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1111 ESEARCH PAPER, SERIES NUMBER 123 JANUARY 1987 UPLANDRICE. INSECTl--1i M gm T -E!R.E OLO U t RTANCE ,AND" ONTzROL J.A. LITSINGER, Al. BARRION, and DANDI SOEKARIA T~ei nternation' Rice Research Instiute P.O.". 16 93, Manila, Philippines

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Page 1: UPLANDRICE. INSECTl--1i M T -E!R.EOLOU t RTANCE ,AND …delphacid.s3.amazonaws.com › 6958.pdf · The significance of abiotic factors in upland rice insect ecology will be apparent

1111 ESEARCH PAPER SERIES NUMBER 123 JANUARY 1987

UPLANDRICE INSECTl--1i M gm

T -EREOLOU t RTANCE

AND ONTzROL

JA LITSINGER Al BARRION andDANDI SOEKARIA

T~einternation Rice Research Instiute PO16 93 Manila Philippines

UPLAND RICE INSECT PESTS THEIR ECOLOGY IMPORTANCE AND CONTROL

JA Litsinger AT Barrion2

and Dandi Soekarna3

ABSTRACI

The diversity of upland rice environments gives rise to a more heterogeneous insect fauna compared with the more homogeneous lowlands A wide array of soil-inhabiting pests - ants termites white grubs crickets root aphids root mealybugs root bugs and wireworms -- common in upland rice cannot tolerate flooding Seedling maggots replace the aquatic whorl maggots as vegetativc foliar pests lnsect-vectored virus diseases arc rare in upland rice Small upland ricefields cause concentrations of the more vagile seed pests during ripening The less stable upland environment -- more restricted growing season smaller area planted greater drought stress - poses greater problems of survival to insects which have overcome them by polyphagy greater longevity off-season dormancy andor dispersal There is no one insect that specializes in upland rice Yield losses to insects however are comparable to those of lowland rice Cultural conrol methods include increased tillage higher seeding rates and crop rotation for soil pests Foliar pests can be minimized by synchronous planting of earlyshymaturing varieties Plant resistance as a method of insect control has iot been greatly exploited because the other breeding objectives of high yield drought tolerance and blast resistance take priority Resistance to stem borers should be a high priority A rich fauna of natural enemies exists but they face even greater problems of survival than the pests The normal low yields of upland rice preclude high levels of insecticide use Seed treatment and baiting are low-cost mnethods as is low-volume sprayi 1g

Entomologist Entomology Department International Rice Research Institute (IRRI) PO Bo- 933 Manila Philippines 2Scnior research assistantEntomology Department IRRI 3Senior researcher Pest and Disease Section Bogor Institute for Food Crops (BORIF) Jalan Cimanggu Kecil 2 Bogor Indonesia

UPLAND RICE INSECT PESTS THEIR ECOLOGY IMPORTANCE AND CONTROL

Upland ricefields are nonpuddled and unbunded without the expectation of impounding water They may be located in many settings - in isolated pockets surrounded by irrigated wetland fields or along steep slopes of recently cleared forest Each enviionment produces a unique composition of insect species

Information on upland rice insect pests is limited and scattered among reports dealing with other rice cultures We found eight reports specifically treating insect problems of upland rice culture (11 59 66 79 116 120 163 176) but none worldwide in scope Most literature focuses on one species or a group of related species Table I lists upland rice insect pests that others have identified and those we have observed Many citations refer to lists of pests with rice as a recorded host or are based on insect habits

Papers concerning upland rice often fail to distinguish between the two ecologically significant aspects ---culture and environment embodied in th term upland rice It is dangerous to extrapolate results of studies of insect pests in well-drained fields where seeds of upland rice

varieties are sown into dry soil (upland rice culture) but in an otherwise lowland environment (fields surrounded by flooded rice) Insects readily fly from the lowland rice-fields to the fields planted in upland rice culture These insects therefore are not necessarily upland rice insects They breed on lowland rice and colonize upland rice We gain greater ecological significance from faunistic records and ecological studies from an upland rice environment where lowland ricefields ar- out of the effective dispersal range of most insect species (tens of kilometers away) In border areas where lowlands and uplands meet studies cannot accurately represent either upland or lowland rice (181)

IPEST (iROUPS

Upland rice insects are more influenced by physical than by biological or socioeconomic parameters I) well-drained soils (lack of prolonged flooding or soil puddling) 2) high probability of drought during crop growth 3) restricted growing season (lengthy nonrice fallow) 4) ricefields interspersed with other crops (diversified flora) and 5) low use of agrochemicals (because of low and unstable yield)

Although a dryland habitat represents an extreme hydrological condition upland rice is host to all but the most aquatic insects (11 176) Whorl maggots caseshyworms water weevils and bloodworms require ponding Many soil and scedling pests are not common in lowland ricefields Deep water rice is established in dry soil and therefore has more in common with upland and rainfed lowland rice than with irrigated rice even though water depths may later reach 1-3 inSecond to deep water rice upland rice represents the most unstable rice environment for foliar insects But upland rice is highly stable for soil insects

The significance of abiotic factors in upland rice insect ecology will be apparent in a discussion of the most prevalent groups of insects attacking upland rice worldwide

Soil -inhabiting pests Well-drained nonpuddled upland rice soils favor pests that pass at least one growth stage underground Soil pests feed on underground plant parts (sown seed or roots) develop entirely in a subterranean habitat and leave only for adult mating and dispersal flights Soil pests include ants (Formicidae) termites (Isoptera) mole crickets (Gryllotalpidae) field crickets (Gryllidae) white grubs and black beetles (Scarabacidac) root aphids (Aphididae) root-feeding mealybugs (Pseudococcidae) root-feeding bugs (Lygacidac Cydnidae) false wireshyworms (Tenebrionidac) wireworms (Elateridae) root weevils (Curculionidae) and soil-inhabiting cutworms (Noctuidac)

A subterranean environment limits mobility particularly in locating food and soil insects have adapted by 1)being long-lived either as individuals (beetles or orthopterans) as colonies olsocial insects (ants or termites) or as dependents on social insects (mealybugs and aphids) and 2) having a wide host range (all species)

Anis Species of -rrestrial ants notably the ubiquitous fire ant Solenopsis geminata and harvester ants Pheidole spp specialize in feeding on ungerminated grass seed Solenopsis readily colonizes disturbed habitats which in turn initially encourage the growth of grasses (39) Colonies of these granivorous ants have a specialized caste that processes seed for food Foraging workers bring seed to

4 IRPS No 123 January 198

Table 1Insect pests of upland rice worldwide

Order Family Species

Isoptera Rhinotermitidae Coptotermesformosanus Shiraki

Termitidae Heterotermesindicola (Wasmann) Anacanthotermesviarum (Koenig) Anacanthotermesrugifrons Mathul et Sen-Sarma Captiterpnesnitobei (Shiraki)Corniterrnesstriatus (lHagen) Procapriterines mnushae Oshima Pericapritermes nigerianus Silvestri (=socialis)Macroternes nutalensis (Haviland)Macrotermes beiiicosus Smcat hman Macroterines gilius (Hagen) Macrotermnes spp Odontoterniesformosanus (Shiraki)Procornitermesarauloi Emerson Procornitermws triacifer (Siivestri) Trinerviterniesgenzinatus Wasmann

[= ebenerianus Sjostedt]MicroceroterynessppMicroterinesspp Synternes nolestits (Burmeister)

Dermaptera Forficulidae Termites unspecifiedDiaperasticuserythrocephalusOlivier

Orthoptera Gryllotalpidae Don lineare (Eschscholtz)GryllotalpaafricanaPalisot de Baeuvois Gryllotalpaorientalis (=africanu) Burmeistera

Gryllidae

Neocurtilla (= Gryllotalpa)hexadacryla (Perty) Scapteriscusdidact)hts(Latreille) Grjllusassinilis(= bimaculatus)(Fabricius)Gryllus (=Liogryllus) bimaculatusde Geer Brachytrupes portentosus (Lichtenstein) Brachytnmpes mnembranaceus DruryPlebeiogrylhsplebejus (Saussure) Teleogryllus testaceus (Walker)Teleogryllus occipitalis (Serville) Loxoblenimus haani Saussure

Pyrgomorphidae

Velarifictorusaspersus(Walker)Velarifictonis sp Euscyrtus concinnus (de Haan)Atractomorpha burri I Bolivar

Tettigoniidae A tractomorphapsittacinapsittacina(de Haan)Conocephahslongipennis(de Haan) Conocephalusrnaculatus(Lethierry) Conoceptahissaltator Saussure Euconocephah s varius (Walker)

Tetrigidae Acrididae

Caulopsiscuspidatus (Scudder) Caulopsis oberthuriScudder Phaneroptera furcifera (Stal) Amphinofus spp Aiolopus thalassinustamulus (Fabricius) Acrida ivilleinsei Dirsch Gesomida mundata zonocera Navas Gonistabicolor (de Haan)Oxyajaponicajaponica (Thunberg)Oxya hya intricata (Stal) Oxyafitscovittata (Marscliall)Oxya velox (Fabricius)Gas frimargus niarmoratusgrandis (Saussure) Xenocatantopshunilishumilis ServilleHierogiyphus banian (Fabricius) ltieroglyphusdaganensis Krauss Hicroglyphus nigrorepletus(1 Bolivar) Hieroglyphus oilziiorosCarl Heteropternisrespondens (Walker)Patangasuccincta (Linnaeus) Stenocatantops splendens (Thunberg) Chondracrisrosea brunneri Uvarov Trigonidium cicindeloides Rambur Locusta migratoriamanilensis (Meyen)Locusta migratoriacapito Saussure

Distribution

Asia Asia Asia Asia Asia Latin America Asia Africa Africa Africa Asia Africa Asia Latin America Latin America Africa

Asia Asia Africa Latin America Asia Africa Latin America Africa Asia Latin America Latin America Asia Asia Asia Africa Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Latin America Asia Latin America Latin America Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Africa Asia Asia Asia Asia Asia Asia Asia Asia Africa

Reference

85 82 190 190 82 85 151 82 85 7 85 85 178 12 82 85 85 195 195 85

193 11 12 157 60 85 195 72 36 186 12 32 59 72 76 84 81 150 195 203 211 81 76 178 b 48 b 76 178 b b b b b 55 178 178 178 65 178 68 81 68 178 178 178 178 178 55 72 178 178 78 178 178 55 55 72 6 82 82 55 54 55 59 130 55 178 55 78 55 82

IRPS No 123 January 1987 5

Table I continued

Order Family Species Distribution Reference

Locusta migratoria migratorioidesReische Africa 33 et Fairmaire

Schistocercagregaria (Forskal) Africa 33 Schistocerca americana (Drury) Latin America 82 195 Valanga nigricornis (Burmeister) Asia 82 Orphulella inticata Scudder Latin America 68

lomoptera Coccidae Pulvinaria sorghicola De Lotto Africa 246 Pulhinariaiceryi (Guerin) Latin America 186 246

Pseudococcidae taennococcus spp Asia 120 Lhorizococcus iu Williams Asia 245 Dysmicoccus boninsis (Kuwana)C Asia Africa 245

Latin AmericaDysmicoccus brevipes (Cockcrell)c Asia Africa 117 203245

Latin AmericaDysmicoccus oryzae (Wirjati)c Asia 245 Geococcus (=Ripersia) oryzae (Kuwana)C Asia 245Brevennia (= tererococcus)rehi (Lindinger) Asia 72 245Nipaccoccus graminis (Maskell) Africa 245 Planococcoides lingnani (Ferris) Asia 245 Pseudococcus saccharicola Takahashi Asia 245 Pseudococcus spp Latin America 60 Saccharicoccus sacchari (Cockerell) Africa Asia 245

Latin America Trionymus ceres Williamsc Asia 119245j rionymus sp Asia b Mealybug unspecified Asia 83 84

Aphididae Capitophorus[= Rhopalosiphumn] prunifoliae Asia 221 (Shinji)

Rhopalosiphum maidis (Fitch) Asia 249 Rhopalosiphum padi (Linnacus) Asia Africa 12 97 249 Rhopalosiphum rufiabdominalis (Sasaki) Asia Africa 66 76 120 195203

Latin America 221229240249 Tetraneura akinire Sasaki Asia 221249Tetraneurabasui ltillcRisLambers Asia 249Terraneura nigriabdominalis (Sasaki) Asia Africa 712 30 76 227229 (=Dryopeia hirsuta)

Tetraneuraradiciola Strand Asia 249 Paracleus ciniciformnis von Heyden Asia 249

[= Forda harukawai Tanaka]Pineus hanikawai (lnouye) Asia 228 Geoica hcifitga (Zehntner) Latin America 76 195 229 249

AsiaGeoica setulosa (Passerini) Asia 249 Anoecia fuliabdominalis (Sasaki) Asia 249 Anoecia comi (Fabricius) Asia 221229249Hysteroneura sctariae (Thomas) Africa Asia 7

Latin America Sipha glyceriae (Kaltenbach) Asia 249 Root aphid unspecified Asia 72 120

Cercopidae Deois schach (Fabricius) Latin America 61 68 194 Sepullia (=Denoplux) nigropunctata Stal Africa 29 Tomaspis ( Deois) fluxtuosa (Walker) Latin America 61 68 194 Tomaspis ( Dcois) flaiopicta (Stal) Latin America 61 61 194 To naspis (Dcois) complcta (Schmidt) Latin Aincrica 68 Tomaspis ( Zulia) entreriana (Berg) Latin America 68 Tomaspis ( Mahanarva) fimbriolata(Stal) Latin America 68 194 Tomaspis ( Aeneohna) spectabilis (Distant) Latin America 219 233 Aeneolamia varia Fciinai Latin America 81 111 Aeneolamia postica (Walker) Latin America 203 Abidama producta Walker Asia 54Delphacidae Sogatella furcifera (Horvath) Asia 59241 Niapanatalugens (Stal) Asia 30 59 72 84 241 Sogatodes oryzicola (Muir) Latin America 6167150Sogatodes cubanus (Crawford) Latin America 150 Sogatodes pusanus (Distant) Asia bPeregrinus maidis (Ashmead) Latin America 186 Laodelphax striatellus (Fallen) Asia 18

Cicadellidae Recilia dorsalis (Motschulsky) Asia 59 241 Nephotettiv nigropictus (Stal) Asia 59241

6 IRPS No 123 January 1987

Table I continued

Order Family

Hemiptera Lygaeidae

Cydnidae

Alydldae

Coreidae Corimelaenidae

Pentatomidae

Thysanoptera Thripidae

Species

Nephotettix virescens (Distant) Nephotettix modulatus Melichar Cofana spectra(Distant)Graphocephala spp Hortensiasimnilis (Walker) Exitianusobscurinervis (Stal)Balclutha spp Draeculacephalaclypeata Osborn Cicadulinabipunctella (Matsumura)Cymoninus turaensis(Paiva) Ninus insigniStal Pachybrachiusnenosus Horvath Caenoblissuspilosus Barber Blissus leucoptens (Say) Blissus spp Dlmnorphoptentscornutus novaeguineae GhauriDimorphopterus similis Slater Paroniuspiratoides (Costa)Pangaeus spp Aethus indicus (Westwood)Geotomnus pygmaeus (Dallas) Stibaropus inolginus Schiodte Cyrornenusbergi FroeschnerCyrtomnenus ciliatus (Palisot de Beauvois) Cyrtonenuscrassus Walker Tominetus spp

Riptortuslinearis (Fabricius) Riptorts spp Leptocorisaacuta (Thunberg)Leptocorisabiguttata (Walker)LeptocorisapalawanensisAhmad Leptocorisaoratorius(Fabricius)Leptocorisachinensis Dallas LeprocorisasolomonensisAhmad Leptocorisaspp Stenocorissouthiwoodi Ahmrad Stenocoris claviformis Ahmad MirperustorridusWestwood aetus trigonus (Thunberg)Alkindus atratusDistant ScaptocorisdivergensFroescher Scaptocoris castaneusPertyScotinopharatarsalis (Vollenhoven) Scotinopharascotti Horvath Eysarcoris (=Stollia) ventralis (Westwood) Dolycorisindicus Stal Macrinajuvenca Burmeister Tanda antiguensis (Westwood) Tantiageifi Schout Tantia perditor (Fabricius) Pygotnenidavaripennis (Westwood) Menida sppNezara viridula (Linnaeus)

Oebaluspoecilus (Dallas) Oebalusypsilon-gnseus (de Geer)Oebalusgrisescens (Sailer) Tibracea limbativentris (Stal)

Aspavia armigera(Fabricius)Diploysfissa Erichson Acrosternum marginatum (Palisot de Beauvois) Acrosternum sppFrankliniellarodeos Pergande Breginatothrips enustos Watson Stenchaetothrips(=Baliothrips)biformis

(Bagnall) (=(Jlloethripsoryzae (Williams)HaplothripspriesnerianusBagnall

Distribution

Asia Africa Asia Latin America Latin America Latin America Latin America Latin America Asia Asia Asia Asia Asia Latin America Asia Asia Africa Asia Latin America Asia Asia Asia Latin America Latin America Latin America Latin America

Asia Africa Asia Asia Asia Asia Asia Asia Asia Africa Africa Africa Asia Latin America Latin America Latin America Asia Asia Asia Asia Africa Latin America Africa Latin America Asia Asia Asia Africa

Latin America Latin America Latin America Latin America

Africa Africa Latin America Latin America Latin America Latin America Asia

Asia

Reference

59 241 217 59 241 6166 61 66 81 150 6166 66 81150 b 178 178 30 83 42 81150 203 30 80 62 178 195 178 178 178 203 203 203 203

55 178 12 72 78 120 214 214 b 59 120178114 178 54 636 82 12 6 12 82 178 81 150 60 is 195 59 120 178 78 36 186 36 186 b 55 12 30 54 55 78 82 120 61 66 81195 61 65 66 195 211 150 195 61 66 81 150 219 233 6 12 82 6 82 186 186 61 66 68 213235

178

7 IRPS No 123 January 1987

Table I continued

Order Family Species Distribution Reference

toleoptera Scarabacidae leteroligus (=Aphonoprocts) 15elcs Billberg Africa 36 Dynastinac (=Ileteroligusmees robustui Prell)

Alissonotum pauper Burnieiser Asia 82 Alissonotuon simile Arrow Asia 209 Dscinetusdubibs (Olivie) Latin America 28 195 233 Devscinetusgagates Burmncster Latin America 20 195 Dyscinctt spp Latin America 60 Eutheola Inimilis Bturmcistr Latin America 20 28 8 1 195 203

233Eutheola identarr(Biurnciter) Latin America 121 203 Heteron)ciwv aw rsonJack Africa 82 Hcreron)chusbtuterculatus Kolbe Africa 82 tleteronichtslicas (Klug) Africa 82 fhteron)chus mosamhicus Peringuey Africa 4 6 48 82 (=Veteronychus orzae Britton)

leteronychts lioderes Redtnbacher Asia 77 82 207 ltcteronchusarator (Fabricius) Africa 82 tleterontchusplebepus (Klug) A frica 57 82Ileteron chus pseuocongonnsis Ferreire Africa 32 Ileteronvchns rogiJrons Fairmaire Africa 82 Heteron chus nrsticus niger (K lug) Africa 82 Heteronvchus spp Africa 17 Lachnosterna longipennis (Blanchard) Asia 82 Lachnosternaspp Asia 120 Ligvros fossator Burmeister Latin America 82 Ligyrus nasutus Burmeister Latin America 203 Ligyrus (=Scarabaeus)ebenus (de Geer) Latin America 20 195

Dynastinac Ligtvros huonilis Burmeister Latin America 60 82 Aaladera castanea (Arrow) Asia 82 Maladera orientalis Motschulsky) Asia 82 Malaaerajaponica Motschulsky Asia 82 Phvlognathusdicynsius (Fabricius) Asia 82

Melolonthinae Exopholis h)ypoletca (Wiedemann) Asia 117 Lerlinosterna[= lolotrichia] consanguinea Asia 122

(Blanchard) Leucopholis rorida (Fabricius) Asia 82 117 Lepidiota blanchardi Dalla Torre Asia 76 82 Leucopholis irrorata (Chevrolat) Asia 76 134 Scrica internpta Walker Asia 76 Holotrichialonipennis (Blanchard) Asia 122 Holotrichia serrata Fabricius Asia 122 Holotrichia leucoph thabna (Wiedemann) Asia 82 Holotrichia seticollis Moser Asia 77 78 Holotrichia pnindanaoana Brenske Asia b Stenoeres spp Latin America 71 195 Atoenius spp Latin America Asia 203 Ph luplg aequata (Bates) Latin America 203 Piyvilophagacaraga Saylor Latin America 203 Phyllophagachiriquina (Bates) Latin America 203 Phyllophagadas)poda (Bates) Latin America 203 Phyllophagaclegans Saylor Latin Ameica 203 Phyllophagahondura Saylor Latin America 203 Phyllophagalatipes (Bates) Latin America 203 Phyllophagamontriesi (Blanchard) Latin America 203 Phyllophaga pamisetis (Bates) Latin America 203 Ph vllophaga sanfosicola Saylor Latin America 203 Phllophagasetifera (Burmeister) Latin America 203 Phyllophaga (=tlolotrichia)helleri Breuske Asia 82 117 159 Phvllophagaspp Latin America 81 150 White grub unspecified Asia 84

Rutclinae Lagochile trigona (Ilost) Latin America 82 Anomala dinidiata var barhata Burmeister Asia 77 78 82 Anomala huteralis(Burmeister) Asia 76 Anoimala lurida (Blanchard) Asia 76 Anomala sulcatula (Burmeister) Asia 76 Anontala varians (Olivier) Asia 82 Anomala antigua (Gyllenhal) Asia 82 Anoinala palida (Fabricius) Asia 82

8 IRPS No 123 January 1987

Table 1 continued

Order Family

Tenebrionidae

Elateridae

Languriidae Chrysomelidae

Chrysomelidae

Coccinellidae Curculionidae

Lepidoptera Lymantriidae

Amatidae

Arctiidae

Limacodidae Eupterotidac Noctuidac

Species

Anomaa volita (Blanchard) Adoretus caliginosus Burmeister Adoretus comprcssus (Weber) Papuana huhneri (Fairmaire) Papuana incrmnis Prell Popillia capricollis Hlope Gonocephalu depressur (Fabricius)Gonocephahm acuiangulum (Fairmaire)Gonocephalum sinphx (Fabricius) Epitragus sallei Champion Anaedus ptnctatissimnts ChampionAeolus spp4griotes spp Agriotes mancus (Say) Conoderts spp Aeoloderma brachmana (Candeze)Wireworni unspecified Anadastus filijf rmis (Fabricius)Dicladispa viridicanca(Kraat z)Trichispasericea (Guerin)Oediopalpastervalis (Weise) Oediopalpa guerini (Italy) Chaetocnema cvlindrica (Baly)Chactocnema denticulata S tephens Chaetocnemabasalis (Baly)Aulacophora sp nr similis Olivier Monolepta caripemne Ialy Monolepta bilasciata (Ilornstedt) Monolepta signata Olivier Cerotoma airojcsciata J acoby Diabrotica adelpha Ilarold Diabrotica balteata Leconte Diabrotica graminLa ltaly Diabrotica speciosa (Germar) Diabrotica limitafa quindecinmunctata(Germar)Diabrotica melanocephala (Fabricius)Diabrotica spp Altica (= Fondia)madagascariensis(Allard)Altica spp Epitrix cuctmeris (Harris) Epitrix spp Oulenjo or pae(Kuwayama) Disonchq (= Donacia) spp Flea beetle unspecified Chnootriba (= htIilachna)sinilis assimilis Mulsant Atactogaster iduriens Walker Hypomeces squtamosus (Fabricius) Tan vinecus discoidalis GyllenhalPhaulosoinusmusculinus (= mnus) Csiki Neobaridia anplitarsis CaseyPsalis pennatula (Fabricius) Laeliasufihsa (Walker) Euproctis irgtncula (Walker)Euproctis minor (Snellen)Euproctis xanthorrhoea (Kollar)Amata sp

Diacrisia obliqua (Walker) Creatonotusgangis Linnacus Latoia (= Parasa) bicolor (Walker)Nisaga simplex Walker Sesamia botanephaga Tams amp Bowden Sesamnia nonagroides botanephagaTfums et Bowden Sesamia calamistis llampson Sesamia inferens (Walker) Spodoptera eridania (Cramer)Spodoptera mauritia acron)ctoides Guenee Spodoptera frugiperda ( ESmith)

Distribution Reference

Asia 82 Asia 82 Asia 82 Asia 82 Asia 82 Asia 78 Asia 82 117 178 Asia 82 117 Africa 82 Latin America 203 Latin America 203 Latin America 195 Latin America 195 Latin America 203 Latin America 195 Asia h Asia Latin America 60 73 117 Asia 200 Africa 36 Africa II Latin America 6166 68 Latin America 6166 68 Asia 225 Latin America 61 66 68 81

Asia 178 Asia 55 Asia 55 Asia h Asia 55 Latin America 203 Latin Amefia 203 Latin America 203 Latin America 203 Latin America 6166 68 Latin Amcrie 195 Latin Amw 195 Latin Aierwca 61 81 Africa 57 Asia 178 Latin Americ 203 Latin America 81 Asia 82 Latin America 195 Asia 120 Africa 6 11 Asia 185 Asia 120 Africa 82 Africa 82 Latin America 66 68 195 233 Asia 117 Asia 117 Asia 117 Asia 117 Asia 117 Asia A S Pradhan

unpubl dataAsia 82 Asia 82 Asia 82 Asia 182 Africa 12 Africa 6 36 Africa 6 8 11 12 36 Asia 59 82 84240 Latin America 150 Asia -0 59 196 Latin America 28 61 66 68 81 150

152195203219233

9 IRPS No 123 January 1987

Table I continued

Order Family Species Distribution Reference

Spodoptera exigua (Hubner) Spodoptera exem pta (Walker) Spodoptera ornithogalli (Guenee)

Africa Africa Latin America

32 217 32 33 36217 150

Mythimna loreyi (Duponchel) Asia 30 Mythimna separata (Walker) Mythimna (- Pseudaletia)latifascia [= adultera

Asia Latin America

178 195

(Schaus)] (Walker)Mythimna (=Pseudaletia) sequax (Fabriclus) Latin America 68 Mythimna rosellinea (Walker) Asia b Mythimna yu (Guenee) Asia b Mocisfrugalis (Fabricius) Asia 30 117 Mocis latipes (Guenee) Latin America 66 68 149 150 203

Platysenta (=Spodoptera) compta (Walker) Asia 219233A Barrion unpubl data

Agrotis ipsilon (Hufnagel) Latin America Africa

33 68 81 150 1)5 203

Agrotis spp Latin America 60 Achaeajanata (Linnaeus) EldanasaccharinaWalker

Asia Africa

117 36

Armyworm unspecified Asia 72 84 Pyralidae Cnaphalocrocis medinalis (Guenee)

Marasmia bilinealisHampson Asia Africa

30 4559 72 84233 36

Marasmia trapezalis(Guenee) Africa 32 217 Marasmia (=Sumia) exigua (Butler) Asia 220 Marasnianmralis (Walker) Asia b Marasiniapatnalis Bradley Marasiniaspp

Asia Asia

b b

Leaf roller unspecified M2liarphaseparatellaRagont

Asia Africa

72 12 57

Maliarphasp Asia b Chilo auricilius Dudgeon Asia b Chilo partellus (Swinhoe) Chilo polychrysus (Meyrick)

Africa Asia

12 92 59 72

Qhilo diffusilineus J de Joannis Africa 6 1229 Chilosuppressalis(Walker)Cibozacconius Blezzinski

Asia Africa

59 72 6 11 12

Cravubus spp Latin America 203 Acigona loftini (Dyar) Latin America 203 Acigona chrysographella(Kollar)Scirpophagaincertulas(Walker)

Asia Asia

82 59 72

Scirpophagainnotata (Walker) Asia 59 200 Scirpophaganiuella (Fabricius) Asia b Elasinopalpuslignosellus (Zeller) Latin America 28 60 61 66 81 150

Diatraeasaccharalis(Fabricius) Latin America 162195203219233 28 66 81 111 150 195

Rupela albincllaCramer Latin America 111

Gelechlidae

Stem borer unspecified Undetermined stem borer nr Maliarpha BrachniaarctraeaMayr

Asia Asia Asia

72 b 55

Brachmi spp Africa 32 Hesperiidae Parnaraguttata Bremer et Grey Asia b

Parnaranaso Fabricius Asia 55 Pelopidasagnaagna (Moore) Pelopidasmathias (Fabricius)

Asia Asia Africa

164 36 155217

Pelopidasconjuncta conjuncta (Herrich-Schaffer) Asia b Borbo fanta Evans Africa 6

Satyridac Melanitisleda ismene Cramer Asia 84213 Mycalesis asophisHew Asia 30

Diptera Cecldomyiidae Mycalesis spp Orseoliaoryzae (Wood-Masn)

Asia Asia

b 72 88 131 181

Orseoliaoryzivora Harris et Gagne Africa 61132218 Chloropidae Steleocerellus (=Mepachymens) ensifer (Thomson)

Oscinella spp Asia Asia

205 205

Gaurax sppChloropsoryzae Matsuntura (= kuwonae Aldrich)

Asia Asia

205 82

Mucldae Atherigonaoryzae Malloch Asia 30 82 117130 Atherigona exigua Stein Asia 82 117 178

10 IRPS No 123 January 1987

Table I continued Order Family

Diopsidae

Agronyzidae

lHymenoptera Formicidae

Species

Atherigona oritentalisSchiner Atherigona indica MallochDiopsis longicornis(= thoracica)Macqurt Diopsis spp Pseudonapnyza asiatica Spencer Pseudonapomyza spicata (Malloch) Agronoza orvzae MunakataAcromyren landoltibalzani(Emery) Acromyrmex landoltifracticornis(Forel)Acromyrnex heyeri Forel)Atta bisphaericaFore Atta capiguaraGoncalves Atta laeLigata (F Smi h1) Atta opacicepsBorgmeierSolenopsisgeminata (abricius)Pheidole sp Pheidolegeton diversus Jerdon Monomorium pharaonis (Linnaeus) Ants unspecified

Distribution Reference

Asia 82 Asia Africa 82205Africa 11 1248 Africa 217 Asia 213 Asia h Asia bLatin America 6168 195 Latin America 61 195 Latin America 61 68 195 Latin America 61 68 195 Latin America 14 61 68 195Latin America 61 68 195 Latin America 61 195 Asia b Asia b Asia 1) Asia b Asia 120

aG orlentalls formerly recorded in Asia as G africana thPhilippines reported in this study CSubterranean root feeders

subterranean nests Foraging ants will feed on germinatedseeds only when the supply of ungerminated seed is low However these ant specialists feed on a wide varietyplants and thus help control grassy weeds (39) This dual

of

pest benefit role extends from tending ants which protecthoneydew-excreting pest species including root aphids and mealybugs (workers repel potential predators and even selectively kill parasitized aphids) to predators of leaf- and stem-feeding rice pests such as leaffolders armywormswhite grubs and stem borers (1 17)

The upland rice environment is particularly well suited to ants because of the lack of flooding Frequent tillage perpetuates grasses and granivorous species that are highlyadapted to constructing new nests in recently tilled fieldsAnt nests however are relatively shallow and tilling with a moldboard plow can destroy them Also upland ricefields are uually small and isolated and a given area in a smallfield has more field borders than it has in a larger fieldallowing high rates of infiltration from more sedentaryspecies such as Plieidole that do not colonize tilled fields but nest in soil under shrubs and trees along field bordersPheidole workers forage into fields from borders and store seed in their nests In slash-and-burn or other no-tillage rice cultures nests would be preserved in the field and possiblylead to high rates of seed predation However granivorous species do not prefer forest habitats

Damage is characterized by reduced usually patchy plantstand (120) Seed-thieving ants greatly prefer rice but they can be selective regarding plants they collect and store Theyharvest more seed than they can consume so patches of germip ing surplus seed (such as Rottboellia) will sproutfrom their nests during the rainy season

In Latin America leaf-cutting ants (Aita spp and Acromyrmex spp) occur only in upland fields and defoliate

young rice plants (47 68) They take leaves to fungal gardensin underground nests where fungi predigest the plant material

Termites Subterranean termites of the family Termitidae which lack symbiotic protozoa to help digest plants are the most frequently mentioned termite pests of upland rice (1185 120) This family cultures fungi in special undergroundcells --- fungal combs made of half-digested plant material (117) The fungi inoculated onto the combs break down plant material into food that termits can digest The termites then consume the fungal combs These grasslandtermites build nests below the plow layer in uplandricefields The colonies attack living plants only when dead plant material is not available They will attack a droughtshystressed crop but prefer older plants having greater cellulose content They will damage newly planted crops where clean culture has removed vegetation

Infestations are worse in deep light-textured soils with low moisture content The first sign of damage is yellowingof older leaves (195) Termites feed on roots (the plantyellows then wilts and finally collapses) and germinatingseed (loss of stand) or move above ground at night to cutseedlings at ground level which they cover with soil for later consumption Soft-bodied termites are highly sensitive to desiccation They live within a self-contained system of airtight chambers and build surface tunnels linei with nudand body secretions (carton) to maintain more than 90 relative humidity Termites can be located by their tunnels

Termites apparently can withstand limited submergence (157) and can be a pest of dry lowland seedbeds (117)perhaps by having nests below perched water tables

Termites appear to do more damage in Latin America where large ricefields are plowed with tractors deprivingtermites of preferred vegetation and forcing them to live on

rice plants In Asia upland ricefields are typically small with Iprge field border areas (ecotones) where termites can forage for preferred food other than growing rice plants All grassland termites do not feed on rice In an upland area in Batangas Province Philippines ricefields are commonly infested by Hosiititernesluzonensis (Oshima) however this termite does not damage rice In the same province though Mrroternesgivus(Hagen)feeds on rice seedlings (178)

In Africa termites may damage rice even though fields are small and surrounded by perennial vegetation These termite species perhaps prefer rice over other hosts

Mole crickets Nymphs and adults are nocturnal and burrow through soil feeding on roots of a wide variety of plants or forage above ground as predators of insects Adults live 3-5 mo and are even cannibalistic Mole crickets prefer low-lyin moist upland soils with high organic matter (II 81) In the lowlands they inhabit rice bunds or nonflooded fields Adults are highly mobile and can leave a flooded field to locate a more suitable habitat Losses show as wilted plants (from root feeding) or reduced stands (120) Damage is more common near field borders where mole crickets relocate after tillage (126)

Fiehl crickets Nymphs and adults of field crickets have similar nocturnal habits and damage rice as much as mole crickets do Piles of weeds removed from fields attract them They make subterranean nests and tunnel through the soil to feed on roots Some species prefer seed to roots Others feed at the base of stems causing deadhearts

Tunnel entrances surrounded by excavated soil are easily seen Young plants are cut at ground level and stored in underground cell Field crickets tre highly polyphagous as are all soil pests (117) Sachan et al (198) reported that field crickets prefer maize to upland rice Like mole crickets field crickets cannot survive in standing water and are therefore more prevalent in upland fields

Whitegrubsandhackbeetles Scarab beetles that feed on living roots as larvae but not as adults are called white grubs or chafers (Melolonthinae Rutellinae) and those that feed as adults but not as larvae on plant crowns are called black beetles (Dynastinac) In the tropics chafers or white grubs have a I-yr life cycle starting with adults emerging from the soil about I mo after the first downpour of the rainy season (134 159) They lay eggs at the same time farmers sow upland rice The rice passes its most susceptible stage and damage is mostly avoided when the white grubs are small After several months the long-lived larvae are large enough for two or three to denude the root system of mature rice (223) This intensity of damage is rare but wilting occurs when root loss is combined with drought stress

The larvae need damp (not saturated or dry) soil to survive and pass the unfavorable dry season 05-2 m underground in aestivation in the moist soil The first heavy rains of the season (20-30 amd) stimulate the grubs to resume activity and after several weeks they develop into

IRPS No 123 January 1987 11

pupae and adults and eventually dig their way to the soil surface and fly to nearby trees to seek food and mates The adults of most white grub species feed only on trees but in some species notably Eutheolahumilis the adults defoliate and cut off stems at ground level (28) Putheola hidentata damages sugarcane fields in Venezuela (121) Floods kill white grubs (208) Grasslands can support large populations and therefore white grubs can be more abundant in newly planted upland ricefields that were previously fallow (81)

Black beetles live up to a year adults can attack a crop at any stage by burrowing into the soil (127) and feeding at the base of stems to cause whiteheads (207) Adults emerge with the early rains and are normally more abundant and do more damage on a young rice crop with small root systems (38 81)

The larvae feed only on dead organic matter in dryland fields and do not attack rice Adults arc highly mobile and although sensitive to flooding can invade a field soon after it drains (115)

Root aplhids Several genera of migratory aphids feed on the roots of upland rice (249) Populations build up more in light-textured soils with high percolation rates Damage has also been reported from dry seed heds of lowland rice (52) Tending ants -- Pheidhle Cren aster Tetramoriuw lsius Tapinoma -- are necessary for root aphids to multiply Aiits harbor aphids in their nests over winter or during unfavorable periods and relocate them on rice plants digging tunnels along the root systems to allow these softshybodied insects to penetrate the soil Most intensive studies on aphid life histories have been done in temperate environments (158 222 229) Root aphids fly to rice plants at the beginning of the rice season and pass through several generations continually developing winged forms to relocate on alternate grassy hosts until winter when they move to perennial hosts usually trees Yield loss occurs mainly through reduced tillering but infested plants become yellow and stunted and in extreme cases wilt (227) Aphid populations build up gradually so damage usually begins in the late vegetative and reproductive growth stages

Root-feeding ineal tugs More than half of the mealybug species or rice feed on roots (245) Being soft bodied mealybugs are not adapted to living underground but they survive because tending ants (the same species that tend root aphids) dig burrows for them and move them from plant to plant Well-drained soils ihelp mcalybugs survive Although six species are recorded as subterranean root pests of rice we found no report citing damage Root-feeding mealybugs are probably more prevalent on perennial grasses than on short-lived annual rice

The most common rice mealyb ig Brevennia rehi is a foliar feeder favored by lack ol rain apparently it is responsible for high yield loss in Northeast India and Bangladesh (N Panda pers comm as stated by IRRI [1061) This report needs confirmation Mealybug damage was only briefly mentioned in another report (156)

12 IRPS No 123 January 1987

Root weevils A few species of upland weevils (IHporneces Donacia Atactogaster) attack rice roots as larvae and feed on stems and leaves as adults Damage ispatchy The larvae are not adapted to submergence and are true upland soil weevils isopposed to the so-called water weevils They rarely become sufficiently abundant to cause damage Adults on rice foliage or the soil surface show that the roots are being attacked by the grubs Root weevils are highly polyphagous (185)

Root heeths Soil-inhabiting false wireworms Gono-cephalumt spp are reported in upland ricefields The adults and larvae normally feed on decaying organic matter but land preparation for upland rice removes these sources The larvae feed on seedling roots and nocturnal adults roam on the soil surface to cut off seedlings at ground level (82) This group of ground beetles is highly omnivorous adults even prey on stem borer larvae (183) True wireworms (Elate-ridae) feed on living roots but rice as a host has been reported Infrequently (117) In Latin America the highlypclyphagous larvae of Diabrotica rootworms are also recorded on rice

Root htus Upland rice seedlings can be attacked by the adults anI nlViphs of chinch bugs Blissusand Caenoblissus and bro n bug Scaptocoris which feed on the roots with theirsucking nouthparts (80 83) Three genera of burrower bugs A tms (eotonhus and Stiharopus -are common iil grasslands in the Philippines and may attack the first crop (rice or maize) planted after the dry season Chinch bugs are most abundant after a dry spell These insects are large and can kill seedlings or stunt and reduce tillering on older plants I hey prefer other hosts and rarely damage rice Adults can also feed oii developing rice grains (81)

(utworms Soil-dwelling noctuid larvae hide under-ground during the day to avoid predation by birds and bccone active at night above grour cutting off you ing plants at ground level (hence tile naiie cvtworn) arvaedrag severed plants into hurrows llie larvae have wide host ranges arid pupatc in the soillgrotis ipsilon occurs worldwide and appears to be the most common cutworm pest of upland environments but is more prevalent inlowland rice particularly in dry seedbeds

IFoliar pests Other groups of insects are adapted to upland environments because they pass one growth stage underground either as egg (in heinrietabolous species) or pupa (in holomnetabolous groups)

Rice seedlibig maggots AitJeriota flies occur in non-fhoodcd environments in Asia and Africa (180) Feeding of larvae one per iller causes deadhearts Flies as opposed to moths only secondarily evolved as pliytophagous pestsfroni a saprophagous Origin Athrnonalarvae secondarihv feed onidecaying tillers whereas larvae of stem borer moths actively Iced on living tillers to cause deadlicarts Seedling maggots will die if they cannot sever a tiller on which to feed

as it decays Plants die under heavy attack whereas lighter infestations cause stunting delayed maturity and ragged leaves (236)

Eggs are laid only on actively growing plants (49) aid damage occurs wiihin the first week after crop emergence(170) Larvae pupate more in stems than in soil Seedling maggot attack is highly seasonal (206) normally peaking 2-3 mo after the onset of the rainy season These pests possiblyaestivatc over the dry season Rice isonly one of their many graminaceous crop and weed hosts

The rice stem maggot (hlorops orzae is adapted to temperate upland and lowland rice growing regions of Asia In Japan and China upland rice grows in mountainous regions a habitat of this chloropid fly The larvae tunnel into tilestems The stem maggot is more damaging during the seedling stage larvac hibernate in grasses

File several dipterans reported as stem maggots in dryshyseeded lowland seedlings inIndia (205) could presumably attack upland rio

Leaf urinrs Icaf miners are common on wheat and maize but are rarely reported on upland rice (213) Three leaf miners are more prevalent on upland than lowland rice inthe Philippines Pvudonalomy a asicanh Vficataand lIAgromrza orzae attack at the seedling stage The larvae develop inside the tunnels they construct in the parenchyma tissue P Vpicata and Aorvzac prefer maizeEehtsine and other grasses to rice spicatacan become abundant on wheat

Leaf beetles Chrysomelid and coccinellid eaf bcetles scrape or otherwise remove leaf tissue as larvae or adults or both This damage accelerates desiccation of plants Tile larvae of sonic species are root feeders Flea beetles Epitrix and (lraetocnetna and rootworms l)iahroticaspp feed on rice foliage only as adults Their root-feeding larvae have hosts other than rice The adults and larvae of the hispa-like Oediolmlpa spp beetles of Brazil and (hmootriha (=EPfishyachra) beetle of Africa feed on upland rice leaves Oealioshypapa spp larvae are leaf miners (mootriha is also pshyvalent in lowland environments (11) Leaf beetles prefer vegetative rice (81)

The rice hispa Dicladipaarnigera is not reported on upland rice and prcers more aquatic habitats In Central Africa however I)cladispm viridicyaiea attacks uplandrice in the vegetative stage as well as lowland seedbeds 7richipasericea is reported on upland rice in West Africa and Madagascar (57)

The rice leaf beetle 1d1ea 01izae riiuech like Chioropsor|rzae is adapted to temperate regions of Ashi and is usually a pest on lowland rice but seldom oi inpland rice in the mountains (82) The adult hibernates on grasses

Armmworms Species of grassland-adapted SpoptloeraPlatysemta and A ithimna larvae create widespread epishydemics They can defoliate ricefields generally in patchesfrom early vegetative growth to harvest (41) Armywornms pupate inthe soil and highly favor upland rice (34) Adults

can disperse long distances to colonize even remote upland areas They become more abundant than their natural enemies particularly after favorable rains following a prolonged drought (196) The drought kills the armyworm and natural enemies (mainly parasites) but the armyworm can recolonize rapidly

The natural increase insoil fertility from mineralization of soil N over the drought period promotes luxuriant plant growth to foster a rapid buildup of these highly fecund species Weeding rice increases tilelikelihood of damage The parasites normally return to control the armyworms but not until after serious damage

Outbreaks on upland rice have been reported in Ghana (3) inzibar (38) Central Africa (36) lPanama (165) Brazil (28) Malaysia (196) and India (167) Defoliation may be severe often leaving only the base of stems Armyworm larvae hide during the day under litter or in soil cracks At night they ascend plants to feed Larvae pupate in the soil

Thrips Thrips frequent upland rice but damage is less than in the lowlands Thrips larvae and adults feed on leaf blades by rasping causing leaf rolling and stippling Thrips prefer vegetative stage rice and their survival increases during drought (heavy rain washes them from foliage) although drought stress usually outweighs thrips damage Drought and thrips cause leaf rolling in upland rice (235) but thrips have been overlooked perhaps becaLse of their small size Thrips are most numerous indry season irrigated rice The combination of favorable plant growth and lack of rainfall is ideal for thrips In contrast the lack of rainfall in upland rice hurts the crop and eventually the thrips whose numbers must decline

In [atin America Frankinilla rodeos attacks panicles before they emerge from the boot calusing sterile grains

Stem hug~s Black bugs are the most common group of rice stem-feeding -lemiptera in Asia The most frequently mentioned species ----Scotinophara coarctaia and Shrida - prefer aquatic habitats and therefore could not occur in upland rice as sometimes reported Upland species in the same gerurs - notably S tarsalis and S scotti -- are morphologically similar and remove sap from stems of grasses and upland rice The upland species are notably more abundant in upland rice planted near forests

In Latin America tile Tibraca ima-large pentatomid tiventris causes deadhearts and whitehcads on upland rice Both adults and nymphs remove sap from internodes of plants more than 3 wk old Tillers are killed by mechanical damage and by the entry of secondary microbial infection This damage is often confused with that of lepidopterous stem borers (233) Eggs are laid on the leaves and stems Nymphs are gregarious This species can also be found on irrigated rice Drought reduces population buildup

Stiem borers Lepidopterous and dipterous stem borers are widely recorded on upland rice but cultural practices and alternate hosts determine abtndance Except for Elasmolmll)uS Iinosellts numbers usually build up toward

IRPS No 123 January 1987 13

the end of the crop growth cycle Therefore late-maturing varieties staggered planting and lack offloodingfavorstem borer buildup Many stem borer species can lie dormant in the stUbbleafter harvest and theirsurvival is encouraged by panicle harvesting (leaving tall stems) and lack of tillage after harvest

Stem borers disperse and lay many eggs Many species are polyphagous however the monophagous MaliarpiasepashyratellaDiopsis Ivmgicornis Scirpophaga incertlas and S innotata can also be abundant on upland rice (hilo Suplressalisdevelops fasterand becomes la rger when reared on upland than on lowland rice (226)

The lesser corn stalk borer lasmopal tt Ittoselhs is semisubterranean and is perhaps the stern borer most adapted to upland rice(129) Highly polyphagous it infests maize peanut and cowpea It attacks seedling rice (one larva can kill tip to four plants before maximum tillering) (195) (hiv and Maliarphastem borers prefer older plants But Scirpophagaspp will attack young plants even in the nursery lasn opalls larvae tunnel into the stems at or below the soil level mausing dcadhcarts Larvae are not found in their turnnels inside sterns beca use vhei disturbed they retreat into cases made from soil particles bound by silk The cases areattached to the tunnel entrance (143) This behavior probably evolved as protection from natural enemies Larvae pupate in the soil The lesser corn stalk borer is more abundant during drought damage is often confused with that of drought (233) This species prefers sandy soils arid its distribution within a field may relate to soil texture (143)

Diatraeasaccharalis aside from the lesser corn stalk borer is frequently a stem borer pest in Latin America According to Teran (233) rice and maize are more preferred hosts than sugarcaine its namesake

Atcitonacrvsograpla is a less known upland rice stem borer in Asia with habits similar to Easinopall s (82) because it is not found in lowlands Acutra has grassy weed alternate hosts As a moth it is often confused with the dark-headed stern borer(hi polur(standstriped stern borer (hi sup)lpsrcais

Other common upland rice stem borers are also reported in the lowlands The dominant species in upland rice vary greatly with location In Uttar lPradesh India (78) and in Japan (123) the pink stem borer Sesamia htfirens is more prevalent hIKenya the principal rice stem borer is Chi partelus

(92) The lowland African sten borers Aialiarpha sepashyratellaDiopsis vngicornis and Sesamia calatnisnis are less abundant Oimupland than on lowland rice(217) but perhaps cause greater yield loss because plant injury is greater if combined with drought css (92) The stalk-eyed stern borer D)ivpsis is more prevalent in wet habitats (6 1I217)

In Central and West Africa Maliarpha and Sesamia calanistis are the dominant stern borers of upland rice (8 9 10 II) (lilo dihlfilineui and Chio partellus inhabit the

14 IRPS No 123 January 1987

upland savannas whereas Chilo zacconio prefers the lowlands (II) Aaliarpha is most abundant in the mangrove swamp habitats (91) coastal regions (16) or rain forest zones (II) of Africa and appears to be the homologue ofScirlpophaga incertulas The yellow stem borr can also be prevalent in upland rice (120 167) The sugarcane borer Eldanasaccharinais so far only a potential pest of uplandrice (91) despite Grist and evers (82) warning

Grasshotl)ers andI locitas Short-horned grasshoppers and locusts -- Locvta Patanrga Schistocerca - layeggs in the soil inhabit grasslands and develop into swarms to seriously damage upland rice from time to time (173) Largenumbers can destroy a field leaving only the stubble In low cmuoroer s giassnoppeirs an cut panicles (173) Farmers may not plant if locust swarms are imminent Prolonged dryweather followed by favorable rain favors development of large swarms (173) Usually breeding in di) grasslands theydisperse toattack crops including upland rice(191) Eggsaresusceptible to desiccation d uring drough t and toinundation during seasonal rains With favorable soil moisture over consecutive seasons natural enemies ca nnot prevent a largepopulation increase leading to migratory swarms (64)

Most grasshoppers on rice are adapted to uplands (107)but lowland rice grasshoppers iervgl(rlhusspp and Ora spp occur occasionally on upland rice (2 82) They lay eggson rice foliage Nymphs are scmiaquatic

The gryllid Lsctrtus coucinnus is a pest of lowland and deep water rice in Bangladesh and Thailand In the Philippines it damages upland rice grown in isolated pockets near lakes The gryllid nymphs and ad ults feed on the central portions of folrage leasing only the midrib and leaf nargins

Leal ler Pvralid moths whose larvae fold leaves to make a feeding shelter occur worldwide (217) but arc appareritly most common in Asia Leaffolde rsattacking rice in Asia are a complex of species (26) (Ctalhahcrcisinedinalis the best-known rice Ia flolde r in Asia may prefer grassy weeds over rice (86) It is a late colonizer of uplandrice U1arasmja eviua a lowland species (220) nreflers rice to grass specres (86) and colonizes upland rice during il early growth stages llarasnria patnalis is the most corn-rnonly encountered species in upland rice in the Philippines Perennial grasses are their alternate hosts and they can survive year-round in upland rice environments The leaf-folders 11arasmia ruralisand Alarasmia spp are least aburndant Dormancy is unknown in leaffolders Cnaplha-ocrocis medhnalis however is known as a migrant (90)Leaffolders remove photosynthetic tissue attack during the flag leaf stage is particularly injurious Popurlations are normally held in check by natural enemies but leaffolders become parlicularly abundant in conditions of high plantfertility and shade

Butterflies Skippers PelopidasInathiasand Borhofanta are more prevalent in African lowland than in uplandhabitats (6) but extensive studies in Japan and China

showed that Pelopidas mathias prefers an upland habitat (155) Parnaragutata seasonally migrates between lowland and upland areas and is equally abundant in both environshyments (164)

The green horned caterpillar Melanitis leda ismene is more abundant in lowlands (212) but our experience in the Philippines suggests that Pelopidas niathiasand 11elanifis leda ismene are often more abundant in uplands than in lowlands Bamboo iaunbuiva spp annually sustains P mathiaspopulations in the Philippine uplands Adults feed on nectar from flowers and migrate to upland rice during the late vegetative stage

iPi erlled h i-1 1d dill L tIC i 7-0 l(become numerous probably because of high egg predationEgg and larval parasitization are often low Butterflies can readily disperse long distances to seek renlote uplandricefields However they have low biotic potential and even it predation rates are low the rarely become abundant Upland environments with their diverse microhabitats can provide more favorable sites than lowland rice plains

tutterfly pests of rice have broad host ranges and become dormant Juring urnfavorable times of the year

IolhagrotslaidopwraThe larvae of many polyshyphagous moths are reported to defoliate upland rice tussock moths lsaispelnnatula Iawlia sUifita luproctisirgtwltaE mnorand P vathorrhoeaslug caterpillar

litotiahicolorhairy caterpillars Nisaga sinp vandA mata spp and wooly bears l)iacrisiaohliqtia and Crvatonotts gangis Many of these polyphgous larvac migrate and are transients in rice including HelhWhis arm iera and 1t zea highly pol phagous species (82)

Gall nidges The Asian rice gall midge Orseoliaorrzae (88 131) and its African counterpart 0 ortzivora(6 11 12 32217) prelerlowland to upland rice Gall midges in rainfed areas are prevalent only during tle rainy season and dispersepoorly Their alternate hosts (wild rices LeersiaIschaetnuan and Pasahmt)are mostlyaquatic grasses and are less likelyto be near upland rice areas In an area of mixed uplandfloating and rainfed lowland rice cultures in northern Thailand upland rice which is ilanted a month earlier was attacked firstbut subsequcent populations were much lower than in rainfed lowla id or floating rice Low numbers of gallmidges passed the dry season orr perennial weeds such as Paspahim (88)

Smot (hid(s Reports of aphids attacking upland rice plants are few (249) Without maize the maize leaf aphidRho alosihhunm maidis attacks upland rice in the Philipshypines The plum peach aphid IIrsteroneuraselariae was reported in Sierra Leerne toinfest leavesand tnriperied grain(7) Siphaglrctriaecaused transverse linrea r necrotic striae iir rice leaves in Italy (172) I-lea v rains held populations in check Rice is not a preferred host for shoot aphids

Leafhoper andI )anthol)pers Cicadellids and delshyphacidsdispersecasilyand readilycolonie upland ricefrom nearby lowland areas A study in Sarawak (241) showed

IRPS No 123 January 1987 15

similar species in upland and lowland rice Nephoiettix The principal vectors of rice virus diseases generally have species dominated The most prevalent species accounting narrow host ranges and alternate hosts in the grass family for 90-95 of the sweep net samples wre N virescens N are mostly annuals dying in the dr3 season Some virus umropictus Nilaparvia hgens Soatellafiircifera Cfana vectors such as BPH can disperse lontg distances but few do

sectraand Recilia dorsalisLowlands had higher popula- Th restriction of upland rice and weed hosts to the rainy tions Katanyukul and Chandaratat (120) monitored hopper season the small area planted to upland rice and the hopper numbers in upland and lowland fields of northern Thaiiand vectors having narrow host ranges all work against virus and found similar results A comparative study of plant- diseases in upland rice hoppers in Fiji showed more brown planthoppe BPH) and In Latin America Sogctdhs o(rzicola and S cuhanus whitebacked planthopper (WBPH) in wetland rice than in transmit hoja blanca virus disease but reports on upland lowland rice and hopperburn was prevalent in lowland rice are rare except in the favored upland areas of Colombia areas with standing water (89) Peru and Venezuela (35 42) S rricolaocurs in Brazil

Hopperburn in upland ricefields in Asia is rare (84) In on upland rice but hojaI blanca has not been reported (67) Batangas Philippines isolated fields of upland rice become S orzicola has a wide host ramgc (10) similar to WBPI- in hopperburned infrequently from WBPH A report from Fiji Asia WBPH however is not known to transmit any virus cites hopperburn from BPH and WBPH (250) In these two disease situations upland rice was grown on highly fertic soil and In Africa pale yellow mosaic virus is endemic to swampy N fertilizer was applied Hoppers respond to better nutrition lowland areas where rice ratoons year-round (19) and is Normally upland rice is grown on poor soil and receives no mechanically vectored mainly by hispid beetles A minor fertilizer vector a flea beetle Chaeiocnema spp is an upland rice

In Asia upland rice ras are normally not far from pest and upland ricetields near endemic areas occasionally lowland ricefields-- a source of hoppers Otherwise hopper contract the virus species with greater polyphagy than BPH or Nelhotettiv Spittle bugs Cercopid nymphs producc a protective virescens would be dominant frothy covering that looks like saliva In Latin America

In Africa leathoppers and planthoppers do not become spittle bugs damage upland rice In parts of Brazil where abundant on upland rice (6 II) probably because lowland upland ricefields are surrounded by pastures adults rice is not widely rlanted nearby The only reported migrating from grasses can damage rice severely and cause hopperburn as from Nilapanrata maeander in breeder complete hopperburn in a young rice crop within a week of plots receiving high N rates in Nigeria (98) Hoppers colonization (194 195) Dispersal to rice occurs even if however are a greter problem in Latin America where pastures are verdant and spittle bugs prefer pasture grasses large grassland areas breed polyphagous species (Gra- to rice (168) Nymphs are rarely abundant enough to phocephala spp Hortensia spp lEvitianus ohscurunervis damage rice Regular rainfall favors their development Balclutha spp and Draeculacephala spp) vhich then (233) disperse to cause hopperburn on seedling rice (68) Rosetto et al (195) reported Graphocep7hala spp and Hortensia spp Seed pests to be equally abundant on upland and lowland rice Alydid and pentatomid bugs are seed pests of upland rice

The leafliopper and planthopper epidemics in Asia that worldwide In Asia Leptocorisa sp prevail Their habits caused severe losses from hopperburn and vijus dicases make them particularly suitable for upland rice 1)thev feed over the past several decades occurred principally on on many grasses (and therefore can survive the earl y wet lowland rice Except in a few isolated cases (160) upland season before rice sets its grain) 2)adults are long-ied and rice has been spared from tungro grassy stunt and ragged mobile allowing them to find isolated plantings 0 rice stunt virus diseases vectorcd by green lcafhopper (GLH) (besides seed bugs only birds and rats appear to have shyand BPI] (123) Virus diseases have occurred in upland rice capability) and 3) adults aestivate in forested areas or where lowland rice was nearby Rice dwarf disease however sugarcane fields during the dry sca son when neither rice nor was first recorded on upland rice in the Philippines (187) grassy weeds are present (I199) Rice seed bugs concentrate Virus diseases can be perpetuated only on livir plants or on small-scale upland riceficlds because they can actively insect vectors and no living rice - plans tooned or search them out The prevalence of groves of trees characshyvolunteer - grows year-round in upland arcas excert in a teristic of upland environments ensures nearby aostivation few with 12 mo of rainfall sites keeping the seed bugs close by

The smaller brown planthopper Laodcelphax striatellu Leptocorisae are uncommon in lowland rice plains occurs in upland rice in Japan (18) It spreads black- because 1) their populations are diluted in a sea of rice streaked dwarf and stripe viruses between winter wheat and planted more or less at the same time (they ca n only feed on barley to lowland rice Early planting of lowland rice rice during the milk to hard dough stages) 2) they lack nurseries spreads these viruses but upland ice has remained aestivation sites and 3) weedy fallow areas are limited uninfected The lowands of Asia are dominated by L oratorius

16 IRPS No 123 January 1987

which lays its eggs hign on tile foliage (199) L acuta and L solonlonlesis oviposit at ground level the former on litter and the latter loose on the ground These egg laying habis explain theirtenvironmental prefcrencecs(214) In Japan the postdiapause adults of L chinensis migrate from moun-tainous areas to upland ricefields during floweiing (1 14)

Pentatornid bugs prevail in Latin America (8182 195) and in Africa pentatomid and coreid bugs arc equally cited (6 82) l)amage is often characterstic of the species (5)rcoris ventralis and Mnida spp are the common

pentatomid seed bugs of Philippine upland rice but they prefer lEchinochva spp grass seed to rice Vearaviritda occurs worldwide and hoften more abundant in the uplands(54) We saw few reports of seed bugs on upland rice but see(d bugs apparently opelrate cqually between by land and upland rice Good dispersal is necessary ior species that call feed for only 2-3 xon a crop and must first find those crops at ripening stages

Other pct species feed n flowers In Mato Grosso Brazil (194) and Santa Cruz Bolivia the panicle weevil Aeo-hariuia amllitarsis attacks rice during flowering (68 233)The larva e rice sten horers adutlts feed at the bases of spikelets caIusing empty grains Also inLatin America thecydnid -lkiituttatralusisa seed bug as anad tilt and nymph (194)

A number of beetles ecd on poilen - coccinellids lialrotia Aulac lhora and Monohpl-a (55) - but becatise riCe is self-pollinating and fcrtiliation occurs before pikelets open they pose no threat to yield

IST ECOLOGY

We now look at characteiistics of the life cycles of upland rice insect pests to lea rn hew each plays an adaptive role in upland rice ecology

Life history strategies Orzasativadoes not tolerate drodgh well so upland rice is highly seasonal normally grown in the wettest months of the year Usually rice is present for less than half of the year The rice-fiee fallow poses serious problems of survival to upland rice insects which have evolved att least four mechanisms to ovrcolne tle cvcl ic-l lack of a hostI) polyphagy 2) longevity 3) dormancy and 4) vagility (dispersal)

Poldt)agy lolyphagy is defined as having hosts of at least two botanical families oligophagy of more than one gents and ionophagy of only one ventus (40) - inour case Orvza Most wild rices however are illi tiC and theieiirc are normally far removed from upland rice hahitat othewise more would be alternate hosts for uplanJ iice insects Thte highly mornophagous rice pests shySciriol)haita hiceritlas Nilal)arvaiahiens Iand Nealo--tettiyvircsctws in Asia and Aalirpha secparatella and 1ioj)sis lngicornis inAfrica - occur on upland rice but

are more abundant in lowland culture They can specialize in rice because they are highly vagile and can attack all growth stages By specializing in rice monophagous rice pests can outcompete related but polyphagous speciesIntensive andI extensive lowland rice culture has favored monophagous species Riqwla alhinella (Cramer) the onlymonophagous species in Latin America is rarely reported on upland ice (I1)

Because of its limited temporal and spatial existence upland rice has not favored the evolution of specialized species Clhang(44)shows evidence that upland rice has only recently been cultured by man from the lowlands thus pests have had little time to fully adapt to the crop All but the lowland-adapted insect pest species have alternate planthosts to rice Root pests with limited mobility are highly polyphagous Pests that specialize in one growth stage eg secdling maggots seed bugs are oligophagous or polyphagous

Most alternat hosts of rice pests are annual grasseswhich also are seasonal Upla_rice is also a host but not the preferred one of the most highly polypliagous spcciessoil pests grass hoppers armvwarrs and Vearaviirielda

The d ry season poses igreat obstacle to ani upland riceinsects survival and only sonic perennial plant hosts are suitable food during the off-seasoin Plolyphagy allows upland rice insects to suLrx-ixe during thC rainy season when plants- both annual and perennial -areactivelygrowing and therefore more nutritious

Loiugevitr The ability of aiinsect to live 2-3 mo without undergoing dormancy helps species sich as root-feedingwhite grbs (as larvae) and secd-feeding bugs (as adtilt) to survive unfavorable periods White grubs arc not highly fecund and their life strategy is to improve survivorship of limited progeny Their subterranean habitat hides them from manv natural enemies lBtut root tissue is not highly nutritious so they ltilst eat great quantities and need a largeinsect biomass to digest it Other soil-inhabiting rootshyfeeding insects have short life cycles but are more fecund

Seed pests need greater longcvi vas adults to locate ahost at early grain dcclopment arnd give sedentary offspring time to develop Few groups of upland rice insects live longin active development Polyphagy and longevity only increase their ability to survive during the rainy seasonOther mechanisms are needed to survive the dry season

orantc In the tropics the nonrice season is the dry season while intemperate areas tile off-+season is winter Many upland rice insects undergo dormancy inlsummer (aestivation) or winter (hibernation) Dormancy is simplyinactivity during winters with prolonged temperatures below 10 C insects find shelter arnd cease movement Activity is resu med with warmer t empcera tu res A deep state of irnactivity termed diapauise occurs in sole species that prepare physiologically (havxing low metabolism and convcrting gonad s to fatty fuel reserves) for the unfavorable season (148) They enter diapaUse inanticipation of unshy

IRPS No 123 January 1987 17

favorable weather and resume activity after favorable Armyworms skippers and locusts actively travel hundreds weather begins usually in response to photoperiod or of kilometers in air fronts such as intertropical convergencethrough a delayed response to a stimulus such as heavy zones to descend onto upland rice fields far from their rainfall (237) Not all individuals of a population have the breeding grounds (31) Several generations can build ip on same threshold for environmental responses for entering or alternate hosts in tile grass family before rice is plantedterminating diapause -- this is seen as a mechanism to These large insects are active fliers Smaller insects such as prevent total mortality if a false cue appears such as a thrips aphids and planthoppers passively migrate long sudden warm spell in winter or rain in the middle of the dry distances by the wind season (154) Otherspecies ean readily travel tens of kilometers - stem

Probably more upland rice insects undL go dormancy borers leaffolders leafhoppes cutworms spittle bugs leaf than we realize Candidates are Atherigona spp root beetles seed bugs ants and termites aphids root weevils crickets leaf beetles thrips and seed The least dispersive species are white grabs mealybugSbugs Insects known to entervarious states of aestivation are seedling maggots crickets weevils and gall midgesleptocorisaspp adults (199) Maliarphascparatellalarvae Each species has evolved a unique set of attributes to (9) Scirpophaga innotata larvae (117) Chilo partelhtsv enable it to survive and adapt to the changing uplandlarvae (21) ScirpopThagainccetulas (179) white grub larvae environments Several evolutionary avenues lead to fitness (134) Patangasuccincia adults (6 82) and Schistocerca (Table 2) but there is no reason to believe one set of gregariaeggs (82) attributes is better than another (224)

Insects known to enter hibernation are LodeIphaDstriatellus nymphs (124) Chtio Tvlpressalis larivae (125) DroughtSesaia inferens larvae (146) but not S calavisis(2) Well-drained rainfed upland soils are subject to drought ifIDiaraea saccharalis larvae (75) Scirpophaga- tcerlttas rains do not fall within 2-3 wk Prolonged drought followedlarae(5 lacchrli s lae (75)saSlOhagasshoperseggs (82)and felo pidas athias pOpae (232) by favorable rains stimulates armyworm and locust outshybreaks but mealybtigs root aphids and thrips become

Lcpmotorisa oratorisand L acuta adults aggregate on numerous onlyafter dry spells for two reasons First heavy trees sugarcane or other shady moist sites L paluwanmsis rain normally kills soft-bodied foliage-feeding thrips and confined to Sulu and Palawan Philippines lives on the mealybugs and soil-inhabiting root aphids and root mealyshygrass Brachiariamutica (Forsk) Stapf in the absence of bugs Drought eliminates this source of mortality Second rice The adults can live for 5 mo When aestivating their the rice plant responds to drought by breaking down rate of metabolism drops and they do not feed They will proteins into soluble N compounds which enter the phloem however puncture plant tissues of their aestivation host in and are taken up by these sap-feeding insects (245) Greater search of water This behavior has led to errors of the host nutrition therefore leads to greater survivorship and range of rice bugs (I) Aestivation quickly terminates with fecundity rain probably meaning aestivation is not deep Small rice area

Some stem borers aestivate as mature larvae in rice Except in the highly mechanized large landholdings of stubble or in straw stacked as livestock feed However Latin America upland ricefieldstypically are patchy within moriality increases proportionally with the duration of the a highly diverse flora The small fields result in higher ratios dry season even if the stubble remains undisturbed If the of perimeter to area than with the typically larger lowland stubble is plowed for a crop following rice few stem borers ricefields Upland ricefields themselves may be intercroppedsurvive Rainfall terminates dormancy but it normally with cereals legumes or root crops Mixed intercroppingtakes several weeks for moths to develop and emerge with a wide variety of species also occ in tribal slash-andshy

Last-instar white grub larvae tunnel 1-2 m deep in the soil burn agriculture (50 74) to construct pupal chambers in which they aestivate The Upland ricefields tend to be small because of the labor larvae are very sensitive to dryness so pupal chambers are needed to clear land in slash-and-burn areas and to till land sealed to coierve body moisture Aestivation terminates to minimize weeds Rice requires more tillage than does with the first heavy rains but as in stem borer 3-4 wk pass maize because rice competes less well with weeds Uplandbefore adults develop rice is more a subsistence than a cash crop and small areas

Insects may respond differently in each region to cold (I ha) can feed a family temperature or drought Biotypes or loal populations may Small fields favor some insect pests such as some ant evolve (56) Local populations of Scirpophagaincertulasor species that can forage from more permanent field borders Patangasuccincta may have become adapted to prevailing Trees provide food and mating sites for white grubs (134) as conditions well as aestivation sites for LetTocorisa spp (199) Field

iagiliti The ability to disperse combined with high border grasses plus tIhose within a ricefield provide alternate fecundity orshort life cycles enable some upland rice pests to food for all but the highly monophagous upland rice better exploit the temporarily favorable upland habitat colonizers and a habitat for natural enemies

0 z

Table 2 Life history patterns of upland rice insects or insect groups

Host range Longevity Dormancy Vagility FertilityMonophagy Oligophagy Polyphagy Low Medium High Yes No Low Dispersive Migratory Low Medium High(Oryza spp) (grasses sedges) (angiosperms) (lt3 wk) (gt2 mo) (lt1 kin) (lt50 eggs) (gt500 eggs)

Ants xTermites x x x

Mole crickets x x x xx x xField crickets x xx

White grubs x x x xx xRoot aphids x x xxMealybugs x

x x x xxSeedling maggots x

x x x x xLeaf beetles

x xx xArmyworms x x xx x

Thrips x x xx xChilo spp x x xx xMaliarpha x x x x

xLocusts x x Xx x xGall midges x x xxiVephotettL iirescens x x xx x x xLeprocorisaspp x

x xLeaftolders x x x

Sogatellafurcifera x

xx x x x x x x

00

IRPS No 123 January 1Q87 19

All this appeals to favor insect pest buildup but the notwithstanding the release of stored nutrients in response contrary is true Loevinsohn (138) showed that insect pest to drought that temporarily benefits philoem feeders populations respond exponentially with the proportion of land devoted to rice up to about 75 of the area then the M POR ANCE I I P1A 1) RWI P1St S

rate of response declines In most upland rico areas tie rice crop occupies less than 50i of the area- therefore the Visits to upland ricefields normally reveal 1kw insect pests potential for population buildup is low This relationship Loevinsohn (138) and Locvinsohn ct al (139) compared may not hold for seed pests that can locate small ricefields insect abundance using annual light trap atchcs in upland

Another outcome of the highly diverse flora and phical rainfed lowland and irrigated lowland sites in the Philipshyenvironment of upland rice isthateach upland ricearca has pines Although only one upland site was studied data a unique composition of insect pests showed equal or fewer insects at the upland site than at the

For example in Tanauan Batangas Philippines the rainfcd lowland sites and fewer than at the irrigated sites main upland rice pests are rice leaffolder(RLF) armvworm (Table 3) M Lltiple regression analysis showed that the key Aythntnaseparata WBPH ants Soli psis gentitataand factors explaining insect abundance among a set of cropping white grub Leucopholis irrorata Sixty kilometers away in intensity variables were the number of rice crops grown per Real Quezon the main pests are seedling maggot Athe- year followed by area devoted to rice Table 3 shows that rigona orae flea beetle (Thacitcmnra hasalis thrips the upland rice site had only I rice crop perym rand that rice Stenchaetothrips hiurnmis and rice bug Leptocorisa was planted in only 20Vi of the area Fertiliter and oratorius In a third Philippine upland rice site in Claveria insecticide use had little bea ring on the trend The armters at Misamis Oriental Mindanao the main pests are ants the upland site applied 60 kg N ha higher than at most Solenopsis gemtinata seedling maggot Alieriona orvzae irrigated sites white grubs Holotrichia in)iUaiuoan t and Lucopholiv Insect occurrence at the upland site was omparable to irrorata stem borers Sesamia ifiArelr (hio auriciliusC that at the Cagayan rainfed lowland site parlicularly in 1982stpessallis Scirpophaga incertulas S innoiata and when drought prevented many farmers fron planting(only

4c ona chrrsurapella root aphids Tetraneura niriah- 40(of the rice area eventually was planted that year) M ore dunrinalisand Rhupaosiphunm riahbdminalis lcaffolders favorable weather occurred in 1980 and 1981 which Marasniapataltis and Chaphalocrocis Medinalis army- supported greater rice areas and consequently more insects worm MIthinna separata and rice bug Leltucorisa The most intensive site for insect pests was South Cotabato oratoriusThe great diversity of insect problems is typical Mindanao where farmers plant 23-25 rice crops per year fer upland rice and makes control efforts more difficult with irrigation The Zaragoa Nutva Ecija site also had particularly regarding breeding for insect resistance high pest abundance and at this site the fields were highly

asynchronous Asynchrony was the third most important Low yield potential Iactor in explaining rice insect abundance Most upland rice is grown on low-nutrient and mineral- Rice cropping intensity in upland areas is bound to toxic soils which when combined with eiratic rainfall remain low Only one rainid upland crop is possible per make upland rice a highly riskycrop for the farmerto invest year (Farmers in Claveria Misamis Oriental Mindanao in costly soil amendments and land management Often Philippines tried asecond upland ricecrop i 1984a ycarof upland rice isgrown far from markets making inputs even favorable rains but no harvest was possible b-cause of more expensive Farmers growing a subsistence crop hosi- drought blast and birds) Rice area isaso bound to remain tate to invest in costly inputs to raise yields low because of the high labor and power rcquirelmenlts to

Even having varieties that could double existing yield prepare the land (because of weeds)and because upland rice potential under present management levels would probably wvill not compete with lowland rice as a cash crop Some not prompt many farmers to use purchased insecticide upland rice sites with prolonged rainy seasons howeverare unless tile risk of crop failure were reduced such as it was planted asynchronously such as the Clacra site with irrigation systems in lowland rice The seasonal abundance of Upland rice insect pests

The Tanauan Batangas site is atypical for upland rice attracted toa light trap wasgrapied for crop year 1980-81 in The soils and rainfall pattern are favorable and because of Tanauan FBatangas (Fig I 2) W13P11 was he most its nearness to Manila farmers have cash resources ftm common rice hopper and its population peaked at the sales of vegetables to purchase tertilier They applied an midgrowth stage as it does in lowland rice D-Vac suction average of 60 kg Nha after particle initiation and obtained samples showed an earlier peak than that from a light trap 25-35 tha yields The high fertility isone reason why RLF indicating emigration W13PI usually emigrate after the armyworms and WBPH developed into large numbers vegetative stage Populations in 1980 reached levels that Farmers however used no insecticide on upland rice caused patches of slight yellowing in some fields WBPH although they did on vegetables probably cause hopperburn in Batangas They immigrate

The poor growing conditions fOr rice generally mean year-round as shown by low levels in the light trap BPH poor growth and low fecundity for insect pests fIeding on it however were scarce in the light trap and not collected with

Table 3 Abundance of rice insect pests and factors in rice cropping intensity of Philippine upland rainfed lowland and irrigated lowland locations where light traps were operated daily for atleast I yr -I

0Rainfed Rainfed lowland Irrigated lowlandupland

Pangasinan Cagayan lloilo Santa Maria Nueva Ecija South Cotabato Tanauan Caaringayan Caaringayan Bangag Bangag Bangag Buray

LagunaCale

Batangas Manaoag 1980-81 Santa Motiica 1982-83 Cabanatuan Zaragoza Avancena NaniamanSolana 1981-82 1982-83 Oton

1980-81 1979-80 Oton 1981-82 1981-82 Koronadal Koronadal1980-81 1979-80 1979-80 1983 1983

Insect (kerosene light trapNephotcttixcatch nolyr)virescens 650 1240 500 1800 330 210 3400 1600 160N nigropictus 210 - 360 18400 5500 2900- 140 90 20 - shy 1700 -Reciliadorsalis 710 - 3200 1700180 890 630 140 170 2400 800Nilaparratalugens 130 690 - - 4200 4600300 520 750 180 60 3300 1600 740 310Sogatellafiircifera 1700 31000 13000 8200310 210 3300 610 270 980 1600 6300 -Cyrrorhinus - 12000 6700600 540 3200 290 160 330 120 410Scirpophaga spp 50 410 620 - - 21000 12000150 180 220 270 30 1000Cnaphalocrocismedinalis 70 70 540 610 270 11000 6100200 140 100 6 60 250Chilo suppressalis 0 60 60 1900 300 200- - 30 20 0 6 3 160 -C auricilius - 0 00 - - 20 3 60 20 3 0Sesania inferens 0 - 80 60 3 80 10 51 -Cropping intensity _ -

Rice crops (noyr) 10 10 12 10 10 10 13Rice area (7c) 17 19 20 20 25 2320 85 85 60 70 40 85 85 90Fertilizer (kg Nha) 60 20 85 80 70 7020 0 0 0 30 30Insecticide (no of 40 60 40 300 04 04 0 03 300 13 13 30 50 40 33 33applicationscrop)

__________

IRPS No 123 January 1987 21

Light trapped Insects Drytnd __ 2(notrap-nght) Suctioned Insects (nom )

50 -)o 9 jO40- SOW ffurciWO Light trp

D-Vc suction

20 to 4

20

Egg porasitizahion (M) 75 60

300 SOOampReOftwciffr

45 30shy

15

-insects (notrop-night) 4 Nileporvata Iuen5

Egg parastization()

Light trapped insects (no trop-nght) Suchoned insects (no m 2 )

6 8 Cytorh~nuls hvidipennii 0

4 05 2

50 0 Insects (noltrop-night)40-ui Ipanh pe g

Moohase 1 Seasonal abundance measured by light trap

Rainfall(etod) and D-Vac suction sampler of whitebacked

to ~ hopper Nilaparvata lugens and lea ffolder60)-t Cnaphalocrocis mnedinalis andplthpe g20 L I II I I I parasites and mtrid predator Ortorhjntus livi-Apr May Jun Jul Aug Sep Oct Nov Dec JaIn Feb Mar dipennis set against cropping pattern daily1 9 80o 19 8 1rainfall and moon phase Tanauan Batangas

1980-81

the D- Vac Cyrtorhinus build up on the crop coterminous are collected in light traps during full moon (109)with WBPH buildup Cyrtorhinus disperses during the Because upland rice shares most of the lowland rice pestsrainy season RLF not readily collected in light traps was and those pest species are generally less impotant to uplandprevalent only during the rice crop rice than lowland rice there is a general belief that insect

Nephotettix virescens was more abundant than Nnigro- pests are less important onpicLus in the light trap collections and on the crop GLH upland than lowland rice Arcview of limited data on upland rice shows an averag of population peaked at the reproductive stage as did zigzag 10-21 yield loss from insect pests on a par with that ofleafhopper (ZLH) Crtorhinus tracked the populations of lowland uee determined by the same methodology (132) planthoppers and leafhoppers Both GLH species were (Table 4) McGuire and Crandall (144) made a similarcollected year-round in the light trap indicating immigra- estimate for Central America Cramer (51) howevertion from nearby irrigated rice areas Possibly because of the estimated 35 yieldlow populations loss in South America based on ainsect number did not correlate with similar pest complex - stem borers and seed bugs shymoon phase although reports indicate that more hoppers attacking irrigated rice in North America This estimate did

22 IRPS No 123 January 1987

Light trapped insects D~~nd (no trap-night )

St plht Suctined insects (nom 2

)

D-Voc suction 1520

1o

0 Lih trap 5 ~105 o5

00 Eggparositizotion()5040 NephOtgtha virneens

30 20

I010 A-Insects (notrop -night)

4 Nephatelh nigropccent i

2 o I vV^J - -- ^ VA____ f _

Light trapped insects (no trap-night) Suctioned insects (no rnE

10 -1208 ReCdO doRsol

6 15

4 10 052 iwxALAiamp 0

Light trapped insects (notrap-night) 2Suc oned nsects (nom )

8 CyrtorhmnusI6o nn]10 i0075

4 05

2 0 0

025

2 Seasonal abundance measured by light trap Roinfall (mmd) Moon phase and D-Vac suction sampler of green leafhoppersNephotettix virescens and N ngropicrus and I zigzag leafhopper Recilia dorsalis their egg 60

parasites and micid predator Cyrtorhinus livi-dipennis set against cropping pattern dailyrainfall and moon phase Tanauan Batangas1980-81

40

5 Lf Apr May Jun

1980 Jul Aug Sp Oct Nov I

-aMar

1981

not account for soil pests and was not based on field data in the absence of pests Carbofuran granules are phytotonicAkinsola (I1) cited mainly data from irrigated rice areas in to rice (239) The prevailing hypothesis is that carbofuranAfrica but estimated yield losses of upland rice between 15 prevents soil bacteria from consuming fertilizer The phytoshyand 30 from insects However losses in Thailand over a tonic effect is through greater availability of fertilizer to the4-yr period ranged from I to 13 averaging 5(106) plant Denitrifying bacteria are known to be more active inYield loss data however should be interpreted with aerobic than anaerobic soils therefore it may not be merecaution (184) The Philippine data were all derived from coincidence that the highest recorded yield losses (22-69)trials on farmers fields under farmers normal agronomic occurred in those trials with 12-30 kg basal Nha On themanagement practices except for Bukidnon and Capiz other hand contro with carbofuran granules may not be aswhere researchers used fertilizer Batangas farmers are good as a seed treatment against seed pests and perhapsatypical because ofhigh levels of fertilizer used The trials in yields would have been higher if a systemic seed treatmentBrazil were carried out at an experiment station but insecticide had been used conditions were similar to those of local farmers The 1979 and 1980 trials in Batangas had two full-

Carbofuran granules were used in every trial as a broad- protection treatments one using carbofuran granules andspectrum insecticide applied basally with the seed An the other using bendiocarb WP as a seed treatment Plotsobjection to using insecticides to measure yield losses isthat protected with bendiocarb yielded higher than those proshyinsecticidescandirectlyorindirectlystimulateplantgrowth tected with carbofuran granules The other trials in the

Table 4 Yield losses to insect pests and yield responses to insecticide in upland rice in the Philippines Brazil and Upper Volta

Yield (tha) Yield loss (5) Location Year Cultivar Protected

with Protected

without Check Total Vegetative Reproductive Ripening

Basal N (kgha) Reference

carbofuran carbofuran Protected No with carbofuran

carbofuran

Philippines Pili Camarines Sur Tanauan Batangas

Pangantucan Bukidnon

Dumarao t-apiz

BrazilGoias Minas Ge~ais

Upper Volta6

1975 1974 1974 1975 1975 1976 1977 1978

1979

1980

1979 1980 1979

1977 1978 1979

Bursiging puti Kinanda Dagge Kinanda Dagge Dagge Dagge Dagge UPL Ri5 Dagge UPL Ri5 Dagge UPL RiS UPL Ri5 UPL Ri5 UPL RiS

IAC47 IAC47 IAC47

14 33 11 36 35 31 30 27 37 26 40 30 40 42 46 35

19 21 09

--

-34 24 24 27 --

29 43 31 42 -

47 25

---

10 28 05 27 27 30 26 29 33 28 33 29 43 23 36 11

15 15 06

29 15 55 25 23

3 13 -7 11

-7 18

3 -7 45 22 69

24 28 35

--

21 -11 -20

4 -

-3

23 7

-2 -23 56

-

-

-__0

6 --

-

-23 14

5 0 8 2

22

-

--

0

-

--18 11

3 -10

4 10

7

--7 -

14 -

-1 6

10 -5

8 9 7

-

--

0 0 0 0 0 0 0 0 0 0 0

30 30 30

12 12 12

99 99

100 100 101 101 102 102 102 102 103 103 104 104 104

66 66 66

Farako-Ba

Mean

1977 1978

IRAT10 IRATIO

38 66

--

40 43

-5 35

21

-

10

--

--

--

- 242 243

Z

tJ

-w

24 IRPS No 123 January 1987

Philippines which had two treatments offering protection In the 9 Philippine trials where yield losses were measuredwith and without carbofuran granules relied on further by growth stage 10 of the yield loss (21) occurred at thecontrol with foliar spray and not on seed dressings The vegetative stage and 517 occurred at each of the reproshycarbofuran-protected plots had an 18yield los compared ductive and ripening stages Ibis result is similar to thewith only 10 with foliar sprayS This yield difference is pattern of yield loss in lowland rice measured by successiveprobably due to differences in protection against sown-seed treatments where the crop is unprotected at each growthpests (foliar spray do not control sown-seed pests) rather stage but proticted during the other growth stagesthan phytotonic effects became basal fertilizer wa used in However the insect complexes r sponsible for vegetativeonly 2 of the 10 comparisons stage yield losses in upland rice are entirely diftfrent than in

The recorded yield losses were not related to yield lowland rice The key vegetative stage pests in lowland ricepotential (Fig 3) One might expect higheryielding crops to arc aquatic whereas those in upland rice are sown--seedhave higher percentage of yield losses but this ws not tile (sown in soil) root and seedling pests This dift rence maycase One reason is that insect damage exacerbates plant explain why agricul torists have overlooked tile ilmportanccinjury from drought stress low yielding fields from d ought of upland rice insect pests Many reports of upland rice pestsstress would show disproportionate differences between in Asia ha Ve focused on major lowland species (46 108)insect-protected and -unprotected treatments Shoot and It isalso surprising that tei vegctative stage of upland rice root pests would be more responsible for losses from would record the highest yield losses becatuseseeral studiesdrought stress than sown-scd pests A second reason have shown that upland rice can readily recover from foliageconcerns seed pests which would have a greater impact on reinoval during that stage (117 152 165 188) Upland ricelow tillering low yielding varieties such as )aggc Low poscsses a hitgh obilitV to compensate for ca rly loss oftillering varieties nnot fill in the space created by removal foliage and will even be stinulated by foliage removal toof seeds oweer 1) R i 5 is high yielding because it tillersP1 produce a higher viel than plants ithout any foliageactively which can close the canopy to provide greater weed removed (63 192 231) [ie reason for this apparentcontrol discrepancy hovce is that yield loss at the vegetative

stage does not octUr frol leat arca loss but fron pestsYield loss () removing sown seed and feeding on devcloping tillers and roots -- damage that cannot be readily conpLtnsatCd for

70 - Measuring yield losses in upland rice cnvironrnents is further complicated by a report from Brazil that greater yield losses occurred in response to better insect control60 - =-09 as Insecticide seed treatments resuled in denser plant stands

(n=21) which in turn created a selectively more favorable environshyment for blast (69) This resuIt was parlticurla rlxexacerbated50 when cirbofuran granules were used in seed furrows Aside

ffrom a greater plant stand ca rbofuran iay have auigmented the N lecl and stimulated blast If so ca rbofuran should be40 - replaced with another insecticide or the seeding or fertilizerI rates decreased

Other methods to determine the impact of upland rice30 shy pests wvcrc to interview farners or to develop single speciesbull bull correlations of population levels and yield loss Feneira (66)2 NIM)st (75(1) Golas farmers reporteddid both in Brazil

20- insect losses in ufpland rice and 251 said those lossesS avcragcd 37(1 [his result is consistent with tie yield losses 0 0 measured by field trials in the same statc(Table4) For every

I0 4 of seedlings removC d by leaf-cutting ants a I 7yield loss is predicted Similarly Ferreira (66) found that ai

0 - average of 53 thrips per panicle before panicle initiation meant 3 times more unfillh d grains

-IO0 I 1 1 I I-101 0 2 3 4 5 6 7

Yield (tho) C)N [ROt IM FI IODS 3 Correlation between upland rice yield and yield lols caused by Upland rice insect pests may be controlled by culturalinsects measured in 21 trials in the Philippines Ivory Coast andBrazil genetic biological and chemical methods

IRPS No 123 January 1987 25

Cultural control dry season plowing will have little effect on them Plowing Crop husbandry to reduce upland insect pest populations when the soil is wet from the first monsoon rains will expose can be classified into practices effective in a single field and soil insects that tunnel close to the soil surface to predatory those effective only at the community level Cultural birds chickens dogs and even man (field crickets nole controls should come first in a pest management program crickets and white grubs a re delicacies in man tra ditional They have broad stable effects as pests have little pos- diets) sibility of overcoming them through selection of biotyps The greatest degree i1 inIscct control fron tillage comes (138) On the other hand some cultu ral controls work only from plowing soon after rice ha rvest when most soil insects with high labor or power inputs and connunity-wilt lie close to the soil surface anrd stern borers are in the rice methods require coordination of iany farniters stuble Whitegrubsare mature and tillage exposesthem to

Pantin time The luxury of being able to shift the time of predators Tirnelv control will prtec the crop following pla ntring upland rice is open onl hr la rniers in areas with a rice and lowr the populationIfor succeedinig years Plowingprolonged rainifall pa ttern lI copping intensity or underthe ice stulble helps it decompose killing sten borer rnechanied land preparatiot Gcnerally planlting as soon larvae inside as the rFiinv season begins will lower populatiois of 1110st [requent tillage destroys ait nests bit not termites insect pests Insect populationsarelo aftcra dlvseoll or Tillage and clean culture howes er rcliove food for winter fallow delayed plaiting lets them build tip on termites which then mtay attack a young rice crop Scarab alIternate hosts that gi vigorotsly with the first seasonial beetles also prefer to miposit in eceitly tilled fields Noshyratiris This first flush iof weds cros lItxuttriartlv with iceCult is dibbling faors the survival oftillace Clre Such rineralied N released during the dr season aid made stils i aels tnd insects thtt pttpate or lay eggs- in the soil

available to the plants with the rains It iliso igrtmitcally Zero tilla hio eerl conserves predators ofthe GIH (37) advanttgeous for the rice ciop to tap this trlLItlaI fertility nt it mcttoi lo lessen the impact o0 soil insects and grow vigorotsly to compete with wrus and to tolerate brotdcasting seed or planting in ftltrrows is prefc ired over pest infestation dihlirig in hills Crickets are large enotlgh to eat seedlings in

Larly plantngs terd to escape seedling maggolts Upland a hill atrd tillering normnally ca iriot fill in tlhe missing space rice in Bitangas is planted in Ma and ine itli the Iirst Oftcn dibbll rd is not well covered and iseasily found by rains Fairners are highly mtotiated to plant early tocscape foraging insets ioIdents and birds drlitght and typlotns ricar harct Arn Atrgtst planting Plan uit sii Incrcatsini sced density protects tile cropwould be evcely attacked by It riim orivzo At tti againrst seed and seelling pests especially ants particularly other Philippiie tipland sites Rea l Qteot antd (laelit if the crop will germinate and ermerge quickly M isainis Oriental seedlig mnaggots arnrige ric boe t se hr trftinll itI India rice intcrclopped witi cotton or farnes have a longer growing season and plnit 2 rin or pigeon pci had lower 6IIt and WBIlII poptrlations than mote after the ortset of the rainy season liaaiangas ftrmners rice alone (211) NfailC arid Urplald ir_ 2re coilltori in the plant ca rly becia use ofa shorter raiinfall period arnd the desi re Philippines he chatngC in pest status folii iniercropping is to lliarvet before the typhoor sea son peaks ltighly locatioll-specific (133) cmianarid the net elfc be either

Early plantings cornbind with early-iatiuring varictics nil bcnelficial ortdetriential (189) For example BIatangias provide higher crp tolerartce for white grubs Aestiatior farmers intercrip Itro in their ipland ricefields A liornshyterminates with the first rains but 5-6 moare ineeded for the nrii iotd-Ahis i (I iiiacius) which norlially feeds resting larvae to develop into beetles emerge and lay eggs on taro ill feed or miarbv rice plants Intercropping and for larvae to reach the danriigii g ird-instar stage probably would not afft0 seed pests A crluanion crop (134 In lataigas this normually iseiough tiite for harvest planted ith rice and haing gleater tolerarnce for root

It Brazil howecver delayed planting is a suggested dariage or being toxic nayact isi trapcrop to root-feeditig cultural control for the lesser cornstalk borer and insect pests Acial-fei i ig pests voultId be most affected by pests favored l drought (66 195) Cropping internsity in intrer ropping Shoot aphids leaf beetles sten borers and Brail is low and delayed plraing places the rice crop in the hopplers would be likely ca ntidates to note effcts On the niost stable rainfall period reducing likelihood old rought othlt halnd the corlipalior crop stch~ sitniaie Iiiv be more

In rilniy arca s rinmirsare roi atl to plant other crops benehited than rice from initecropping ( 137) before rice and althNgh it woultl be desirabh to avoid 11echi Weeuling luring the first month after clop infestations by planting rice cuatl lariners lack the labor cstablishent will force pcts such is anyvworris which and time to do so ol a low priority crp prefer grassy cetds ontor tle vorling rice Soil iampsecs will

77age Uplild rice Stils ae light textured and Call be mvnye to the rice crop Cleani citrltirc also lay lore territes plowed duiring the d rv seasor notl only to grin time for to attack I u cetincti inrice ceds i ay he le optinal planting with lie fitst Inilsoon riiains but also to desiccate solition 13) soil insects arid weeds such is ntsedge that foim rhizomes Irtlizr Itneatsing N Iciiliier oin hlwland rice has Htowever becta use most soil insects lie below die plow layer fAtwvored Irigher populains of pliithoppeiv Imlhoppers

26 IRPS No 123 January 1987

and leaffolde rs However because of the need to temper fertilizer use on upland rice o prevent blast fIrtilizer rates will be lower than is comonia r for lowlad rice causing relatixely less hopper and IaIfTfoide r population buildup However inupland rice areas without a history of blast such Is Batangas farmers apply high rates of N and conscqtrently the crop has high infcstation levels of WBPI]and RIF The fhrmrcrs leec insect buildup by splitting the fertili11r in0to two to three applicatiois bit do not applV fertilizer at the vegetati estage because this prodrces a tall crop which illeasily lodge during typhoons Greater crop fertility leads to greater insect survival (better nutrition) larger insects (to cat no re foliage) arid greater fec titd t (to lav nmre eggs) (128 145 171) Not all insect species respondequally hovr Increasing rates (if N l anrid K increased Dialraa s-acchralis and (iaeiurema flea beetle but reduced thrips (66) Zn rcdLuced deadheart density from the lesser cornIstalk borer

Fnldingi Although most Upland rice areas do nIot hawxcaccess to wa ter floodiig the fields is rRrrnriierdcd in areas of Brazil or root bug and white gib control ( 195)

Crop rotatioti Tillage before planting a crop after rc harvest will control stern boreis and other pests renain n the stubble arid will unearth soil pests Planting a non-gratriiaceous crop after rice is recommncrded for termite control (66) Crop rotation is less nttdcd in upland rice culture than in lowlands because atcrop-free period is assured even after harvest of crops following rice

t1ulchilt iryl plastic irli irig designed for weedcoitrol ii upland rice inJapan resulted in ntr highanri liy infestation of (ihilo sulytresvais (93 9-1 95) NIulched Upland rice grew more luxuriantlv and attracted more (Avipositing mothis than u lched rice Also the rice plant and stein borers imatoured 10rC quickly and a partialgeneratiori af vouing ista iNoverwintcred in the stubbleLotili--malitrt varieti Quickly maturing crops reduce the tmbCr of pCst generations that Calt build up Rapid crop establishment olan ca rl v-natu ring vaitvty if)pa 11icLa r will lessen white gib and stem borer damiage White grub larvae intle last iistar are highly destIruCtive so aii early-iiatri ng variety will escape serious root loss Stem borers build 1ip slowlyx in rice to ecormic abundant in a late-Maturing crop anrd caiuse whi tlieads Numibers of somespecies increase exponeitially with each succeeding genera-ti so a xarietyexhat atures I trowh earier thian another will hac less insect damage Early-naturing vaiicties however will not lessen seed pest daiage either ii the soil or oina standing crop

Strwhoi hAIJttam Inrscts tcd irig on thlie aerial portions of rice plats disperse from carly to late plantedfields AS iiiibCirs increase exponentially with eachinsect

gtscnrattm e eiglbrig f rmc~s1(uil upp wh rc generation pests build tIP where neighboring fihrTC e9r1sstagger their plantings II Batangas farmers plant uplandrice within I iro use 120-d s-arieties and plant at the start of the rainy season Ini Misarnis Oriental laricrs stagger their

plantings up to 3 mo apart and use a set of atieties that mature in 4-7 mo Insect pest danage particularly from stem borers is greater in Misarmis Oriental Also seedling maggots damage rice there because rainfall favors frequent planting of maic In Batangas rice escapes seedling maggots bCca usC of distinct drs season and synchron0us early planting with the onset of ains

Therefore combinations of cultural practices ea rlyplanting synchronous planing crop rotation and earlyshymaturing vaneties protect the rice crop against most insect pests

Ilmant resistance Beca tse upland rice has few specialized insect pests and many insect types attack the crop worldwide regional breeding and strong internatiornal cooperation are required

Sown-sced pests arc economically controlled by insectishycide seed treatment and should the relore receive lowxpriori ty inbreeding objectives

Root pests are normally cost ly to control with insecticides but finding resista t sonres mayvbe di fficult Tanaka (229) for exa -ple fi led to find varieties resistant to root aphids Selecting for large root bioniass perhaps should be the strategy for pests sLuch as Whlite grubs that renliove roots Fainter(174) howcvcr cites varietiesof crops tolerant ofor resistant t(i root pests Some sorghun varieties are high in cyanide so rice sorghun might incorporate broadshyspcctruri chemical resistance

Seedling vigor and drought tolerance are high prioritybreeding objectives so tupland rices should have large r root systernsand be able to tolerate ligherlevels of root loss Sonie root loss inolder plants is beneficial if it stimulates new root developnrut and therefore crihances intake of soil nutrients (53) The rice seedling inaggot rtiLotia spp may be

controlled thirough resistant aittics A hi rgc breeding effort on sorghniu has had fair success inideveloping arreties esrstait to Asoccata Rodani (251) A field trial comparing 10 upland ricc varieties showed differences aniong varieties va ring from II to 4(- damaged tilleis (215) Hovver resistarce to 1Iill-0r gtia species may rot cross ox-rti otier

Shiraki (210) reported that upland i-ice stens wcrc harder than lows-land rice stemsperhaps becaIusc (f a higher silicoricontent and thus were more resistant to yellow stern borer hc tortia Iow tillering (f uplaid iice xarneties prodtres a

higher percentage of infested tillers than would occor in higher tillering lowland iice under a siiilar egg density The corollary therefore is to select for high tilleriig upland rices

Stemii bor amcstheh main targes dobrecders inAfrica (571 204 )but nticl oifth e sc reen irig is dotie unde r lo wland co d t osrm s wih A h r~ o l pe i s n acorditions (11 96 218) As with lthrinrnwspecies mayhave to be dealt with independently however initial results show cross resistance between African and Asian species(92)

In Brazil screening Ior insect resistance focttses oil lou I pest problems Varieties tolerant of lasmolwalpus ligi-sellus and resistant to )iatraea saccharalis have been identified (70 153) Current work also emphasies 7iraca limativetntris and Thmialspis (=Iosis) flavouicta (66) These insect pests Ilt onlV citise IIigh yield losses tihey also ire difficult to control chemically ofr by other ilealls

ManV upland rice vaieties aic early maturing and are resistant to pests such as gall midge (I Xl ) Early mIturity is often a highly desirable trait to esuilpe pest buildup but it is not true resista nce bca usc ilthose va reties are plinted late they will be itsdamaged as ain early planted susc ible variety

P~erhaps tile cosiiiopolitall of all upland rice pestsimost are the sced pests (ripenling stagc) which pose a particular problem beta use scuds re ised IsIO for hitnit4ans Ayltoxic substatice introd tcld geticticllyilmay also be toxic to nal iniless it caln he placed old in thfe IdhuffSelectmi fo hairiness or lotng wns kill mIlct sthilrohleiiis dutilig threshing pa rtictfarl maitial threshinfl [hireshers ill complain of itcfimin atid itteascd (fust levels

Attempts itIRRI to fitid tcistatncc to the rice bug plorisa have ilotbeCnl prmitisilg lIIetIattonlids iorc

through the lenilna or palca hillhairs sedstVIor seeds wilh awns that deter bugs promote complaints from threshers Seed bugs liptocorisa and Stcnocoris enter the seed through the opening between tile lemna and palC but varieties with narrow openings are diifficult to mill

Breeding priorities should concentrate ott inotCI gruLtLIps that are most difficult to c nttrol by other meains Steil) horers appear to be the most widespread group that sigiiificantly dalag s rice aind is hard to manage by other Ilnca us

Biological control Very fe stutdic of mitatt~l eneties of rice pests in upland envirollnletits wee in tile wefouiid literature Therefore have not compiled a list of species Ve present a list of natural enemies iOtid in upland ricefields tuic latialln Batangas for some conitno iOldiar pest species ([itble 5)

Muich ofl thc lollowinle itfor ittiol cotlliCs rol Stutdies in the lowlatnds or roi crops otithcr thlin tice

Qu(ailtr o 11ll t11t( Nat trrtf lii hcoeits ight Ile used agaitist ile social insects tilt and termites Sollie predator specice live ill theirf nests n ilnic theirnllt ip-pearanIlCe aMRidbehavior IJtrouMdCing a mote aggressise ant species to displace pest ant codonies is a possibility (169) Patihogens as potential natural encnies have reccived little atteition (I13) Natural ncnuics asa group however hac not been toIl iporta it in Iegulating aintshown be and ternite numbers

The nocturnal behavior aid subteitmien habitat of lole crickets and field crickets protect them from many ntllral enemies There are only limited records of natural cnelics for these groups of pests some pathogens (87) but mainly

IRPS No 123 JattuIny I 271987

parasites - scelionids on eggsand sphecids on ilymphs and adults Sphecid wasps Larra carutmaria (SImitI hI L huzwuensis Rohwer and lsaltiuhta Willi ams speciali in mole crickets pa ralyzing them nd dragging the ninto thcir nests as food lot their voting ()tier sphCcid species Lins aurulenoa (Iabricius) lot maiac (Ashmead ) 1 lhshylssellaus (Smith) and Ml lal)oriosu (Snithf specialie in field clickets

White grUbsO the other hand hae Icti scriuns enuigh to have been extensively studied for cotttroi 1W nati al enemies( 112 140 141) Scoliid parasites C(ajsoMcispp have beetn intrtduced itto tilePhilippites for control of Ielxl ]udisiro la The rcsearchcts w Intttrdced themle during ill era claimed that the white gttionthreak as controlled (142) howevcr it isnormal for pest epidemics to subside and there was no diect evidence to indictte that tfe parasite was responsible A tmore promisitg titctllod might he to identify iso ltcttass prt~f tcartd apply hiactelaln fingal pathogens to tile soil Ihe lalattese beetle in North America has hcci coitrolled hv tite conmnleciall aailable hactcrial prcpa ration itacihlpu pav ofr milksdiscas (71) Spote prtyefott the soil itloted ilrtier reitiain actic fr Veas

Root aphid sardtnctly bugs readilslic tilib to predatots and parlsites once their tetndine ants are controlled The larvac and adtilts of a coccinellid beetle 1Scynms sp prey on llriadlouais in slash and burn upland rice Baits treated with insecticide may be used to control tending ants by killing their ourig (244)

Root-tkcditig blugs false wireoris wireorms and root wccils aic r ofsuch little ifilpoltatice that thenomttll role of liatural Cticniics hias tiot beei asscssed A possibility hoevrl tiglt he tit ItaIIitic niaCIMtodcs lintt1g11en Itese soil-itlhlfbitine mCtitodcs W5uhtfhave to be imiass Produiced

large miltibcr of niatltrll cnemics ircknown ft CltIoriS IC IpblCeii is thtt clttvortnlls ofonlie the crop soott alter land ptel)tritiOti a1tithe bcgit iniig of the raittv

Citsoln hfien tiatllral leniiiy populations aric lo Colltloil WOtuld iii ole mass prodtticing andirclcaslng key spccies

le egg stage is the fpirt (olthe Cedlitgt Ialot life cycle tI hich to f)ctis ftor tittlril elctit cottntrol hplraites o1 pleditors ttsit elleggs highly expoIsYCl I Sgg pailsites or predators stolld probahly e ittass andtbe produced released Littlh i toruialioll is a ailable ott what species to trV o ho to Iniss podLice thiCIi A etiloplhid wasp Icmihlaol sp effctiicly regushy

lates the leatf lcloalapolm2a spioanoiiitr heat andc rice antid should be collsctvcd

leaf beetles whCllose e am] eggs ate laid in the grutindhirO ould be motC difficutlt to control by niatulral enelV Mnll ipuhltiOn But species sUici IS c_gsthispid beetles ith

and larVle oilleascs ate vtilcratble to iparaites and pledators

Natural enemisc of tlirips otl rice are little known

28 IRPS No 123 January 1987

Table S Parasites recorded from upland rice insect pests in Tanauan Batangas Philippines 1977-84

Order Family Species Host stage

I lymttenoptera Braconidae Leaffolders Cnaphalocrocis Marasinia spy

Cardiochilesphilipplinensis Ashmead a Larva

Cotesia angustillasis(Gahan)Cotesia or cypris (Nixon)

Larva Larva

Cotesia or taeniaticornis(Nixon) Larva Cilonusinotakatae Munakata Chelonits SppAlacrocetrts philippinensis (Ashtmead)

Egg to larva Egg to larva Larva

Iclineumonidac Orgilts spp Isclutojppa hireator(Fabricius)

Larva Larva

Itoplectis naraigav(Ashmcad) Larva Tenelhtcha philippiensis (Ashmead) Larva Tenelchha stangli (Ashmead) a

irichomma cnaphalocrosis Uchida Larva Larva

Bethylidac Elasmidac

Goniozms or trianguli r Kieffer lasmtusalhopictus Crawford

Larva Larva

Diptera

Encyrtidac Chalcididac

Tachinidac

Elasmtus spp Copidosomopsisnacohiae (Fidy) Brach nteria evcarinata Gahan Biachyieria lasus (Walker) Ztgohothria ciliata (Wulp)

Larva Egg to larva Pupa Pupa Larva

Argyrophylx nigrotihialis(Blaranov) Larva

l-ymenoptera Braconidac Yellow stern borer Scirpopiaga incertulas

Braco chinensisSzeplegeti Larva CIthelotis spp Chelonus niunakatae Munakata Cotesi flaripes Cameron Stenoracon nicerilli (Bingham) a

Larva Egg to larva Larva Larva

Ichneumonidac Tropobracon schoenobii (Viereck) Amatroinorpiaaccepta metathoracica(Ashmiead)Frihorus sinicus (I lolmgren)

Larva Larva Larva

Ischnojoppaluteator (Fabricius) Larva Isotima nr dammeratiti Rohwer Larva Tenelucha philippinensis (Ashmead) a

Larva Tenehicha stangli (Ashmcad)Trichomma enaphalocrosis Uchida

Larva Lara

Chalcididac Eulophidac Trichogranmmatidae

Xanthopimnpla stemmator (Thunberg) Charops hrachYpterumn Gupta and Malteswary Brachtm eria sppTetrasticttus schoenohii Ferriere Trichorigranm chihunis Ishi

Larva to pupa Larva Pupa Egg Egg

Diptera

Scelionidac Pteromalidae Tachinidae

Tt ichograintnta aponiicuitt (Ashniead) Telenomits rowani (Gahan) a

Trichoinalopsis apattelctena (Crawford) Jeirhaea spp

Egg Fgg Egg pupa Larva

Z4gobothria ciliata (Wulp) Larva

Ilymenoptera lecneumonirlac Gol fringed borer (hilo auricilius

Trichomm enoptalocrosisUchida Larva

Ieioenoptera Braconidae

Aantholimlh steiniator (Thtinberg) a

Pink stet Ioret Sesamia inferens StoInacoil nicrilhi (Bingh am)

Loarva to pupa

larva

Diptera Ilyinenoptera

Armvtnyorms Mythimna separata Spodoptera mauritia acronyetoidesTachinidae Zvgohothriaatropiora( Robinea i -Desvoid y)Eulophidae It-phctrtiscitapadoe (Ashimead)

Larva Larva

(halcididae Braconidac

Brachpin ria spip (oesia spp

Larva larva

IHyntenoptera I)iptera

Rice skipper Ielop ida (halcididae Tachinidae

math ias aidgreen iortedci ciliarNIelan itis eda Brat lhincria sp iii matuqitatt Cameron Irgyrolhirlaxi trotlhiais(marumtv)a1

ismtne Larva Larva

Diptera Ilymenoptera

Tachnidae lelineutionidae

fuown se tilo per Mois frugalisIrgtrophrlax 1igrothiahiv 3 a rat ov) anthopimpa pmnctata (Iabricius)

Larva Larva to pupa

aMost dominant

IRPS No 123 January 1987 29

Normally predatory thrips prey on herbivorous thrips strepsiptera on both Spiders dominate tilenymphaladultNatural enemies of stem bugs --Scolinoplara and Tibraa predators Spittle bugs move to rice from pasture grasses - are in order of importance egg par~isites and predators and biocontrol would be successful only ifcarried out before fungal pathogens and nymphal adult parasites Egg tiley immigrate into ricefields parasites could either be introduced fron other areas or Biological control tactics for hemipterous seed bugs mass produced and released ould follow those outlined for stein bugs

Armyworms are normally lwld in hcck by the activities Coml)arison of environnewtns Natural enemies are of egg and larval parasites When these natural enemies fail perhaps faced with even greaterconstraints than insect pestsusually because of drought armyworns become epidemic to survive ind reproduce in upland environments TheyIt may be worthwhile therefore to release parasites during have adapted ani survived by having wide host ranges the the rainy months following a drought Virus diseases of ability to aestivate or hibernate over unfavorable seasonslarvae would beanotheravnue to exploreforarnyworms and the ability to disperse or by other mechanisms

Parasites have ben traditionallyconsidered for biolegical Comparing the most dominant and therefore the most control attempts against stein borers usually by rekasing adapted species of natural enemies collected in various exotic species( 166) This approach has not met with success environments in the Philippines we can understand how and new avenues should be explored The role of predators different rice environments are arid pathogens is little inderstood and deserves greater Anagqrms olptahiisa mymarid planthopper egg parasiteattention Egg parasitism rates are normally reasonably is cquallyadapted to all Philippine rice environments (Tablehigh but should be supplemented with effective egg 6) It is selective for planthoppers and even parasitizes eggspredators Orthopterans are potential egg predators of the n-iize planthopper Peregrinitsmaidis itt upland areas Metioche vittaticollis (Sta ) Anavipha nifiwnnis (Saus- Alavohis also parasitizes maize planthopper eggs but is sure) and Conocephahs hongipnni (de i-laan) feed on most prevalent on BPH in the irrigated wetlands (onashystem borer eggs etioche and Anaxil)ha specialize in eggs tocerus spp like A optalhilis on BPH and WBPI Iattacks with no hair such as Chilo spp The omnivorous (ono- the eggs of GLH in ell rie environments It is also found cephahsspecializes in eggs cov red with hairs it is a proven parasitizing the eggs ef the white rice licahopper (oimna egg predator of Scirlouphaga spp but has the discouraging spetra Oirosita naias a trichogrammatid egg parasitehabit of eating rice guin (65) attacks BIPH and WBPH in the wetlands and has no other

Grasshoppers have egg (scelionid) and nymphalad ult host A related species 0 aesopi specializes in G1-H in (nemestrinid tachinid and sarcophagid) v irasites Among wetland environments Dryinid strepsipt-an and pipunshypathogens protozoans have been recorded most frequently culid nymphal and adult parasites of planthoppers and Control of locust species by natural enemies could con- leafhoppers also specialize either in hopper species or centrate on their habitual breeding areas environments Among the dryinids Pseuhdgonatoipsflavi-

Leaffolders have rich complexes of natural enemies some feimur prefers the lowlands and BPH while P nuch has no adapted to upland rice (22 27) ranging from egg predators environmental prefirences but is adapted only to WBPH (gryllids coccinellids) to larval parasites (braconids ichncu- The main d ryinid species on GLH aptogonatop)usspp is monids) to larval prcdators (ants and carabids) to larval most dominant in rainfed lowlands and also attacks other pathogeiis (vira and fungal diseases) Efforts to augment leafhoppers within Amnrasca Cicadulinaand Bachaltuathese natural enemies should focus on the egg predators Strepsiptera are more dominant on plarithoppers than (rearing releasing)and viral or fungal pathogens (culturing lealhoppers The Elenchus species in the Philippines attack spraying) all species of planthoppers within the genera Sogawilla

Parasitization of the large conspicuous larvae of Sogatodels NilaparsataHarualia and Opiconsiva Elenshyskippers green horned caterpillars and other polyphagous cusrvasuntaisuiis adapted to the major rice environments Lepidoptera by tachinids and chalcids is normally low ELg but prefers BPH in rainfed arid WBPIH in wetland environshyparasites and predators are perhaps the key to their control ments Ilalictoph(utsmtnroeiand HVpectrus attack G LH

Gall midges are parasitized by pterormalids and platy- and COfimravcctra respectivelygasterids that aestivate within their larval host between Pipunculids are mainly parasitic oii lealhoppers Each seasons Shoot aphids are preyed upon by coccinellids species has a unique environmental preference Tomosshysyrphids and chrysopids varireIa oraetoraprefers Ne7 otetltix nigropictrs and

Planthoppers arid leafhoppers have egg parasites and Pilunwulhtsjavanensisattacks Dehoceplhahls spp which predators as well as nymphialad nlt parasitesand predators are more abundant in upland than irrigated lowland their rich complex of natural enemies includes fungal environments T suhvirescensattacks Nephotettix virescens pathogens The principal egg parasites are myrnarids and and prefers the lowlands Pilpuiulus nttillattls is an trichogramniatids Egg predators are norinally mirid plant upland rice parasite arid attacks only Nephotetix speciesbugs that suck out the yolk Nymphal ad nIt parasites are More than 17 recorded species of leaffolder parasites in dryinids on planthoppers pipunculids on lealhoppers and the Philippines attack species of C(naplwocrocis and

30 IRPS No 123 January 1987

Table 6 Dominant natural enemies of rice pests in upland rainfed lowland and irrigated lowland rice environments of the Philippines

Natural enemy

Host stage Species

Brown planthop-Parasite Egg

MymaridaeAnagrus optabilis(Perkins) Anagrusflaveolus Waterhouse

TrichogrammatidacOligosita naiasGirault

NymphAdult Dryinidac

Pseudogotatopuslaiifemttr Esaki et Hashimoto Elenchidae

Elenchus yasurnatsui Kifune and Hirashima

Environment

Upand Rainfed lowland Irrigated lowland

Ailaparata lugens

x x X x

x

x x

x X X Whitebacked planthoppers Sogatella furcifera and Sogatodes pusanusEgg

MymaridaeAnagrus optabilis(Perkins)

TrichogrammatidacOl~gositanaias Girault

NymphAdult Dryinidae

Pseudogonatopusnudus Perkins Elenchidae

Elenchus yasutatsui Kifune et Hirashima

Parasite Egg

MymaridaeGonatocerus spp

TrichogrammatidaeOligositaaesopi Girault

NymphAdult Pipunculidac

Tomosvaryella subviresccns Loew Tomosvaryella oryzaetora (Koizumi) Pipunclusmutillatus (de Meijere)

DryinidaeIlaplogonatopusspp

Egg Copidosomopsisnacoleiae (Eady)

Larva BraLonidae

Cardiochiles philipphnensisAshnead lchneumonidac

Trichiomnia cnaphalocrosisUchida Temelucha stangli (Ashmead)

Pupa Chalcididae

BrachymeriaexcarinataGahan Brachymerialasus (Walker) Brachymeria nr tachardiaegroup

Parasite Egg

Scelionidae Telenomus rowani (Gahan)

EulophidacTettasticusschoenobhi Ferriere

Larva Braconidae

Stenobraconnicevillei (Bingham) Iclhncurnonidac

Temelucha phiiippinensis(Ashinead)Temelucha stangli (Ashmead)

x x x

x x

X x X

x x Green leafhopper Nephotettix virescens

X x x

x x

X X X X

X

X Rice leaffolders CnaphalocrocisMarasmia

x X x

X

X X

x x x x x X x x x

Yellow stem borer Scirpophaga incertulas

X X x

x

x

x X XAinauromorphaaccepta metathoracica(Ashmead) x

continued on next page

IRPS No 123 January 1987 31

Table 6 cntinued

Natural enemy Environment

Host stage Species Upland Rainfed lowland Irrigated lowland

Larva-pupa Gold fringed borer Chilo attriciiuslchneumonidae

Xanthopimpla stemmator (Thunberg) x aPupa x

Pteromalidae Trichoinalopsis apanteloctena (Crawford) ax

Pink stem borerb Sesamnia inferensParasite

Larva Braconidae

Stenobracon nicevillei (Bingham) x b b Armyworms Mvthinna SpodopteraLarva

Tachinidac Zygobothriaarroptvora (Robineau-Desvoidy) x x

Braconidac Cotesia spp

X Skipper Pelopidas mathias and green horned caterpillar Melanitis leda ismeneF-gg

TrichogramrnatidacTrichogrannaspp x

Scelionidae Undetermined Scelionid

x Larva

Tachinidae Argyroph lax nitrotibialis (Baranov) X x

Pteromalidae Trichomnalopsis apanteloctena (Crawford)

X Predator

CoenagrionidaeAgriocnenis femina femina Brauer x xIschnura senegalensis tambur

Miridae CrtorhinsliidipennisReuter x x

Nabidac Stcnonabis tagalicus (Stal) x

VeliidaeAlicrovelia douglasi atrolineata (Bergroth) x

Coccinellidae Micraspis crocea (Mulsant) X x x

Vespidae Eumenes campaniformis (Fabricius) x

Formicidac Diaecamna spp xOdontoponera transversa (F Smith) x

Carabidae Chaenius spp OphioneaW- aJasciataSchmidt-Goebel

x

Ophionea ishii ishii Habu x x X

Gryllidav Anaxipta longipennis (Saussur-) x xMetioche ittaticollis (Sial) x x x

Tettigoniidae Conocephalus longipennis (de Haan) x x x

aNo rearing made on rainfed lowland bNo rearing made on rainfed lowland or Irrigated lowland

MrasmiaAmong the Laval parasites the braconid Cardio- lowlands and parasitizes oriental maize borer OsIriniachilesphilijpphensis is most adapted to an upland environ- furnacalis (Guenee) larvae The leaffolder pupal parasitesment It also parasitizes Hydelepta indicata a common not only occur equally in all environments but also haveleaffolder of legumes The ichneumonids Trichomma cna- wvide host ranges- parasitizing species of OsIrinia Marucaphalocrosis and Temehcha stangli occur more in the Hydelepta and Honiona - in maize and legumeslowlands T stangli is most adapted to irrigated lowlands The principal egg parasite of yeltow stem borer (YSB)and also parasitizes Chilo and Scirpophaga stem borer Telenomus rowani is widely adapted to all environmentslarvae T cnaphalocrosis is most prevalent in rainfed and has an unusual alternate host -Tabanus eggs (25)

32 IRPS No 123 January 1987

Tetrastictu schoenobii specializes only in YSB and occurs mainly in irrigated lowlands -- further proof that YSB is most adapted to the wetter environments

Larval parasites of YSB are highly environment-specific The braconid Stenohracon nicevillei occurs only in upland rice but also attacks the pink stem borer (PSB) Tieinchwa phiiiplPwnsis an ichneumonid occurs in upland rice and also parasitizes the larvae of (7hiu stqpresaisinatro-morpha accepta melaihoracica like 7 stangi is mnost adapted to the irrigated lowlands

The dark-headed stem borer(l)HSB) (ih pochit3sus was not collected in all rice environments and the gold fringed borer Chilo auricilius nd PSB were collected only in upland rice Xaantiholiplastenrnaloran ichneumonid larval parasite appears to be widely adapted whereas TrihomaslopisapanitI)cteiiaa pteromalid pupal parasite occurs mostly in wetlands and also parasitizes larvae of Chilo suppressais and Pelhuidas mathias

Armyworms Myit hinin and Spodujipera have two main larval parasites - a tachinid Zrgohothriaatropivora in rainfed environments and a braconid Cotesia spp in irrigated lowlands

The butterfly pests Pelopidas and 4elanitishave tricho-grammatid egg parasites found only in irrigated lowland environments

Two larval parasites show environmental preferences --Argrroph av nigrotihialis a tachinid in rainfed habitats and Trichonialopsisapantelkcnela in irrigated ricefields A limgrotibialisalsoattacks the sweet potato hornwormAgrius convolvtdi and T apanteloctena parasitizes DH S1B

Predators as a group show more distinct environmental preferences The aquatic damselflies Coenagrionidae are most abundant in the lowlands Cvriorhinus is also most adapted to the lowlands The nabid Stenonabis tagalicus occurs only in the uplands and isalso prevalent on legumes The coccinellid lady beetles wasps and ants are more prevalent in upland areas AlicrasIpis cit-ea larvae prey ona variety of aphids attacking legumes and maize Etunienes campanonia vespid mud wasp makes nests in trees and is therefore most adapted to the more botanically diverse uplands Soil-dwelling ants cannot tolerate flooding

Upland environments are habitats to arboreal carabid beetles Three species of Chlaeniusbeetles prey on leaffolder larvae and are perhaps more important than parasites (27)

The gryllids and tettigoniids are widely adapted egg predators not only on rice but also on maize and legumes They feed on eggs of most insects that are laid on leaves

The spidercommunity of upland rice environments isrich in species At one site (Batangas Philippines) 31 species have been recorded (23) Of the 176 spider species recorded in Philippine ricefields about one-half (82) occur in upland rice The spiderspecies of upland rice environments overlap more with those of rainfed lowland than with those of irrigated lowland rice (24) The same study showed that of the three environments irrigated rice has (ie greatest spider species diversity foiiowed by rainfed lowland and upland

However of the 10 most prevalent ricefield spiders 9 were abundant in upland rice showing wide adaptation All three spider guilds -- orb-web space-web and huting spiders shywere prevalent in upland ricefields There were however differences in environmental prelerences for some spider species

Of the orb-wcavers Tetragniathamanlihulatawas partishycila rlv abundant 7 japonica was low and Ltucatge dehcorata was absent in upland rice

As in other environments Aiypeiwnaifirtosana was the most dominant space-web spider however its relative numbers were lowerthan for lowland sites Among hunting spiders Ox|opes javanus and Lcosa leatcost fgla were more abundant and Lvcosa lpseudoanmnlata was less abundant in upland compared with lowland rice envishyroniments

Extensive rearing of rice insect pests in upland rainfed lowland and irrigated sites over an 8-yr period in the Philippines provided insight into the effectiveness of parasites as natural enemies (Table 7 8) Most of toe data are from large sample sizes taken over at least one crop which overcomes the pitfalls expressed by Van Driesche (238)

VSB egg parasitization was surprisingly similar across all environments Larval parasitization was highly dynamic among sites and even years within the same site but showed somewhat lowerlevels in uplandsthan in irrigated wetlands This would have been predicted for egg parasites as well since YSB is adapted to the lowlands and so must be its parasites This isevidence that Telenomus rowani is as good at dispersing as its main hosts Trowani clings onto the anal tufts of female YSB and parasitizes eggs as they are laid This phoresy may explain its wide adaptation The more specialized lowland YSB larval parasites therefore appear more effective than their upland counterparts with more alternate hosts

Low collection levels of PSB larvae especially in the wetlands make comparisons between environments difficult Low levels of larval parasifization were recorded in both rainfed environments Except in the Cagayan rainfed wetland site in 1980 low levels ef larval parasitization were evident at irrigated and rainfed sitesalike No differences by environment were evident Higher levels of parasitization occurred in Sarawak during a 1967 outbreak also from tachinids Rothschild (197) indicated that higher parasishytization occurred in upland rice areas

Skipper and green horned caterpillar larval parasitization rates were generally low and showed no environmental effect htighest activity occurred in Cagayan in 1980 (48 parasitization) Although larval parasitization levels of RLF were variable year to year activity appeared greater in the irrigated lowlands than in the uplands Again the highest levels were recorded in Cagayan in 1980 Cagayan has extensive grasslands and fallow around the rice areas and a relatively long rainfall period Rainfall in 1930 was good for crop growth and apparently for increase of larval parasites

IRPS No 123 January 1987 33

Table 7 Parasitization rates of rice planthoppers and green leafhopper in upland rainfed lowland and irrigated lowland environments Philipshypines 1976-840

Environment and site Year Egg parasitization () Nymphaladult parasitization ()

BPH W13P GLH BPII WBPH GLIH

UplandBatangas

Rainfed lowland

1976 1979 1980 1982

15 17 12

2(47)8(39)

4(51)

9(1800)14 (212)

12(304)

8 (101)6 (127)

9(116)

Iloilo

Cagayan

1976 1977 1978 1981 1980 1981 1982

0 (69)

11

44 48 40

-

-

46 55 44

0 (471)

11

52 56 49

0 (200)9 (214)

10 (1259)8(235)6 (900) 3(800)

-7 (462)

11 (270)3 (313)5 (900) 1 (800)

4 (600)10 (725)10 (362)12(870)5 (900) 3 (800)

Pangasinan

Irrigatedlowland

1984 1976 1978 1979 1982

13 (436) 13(379) 14 (457)

6(14)7 (197)

17(198) 11 (486) 18 (275)

3 (81)10 (274)3 (116)

16(871) 19(88)

28(211)9 (764)5 (647)7 (1204) 8(305)

Laguna 1977 11 7 21 1978 31 1979 14 4 4 28 1980 7 8 20 1981 1983

56 7(278)

26 16 (1650)

16 27(1921)

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R I Apostol unpublished 1lo1lo and Pangasinan 136 Cagayan BCanapi unpublished Liuna 175 I C Pantua unpublished 118 234 43 Carino and Shepard unpublished BPH = brown planthopper Nila parvara htgens W131Pi =whitebacked planthopper Sogatelia furcifera GLH = green leafhopper Nephotettrix virescens Figiires in parentheses are sample sizes

Tauher et al (230) reported that internal external and genetic factors influence the seasonal activity of parasitoids

Hopper egg parasites but not nymphaladult parasites were also parlicularly abundant at the Cagayan site possibly the egg parasites have more alternative hopper prey in the weedy areas than nymphaladult paisites

The seasonal dynamics of hopper egg parasites can be seen for upland rice in Batangas(Fig I 2) and for rainfed lowland lice in Pangasinan( 135)and Cagayan( 105) Levels of BPHWBPI-L and GLH egg parasitization were corn-parable in upland and irrigated lowland environments Except for some high rates of paraMsitization early in the season the parasitization rate tended to be steady in all envilonments However hopper nymphaladult parasi-tizution levels were more similar in the rainfed environments and lower than in irrigated rice

Overall parasites do not appear highly effective by themselves against upland or even lowland pests Among natural enemies parasites have attracted the most attention because they can be more readily assesscd

Predators and pathogens lend themselves better to management practices Predators the most important group of natural enemies in riccfields are difficult to quantify On the other hand pathogens appear less im-portant but can be readily cultured and disseminated

The natural enemy commulnity of upland rice is rich in species and differs significantly from that of lowland rice

These beneficial organisms must be conserved by applying insecticides judiciously particularly sprays Seed or soil placement of chemicals minimizes exposure to natural enemies The strategy to derive the greatest benefit from natural enemies isto allow the greatest number of beneficial species to thrive Then perhaps at least one species will be effective against each pest at any tine overcoming the variable abundance of each species within and between years

Programs to introduce exotic species or mass produce indigenous ones are ambitious and expensive because they require trained people on a sustained not ad hoc basis

Chemical control Insecticides offer rapid and efficient control of upland insect pests (190) but-should be used only after other control measures have been considered Insecticides are rarely used on upland rice because of cost (32) Upland rice yields are normally too low to justify the expenditure Also most upland rice is grown as a cash crop Spraying upland rice is more difficult than lowland rice because water is less accessible Government subsidy programs to provide pestishycides to tribal peoples for upland rice production have created problems of toxicity to humans because isolated people have had no experience to be able to handle pesticides safely

However ins-eticide use has beenjustified when a crop is

0Z

Table 8 Parasitization rates of lepidopterous pests in upland rainfed lowland and irrigated lowland environments Philippines 1976-84a

Parasitization (5)

Environment and site Year YSB GFB PSB Earcutting Swarming Skipper Greenborned RLF Brown larva larva caterpillar caterpillar caterpillar caterpillar larva semilooperEgg Larva larva larva larva larva larva

UplandBatangas 1977 - 0 (57) - 0 (63) - 0 (71) - 0 (106) 11(142) 2 (101)1978 43 (4697) 8(451) 5 (38) 3 (40) 6 (61) 2(58) 15 (57) 13 (129) 13 (1286) 4 (30)1979 59(4135) 12(320) 13 (11) 5(28) 9(45) 6(46) 7(23) 8(160) 9(1152) 8(26)

Rainfed lowland Iloilo 1976 - 0 (10) 0(10) 0 (7) 0 (19) 2 13) 15 (180) 5 (76)1978 0 (54) 0 (3) 0 (17) 9 (64) 4 (78) 3 (73) 0 (121) 0 (191)1979 64 (2350) 16 (84) 6 (17) 0(6) 18 (68) 25 (104)

1981 56 (1322) 18(304) 0(5) 12(51) 18(138)Pangasinan 1976 60 (7 7 )b 62(82) 0(111) 5 (39)

1978 58 (53)b 5(51) Cagayan 1979 59(713) 8 (101) 6 (34) 0(7)

1980 0(11) 4 (28) 15 (286)

62 ( 48(177) 61(140)1981 0(5) 12 (2) 13(45) 14 (92)

Irrigatedlowland Laguna 1974 62 8

1974-75 50 41 1975 43 72 1978 96 (623) 21(24) 33 (60)1979 83(2017) 18(110) 8(62) 11 (89) 22(400)1980 78 (1625) 20(75) 0 (16) 4 (72)1983 60

9 (80) 7 (270) 30

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R F Apostol unpublished Iloilo and Pangasnan 136 Cagayan B Canapi unpublished Laguna 175 P CPantua unpublished 118 234 143 Carino and Shepard unpublished YSB = yellow stem borer ScirpophagaincertulasGFB = gold fringed borer Chilo auricilius PSB = pink stem borerSesamia inferens earcutting caterpillar Alythimna separata swarming caterpillar Spcdoptera mauritia acronyctoides skipper Pelopidas mathias reen horned caterpillar Melanitls ledaismene RLF = rice leaffoldersCnaphalocrocismedinalis and Marasmiaspp brown semilooper Mocisfrugalis Figures in parentheses are sample sizes Egg masses

IRPS No 123 January 1987 35

Table 9 Effect of seeding rate and insecticide on plant density and Yield losses from insect pests in upland rice often areyield of Dagge upland rice Batangas Philippines 1976a comparable to those of lowland rainfed or irrigated rice

Seeding rate Insecticide Dosage Plant density Yield (0-Y30o) but usually from different pests The principal (kg aiha) seed treatment (kg aiha) (noIra-row) (tha) groups are soil pests feeding on sown seed and rootsat 14 DE followed by seedling pests These groups have largely gone

50 None - 35 a 20 a unnoticed leading researchers to conclude that stem borers50 Carbofuran F 10 38 ab 23 a and seed bugs are the most important pests (17) Yield loss 10 beCabofuranF100 None - 4551 c 23 a trials that pinpoint major pests need to includc seed

100 Dieldrin WP 10 52 cd 22 a treatments Granules applied in the furrow may not give150 None - 59 d 22 a adequate protection from many seed pests

aAv of 8 fields DE = days after crop emergence In a column means The floristic and pedological diversity of upland rice followed by a common letter are not significantly different at the environments is matched by the diversity of arthropods5level by D)MRT F = flowable Wi = wettable powder both pests and their natural enemies Irrigation has homoshy

genized lowland rice environments Flooded soils tend tothreatened by epidemics of locusts or armyworms New have the same properties and to eliminate soil pests The technology stressing efficiency has minimized insecticide spatial and temporal dominance of cultivated rice has cost benefited specialist pests

Seed treatment is normally inexpensive and can be Upland rice has no specialist arthropod species The economicallyjustified in -nanycases(66) Treating seed with monophagous pests on upland rice prefer the lowlands systemic insecticide (216) but not chlorinated hydrocarbons where rice evolved Rice in the uplands exists only because (247) is most effective in controlling seedling maggots of man and even though lowland rice predates upland riceFoliar sprays have to be repeated to obtain control and specialist species evolved Wheat maize and sorghum do often are washed off by frequent early season rains not have monophagous pests leading us to conclude that Granular insecticides are effective but required dosages are there is no evolutionary advantage for inssects to have too expensive to justify except under heavy infestation narrow host ranges in upland graminaceae(247) When directed against first-instar larvae banding Pests have adapted to the uplands not only by having granules in seed furrows (177) can be effective against white wider plant host ranges but also by having long life cyclesgrubs at low dosages (025 kg aii ha) (134) Seed treatments undergoing dormancy or being dispersive Each pest has a are not promising for white grub control (248) unique life history strategy suited to highly unpredictable

Seed treatment is also effective ainst ants and may be environments In agivenyear someinsect specics an better cheaper than increasing the seeding rate In Batangas exploit upland rice as a food source but no pest is a senious treating 50 kg seed ha was as effective as increasing the one every year on upland rice seeding rate to 100 kgha and using untreated seeds (Table The low yielding potential of upland rice means less 9) In Brazil seed treatments control termites and other soil quantity and lower quality of food for pests to exploit but insects including the lessercornstalk borer(66) Domiciano also less incentive for farmers to undertake control (58) however found seed treatments inconsistent in their The most effective control would be early and synchroshyeffect against high populations of lesser corn stalk borer nous planting of early-maturing upland rice varieties to

Baiting can be an inexpensive way of controlling ants prevent insect pest buildup Breeding for insect resistance seedling maggot flies mole crickets and field crickets Baits should initially focus on stem borers because they are verymade of locally available material and impregnated with difficult to control by other means Biocontrol efforts insecticide can be sparingly distributed taking adantage of should first stress conservation of natural enemies preshythe pests ability to disperse and encounter bait sites dators and pathogens rather than parasites Efficient

The new low-volume hand-held controlled-droplet chemical control includes seed treatment or baiting for seed sprayers may offer an advantage over high-volume knap- seedling and some root pests and using controlled dropletsack sprayes for controlling foliar pests Upland rice sprayers for spot treatments against foliar pests and seed canopies are more open than those in lowland rice and bugs when economic thresholds are reached better droplet penetration should result Controlled droplet applicators would be ideal for seedbugs and defoliators

CONCLUSION

Upland rice is attacked by a wider array of insect pests than is lowland rice mainly beca use of the addition of soil pests Generally population levels of lowland rice pests are lower in upland than lowland rice leading many people to conclude that insect pests are not important on upland rice

36 IRPS No 123 January 1987

REFERENCES CITED

1Abraham C C and K S R Mony 1978 Occurrence ofLeptocorisa acuta Fabr (Coreidae Hemiptera) as a pest of nutmeg trees J Bombay Nat ttistSoc 74553

2 Agyen-Sampong M 1975 HlierogrlphusdiaganensisKrauss (Orthop-tera Acrididae) a new pest of rice in Northern Ghana Ghana J Agric Sci 8249-253

3 Agyen-Samnpong M 1976 Varietal resistance and other inle-ratcd pest control components as factors of rice varietal improvement in West Africa Prcsented at the Second varietal improvement serinar 14-18 Sep 1976 Bouak Ivory Coast West Africa Rice Development Association Monrovia Liberia 18 p

4 Agyen-Sampong M 1977 Outbreak of Ihccrorrdnisorae in rice seedlings near Rokupr Sierra Leone lit Rice Res Newsl 2(6)16-17

5 Agyen-Sampong M 1980 Dirty panicles and rice yield reduction caused by hugs IntRice Res Newsl 5(l)11-12

6 Agyen-Sampong M 1982 The major pest problems of irrigated upland and mangrove swanip rice ecosystems ir the humid tropical Guinea savanna and Salel climate zones Pages 318-328 in Pro-ceedings training course cn concepts techniques and application of irtegrated pest management in rice inWest Africa 10-20 Jan 1982 Fendall Liberia West Africa Rice l)evelopioent Association Monrovia Liberia 505 p

7 Akibo-Betts V I and S A Raymundo 1978 Aphids as rice pests in Sierra Leone lit Rice Res Newsl 3(6)15-16

8 Akinsola F A 1975 Present status of different rice stemborers in parts of Nigeria iicL Entomo Newsl 328

9 Akinsola FA 1979 Ihe biology and ecology of rice stemborers in Nigeria Ph 1) thesis University ol lbadan Nigeria 217 p

10 Akinsola E A 1979 Guidelines of integrated management of insect pests of upland rice in West Africa Presented at the Seminar on integrated management of rice diseases and insect pests at Bobo-Dioulasso Upper Volta 17-22 Sep 1979 West Africa Rice Develop-ment Association Monrovia Liberia

II Akinsola IA 1984 Insect pests of upland rice in Africa Pages 301-305 in An overview of upland rice research Proceedings of the 1982 lIouak Iwsrv Coast upland rice workshop International Rice Research Institute 13O Box 933 Manila Philippines

12 Alam M S V TJohn and Kaung Zan 1985 Insect pests and diseases of rice in Africa Pages 67-82 in Rice improvement in Eastern Central and Southern Africa proceedings of the international rice workshop at Iusaka Zambia 9-19 April 1984

13 Alticri M A A van Schoonhovcn and J D Doll 1977 The ecological role of weeds in insect pest management systems a review illustiated with bean (Phaseohsvulgaris L)cropping systems PANS (Pest Artie News Summ) 23195-205

14 Anmante F 1967 The leaf-cutting ant Atia capiguaraa pest of pastures [in Portuguese] Biol6gico (Sao Paulo) 33113-120 (Rev Appl Entomol A56207)

15Andrade A C and D Puzzi 1951 Preliminary results of experi-merits on the control of Sculptcoris iastantus on sugarcane [in Portuguese] Bioulgico (Sao Paulo) 1744-49 (Rev Appl Entomol A41421422)

16 Appert J 1970 Maliarplia separaelha (rice stem borer) new observations of entomological problems of rice in Madagascar [in French] Agron Trop (Paris) 25329-334

17 Ariadeau Mand Z Harahap 1986 Relevant upland rice breeding objectives Pages 189-197 in Progress in upland rice research Proceedings of the 1985 Jakarta conference International Rice Research Institute 1O Box 933 Manila Philippines

18Asar S and Kisimoto 1975 Seasonal occurrence of the smaller brown planthopper in upland rice [in Japanese] Proc Kanto-Tosan Plant Prot Soc 2228

19 Bakker W 1974 Characterization and ecological aspects of rice yellow mottle virus in Kenya Meded Landbouwhogesh Wage-ningen 152 p

20 Balut F F 1970 White grubs attacking rice Oryza ativa L [in Portuguese] Biol6gico (So Paulo) 36321-322

21 Banerjee S N and L M Pramanik 1967 The lepidopterous stalk borers of rice and their life cycles in the tropics pages 103-124 in The major insect pests ofthe rice plant Proceedings ofa symposium at the International Rice Research Institute September 1964 The Johns Hopkins Press Baltimore Maryland USA

22 Barrion A T and J A Litsinger 1980 Ants a natural enemy of cliaiarac in dryland rice IntRice Res Newsl

5(4)22-23 23 Barrion A T and J A litsinger 1981 The spider fauna of

Philippine rice agroecosystem 1Dryland Philipp Entomnol 5139-166

24 Barrion A T and 1 A Iitsinger 1984 The spider fauna of Philippine rice agroccosystcms II Welland Philipp Eiitomol 611-37

25 Barrion A T1 and J A Litsinger 1984 lihanus (Diptera Tabanidae) eggs an alternative host of rice stei borer (SI3) egg parasite Telenoonis digns (Iliymenoptcra Scelionidae) lit Rice Res Newvsl 9(6)19

26 Barrion A T and J A Litsinger 1985 Identilication of rice leaffolders by wing markings Int Rice Res Newsl 10( 1)24

27 Barrion A T and J A Litsinger 1985 Chctiosspp (Coleoptera Carabidae) a leaffolder ILF)predator lit Rice Res Newsl 10(1)21

28 Bertels A 1970 Insect pests of rice and theircontrol [in Portuguese] Pelotas Brazil Instituto de Expcrimentacao Agropccuararias do Sul Circular 43 24 p

29 Bonzi S M 1982 Ctilo (flisilineus J de Joannis (Lepidoptera Pyralidae) stem borer of irrigated cereals in Upper Volta LioFrench] Agron Trop Riu Rizic Cult Vivrieres Trop 37207-209

30 Bourke T V T L Fenner J N LStihick G Baker EHassan D F OSullivan and C S li 1977 Insect pest survey for the year ending 30th June 1969 Papta New Guinea )epartment of Agrishyculture Stock and Fisheries Port Moresby 57 p

31 Bowden 1 1973 Migration of pests in the tropics Meded Rijksfac Landbouwwet Gent 38785-796

32 Breniere J 1973 The main rice parasites in West Africa and their control Pages 27-42 in Proceedings seminar on plant protection for the rice crop West Africa Rice Development Association Monrovia Liberia

33 Breniere J 1976 The principal insect pests of rice in West Africa and their control West Africa Rice Development Association Monrovia Liberia 52 p

34 Brown ES 1962 The African arinyworn Spoiolptera eXI)Ina (Walker) (Lepidoptera Noctuidae) a review of the literature Cornshymonwealth Institute of Entomology London UK57 p

35 Brown F B 1968 Upland rice in Latin America FAO-International Rice Commission 12th Session of the Working Party on Rice Production and Protection 2-14 Sept Peradeniya Sri lanka Food and Agriculture Organization Rome Italy 6 p

36 Buyckx F J 1 1962 Disease and insect pests of crops inCongo Rwanda and Burundi [iil French] Publications dc LInstilut National pour lestude Agrononiquedu Congo M Weissenbruck S A Brussels Belgium 708 p

37 Cariiio F 0 1981 Role of natural enentics inpopulation suppression and pest management of green rice lealhoppers Ph D thesis University ofthe Philippines at Los Bafitos PhilippinesLaguna 136 -

38 Carpente A J 1983 The development of rice cultivation and extension of vegetable productin in Zanzibar Final Report Vol II1 October 1983 Food and Agricultme Organization Ronte Italy

39 Carroll C P and S1 Risch 1983 Tropical annual cropping systems ant ecology Environ Manage 751-57

4(0 Cates R G 198 1 Host plant predictability and the feeding patterns of monophagous oligophagous and polyphagous insect herbivores Oecologia (Berl) 48319-326

IRPS No 123 January 1987 37

41 Cendcafia S M and F B Caora 1967 Insect pests of rice in the 62 Ene 1 C 1981 I)iiorldhol)ierusihilis Slater (licteroplera Philippines Pages 591-616 in The najor insect pests ofthe rice plant lygacidue Illissinac) a new rice pest front Nigeria lit Rice Res Proceedings of a symposium at the International Rice Research Newsl N3)15 Institute September 1964 The Johns Hopkins Press Baltimore 63 Fugenio T S 1965 rhe effect oi e-vCgetation on rice production Maryland USA 729 p and the possibility of overcoming it by leaf pruning MS thesis

42 Centro Ihternacional de Agricklittira Tropical 1984 Upland rice in University ol the Philippines at Los Baijos ILaguna Ihilippins 69 p Iatin Anerica Pages 93-119 hi An os ervies of uplaind rice research 04 Farrow R A 1975 The Alrican nmigratory locust ii is main Proceedings of the 1982 Boinakamp Ivory Coast upland rice workshop outbreak area of the middle Niger qlinlitatise studies of solitary International Rice Research Institute P0 Blox 933 Manila populations in relation to environmental factors Locusla No II Philippines 198 p

43 Chandra G 1979 Taxonomy ad hn riononlics ofthe imect parasites of 65 Fennal R G 1943 Food crop investigation Windward and rice Icalloppers and planthoppers in the Pihilippines and their Ieeward Islands First report Report onl work dunring the periodiiiportairce iii natural biol)gical cointrol Philipp Fntoltol 4119-139 April 1942-April 1943 Imperial College of Tropical Agriculture

44 Chang T T 1976 The originii colut ion enlIivation dissemination Trinidad 16 p (Rev of Appl Fintornol A3222-25) and diersificition of Asian and African rices -uphytica 25425-44 1 66 Ferreira F 1983 Integrated pest ritaageiret of upland rice [ill

45 Chatierjee P B 1979 Rice leaffolder attacks in India lit Rice Res Portuguese English summary] Pages 323-341 in Upland riceculttre Newsl 4(3)21 factors affecting production E Ferreira T Yamada and F-

46 Cheng Cbien-pan 19(4 Upland rice in Taiwan International Rice Malmolta eds Institutoda Potassae Foslato Institnto Internacional Cnmmission Wiorking Party oit Rice Production ard Protection I()th da Potassa Piracicaha Sao Paulo lraid Meeting 3-10 March 1964 Manila Philippines Food and Agricul- 67 Ferreira E arid F dl Silva Martins 1979 Occurrence of ture Organiaion Rome Italy 5 p Sogatohs orvziola (Muir) inupland rice in Goias Ilrail lit Rice

47 Chcrrett 1 M and 1) 1Peregrine 1976 A review of the status of Rcs Ncwssl 4(4)18 leaf-cutting ants and their control Ann Appl Biol 84124-128 68 Ferreira E and 1F da Silva Martiris 1984 Insect pestof rice ill

48 Chipeta F M 1974 Pests of rice in Malawi Pages 90-95 in Brazil and their control Jill Portugucsej FMIlRAtPA-CNPAF Proceedings Sith eastern Africa cereals research conference 10-15 Goiania Brmiil 67 p March 1974 DRB Mialda ed Ministry of Agriculture lilongwe 69 Ferreira FI F ca Silha Miartins S S Nero and 1 It P de Malawi Carvallio 1982 M tltiplc regression analysis todeterimine the effect of

49 Chu Yau-i 1979 The d atiaging period of the rice seedling fly insect populatiois ott upland rice yield [inIPortuguese Pesqui (Atlterigotiavxigu Stcin) (NItiscide Diptera) oil cori) at Fast Java Agropecu Bras 17671-675 Indonesia Plant Prot Bill 21403413 70 Ferreira F 1F da Silva Martins and F 1PZimmermann 1979

50 Conklin H C 1957 IHaninoo agriculture a report on aln integral Rice varietal resistance to tle lesser corn stalk borer [in Port uguese] systent of shi fti rig cult ivationi in the Pltilippincs Food and Agriculture Pesqui Agropccu Biras 14317-321 Organization Rome Italy 209 p 71 Fleming W F 1968 Biological control of lie hapanese beetle U S

51 Cramer H II 1967 Plant proection and world crop production Dep Agric RLs Sers [ech Bull 1383 78 p Pilaniensclhutz-Nachr 201-524 72 Food and Agriculture Organiation of the United Nations 1982

52 l)ani R C and Majunidar 1978 Prelininary observations on tie Country report surmimary Pages 27-37 in Report on the first incidence of root aphids on differeit rice cultnrcs Sci Cult 4488-89 FAOIUN)IPhail and regional training coiuirse in iniproved ctltural

53 Davidson R I 1979 lffects of root feeding on foliage yield Pages practices for upland rice 5-23 Oct 1981 Northern Region Agrculturd117-120 in T K Ci osb and R B Pottinger eds Proceedings of the Developririt Center Chiang Mai Food and Agricuilture Organiiashy2nd Australian conlerence on grassland invertebrate ecology Govt tion Rome Italy 37 n Printer Wellington New Zealand 73 Franco do Ariaral S 195 1 Residual action of insecticides in turfy

54 l)ean G 1W 1976 Rice insect pests inlaos Int Rice Res Newsl soil--the results of a field experiment [in Portuguese] Biol6gico (Sao 1(2)15 Paulo) 17209-214 (Rev Appl Entomol A41423)

55 I)ean G J W 1978 Insect pests of rice in Laos PANS 24(3)280-289 74 Freeman ) 1970 Report onithe Iban london School of -coroiics 56 )ingle H 1984 Behavior genes and lifc histories conplex adapta- Monographs on Social Anthropology No 41 Athlane Press

tions to uncertain environments Pages 169-194 in A new ecology London UK 317 p novel approaches to interactive systems P W Price C N Slobod- 75 Fuchs T W1 A Harding and 1W Smithlr 1979 Induction and chikoff and W S Gaud cdsl ohn Wiley New York USA termination oldiapause in the sugarcane borer Ann Flitonlol Soc

57 Dobelmann-JP and M Falais 1968 Upland rice cultivation ill tie Am 72271-274 North-western part of Madagascar in 1966-1967 [in French English 76 (hbriel B1) 1975 A review tf the major insect pests of some tpland summary] Trop Agron 2313-21 crops in tlc Philippines Pages 86-111 in Ad Htoc panel of experts on

58 Doniiciano N L 1979 Chemical control of tie lesser corn stalk pest disease arnd weed problems in some rainfed crops Bangkok (lolsonno~paloIgnowslhts) (Zeller 1848) (I-epid(ptera Ihycitid ae) Thailand Food aid Agriculture Organization Roire Italy

on rice in Ponta Gressa Parara 1977-1978 [in Portuguese] Pages 77 Garg D K and N K Shal 1983 White grub outbreak on rainfed 291-303 in Anais 1 Reuniao de tecnicos cmiirizicultura do estado de dryland rice in Uttar Pradeslh Int Rice Res Ncwsl 81(4)16-17 Sao Paulo-Campinas Cati Brazil 78 Garg I) K and IP Tandon 1982 Major insect pests of rice on hilly

59 Dunsmore 1 R 1970 Investigations on tlie varieties pests and tracts of Uttar Pradesh India lilt Rice Res Newsl 7(11)11-12diseases of upland rice (hill padi) in Sarawak Malaysia Int Rice 79 Garrity 1) P 1983 Upland rice research an international bibliograpy Comm Newsl 1929-35 1975-1982 Iternational Rice Research hnstitnte PO Box 933

60 Elias 1R1965 The new decline of rice seedlings is not caused by Manila Philippines 121 p parasites [in Portuguese] Biol6gico (Sao Paulo) 313-6 80 Ghauri M S K 1982 A new subspecies of Iinorh)rone bull

61 Empresa Brasiliera de Pesquisa Agropecuaria 1984 Upland rice itt cornlus Slater (Hemiptera Iygaeidae) front Papua New Guinea on Brazil Pages 121-134 it Av overview of upland rice research rice and carpet grass Ihll lEntoniol Res 72133-137 Proceedings ofthe 1982 Blouak Ivory Coast Upland rice workshop 81 Gonzales F 0 PArregoces R I ihernandef and OT Parada International Rice Research Institute PO Box 933 Manila 1983 Insect and imite pests of rice arid their control in Latin America Philippines [in Spanish] la Federacion Nacional de Arrocerts (Fedearroz)

Bogota Colombia 60 n

38 IRPS No 123 January 1987

82 Grist D H and R J A W Lever 1969 Pests of rice Longmans London UK520 p

83 Hale P R 1979 Two subterranean pests of upland rice in Papua New Guirnca IntRice Res Newsl 4(2)16-17

84 Hale P R and J T Hale 1975 Insect pests of rice in the New Guinea Islands Rice Entomol Newsl 34

85 Harris W V 1969 Termites as pests of crops and trees Common-wealth Institute of Entomology London UK 41 p

86 Hattori 1 1968 A method to distinguish between grass leaf roller (Ctaplha rocis medinalisGuenee) and the rice leaf roller (Susurtia exigua Butler) [in Japanese] Plant Prot (Japan) 23167-170

87 Henry I E and 1A Onsager 1984 Experimental control of the mormon cricket Iahrosshnph by NJsena locustae(Aficrospora Microsporidae) a protozoan parasite of grasshoppers (Ort Acri-didae) Entomophaga 27197-201

88 Hidaka 1 P Vungsilabulr and SKadkao 1974 Studies on ecology and control of the rice gall midge in Thailand Tech Bull 6 TropicalAgricultural Research Center Tokyo Japan 113 p

89 Hincklcy A 1) 1963 Ecology and control of rice planthoppers in Fiji Bull Entomrol Res 53467 481

90 Hirao J 1982 Ecology and chtInical control of the rice leaffolder Jpn Pestic Inf 4114-17

91 Ho DT 1984 Sten borer incifience in rice ecosystems in KenyaEast Africa Pages 307-320 in A overview of upland rice research Proceedings of the 1982 Bouac Ivory Coast upland rice workshop International Rice Research Institute PO Box 933 Manila Philippines

92 Ho )T J 1Njokah and J GKibuka 1983 Studies on rice sten borers in Kenya with emphasis on ifaliarpiaipiraellRag Insect Sci Appl 465-73

93 Ichihara I T Ito A NIatsuo and M Sawada 1971 Chemical control of the rice striped stem borer in upland rice grown in plastic mulch [iii Japanese] Proc Kanto-Tosan Plant Prot Soc 1882

94 IclhiharaIand A Matsuo 1969 Damage caused by rice striped stett borer to upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1686-87

95 Ichhara I atid A MIatsun11970 Conside ration oft lie extraordinary prevalence of rice striped stem borer in upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1778-79

96 Institut de Recherches Agrolnliqes Iropicale et des Cultures Vivrieres 1978 Insect resistance Pages 12-13 in Upland rice 1978 atnual report Bouak Ivory Coast 27 p

97 International Institute for Tropical Agriculture 1983 Aphids itt rice roots I ITA Res Briefs 4(1-3)4-5

98 International Institite fi Tropical Agriculture 1983 Hopperburn in rice caused by brown plathopper IITA Res Briefs 4(l-3)6-7

99 International Rice Research Institute 1975 Annual report for 1974 PO Box 933 Manila Philippines p 216

100 International Rice Restarch Institute 1976 Annual repiort for 1975 PO Box 933 Manila Philippines p 228

101 International Rice Research Institute 1979 Cropping systems program Component technology development and evaluation Page 404 i Annual report for 1978 11O Box 933 Manila Philippines

102 International Rice Research Institute Department of Entomology 1981) Insecticid evaluation 1979 Tables 73 74 PO Box 933Manila Philipes

103 International Rice Research Institute Department of Entonmology 1981 Insecticide evalomtion 1980 Table 81 POBox 933 Manila Philippines

104 International Rice Research Institute 1981 Report of a meeting of Asian rice-based cropping systems entomologists Eleventh cropping systems working group meeting 18-22 May 1981 PO Box 933 Manila Philippines p 132 133 144

105 International Rice Research Institute 1984 Cropping systems program Pages 368-373 in Annual report for 1983 PO Box 933 Manila Philippines

106 International Rice Research Institutc 1984 Upland rice in Asia Pages 45-68 in An overview of upland rice research Proceedings of the 1982 Bouakamp Ivory Coast upland rice workiiop loternational Rice Research Institute 11O Box 933 Manila Philippines

107 Irshad M 1977 Grasshoppers in rice in Pakistan lit Rice Res Newsl 2(3)8-9

108 Ito R and TOno 1964 Upland lice its culture and varieties in Japan lIternational Rice Cormmission loth Meeting of the Working Party on Rice Production and Protection 3-10 March 1964 Manila Philippines Food and Agriculture Organization Rome Italy 12 p

109 Jeffrey M I and V A Dyck 1983 Effect of moonlight on lightshytrap catches of brown planthopper lit Rice Res Newsl 8(6) 13

110 Jennings P R and T Alicia Pineda 1983 Management ofthe rice plnthoppers Sogatodsorivi(coIl in Latin America Presented at the meeting on stable plant resistance FAO and lIndocsian I)inectorate of Food Crops Protection Bali lndonesia 29 Novcniber-2 l)ccemshyber 9 p

IllJimenez Gomez J A 1979 Rice insect pests i lla ios ()riCntales lil Spanish] Pages 99-103 in Rice ciltivation Instituto (olotibiatto Agropecuario Compndio No 29

112 Jongeleen F JJ Soesanto and A Mockidjo 1979 Studies oti pathogens of two species of Scarabacidac froinIndonesia Z Angew Enlomnol 87412-417

113 Jouvenaz 1) PC S lofgren and W A Baiks 1981 Biological control of imported fire antsa review of cirrent knowledge Bull Entomol Soc Am 27203-208

114 Kainoh Y K Shiniiu S MarUani Y Taiaki 1980 lost finding behavior of tie rice bu lplotori+JchinvittjI)allas (liciniptera Corcidac) with special reference to diet pattern ol aggregation and feeding on the rice plant Appl Entomol Zool 15225-233

115 Kakkar K 0 P Sharma and P I Sharma 1977 Note on Ihteronyrchus liodresRedt (Coleopicra I)ynastidae) as asporadic pest of paddy and maie inHimachal Pradesh Corr Agric 156-57

116 Kalode M131 79 Major insect pest problems and their control in upland rice (Unpublishcd mineogriph CWI oJNaaiat lni Chandaratat (198 1)original not seen)

117 Kalshoven L i 98 I The pests of crops in I ldoniesia (translated by P A vatn der Laan) 1 T Ichtiar B3aru-vati Iloeve lakarta Indonesia 701 p

118 Kamal N Q 1981 Suppression of whitebacked planto ner Sogatella fircaifra (Horvath) aLId rice lealfolder (nhal -cis licdb lis (Guenee) populations by natural cnclmie hlI) tiesis Grcgorio Araneta University Foundation Victoncta Park Metto Manila Philippines 179 Ip

119 Karim A N M Reaul 1975 A species ofliealy bug Trioni i70s ceres new oii rice in the Philippines Rice Entoniol Newsl 247

120 Katanyukul Wand C Chandaratat 1981 Insect pestsof uplaid rice and their control Report firstFAOUNlFi Tliailand training course on improved cultural practices for tplad rice Chiang Mai Thailand 5-23 Oct 1981 Food and Agriculture Organiation Rome Italy 22 p

121 Kern F 1955 lhletwioa(I)uninemts) bhhit Biturm(Coleoptera) a new pest of sugarcane inVenezuela [inSpanish with Etnglish sunmary] Agron Trop (Maracay) 527-29 (Rev Appl Eitoniol A4572)

122 Khan K M and S Ghai 1974 White grubs ald their control in India Pesticides (1Bombay) 819-25

123 Kiritani K 1975 Integrated pest management of rice pests Pages 66-70 itt Proceedings platning workshop ott cooperative hIeld research in pest managementP SMotooka ed East-West Center Honololu Hawaii USA

124 Kisimoto R 1967 Ecology of insect vectors forecasting and chemical control Pages 243-255 in The virus diseases of tle rice plant Proceedings of a symposium atthe International Rice Research Institute April 1967 The Johns Hopkins Press altimore Maryshyland USA

125 Kisimoto R and V A Dyck 1976 Climate and rice insects Pages 367-391 in Climate and rice International Rice Research Institute PO Box 933 Manila Philippines

126 Kureha S S Kobayashi SAsari and T Ishihara 1977 Life history and control of the mole cricket Grrllotalpa africana Palisot de Beauvois [in Japanese English translation 7520 1RRI library] Proc Kanto-Tosan Plant Prot Soc 21192-197

127 Kushwaha K S 1981 The black beetle as a pest of rice in Haryana India Int Rice Res Newsl 6(4)14

128 Lawani S M 1982 A review of tile effects of various agronomic practices on cereal stem borers Trop Pest Manage 28266-276

129 Leuck D B 1966 Biology of the lesser cornstalk borer in South Georgia J Econ Entomol 59 797-801

130 lever R J A W 1955 Insect pests of the rice plant in Malaya Int Rice Comm Newsl 1612-21

131 Li Ching-Sing and Shin-Foon Chiu 1951 The study ofthe rice gall midge Iachydiplosis orvza Wood Mason [in Chinese with English summary] 1Taiwan Agric Res 21-13

132 Iitsinger J A 1984 Assesament of the need-based application for rice Proceedings TechnologyTransfer Workshop 15-16 March 1984 International Rice Research Institute 1PO Box 933 Manila Philippines 6 p

133 Litsinger J A 1984 Environmental diversity and insect pest abundance with reference to the Pacific Pages 285-307 in Proceedings sub-regional training course on methods of controlling diseases insects and other pests of plants in the South Pacific 4-20 Oct 1982 Vaini Kingdom of Tonga Nukualofa

134 Litsinger J A RF Apostol and M BObusan 1983 White grub Leucopholis irrorata (Coleoptera Scarabaeidae) pest status popula-tion dynamics and chemical conllol in arice-maize cropping pattern in the Philippines J Econ Entoinol 761133-1138

135 Litsinger J A C de la Cruz F Raymundo A T Barrion M D Lumaban J P Bandong M S Venugopal F Paragna and E Balete 1982 Insect pests and insecticide reponse of dry-seeded bunded rice Pages 179-193 in Report of workshop on cropping systems research in Asia International Rice Research Institute Los Bafos Philippine

136 Litsinger J A M 1) Luinaban J P Bandong A T Barrion W Katanyukul and N Panda 1978 Insect pest management in rainfcd lowland rice agroecosystms of tropical Asia International Rice Research Conference 17-21 April 1978 International Rice Research Institute Los Bafios Philippines

137 Litsinger J A and K Moody 1976 Integrated pest management in multiple cropping systems Pages 293-317 in R I Papendick P A Sanchez and G B Triplett eds Multiple cropping American Society of Agronomy Madison Wisconsin USA

138 Loevinsohn M E 1984 The ecology and control of rice pests in relation to the intensity and synchrony of cultivation Ph D thesis University of London 36) p

139 Loevinsohn M E J A Litsinger N Panda M A Austria R F Apostol C Andrion and J Bandong 1982 A basis for predicting rice insect pest abundance in cropping systems Paper presented at the 1982 International rice research conference on rice and cropping systems 19-23 April 1982 International Rice Research Institute Los Bafios Iaguna Philippines 25 p

140 Lopez A W 1930 Repot of the Entomology Department Pages 145-172 in Annual report of the Philippine Sugar Association 1929-30 Manila

141 Lopez AW 1931 Entomology Department Pages 227-266 in Annual report of the director or research 1930-31 Philippine Sugar Association Manila

142 Lopez A W 1932 Entomology Deparment Pages 252-279 in Annual report of the Philippine Sugar Association 1931-32 Manila

143 Lungibill P and G G Ainslee 1917 The lessercornstalk borer US Dep Agric Agric Res Serv Tech Bull 539 27 p

IRPS No 123 January 1987 39

144 McGuire J U Jr and BS Crandall 1967 Survey of insect pests and plant diseases ofselected food crops of Mexico Central America and Panama U SDepartment of Agriculture International Agriculshytural Development Service Agricultuial Researrh Service October 1967 11 p

145 McNeill Sand T R ESouthwood 1978 The role of nitrogen in the development of insectplant relations Pages 77-9 in lliochemical aspects of plant and animal coevolution J B Ilarbornu ed Academic Press London UK

146 Maebara H 1955 Spatial distribution of lastinstiarlanaeof purplish stem borer (Sesamia iqferens Walker) hibernating in an upland rice field Bull Kagoshima Univ Fac Agric 438-41

147 Mallick E H and N G Hajra 1977 Response of rice plant to defoliation treatments at different developmental stages Indian Agric 2151-56

148 Mansingh A 1971 Physiological classification of dormancies in insects Canl Entomol 103983-10)9

149 Martell C G 1974 Catalogue of plant feeding insects of Mexico [in Spanish] Fitofilo 27(69)1-176

150 Martinez 0 J 1982 Principal rice pests in Ilanos orientalis [in Spanish] P~ages 80-94 in Curso deact nalization de conocimientos en arroz Villaniencis Instituto (olonmbiano Agropecuario

151 Martins J F da Silva F Ferrcira A S Prabhu F 1 P Zimmermann 1980 Use of pesticides for the control of the main subterranean upland rice pests esqui Agropecu Bras 1553-u

152 Martins I F da Silva F Fe eira rod BdamSilveira Pinheiro I Q2 Damage simulation caused by defoliators of upland rice [in Portuguese with English summary] Pesqui Agropecu Bras 171113-1119

153 Martins 1F da Silva N(iyuen van Tan and 1Ida Silveira Pinheiro 1981 Stem borer resistance in upland rice associated with plant morphological characters [in Portuuce Iesqui Agropecu Bras 16187-192

154 Masaki S 1980 Sumnmer diapause Ann Rev Entomiol 251-25 155 Masuzawa T 11 Suwa and F Nakasoji 1983 l)ifkleences of

oviposition preferencc and survival rate of two skipper butterflies Parnara guttata and PIchlpidai iathiais (Iepidoptera HIesperiidac) on rice plant and cogon grass New Entomol 3247-56

156 Maurya D M and C P Vaish 1984 Upland rice in India Pages 69-92 in An overview oftpland rice research Proceedings ofthe 1982 Bouakamp Ivory Coast upland rice workshop International Rice Research Institute IO Box 933 Mnila Philippines

157 Menon M G R R N Katiyai S Kuar and S S Misra 1970 Termites attacking the paddy crop Sci Cult 36294-295

158 Miyahara Y and 1 Nishikubo 1967 Behavior of young nymphs of the red rice root aphid Iholahsipiun ri liabhdontinalisSasaki in relation to movement from the leaf to the roots of upland rice [in Japanese] Pioc Assoc Plant Prot Kyushu 1363-66

159 Mo ljoa Tjien 1952 White grub of upland rice (Holotrichiahelleri Brsk) [in Indonesian with English summary] Contrib Agric Res Stn Bogor No 13134 p

160 Mochida 0 and T Suryana 1979 The occurrence of the brown planthopper and ragged stunt on lowland and upland rice ill Central Java and logyakarta in tile late of January 1979 Entomnolgy Survey Report No 15 Central Research Institute of Agriculture (CRIA) Sukamandi Indonesia I February 1979 9 p

161Monet C 1979 Varietal resistance to upland rice pests [inI French] Pages 35-39 in Annual report 1977 Institut des Savanes Department des Cultures Vivrieres Bouak6 Ivory Coast

162 Monte 0 1942 A caterpillar ofrice-fields[in Portuguese] Biol6gico (Sao Paulo) 8161-163 (Rev Appl Entomol A31414)

163 Munroe I) D 1979 Pest management strategy for hill rice in four resettlement areas in Sarawak Malaysia Pages 89-102 i1 Guidelines for integrated control of rice insect pests Food and Agriculture Organization Plant Production and Protection Paper 14 Rome Italy 115 p

40 IRPS No 123 January 1987

164 Nakasuji F M Ishii and Jen-Zon Ito 1984 Rice skippers in Taiwan and their life histories ZZrC (Taipei) Tech Bull 82 10 p

165 Navas I) 1976 Fall armyworum in rice Pages 99-106 il Ecological animal control by habitat management Proc Tall Timbers Con-ference 28 Feb-I Mar 1974 Fall Timbers Res Stn Misc Publ 6 Tallahassee Horida USA

166 Nickel 1 1 1961 Biological control of rice stem borer A feasibility study Tcch Bull 2 International Rice Rcarch Institute PO Box 933 Manila Philippines IIl p

167 Nigan P M and R A Vernia 1985 Insect pests of uplaild rice in Uttar Pradesh Int Rice Res Newsl 10(6)22

168 Nilakhe S S 1985 -cological observations on spittle bugs with emphasis on their occirence in rice Pcsqtui Agropecu Bras 20407-414

169 Nixon G EJ 195 1 lie association of ats with aphids and coccids Commonwealth Iusti lcte of Entonology Iondon UK 36 p

170 Nurdin F an Ishak Manti 1985 Study oit seedling fly of upland rice Interlational Upland Rice Moitoring Tour Report 26 lebruary-2 MarLI Sukaranti West Sumatra

171 Oka 1 N 1979 Ciltural conttol of the brown planthopper Pages 357-369 in Brown planthopper threat to rice production in Asia International Rice Research Institutc 1p0 Box 933 Manila Philippines

172 Ohni M and A Vilnaii 1975 Rice insect pests of Italy biology of Sipha grcria (Kaltenbach) and its control [in Italian] Riso (Milan) 2459-71

173 Otanes F Q atnd 1 I Sison 194 1 Pests of rice Philipp J Agric 12211-259

174 Painter R I 1951 Insect icsistance in crop plants University Press of Kansits Lawrence Kansas USA 520 p

175 Pantua P C and 1A litsinger 1981 Arthropod abundance in continuous vs biannual rice cropping systeis Int Rice Res Ncwsl 6(2)20-21

176 Pathak M ) and V A l)yck 1975 Sttudics ott insect pests of upland rice Pages 186-191 in Major research in upland rice International Rice Research Institute PO Box 933 Ilanila Philippines

177 Patil B R B M IHasabe and 1) S Ajri 198 I Chemical control of white gritb lIditriihinhorraia Fabr intfesting groundntut paddy atnd sorghum Indian 1Entomol 43232-236

178 Pawar A 1) 1974 (ollection and identification of insect pests of rice rlerminal repirt lInternational Rice Research Institute PO Box 933 Manila Philippines 72 p (tnimco)

179 IerrajU A aid A S Reedy 1965 Observations ott diapausc itt tie steti borer ohrice Rice News Teller (lai-Mar)9-13

180 Pont A C 1972 A review of the Oriental species of theri ona Rondani (I)iptera Muscidac) of econtomic importance Pages 27-102 in Control of sorglium shoot fly M G lomtani and W R Young eds Oxford atd IH1 Ptthl Co New I)clhi India 324 p

181 Prasad S C 1PChand aid M 1PSingh 1983 Incidence of rice gall midge in the upland plateau region of Hihar Oryza 20235-236

182 Rai P S 198 0 Nisaga simpex damage to rice in the hill tracts of South India Int Rice kles News[ 5(I)12

183 Rathorc Y S 1969 Gonocplhahli dcprssunt Fabr as itpredator of (tilo zonwIlhis (Swinhoc) and other moths in field cage at Patnagar Uttar Pradesh Indian 1Entotnol 3127

184 Reed W 1983 Crop losses causcd by insect pists in the developing world Proceedings 10th Internatial Congress of Plant Protection lrighton UK 174-80

185 Reedy M V 1981 Ateactogasterinhens(Walker)a new root-grub pest of tipland paddy and inaize Curr Sci 501076

186 Reveno M A 1968 Catalogue of the insect pests of economic crops in Colombia [in Spaiish] Asociacion Latino Americana de Entomo-logia (AIAE) 129-33

187 Reyes G M 1957 Rice dwarf disease in the Philippines Food and Agriculture Organiiation Plant Prot Bull 617-19

188 Rice S E A A Grigarick and M 0 Way 1982 Effect of leaf and panicle feeding byarmyworm (Lepidoptera Noctuidae) larvae on rice grain yield Environ Entomol 7593-595

189 Risch S J 1) Andow and M A Altieri 1983 Agioccosystem diversity arnd pest control data tentative conclusions and nizw research dirctions Environ Entomol 12625-629

190 Rivera C T 1956 Three organic insecticides in thecontrol of insects affecting upland rice Philipp Agric 39465-472

191 Roffrey 1 1972 Locusta outbreaks in the Philippines Acrida 1177-188

192 Rojas R 1) 1941 The effect of pruning at varying ages upon the yield of Ramairio Phipp Agric 29851-860

193 Roonwal M i 1979 Ternite life and termite control in tropical South Asia Scientilic Pnbl odhpr India 139 p

194 Rosetto C1 F da Silva Martins N C Schmidt and L EAzzini 1978 Damage caused by pasturespittle bugs 4Dvoi)lavopictaandDJ schach) on rice lBragantia Nota 6 (Matcli)25-27

195 Rosztto C1- S S Neto I- Amantc I) Iink 1Garcia-Vieira D M de SoLza N V Bansa t and A M de Olivcira 1971 Rice pests of Brazil [in Portuguese] Pages 149-238 inrFA()-International Rice Commission 2nd meeting of the Rice Committee for the Americas 6-11 l)ecember 1971 Pelotas Rio Grande do Sul Brazil

196 Rothschild G H L 1969 Obseratiois on the armyworm Spodoptera matritia acronrctoiiles Gn (Lep Noctuidae) Bull Entomol Res 59143-160

197 Rothschild G 1- L 1974 Parasites of the armworm Spolopiera mauritiaacroncitoihes in Malaysia Fntornopliaga 19293-299

198 Sachan i N S K Gangwar and 1N Katujar 1980 Field crickets damaging cages in North Eastern IHill Region Indian J Entomol 42526-527

199 Sands 1) P A 1977 The biology and ecology of Lepocorisa (Hemiptera Alydidac) in Papua New Guinea Res Bull 18 Departshyment of Primary Industry Port Moresby Papua New Guinea 104 p

200 Sarawak 1968 titomology division Pages 97-106 in Report of the research branch of the Department of Altriculture 1966 Kuching (Rev AIppl Eintomol A56603)

201 Satpathy JM M S Das and K Naik 1977 Effect of multiple and mixed cropping on the incidence of scine important pestsI Entomol Res 178-85

202 Sauer IHF G 1939 Notes Ott Elasmpnlpualtsligniosellths Zeller (Lep Pyr) a serious pest ofcereals in Sao Paolo [in Portuguese] Arq Inst Biol 10199-206 (Rev Appl Entotnol A28378)

203 Satinders 1L A 1 SKing and C L Vargas S 1983 Crop pests of Central Anerica with t bibliography [in Spanish] Centro Agroshynotnico Tropical de Investigacion y Ensenanza Turrialba Costa Rica (CATIE) Tec Bol 9 90 p

204 Sauphanor B 1985 Some factors of upland rice tolerttce to stem borers in West Africa Inseu Sci Appl 6429434

205 Sen P atnd S Chakravorty 1976 Stentflics in Haritighata Nadia West Bengal India [it Rice Res Newsl 1(12) 15

206 Senapai B and 1M Satpathy 1983 The seedling fly Atherigona orrzae Malloch is a pest of rice in Orissa (India) J Entomol Res 777-79

207 Shhjahan M S Alatn M Ah nad and BN Islam 1983 A new scarabaeid beetle pest of dryland rice in Bangladesh Int Rice Res Ncwsl 8(I)16-17

208 Sharma G L S C Modgol and R C Gautam 1979 Effect of depth of submergence ott incidence of bacterial blight and white grub infestation in transplanted rice Int Rice Res Newsl 4(6)12-13

209 Sharma S 1985 Scarabaeid beetle outbreak itt upland rice Int Rice Res Newsl 1(1(l)24

210 Shiraki T 1917 The paddy borer Schoenobhits incertulas Walker [achaku Agric Exp Stn Formosa 256 p

211 Silva A de Brito and 11P Magalhaes 1981 Insect pests of rice [in Portuguese] Circular Tecnica No 22 EMBRAPA Centro de Pesquisa t ecuaria de Tropico Unido (CPAJU) Belem Brazil 14 p

212 Singh S A 1979 Emergence of rice leaf butterfly as a rice pest in Manipur India int Rice Res Newsl 4(4)16-17

213 SinghS A 1983 Changing trends of insect pestsof rice in Manipur Int Rice Res Newsl 8(f)12

214 Siwi S S and P H van Doesburg 1984 Lenovorisa Latreille in Indonesia (Heteroptera Coreidae Alydinae) Zool Meded (Leiden) 58117-129

215 Soejitno J 1977 Reaction of some upland rice varieties toseedling fly eltherikonaexigua Stein [in Indonesian] Siri Hana I Penyakit 630-35 Lembga I osat Fenclilian Pertanian Bogor Indonesia

216 Socnardi 1967 Insect pests of rice in Indonesia Pages675-683 in The major inse pests of thu rice plant Proceedings of a symposium at the Internatital Rice Rec Wi itt S pternber 1964 The Johns Hopkins Press Baltimore Maryland USA

217 Soto P E A T Petcz and I W Buddenhagen 1976 Survey of insect pests and diseases of rice in different ecological zones in Nigeria Rice Entomol Ncwsl 435-46

218 Soto 11E and Z Ziddiqi 1976 Screening for resistance to African rice insect pests Varietal improsetnent sCmi n r Bouak Ivory Coast 13-17 Sept 1976 West Africa Rice l)evelopment Association Monrovia Iiberia

219 Squire F A 1972 Entomological problems in Bolivia PANS (PestArtic News Sumr) 18249-268

220 Stapely 1 II 1978 The rice leaf roller in the Solomon Islands Int Rice Res Newsl 3(l)13

221 Suenaga H 1937 On aphids attacking the roots of rice Oyo-I)oubts Zasshi 9150-152 [in Japanese] (Rev Appl Entomol A25745)

222 Suenaga H 1952 On the seasonal abundance of three subterranean aphids on upland rice Bull Kyushu Agric Exp Sti 1249-262

223 Sutynso Yas and Dandi Soekaria 1964 Beberapa hana yang ttenperang tana man padi tanah kering [in Indonesian] Rapat kerja Padi Tanah Kering 6 sd 8 Juily 1964 Kaliurang 9 p

224 Tallamy I) W and R F Denno 1983 Alternative life history patterns in risky environments in example from lace bugs Pages 129-147 in Insect life history patterns R F l)cnio and If Dingle eds Spring-Verlag 225 p

225 Tamura 1 1939 Observations on some insect pests of field crops [in Japanese] 1 Plant Prot 26414-420 (Rev Appl EntomolA27-556)

226 Tamura and Suzuki 1964 Influences of the early-transplanting practice in rice culture upon development of the rice stem borer Chilo siql)ressalisWalker Bull Ilokuriku Natil Agric Exp Stn 761-94

227 Tan J P 1924 The rice root aphid Drowhi ihirsoia Philipp Agric 13277-288

228 Tanaka 1 1957 Taxonomy and distribution of some subterranean aphids injurious to upland rice in Japan with description of a new species [in Japanese English summary] Botyu-Kagaku 22168-176

229 Tanaka T 1961 The rice root aphids their ecology and control Spec [full College of Agriculture Utsunomiya University Japan F3 p

230 Tauber M IC A Tauber R Nechols and 1Obrycki 1983 Seasonal activity of parasitoids control by external internal and genetic factors Pages 87-108 itt I)iapause and life cycle strategies in insects V K Brown and I Ilodek eds )r W Junk The Hague Netherlands

231 Taylor W F 1972 Effects of artificial defoliation (simulating pest damage) on varieties of upland rice Exp Agric 879-83

232 feotia T P S and S Natud 1966 Biotoomics of the rice skipper Parnaraniathiis Fabricius (Lepidoptera Hesperiidac) Indian J Entomol 28181-186

233 Teran F 0 1971 Rice pests of Santa Cruz [in Spanish Ministerio de Agricultura Sita Cruz Bolivia Bull 45 8 p

234 Than Itun 1976 Population dynamics of the yellow rice borer Tryporza itcertulas (Walker) and its damage to the rice plant MS thesis University of the Philippines at Ios Bafios Laguna Philippines 92 p

235 Uichanco L B 1911 Pests Pages 55-74 it Rice Philipp Agric 11-200

IRPS No 123 January 1987 41

236 van der Laan P A 195 1 Life history of the rice seedling fly Contrib Gen Agric Res Stn Boor Indonesia 118 15 p

237 van der Laan 11A 1959 Correlation between rainfall in the dry season and the occurrence of the white rice borer (Scirpolhaga innotala WIk) in lava Entomol Fxp Appl 212-20

238 Van Driesche R G 1983 Meaning of percent parasitismin studies of insect parasitism Fnviron Entomol 121 I1- 122

239 Venugopal M S and A litsingcr 1984 Fffect ofcarbofuran on rice growth Prot Ecol 7313-317

240 Vietnam Directorate of Rural Affairs 1962 Annual work progress report on crop improvement program of rice sugarcane vegetable and field crops (lor the period from July 1961 to June 1962) (11) 299 p (Rev Appl Entomol A51526-527)

241 Wan M T K 1972 Observations on rice leaf- and planthoppers in Sarawak (Malaysian Borneo) Malay Agric 148308-335

242 West Africa Rice I)esclopnient Association 1978 Annual research report Vol 2 Statistics Monrovia Liberia

243 West Africa Rice Development Association 1979 Annual research report p47 Monrovia Liberia

244 Williams ) F and CS lofgren 1981 Fli I illy l-468 anew bait toxicant for control of the red imported lire at Fla Entomol 64472477

245 Williams I) 1 1970 The mealybugs OFlomoptera Coccoidea Iscudococcidac) of sugar-cane rice and sorghum Bull Entomol Res 0109-188

246 Williams D) 1 1982 Pulinaria icervi (Signoret) and its allies ott sugar cane and other grasses Btll Entoinol Res 72111-117

247 Wirjosuiardjo S 1976 Study ofchenmical cont rot of insect pests ott upland rice Imiu Pertanian (Jogjakarta) 1323-329

248 Wirjosuhardjo S and N Wirarta 1976 Chemical control of the white grub tIolorithiatlltriin upland rice in M uneng Irobolinggo Indonesia) [in Indonesian] Imut Pertanian (Jogjakarta) 1330-332

249 Yano K T Miyake and V F Eastop 1983 The biology and economic importance of rice aphids (Henuiptera Aphid idae) a review Bull Entomol Res 73539-566

250 Yelp D and P Rhodes 19(4 lhe success of drilled dryland rice iii the Fiji Islands International Rice Commission Working Party on Rice Production and Protection Manila Philippines 3-10 March1964 Food and Agriculture Organization Rome Italy 13 p

251 Young W R 1981 Fifty-five years of research on the sorghum shootfly Insect Sci Appi 23-9

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axial-flow pump

Page 2: UPLANDRICE. INSECTl--1i M T -E!R.EOLOU t RTANCE ,AND …delphacid.s3.amazonaws.com › 6958.pdf · The significance of abiotic factors in upland rice insect ecology will be apparent

UPLAND RICE INSECT PESTS THEIR ECOLOGY IMPORTANCE AND CONTROL

JA Litsinger AT Barrion2

and Dandi Soekarna3

ABSTRACI

The diversity of upland rice environments gives rise to a more heterogeneous insect fauna compared with the more homogeneous lowlands A wide array of soil-inhabiting pests - ants termites white grubs crickets root aphids root mealybugs root bugs and wireworms -- common in upland rice cannot tolerate flooding Seedling maggots replace the aquatic whorl maggots as vegetativc foliar pests lnsect-vectored virus diseases arc rare in upland rice Small upland ricefields cause concentrations of the more vagile seed pests during ripening The less stable upland environment -- more restricted growing season smaller area planted greater drought stress - poses greater problems of survival to insects which have overcome them by polyphagy greater longevity off-season dormancy andor dispersal There is no one insect that specializes in upland rice Yield losses to insects however are comparable to those of lowland rice Cultural conrol methods include increased tillage higher seeding rates and crop rotation for soil pests Foliar pests can be minimized by synchronous planting of earlyshymaturing varieties Plant resistance as a method of insect control has iot been greatly exploited because the other breeding objectives of high yield drought tolerance and blast resistance take priority Resistance to stem borers should be a high priority A rich fauna of natural enemies exists but they face even greater problems of survival than the pests The normal low yields of upland rice preclude high levels of insecticide use Seed treatment and baiting are low-cost mnethods as is low-volume sprayi 1g

Entomologist Entomology Department International Rice Research Institute (IRRI) PO Bo- 933 Manila Philippines 2Scnior research assistantEntomology Department IRRI 3Senior researcher Pest and Disease Section Bogor Institute for Food Crops (BORIF) Jalan Cimanggu Kecil 2 Bogor Indonesia

UPLAND RICE INSECT PESTS THEIR ECOLOGY IMPORTANCE AND CONTROL

Upland ricefields are nonpuddled and unbunded without the expectation of impounding water They may be located in many settings - in isolated pockets surrounded by irrigated wetland fields or along steep slopes of recently cleared forest Each enviionment produces a unique composition of insect species

Information on upland rice insect pests is limited and scattered among reports dealing with other rice cultures We found eight reports specifically treating insect problems of upland rice culture (11 59 66 79 116 120 163 176) but none worldwide in scope Most literature focuses on one species or a group of related species Table I lists upland rice insect pests that others have identified and those we have observed Many citations refer to lists of pests with rice as a recorded host or are based on insect habits

Papers concerning upland rice often fail to distinguish between the two ecologically significant aspects ---culture and environment embodied in th term upland rice It is dangerous to extrapolate results of studies of insect pests in well-drained fields where seeds of upland rice

varieties are sown into dry soil (upland rice culture) but in an otherwise lowland environment (fields surrounded by flooded rice) Insects readily fly from the lowland rice-fields to the fields planted in upland rice culture These insects therefore are not necessarily upland rice insects They breed on lowland rice and colonize upland rice We gain greater ecological significance from faunistic records and ecological studies from an upland rice environment where lowland ricefields ar- out of the effective dispersal range of most insect species (tens of kilometers away) In border areas where lowlands and uplands meet studies cannot accurately represent either upland or lowland rice (181)

IPEST (iROUPS

Upland rice insects are more influenced by physical than by biological or socioeconomic parameters I) well-drained soils (lack of prolonged flooding or soil puddling) 2) high probability of drought during crop growth 3) restricted growing season (lengthy nonrice fallow) 4) ricefields interspersed with other crops (diversified flora) and 5) low use of agrochemicals (because of low and unstable yield)

Although a dryland habitat represents an extreme hydrological condition upland rice is host to all but the most aquatic insects (11 176) Whorl maggots caseshyworms water weevils and bloodworms require ponding Many soil and scedling pests are not common in lowland ricefields Deep water rice is established in dry soil and therefore has more in common with upland and rainfed lowland rice than with irrigated rice even though water depths may later reach 1-3 inSecond to deep water rice upland rice represents the most unstable rice environment for foliar insects But upland rice is highly stable for soil insects

The significance of abiotic factors in upland rice insect ecology will be apparent in a discussion of the most prevalent groups of insects attacking upland rice worldwide

Soil -inhabiting pests Well-drained nonpuddled upland rice soils favor pests that pass at least one growth stage underground Soil pests feed on underground plant parts (sown seed or roots) develop entirely in a subterranean habitat and leave only for adult mating and dispersal flights Soil pests include ants (Formicidae) termites (Isoptera) mole crickets (Gryllotalpidae) field crickets (Gryllidae) white grubs and black beetles (Scarabacidac) root aphids (Aphididae) root-feeding mealybugs (Pseudococcidae) root-feeding bugs (Lygacidac Cydnidae) false wireshyworms (Tenebrionidac) wireworms (Elateridae) root weevils (Curculionidae) and soil-inhabiting cutworms (Noctuidac)

A subterranean environment limits mobility particularly in locating food and soil insects have adapted by 1)being long-lived either as individuals (beetles or orthopterans) as colonies olsocial insects (ants or termites) or as dependents on social insects (mealybugs and aphids) and 2) having a wide host range (all species)

Anis Species of -rrestrial ants notably the ubiquitous fire ant Solenopsis geminata and harvester ants Pheidole spp specialize in feeding on ungerminated grass seed Solenopsis readily colonizes disturbed habitats which in turn initially encourage the growth of grasses (39) Colonies of these granivorous ants have a specialized caste that processes seed for food Foraging workers bring seed to

4 IRPS No 123 January 198

Table 1Insect pests of upland rice worldwide

Order Family Species

Isoptera Rhinotermitidae Coptotermesformosanus Shiraki

Termitidae Heterotermesindicola (Wasmann) Anacanthotermesviarum (Koenig) Anacanthotermesrugifrons Mathul et Sen-Sarma Captiterpnesnitobei (Shiraki)Corniterrnesstriatus (lHagen) Procapriterines mnushae Oshima Pericapritermes nigerianus Silvestri (=socialis)Macroternes nutalensis (Haviland)Macrotermes beiiicosus Smcat hman Macroterines gilius (Hagen) Macrotermnes spp Odontoterniesformosanus (Shiraki)Procornitermesarauloi Emerson Procornitermws triacifer (Siivestri) Trinerviterniesgenzinatus Wasmann

[= ebenerianus Sjostedt]MicroceroterynessppMicroterinesspp Synternes nolestits (Burmeister)

Dermaptera Forficulidae Termites unspecifiedDiaperasticuserythrocephalusOlivier

Orthoptera Gryllotalpidae Don lineare (Eschscholtz)GryllotalpaafricanaPalisot de Baeuvois Gryllotalpaorientalis (=africanu) Burmeistera

Gryllidae

Neocurtilla (= Gryllotalpa)hexadacryla (Perty) Scapteriscusdidact)hts(Latreille) Grjllusassinilis(= bimaculatus)(Fabricius)Gryllus (=Liogryllus) bimaculatusde Geer Brachytrupes portentosus (Lichtenstein) Brachytnmpes mnembranaceus DruryPlebeiogrylhsplebejus (Saussure) Teleogryllus testaceus (Walker)Teleogryllus occipitalis (Serville) Loxoblenimus haani Saussure

Pyrgomorphidae

Velarifictorusaspersus(Walker)Velarifictonis sp Euscyrtus concinnus (de Haan)Atractomorpha burri I Bolivar

Tettigoniidae A tractomorphapsittacinapsittacina(de Haan)Conocephahslongipennis(de Haan) Conocephalusrnaculatus(Lethierry) Conoceptahissaltator Saussure Euconocephah s varius (Walker)

Tetrigidae Acrididae

Caulopsiscuspidatus (Scudder) Caulopsis oberthuriScudder Phaneroptera furcifera (Stal) Amphinofus spp Aiolopus thalassinustamulus (Fabricius) Acrida ivilleinsei Dirsch Gesomida mundata zonocera Navas Gonistabicolor (de Haan)Oxyajaponicajaponica (Thunberg)Oxya hya intricata (Stal) Oxyafitscovittata (Marscliall)Oxya velox (Fabricius)Gas frimargus niarmoratusgrandis (Saussure) Xenocatantopshunilishumilis ServilleHierogiyphus banian (Fabricius) ltieroglyphusdaganensis Krauss Hicroglyphus nigrorepletus(1 Bolivar) Hieroglyphus oilziiorosCarl Heteropternisrespondens (Walker)Patangasuccincta (Linnaeus) Stenocatantops splendens (Thunberg) Chondracrisrosea brunneri Uvarov Trigonidium cicindeloides Rambur Locusta migratoriamanilensis (Meyen)Locusta migratoriacapito Saussure

Distribution

Asia Asia Asia Asia Asia Latin America Asia Africa Africa Africa Asia Africa Asia Latin America Latin America Africa

Asia Asia Africa Latin America Asia Africa Latin America Africa Asia Latin America Latin America Asia Asia Asia Africa Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Latin America Asia Latin America Latin America Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Africa Asia Asia Asia Asia Asia Asia Asia Asia Africa

Reference

85 82 190 190 82 85 151 82 85 7 85 85 178 12 82 85 85 195 195 85

193 11 12 157 60 85 195 72 36 186 12 32 59 72 76 84 81 150 195 203 211 81 76 178 b 48 b 76 178 b b b b b 55 178 178 178 65 178 68 81 68 178 178 178 178 178 55 72 178 178 78 178 178 55 55 72 6 82 82 55 54 55 59 130 55 178 55 78 55 82

IRPS No 123 January 1987 5

Table I continued

Order Family Species Distribution Reference

Locusta migratoria migratorioidesReische Africa 33 et Fairmaire

Schistocercagregaria (Forskal) Africa 33 Schistocerca americana (Drury) Latin America 82 195 Valanga nigricornis (Burmeister) Asia 82 Orphulella inticata Scudder Latin America 68

lomoptera Coccidae Pulvinaria sorghicola De Lotto Africa 246 Pulhinariaiceryi (Guerin) Latin America 186 246

Pseudococcidae taennococcus spp Asia 120 Lhorizococcus iu Williams Asia 245 Dysmicoccus boninsis (Kuwana)C Asia Africa 245

Latin AmericaDysmicoccus brevipes (Cockcrell)c Asia Africa 117 203245

Latin AmericaDysmicoccus oryzae (Wirjati)c Asia 245 Geococcus (=Ripersia) oryzae (Kuwana)C Asia 245Brevennia (= tererococcus)rehi (Lindinger) Asia 72 245Nipaccoccus graminis (Maskell) Africa 245 Planococcoides lingnani (Ferris) Asia 245 Pseudococcus saccharicola Takahashi Asia 245 Pseudococcus spp Latin America 60 Saccharicoccus sacchari (Cockerell) Africa Asia 245

Latin America Trionymus ceres Williamsc Asia 119245j rionymus sp Asia b Mealybug unspecified Asia 83 84

Aphididae Capitophorus[= Rhopalosiphumn] prunifoliae Asia 221 (Shinji)

Rhopalosiphum maidis (Fitch) Asia 249 Rhopalosiphum padi (Linnacus) Asia Africa 12 97 249 Rhopalosiphum rufiabdominalis (Sasaki) Asia Africa 66 76 120 195203

Latin America 221229240249 Tetraneura akinire Sasaki Asia 221249Tetraneurabasui ltillcRisLambers Asia 249Terraneura nigriabdominalis (Sasaki) Asia Africa 712 30 76 227229 (=Dryopeia hirsuta)

Tetraneuraradiciola Strand Asia 249 Paracleus ciniciformnis von Heyden Asia 249

[= Forda harukawai Tanaka]Pineus hanikawai (lnouye) Asia 228 Geoica hcifitga (Zehntner) Latin America 76 195 229 249

AsiaGeoica setulosa (Passerini) Asia 249 Anoecia fuliabdominalis (Sasaki) Asia 249 Anoecia comi (Fabricius) Asia 221229249Hysteroneura sctariae (Thomas) Africa Asia 7

Latin America Sipha glyceriae (Kaltenbach) Asia 249 Root aphid unspecified Asia 72 120

Cercopidae Deois schach (Fabricius) Latin America 61 68 194 Sepullia (=Denoplux) nigropunctata Stal Africa 29 Tomaspis ( Deois) fluxtuosa (Walker) Latin America 61 68 194 Tomaspis ( Dcois) flaiopicta (Stal) Latin America 61 61 194 To naspis (Dcois) complcta (Schmidt) Latin Aincrica 68 Tomaspis ( Zulia) entreriana (Berg) Latin America 68 Tomaspis ( Mahanarva) fimbriolata(Stal) Latin America 68 194 Tomaspis ( Aeneohna) spectabilis (Distant) Latin America 219 233 Aeneolamia varia Fciinai Latin America 81 111 Aeneolamia postica (Walker) Latin America 203 Abidama producta Walker Asia 54Delphacidae Sogatella furcifera (Horvath) Asia 59241 Niapanatalugens (Stal) Asia 30 59 72 84 241 Sogatodes oryzicola (Muir) Latin America 6167150Sogatodes cubanus (Crawford) Latin America 150 Sogatodes pusanus (Distant) Asia bPeregrinus maidis (Ashmead) Latin America 186 Laodelphax striatellus (Fallen) Asia 18

Cicadellidae Recilia dorsalis (Motschulsky) Asia 59 241 Nephotettiv nigropictus (Stal) Asia 59241

6 IRPS No 123 January 1987

Table I continued

Order Family

Hemiptera Lygaeidae

Cydnidae

Alydldae

Coreidae Corimelaenidae

Pentatomidae

Thysanoptera Thripidae

Species

Nephotettix virescens (Distant) Nephotettix modulatus Melichar Cofana spectra(Distant)Graphocephala spp Hortensiasimnilis (Walker) Exitianusobscurinervis (Stal)Balclutha spp Draeculacephalaclypeata Osborn Cicadulinabipunctella (Matsumura)Cymoninus turaensis(Paiva) Ninus insigniStal Pachybrachiusnenosus Horvath Caenoblissuspilosus Barber Blissus leucoptens (Say) Blissus spp Dlmnorphoptentscornutus novaeguineae GhauriDimorphopterus similis Slater Paroniuspiratoides (Costa)Pangaeus spp Aethus indicus (Westwood)Geotomnus pygmaeus (Dallas) Stibaropus inolginus Schiodte Cyrornenusbergi FroeschnerCyrtomnenus ciliatus (Palisot de Beauvois) Cyrtonenuscrassus Walker Tominetus spp

Riptortuslinearis (Fabricius) Riptorts spp Leptocorisaacuta (Thunberg)Leptocorisabiguttata (Walker)LeptocorisapalawanensisAhmad Leptocorisaoratorius(Fabricius)Leptocorisachinensis Dallas LeprocorisasolomonensisAhmad Leptocorisaspp Stenocorissouthiwoodi Ahmrad Stenocoris claviformis Ahmad MirperustorridusWestwood aetus trigonus (Thunberg)Alkindus atratusDistant ScaptocorisdivergensFroescher Scaptocoris castaneusPertyScotinopharatarsalis (Vollenhoven) Scotinopharascotti Horvath Eysarcoris (=Stollia) ventralis (Westwood) Dolycorisindicus Stal Macrinajuvenca Burmeister Tanda antiguensis (Westwood) Tantiageifi Schout Tantia perditor (Fabricius) Pygotnenidavaripennis (Westwood) Menida sppNezara viridula (Linnaeus)

Oebaluspoecilus (Dallas) Oebalusypsilon-gnseus (de Geer)Oebalusgrisescens (Sailer) Tibracea limbativentris (Stal)

Aspavia armigera(Fabricius)Diploysfissa Erichson Acrosternum marginatum (Palisot de Beauvois) Acrosternum sppFrankliniellarodeos Pergande Breginatothrips enustos Watson Stenchaetothrips(=Baliothrips)biformis

(Bagnall) (=(Jlloethripsoryzae (Williams)HaplothripspriesnerianusBagnall

Distribution

Asia Africa Asia Latin America Latin America Latin America Latin America Latin America Asia Asia Asia Asia Asia Latin America Asia Asia Africa Asia Latin America Asia Asia Asia Latin America Latin America Latin America Latin America

Asia Africa Asia Asia Asia Asia Asia Asia Asia Africa Africa Africa Asia Latin America Latin America Latin America Asia Asia Asia Asia Africa Latin America Africa Latin America Asia Asia Asia Africa

Latin America Latin America Latin America Latin America

Africa Africa Latin America Latin America Latin America Latin America Asia

Asia

Reference

59 241 217 59 241 6166 61 66 81 150 6166 66 81150 b 178 178 30 83 42 81150 203 30 80 62 178 195 178 178 178 203 203 203 203

55 178 12 72 78 120 214 214 b 59 120178114 178 54 636 82 12 6 12 82 178 81 150 60 is 195 59 120 178 78 36 186 36 186 b 55 12 30 54 55 78 82 120 61 66 81195 61 65 66 195 211 150 195 61 66 81 150 219 233 6 12 82 6 82 186 186 61 66 68 213235

178

7 IRPS No 123 January 1987

Table I continued

Order Family Species Distribution Reference

toleoptera Scarabacidae leteroligus (=Aphonoprocts) 15elcs Billberg Africa 36 Dynastinac (=Ileteroligusmees robustui Prell)

Alissonotum pauper Burnieiser Asia 82 Alissonotuon simile Arrow Asia 209 Dscinetusdubibs (Olivie) Latin America 28 195 233 Devscinetusgagates Burmncster Latin America 20 195 Dyscinctt spp Latin America 60 Eutheola Inimilis Bturmcistr Latin America 20 28 8 1 195 203

233Eutheola identarr(Biurnciter) Latin America 121 203 Heteron)ciwv aw rsonJack Africa 82 Hcreron)chusbtuterculatus Kolbe Africa 82 tleteronichtslicas (Klug) Africa 82 fhteron)chus mosamhicus Peringuey Africa 4 6 48 82 (=Veteronychus orzae Britton)

leteronychts lioderes Redtnbacher Asia 77 82 207 ltcteronchusarator (Fabricius) Africa 82 tleterontchusplebepus (Klug) A frica 57 82Ileteron chus pseuocongonnsis Ferreire Africa 32 Ileteronvchns rogiJrons Fairmaire Africa 82 Heteron chus nrsticus niger (K lug) Africa 82 Heteronvchus spp Africa 17 Lachnosterna longipennis (Blanchard) Asia 82 Lachnosternaspp Asia 120 Ligvros fossator Burmeister Latin America 82 Ligyrus nasutus Burmeister Latin America 203 Ligyrus (=Scarabaeus)ebenus (de Geer) Latin America 20 195

Dynastinac Ligtvros huonilis Burmeister Latin America 60 82 Aaladera castanea (Arrow) Asia 82 Maladera orientalis Motschulsky) Asia 82 Malaaerajaponica Motschulsky Asia 82 Phvlognathusdicynsius (Fabricius) Asia 82

Melolonthinae Exopholis h)ypoletca (Wiedemann) Asia 117 Lerlinosterna[= lolotrichia] consanguinea Asia 122

(Blanchard) Leucopholis rorida (Fabricius) Asia 82 117 Lepidiota blanchardi Dalla Torre Asia 76 82 Leucopholis irrorata (Chevrolat) Asia 76 134 Scrica internpta Walker Asia 76 Holotrichialonipennis (Blanchard) Asia 122 Holotrichia serrata Fabricius Asia 122 Holotrichia leucoph thabna (Wiedemann) Asia 82 Holotrichia seticollis Moser Asia 77 78 Holotrichia pnindanaoana Brenske Asia b Stenoeres spp Latin America 71 195 Atoenius spp Latin America Asia 203 Ph luplg aequata (Bates) Latin America 203 Piyvilophagacaraga Saylor Latin America 203 Phyllophagachiriquina (Bates) Latin America 203 Phyllophagadas)poda (Bates) Latin America 203 Phyllophagaclegans Saylor Latin Ameica 203 Phyllophagahondura Saylor Latin America 203 Phyllophagalatipes (Bates) Latin America 203 Phyllophagamontriesi (Blanchard) Latin America 203 Phyllophaga pamisetis (Bates) Latin America 203 Ph vllophaga sanfosicola Saylor Latin America 203 Phllophagasetifera (Burmeister) Latin America 203 Phyllophaga (=tlolotrichia)helleri Breuske Asia 82 117 159 Phvllophagaspp Latin America 81 150 White grub unspecified Asia 84

Rutclinae Lagochile trigona (Ilost) Latin America 82 Anomala dinidiata var barhata Burmeister Asia 77 78 82 Anomala huteralis(Burmeister) Asia 76 Anoimala lurida (Blanchard) Asia 76 Anomala sulcatula (Burmeister) Asia 76 Anontala varians (Olivier) Asia 82 Anomala antigua (Gyllenhal) Asia 82 Anoinala palida (Fabricius) Asia 82

8 IRPS No 123 January 1987

Table 1 continued

Order Family

Tenebrionidae

Elateridae

Languriidae Chrysomelidae

Chrysomelidae

Coccinellidae Curculionidae

Lepidoptera Lymantriidae

Amatidae

Arctiidae

Limacodidae Eupterotidac Noctuidac

Species

Anomaa volita (Blanchard) Adoretus caliginosus Burmeister Adoretus comprcssus (Weber) Papuana huhneri (Fairmaire) Papuana incrmnis Prell Popillia capricollis Hlope Gonocephalu depressur (Fabricius)Gonocephahm acuiangulum (Fairmaire)Gonocephalum sinphx (Fabricius) Epitragus sallei Champion Anaedus ptnctatissimnts ChampionAeolus spp4griotes spp Agriotes mancus (Say) Conoderts spp Aeoloderma brachmana (Candeze)Wireworni unspecified Anadastus filijf rmis (Fabricius)Dicladispa viridicanca(Kraat z)Trichispasericea (Guerin)Oediopalpastervalis (Weise) Oediopalpa guerini (Italy) Chaetocnema cvlindrica (Baly)Chactocnema denticulata S tephens Chaetocnemabasalis (Baly)Aulacophora sp nr similis Olivier Monolepta caripemne Ialy Monolepta bilasciata (Ilornstedt) Monolepta signata Olivier Cerotoma airojcsciata J acoby Diabrotica adelpha Ilarold Diabrotica balteata Leconte Diabrotica graminLa ltaly Diabrotica speciosa (Germar) Diabrotica limitafa quindecinmunctata(Germar)Diabrotica melanocephala (Fabricius)Diabrotica spp Altica (= Fondia)madagascariensis(Allard)Altica spp Epitrix cuctmeris (Harris) Epitrix spp Oulenjo or pae(Kuwayama) Disonchq (= Donacia) spp Flea beetle unspecified Chnootriba (= htIilachna)sinilis assimilis Mulsant Atactogaster iduriens Walker Hypomeces squtamosus (Fabricius) Tan vinecus discoidalis GyllenhalPhaulosoinusmusculinus (= mnus) Csiki Neobaridia anplitarsis CaseyPsalis pennatula (Fabricius) Laeliasufihsa (Walker) Euproctis irgtncula (Walker)Euproctis minor (Snellen)Euproctis xanthorrhoea (Kollar)Amata sp

Diacrisia obliqua (Walker) Creatonotusgangis Linnacus Latoia (= Parasa) bicolor (Walker)Nisaga simplex Walker Sesamia botanephaga Tams amp Bowden Sesamnia nonagroides botanephagaTfums et Bowden Sesamia calamistis llampson Sesamia inferens (Walker) Spodoptera eridania (Cramer)Spodoptera mauritia acron)ctoides Guenee Spodoptera frugiperda ( ESmith)

Distribution Reference

Asia 82 Asia 82 Asia 82 Asia 82 Asia 82 Asia 78 Asia 82 117 178 Asia 82 117 Africa 82 Latin America 203 Latin America 203 Latin America 195 Latin America 195 Latin America 203 Latin America 195 Asia h Asia Latin America 60 73 117 Asia 200 Africa 36 Africa II Latin America 6166 68 Latin America 6166 68 Asia 225 Latin America 61 66 68 81

Asia 178 Asia 55 Asia 55 Asia h Asia 55 Latin America 203 Latin Amefia 203 Latin America 203 Latin America 203 Latin America 6166 68 Latin Amcrie 195 Latin Amw 195 Latin Aierwca 61 81 Africa 57 Asia 178 Latin Americ 203 Latin America 81 Asia 82 Latin America 195 Asia 120 Africa 6 11 Asia 185 Asia 120 Africa 82 Africa 82 Latin America 66 68 195 233 Asia 117 Asia 117 Asia 117 Asia 117 Asia 117 Asia A S Pradhan

unpubl dataAsia 82 Asia 82 Asia 82 Asia 182 Africa 12 Africa 6 36 Africa 6 8 11 12 36 Asia 59 82 84240 Latin America 150 Asia -0 59 196 Latin America 28 61 66 68 81 150

152195203219233

9 IRPS No 123 January 1987

Table I continued

Order Family Species Distribution Reference

Spodoptera exigua (Hubner) Spodoptera exem pta (Walker) Spodoptera ornithogalli (Guenee)

Africa Africa Latin America

32 217 32 33 36217 150

Mythimna loreyi (Duponchel) Asia 30 Mythimna separata (Walker) Mythimna (- Pseudaletia)latifascia [= adultera

Asia Latin America

178 195

(Schaus)] (Walker)Mythimna (=Pseudaletia) sequax (Fabriclus) Latin America 68 Mythimna rosellinea (Walker) Asia b Mythimna yu (Guenee) Asia b Mocisfrugalis (Fabricius) Asia 30 117 Mocis latipes (Guenee) Latin America 66 68 149 150 203

Platysenta (=Spodoptera) compta (Walker) Asia 219233A Barrion unpubl data

Agrotis ipsilon (Hufnagel) Latin America Africa

33 68 81 150 1)5 203

Agrotis spp Latin America 60 Achaeajanata (Linnaeus) EldanasaccharinaWalker

Asia Africa

117 36

Armyworm unspecified Asia 72 84 Pyralidae Cnaphalocrocis medinalis (Guenee)

Marasmia bilinealisHampson Asia Africa

30 4559 72 84233 36

Marasmia trapezalis(Guenee) Africa 32 217 Marasmia (=Sumia) exigua (Butler) Asia 220 Marasnianmralis (Walker) Asia b Marasiniapatnalis Bradley Marasiniaspp

Asia Asia

b b

Leaf roller unspecified M2liarphaseparatellaRagont

Asia Africa

72 12 57

Maliarphasp Asia b Chilo auricilius Dudgeon Asia b Chilo partellus (Swinhoe) Chilo polychrysus (Meyrick)

Africa Asia

12 92 59 72

Qhilo diffusilineus J de Joannis Africa 6 1229 Chilosuppressalis(Walker)Cibozacconius Blezzinski

Asia Africa

59 72 6 11 12

Cravubus spp Latin America 203 Acigona loftini (Dyar) Latin America 203 Acigona chrysographella(Kollar)Scirpophagaincertulas(Walker)

Asia Asia

82 59 72

Scirpophagainnotata (Walker) Asia 59 200 Scirpophaganiuella (Fabricius) Asia b Elasinopalpuslignosellus (Zeller) Latin America 28 60 61 66 81 150

Diatraeasaccharalis(Fabricius) Latin America 162195203219233 28 66 81 111 150 195

Rupela albincllaCramer Latin America 111

Gelechlidae

Stem borer unspecified Undetermined stem borer nr Maliarpha BrachniaarctraeaMayr

Asia Asia Asia

72 b 55

Brachmi spp Africa 32 Hesperiidae Parnaraguttata Bremer et Grey Asia b

Parnaranaso Fabricius Asia 55 Pelopidasagnaagna (Moore) Pelopidasmathias (Fabricius)

Asia Asia Africa

164 36 155217

Pelopidasconjuncta conjuncta (Herrich-Schaffer) Asia b Borbo fanta Evans Africa 6

Satyridac Melanitisleda ismene Cramer Asia 84213 Mycalesis asophisHew Asia 30

Diptera Cecldomyiidae Mycalesis spp Orseoliaoryzae (Wood-Masn)

Asia Asia

b 72 88 131 181

Orseoliaoryzivora Harris et Gagne Africa 61132218 Chloropidae Steleocerellus (=Mepachymens) ensifer (Thomson)

Oscinella spp Asia Asia

205 205

Gaurax sppChloropsoryzae Matsuntura (= kuwonae Aldrich)

Asia Asia

205 82

Mucldae Atherigonaoryzae Malloch Asia 30 82 117130 Atherigona exigua Stein Asia 82 117 178

10 IRPS No 123 January 1987

Table I continued Order Family

Diopsidae

Agronyzidae

lHymenoptera Formicidae

Species

Atherigona oritentalisSchiner Atherigona indica MallochDiopsis longicornis(= thoracica)Macqurt Diopsis spp Pseudonapnyza asiatica Spencer Pseudonapomyza spicata (Malloch) Agronoza orvzae MunakataAcromyren landoltibalzani(Emery) Acromyrmex landoltifracticornis(Forel)Acromyrnex heyeri Forel)Atta bisphaericaFore Atta capiguaraGoncalves Atta laeLigata (F Smi h1) Atta opacicepsBorgmeierSolenopsisgeminata (abricius)Pheidole sp Pheidolegeton diversus Jerdon Monomorium pharaonis (Linnaeus) Ants unspecified

Distribution Reference

Asia 82 Asia Africa 82205Africa 11 1248 Africa 217 Asia 213 Asia h Asia bLatin America 6168 195 Latin America 61 195 Latin America 61 68 195 Latin America 61 68 195 Latin America 14 61 68 195Latin America 61 68 195 Latin America 61 195 Asia b Asia b Asia 1) Asia b Asia 120

aG orlentalls formerly recorded in Asia as G africana thPhilippines reported in this study CSubterranean root feeders

subterranean nests Foraging ants will feed on germinatedseeds only when the supply of ungerminated seed is low However these ant specialists feed on a wide varietyplants and thus help control grassy weeds (39) This dual

of

pest benefit role extends from tending ants which protecthoneydew-excreting pest species including root aphids and mealybugs (workers repel potential predators and even selectively kill parasitized aphids) to predators of leaf- and stem-feeding rice pests such as leaffolders armywormswhite grubs and stem borers (1 17)

The upland rice environment is particularly well suited to ants because of the lack of flooding Frequent tillage perpetuates grasses and granivorous species that are highlyadapted to constructing new nests in recently tilled fieldsAnt nests however are relatively shallow and tilling with a moldboard plow can destroy them Also upland ricefields are uually small and isolated and a given area in a smallfield has more field borders than it has in a larger fieldallowing high rates of infiltration from more sedentaryspecies such as Plieidole that do not colonize tilled fields but nest in soil under shrubs and trees along field bordersPheidole workers forage into fields from borders and store seed in their nests In slash-and-burn or other no-tillage rice cultures nests would be preserved in the field and possiblylead to high rates of seed predation However granivorous species do not prefer forest habitats

Damage is characterized by reduced usually patchy plantstand (120) Seed-thieving ants greatly prefer rice but they can be selective regarding plants they collect and store Theyharvest more seed than they can consume so patches of germip ing surplus seed (such as Rottboellia) will sproutfrom their nests during the rainy season

In Latin America leaf-cutting ants (Aita spp and Acromyrmex spp) occur only in upland fields and defoliate

young rice plants (47 68) They take leaves to fungal gardensin underground nests where fungi predigest the plant material

Termites Subterranean termites of the family Termitidae which lack symbiotic protozoa to help digest plants are the most frequently mentioned termite pests of upland rice (1185 120) This family cultures fungi in special undergroundcells --- fungal combs made of half-digested plant material (117) The fungi inoculated onto the combs break down plant material into food that termits can digest The termites then consume the fungal combs These grasslandtermites build nests below the plow layer in uplandricefields The colonies attack living plants only when dead plant material is not available They will attack a droughtshystressed crop but prefer older plants having greater cellulose content They will damage newly planted crops where clean culture has removed vegetation

Infestations are worse in deep light-textured soils with low moisture content The first sign of damage is yellowingof older leaves (195) Termites feed on roots (the plantyellows then wilts and finally collapses) and germinatingseed (loss of stand) or move above ground at night to cutseedlings at ground level which they cover with soil for later consumption Soft-bodied termites are highly sensitive to desiccation They live within a self-contained system of airtight chambers and build surface tunnels linei with nudand body secretions (carton) to maintain more than 90 relative humidity Termites can be located by their tunnels

Termites apparently can withstand limited submergence (157) and can be a pest of dry lowland seedbeds (117)perhaps by having nests below perched water tables

Termites appear to do more damage in Latin America where large ricefields are plowed with tractors deprivingtermites of preferred vegetation and forcing them to live on

rice plants In Asia upland ricefields are typically small with Iprge field border areas (ecotones) where termites can forage for preferred food other than growing rice plants All grassland termites do not feed on rice In an upland area in Batangas Province Philippines ricefields are commonly infested by Hosiititernesluzonensis (Oshima) however this termite does not damage rice In the same province though Mrroternesgivus(Hagen)feeds on rice seedlings (178)

In Africa termites may damage rice even though fields are small and surrounded by perennial vegetation These termite species perhaps prefer rice over other hosts

Mole crickets Nymphs and adults are nocturnal and burrow through soil feeding on roots of a wide variety of plants or forage above ground as predators of insects Adults live 3-5 mo and are even cannibalistic Mole crickets prefer low-lyin moist upland soils with high organic matter (II 81) In the lowlands they inhabit rice bunds or nonflooded fields Adults are highly mobile and can leave a flooded field to locate a more suitable habitat Losses show as wilted plants (from root feeding) or reduced stands (120) Damage is more common near field borders where mole crickets relocate after tillage (126)

Fiehl crickets Nymphs and adults of field crickets have similar nocturnal habits and damage rice as much as mole crickets do Piles of weeds removed from fields attract them They make subterranean nests and tunnel through the soil to feed on roots Some species prefer seed to roots Others feed at the base of stems causing deadhearts

Tunnel entrances surrounded by excavated soil are easily seen Young plants are cut at ground level and stored in underground cell Field crickets tre highly polyphagous as are all soil pests (117) Sachan et al (198) reported that field crickets prefer maize to upland rice Like mole crickets field crickets cannot survive in standing water and are therefore more prevalent in upland fields

Whitegrubsandhackbeetles Scarab beetles that feed on living roots as larvae but not as adults are called white grubs or chafers (Melolonthinae Rutellinae) and those that feed as adults but not as larvae on plant crowns are called black beetles (Dynastinac) In the tropics chafers or white grubs have a I-yr life cycle starting with adults emerging from the soil about I mo after the first downpour of the rainy season (134 159) They lay eggs at the same time farmers sow upland rice The rice passes its most susceptible stage and damage is mostly avoided when the white grubs are small After several months the long-lived larvae are large enough for two or three to denude the root system of mature rice (223) This intensity of damage is rare but wilting occurs when root loss is combined with drought stress

The larvae need damp (not saturated or dry) soil to survive and pass the unfavorable dry season 05-2 m underground in aestivation in the moist soil The first heavy rains of the season (20-30 amd) stimulate the grubs to resume activity and after several weeks they develop into

IRPS No 123 January 1987 11

pupae and adults and eventually dig their way to the soil surface and fly to nearby trees to seek food and mates The adults of most white grub species feed only on trees but in some species notably Eutheolahumilis the adults defoliate and cut off stems at ground level (28) Putheola hidentata damages sugarcane fields in Venezuela (121) Floods kill white grubs (208) Grasslands can support large populations and therefore white grubs can be more abundant in newly planted upland ricefields that were previously fallow (81)

Black beetles live up to a year adults can attack a crop at any stage by burrowing into the soil (127) and feeding at the base of stems to cause whiteheads (207) Adults emerge with the early rains and are normally more abundant and do more damage on a young rice crop with small root systems (38 81)

The larvae feed only on dead organic matter in dryland fields and do not attack rice Adults arc highly mobile and although sensitive to flooding can invade a field soon after it drains (115)

Root aplhids Several genera of migratory aphids feed on the roots of upland rice (249) Populations build up more in light-textured soils with high percolation rates Damage has also been reported from dry seed heds of lowland rice (52) Tending ants -- Pheidhle Cren aster Tetramoriuw lsius Tapinoma -- are necessary for root aphids to multiply Aiits harbor aphids in their nests over winter or during unfavorable periods and relocate them on rice plants digging tunnels along the root systems to allow these softshybodied insects to penetrate the soil Most intensive studies on aphid life histories have been done in temperate environments (158 222 229) Root aphids fly to rice plants at the beginning of the rice season and pass through several generations continually developing winged forms to relocate on alternate grassy hosts until winter when they move to perennial hosts usually trees Yield loss occurs mainly through reduced tillering but infested plants become yellow and stunted and in extreme cases wilt (227) Aphid populations build up gradually so damage usually begins in the late vegetative and reproductive growth stages

Root-feeding ineal tugs More than half of the mealybug species or rice feed on roots (245) Being soft bodied mealybugs are not adapted to living underground but they survive because tending ants (the same species that tend root aphids) dig burrows for them and move them from plant to plant Well-drained soils ihelp mcalybugs survive Although six species are recorded as subterranean root pests of rice we found no report citing damage Root-feeding mealybugs are probably more prevalent on perennial grasses than on short-lived annual rice

The most common rice mealyb ig Brevennia rehi is a foliar feeder favored by lack ol rain apparently it is responsible for high yield loss in Northeast India and Bangladesh (N Panda pers comm as stated by IRRI [1061) This report needs confirmation Mealybug damage was only briefly mentioned in another report (156)

12 IRPS No 123 January 1987

Root weevils A few species of upland weevils (IHporneces Donacia Atactogaster) attack rice roots as larvae and feed on stems and leaves as adults Damage ispatchy The larvae are not adapted to submergence and are true upland soil weevils isopposed to the so-called water weevils They rarely become sufficiently abundant to cause damage Adults on rice foliage or the soil surface show that the roots are being attacked by the grubs Root weevils are highly polyphagous (185)

Root heeths Soil-inhabiting false wireworms Gono-cephalumt spp are reported in upland ricefields The adults and larvae normally feed on decaying organic matter but land preparation for upland rice removes these sources The larvae feed on seedling roots and nocturnal adults roam on the soil surface to cut off seedlings at ground level (82) This group of ground beetles is highly omnivorous adults even prey on stem borer larvae (183) True wireworms (Elate-ridae) feed on living roots but rice as a host has been reported Infrequently (117) In Latin America the highlypclyphagous larvae of Diabrotica rootworms are also recorded on rice

Root htus Upland rice seedlings can be attacked by the adults anI nlViphs of chinch bugs Blissusand Caenoblissus and bro n bug Scaptocoris which feed on the roots with theirsucking nouthparts (80 83) Three genera of burrower bugs A tms (eotonhus and Stiharopus -are common iil grasslands in the Philippines and may attack the first crop (rice or maize) planted after the dry season Chinch bugs are most abundant after a dry spell These insects are large and can kill seedlings or stunt and reduce tillering on older plants I hey prefer other hosts and rarely damage rice Adults can also feed oii developing rice grains (81)

(utworms Soil-dwelling noctuid larvae hide under-ground during the day to avoid predation by birds and bccone active at night above grour cutting off you ing plants at ground level (hence tile naiie cvtworn) arvaedrag severed plants into hurrows llie larvae have wide host ranges arid pupatc in the soillgrotis ipsilon occurs worldwide and appears to be the most common cutworm pest of upland environments but is more prevalent inlowland rice particularly in dry seedbeds

IFoliar pests Other groups of insects are adapted to upland environments because they pass one growth stage underground either as egg (in heinrietabolous species) or pupa (in holomnetabolous groups)

Rice seedlibig maggots AitJeriota flies occur in non-fhoodcd environments in Asia and Africa (180) Feeding of larvae one per iller causes deadhearts Flies as opposed to moths only secondarily evolved as pliytophagous pestsfroni a saprophagous Origin Athrnonalarvae secondarihv feed onidecaying tillers whereas larvae of stem borer moths actively Iced on living tillers to cause deadlicarts Seedling maggots will die if they cannot sever a tiller on which to feed

as it decays Plants die under heavy attack whereas lighter infestations cause stunting delayed maturity and ragged leaves (236)

Eggs are laid only on actively growing plants (49) aid damage occurs wiihin the first week after crop emergence(170) Larvae pupate more in stems than in soil Seedling maggot attack is highly seasonal (206) normally peaking 2-3 mo after the onset of the rainy season These pests possiblyaestivatc over the dry season Rice isonly one of their many graminaceous crop and weed hosts

The rice stem maggot (hlorops orzae is adapted to temperate upland and lowland rice growing regions of Asia In Japan and China upland rice grows in mountainous regions a habitat of this chloropid fly The larvae tunnel into tilestems The stem maggot is more damaging during the seedling stage larvac hibernate in grasses

File several dipterans reported as stem maggots in dryshyseeded lowland seedlings inIndia (205) could presumably attack upland rio

Leaf urinrs Icaf miners are common on wheat and maize but are rarely reported on upland rice (213) Three leaf miners are more prevalent on upland than lowland rice inthe Philippines Pvudonalomy a asicanh Vficataand lIAgromrza orzae attack at the seedling stage The larvae develop inside the tunnels they construct in the parenchyma tissue P Vpicata and Aorvzac prefer maizeEehtsine and other grasses to rice spicatacan become abundant on wheat

Leaf beetles Chrysomelid and coccinellid eaf bcetles scrape or otherwise remove leaf tissue as larvae or adults or both This damage accelerates desiccation of plants Tile larvae of sonic species are root feeders Flea beetles Epitrix and (lraetocnetna and rootworms l)iahroticaspp feed on rice foliage only as adults Their root-feeding larvae have hosts other than rice The adults and larvae of the hispa-like Oediolmlpa spp beetles of Brazil and (hmootriha (=EPfishyachra) beetle of Africa feed on upland rice leaves Oealioshypapa spp larvae are leaf miners (mootriha is also pshyvalent in lowland environments (11) Leaf beetles prefer vegetative rice (81)

The rice hispa Dicladipaarnigera is not reported on upland rice and prcers more aquatic habitats In Central Africa however I)cladispm viridicyaiea attacks uplandrice in the vegetative stage as well as lowland seedbeds 7richipasericea is reported on upland rice in West Africa and Madagascar (57)

The rice leaf beetle 1d1ea 01izae riiuech like Chioropsor|rzae is adapted to temperate regions of Ashi and is usually a pest on lowland rice but seldom oi inpland rice in the mountains (82) The adult hibernates on grasses

Armmworms Species of grassland-adapted SpoptloeraPlatysemta and A ithimna larvae create widespread epishydemics They can defoliate ricefields generally in patchesfrom early vegetative growth to harvest (41) Armywornms pupate inthe soil and highly favor upland rice (34) Adults

can disperse long distances to colonize even remote upland areas They become more abundant than their natural enemies particularly after favorable rains following a prolonged drought (196) The drought kills the armyworm and natural enemies (mainly parasites) but the armyworm can recolonize rapidly

The natural increase insoil fertility from mineralization of soil N over the drought period promotes luxuriant plant growth to foster a rapid buildup of these highly fecund species Weeding rice increases tilelikelihood of damage The parasites normally return to control the armyworms but not until after serious damage

Outbreaks on upland rice have been reported in Ghana (3) inzibar (38) Central Africa (36) lPanama (165) Brazil (28) Malaysia (196) and India (167) Defoliation may be severe often leaving only the base of stems Armyworm larvae hide during the day under litter or in soil cracks At night they ascend plants to feed Larvae pupate in the soil

Thrips Thrips frequent upland rice but damage is less than in the lowlands Thrips larvae and adults feed on leaf blades by rasping causing leaf rolling and stippling Thrips prefer vegetative stage rice and their survival increases during drought (heavy rain washes them from foliage) although drought stress usually outweighs thrips damage Drought and thrips cause leaf rolling in upland rice (235) but thrips have been overlooked perhaps becaLse of their small size Thrips are most numerous indry season irrigated rice The combination of favorable plant growth and lack of rainfall is ideal for thrips In contrast the lack of rainfall in upland rice hurts the crop and eventually the thrips whose numbers must decline

In [atin America Frankinilla rodeos attacks panicles before they emerge from the boot calusing sterile grains

Stem hug~s Black bugs are the most common group of rice stem-feeding -lemiptera in Asia The most frequently mentioned species ----Scotinophara coarctaia and Shrida - prefer aquatic habitats and therefore could not occur in upland rice as sometimes reported Upland species in the same gerurs - notably S tarsalis and S scotti -- are morphologically similar and remove sap from stems of grasses and upland rice The upland species are notably more abundant in upland rice planted near forests

In Latin America tile Tibraca ima-large pentatomid tiventris causes deadhearts and whitehcads on upland rice Both adults and nymphs remove sap from internodes of plants more than 3 wk old Tillers are killed by mechanical damage and by the entry of secondary microbial infection This damage is often confused with that of lepidopterous stem borers (233) Eggs are laid on the leaves and stems Nymphs are gregarious This species can also be found on irrigated rice Drought reduces population buildup

Stiem borers Lepidopterous and dipterous stem borers are widely recorded on upland rice but cultural practices and alternate hosts determine abtndance Except for Elasmolmll)uS Iinosellts numbers usually build up toward

IRPS No 123 January 1987 13

the end of the crop growth cycle Therefore late-maturing varieties staggered planting and lack offloodingfavorstem borer buildup Many stem borer species can lie dormant in the stUbbleafter harvest and theirsurvival is encouraged by panicle harvesting (leaving tall stems) and lack of tillage after harvest

Stem borers disperse and lay many eggs Many species are polyphagous however the monophagous MaliarpiasepashyratellaDiopsis Ivmgicornis Scirpophaga incertlas and S innotata can also be abundant on upland rice (hilo Suplressalisdevelops fasterand becomes la rger when reared on upland than on lowland rice (226)

The lesser corn stalk borer lasmopal tt Ittoselhs is semisubterranean and is perhaps the stern borer most adapted to upland rice(129) Highly polyphagous it infests maize peanut and cowpea It attacks seedling rice (one larva can kill tip to four plants before maximum tillering) (195) (hiv and Maliarphastem borers prefer older plants But Scirpophagaspp will attack young plants even in the nursery lasn opalls larvae tunnel into the stems at or below the soil level mausing dcadhcarts Larvae are not found in their turnnels inside sterns beca use vhei disturbed they retreat into cases made from soil particles bound by silk The cases areattached to the tunnel entrance (143) This behavior probably evolved as protection from natural enemies Larvae pupate in the soil The lesser corn stalk borer is more abundant during drought damage is often confused with that of drought (233) This species prefers sandy soils arid its distribution within a field may relate to soil texture (143)

Diatraeasaccharalis aside from the lesser corn stalk borer is frequently a stem borer pest in Latin America According to Teran (233) rice and maize are more preferred hosts than sugarcaine its namesake

Atcitonacrvsograpla is a less known upland rice stem borer in Asia with habits similar to Easinopall s (82) because it is not found in lowlands Acutra has grassy weed alternate hosts As a moth it is often confused with the dark-headed stern borer(hi polur(standstriped stern borer (hi sup)lpsrcais

Other common upland rice stem borers are also reported in the lowlands The dominant species in upland rice vary greatly with location In Uttar lPradesh India (78) and in Japan (123) the pink stem borer Sesamia htfirens is more prevalent hIKenya the principal rice stem borer is Chi partelus

(92) The lowland African sten borers Aialiarpha sepashyratellaDiopsis vngicornis and Sesamia calatnisnis are less abundant Oimupland than on lowland rice(217) but perhaps cause greater yield loss because plant injury is greater if combined with drought css (92) The stalk-eyed stern borer D)ivpsis is more prevalent in wet habitats (6 1I217)

In Central and West Africa Maliarpha and Sesamia calanistis are the dominant stern borers of upland rice (8 9 10 II) (lilo dihlfilineui and Chio partellus inhabit the

14 IRPS No 123 January 1987

upland savannas whereas Chilo zacconio prefers the lowlands (II) Aaliarpha is most abundant in the mangrove swamp habitats (91) coastal regions (16) or rain forest zones (II) of Africa and appears to be the homologue ofScirlpophaga incertulas The yellow stem borr can also be prevalent in upland rice (120 167) The sugarcane borer Eldanasaccharinais so far only a potential pest of uplandrice (91) despite Grist and evers (82) warning

Grasshotl)ers andI locitas Short-horned grasshoppers and locusts -- Locvta Patanrga Schistocerca - layeggs in the soil inhabit grasslands and develop into swarms to seriously damage upland rice from time to time (173) Largenumbers can destroy a field leaving only the stubble In low cmuoroer s giassnoppeirs an cut panicles (173) Farmers may not plant if locust swarms are imminent Prolonged dryweather followed by favorable rain favors development of large swarms (173) Usually breeding in di) grasslands theydisperse toattack crops including upland rice(191) Eggsaresusceptible to desiccation d uring drough t and toinundation during seasonal rains With favorable soil moisture over consecutive seasons natural enemies ca nnot prevent a largepopulation increase leading to migratory swarms (64)

Most grasshoppers on rice are adapted to uplands (107)but lowland rice grasshoppers iervgl(rlhusspp and Ora spp occur occasionally on upland rice (2 82) They lay eggson rice foliage Nymphs are scmiaquatic

The gryllid Lsctrtus coucinnus is a pest of lowland and deep water rice in Bangladesh and Thailand In the Philippines it damages upland rice grown in isolated pockets near lakes The gryllid nymphs and ad ults feed on the central portions of folrage leasing only the midrib and leaf nargins

Leal ler Pvralid moths whose larvae fold leaves to make a feeding shelter occur worldwide (217) but arc appareritly most common in Asia Leaffolde rsattacking rice in Asia are a complex of species (26) (Ctalhahcrcisinedinalis the best-known rice Ia flolde r in Asia may prefer grassy weeds over rice (86) It is a late colonizer of uplandrice U1arasmja eviua a lowland species (220) nreflers rice to grass specres (86) and colonizes upland rice during il early growth stages llarasnria patnalis is the most corn-rnonly encountered species in upland rice in the Philippines Perennial grasses are their alternate hosts and they can survive year-round in upland rice environments The leaf-folders 11arasmia ruralisand Alarasmia spp are least aburndant Dormancy is unknown in leaffolders Cnaplha-ocrocis medhnalis however is known as a migrant (90)Leaffolders remove photosynthetic tissue attack during the flag leaf stage is particularly injurious Popurlations are normally held in check by natural enemies but leaffolders become parlicularly abundant in conditions of high plantfertility and shade

Butterflies Skippers PelopidasInathiasand Borhofanta are more prevalent in African lowland than in uplandhabitats (6) but extensive studies in Japan and China

showed that Pelopidas mathias prefers an upland habitat (155) Parnaragutata seasonally migrates between lowland and upland areas and is equally abundant in both environshyments (164)

The green horned caterpillar Melanitis leda ismene is more abundant in lowlands (212) but our experience in the Philippines suggests that Pelopidas niathiasand 11elanifis leda ismene are often more abundant in uplands than in lowlands Bamboo iaunbuiva spp annually sustains P mathiaspopulations in the Philippine uplands Adults feed on nectar from flowers and migrate to upland rice during the late vegetative stage

iPi erlled h i-1 1d dill L tIC i 7-0 l(become numerous probably because of high egg predationEgg and larval parasitization are often low Butterflies can readily disperse long distances to seek renlote uplandricefields However they have low biotic potential and even it predation rates are low the rarely become abundant Upland environments with their diverse microhabitats can provide more favorable sites than lowland rice plains

tutterfly pests of rice have broad host ranges and become dormant Juring urnfavorable times of the year

IolhagrotslaidopwraThe larvae of many polyshyphagous moths are reported to defoliate upland rice tussock moths lsaispelnnatula Iawlia sUifita luproctisirgtwltaE mnorand P vathorrhoeaslug caterpillar

litotiahicolorhairy caterpillars Nisaga sinp vandA mata spp and wooly bears l)iacrisiaohliqtia and Crvatonotts gangis Many of these polyphgous larvac migrate and are transients in rice including HelhWhis arm iera and 1t zea highly pol phagous species (82)

Gall nidges The Asian rice gall midge Orseoliaorrzae (88 131) and its African counterpart 0 ortzivora(6 11 12 32217) prelerlowland to upland rice Gall midges in rainfed areas are prevalent only during tle rainy season and dispersepoorly Their alternate hosts (wild rices LeersiaIschaetnuan and Pasahmt)are mostlyaquatic grasses and are less likelyto be near upland rice areas In an area of mixed uplandfloating and rainfed lowland rice cultures in northern Thailand upland rice which is ilanted a month earlier was attacked firstbut subsequcent populations were much lower than in rainfed lowla id or floating rice Low numbers of gallmidges passed the dry season orr perennial weeds such as Paspahim (88)

Smot (hid(s Reports of aphids attacking upland rice plants are few (249) Without maize the maize leaf aphidRho alosihhunm maidis attacks upland rice in the Philipshypines The plum peach aphid IIrsteroneuraselariae was reported in Sierra Leerne toinfest leavesand tnriperied grain(7) Siphaglrctriaecaused transverse linrea r necrotic striae iir rice leaves in Italy (172) I-lea v rains held populations in check Rice is not a preferred host for shoot aphids

Leafhoper andI )anthol)pers Cicadellids and delshyphacidsdispersecasilyand readilycolonie upland ricefrom nearby lowland areas A study in Sarawak (241) showed

IRPS No 123 January 1987 15

similar species in upland and lowland rice Nephoiettix The principal vectors of rice virus diseases generally have species dominated The most prevalent species accounting narrow host ranges and alternate hosts in the grass family for 90-95 of the sweep net samples wre N virescens N are mostly annuals dying in the dr3 season Some virus umropictus Nilaparvia hgens Soatellafiircifera Cfana vectors such as BPH can disperse lontg distances but few do

sectraand Recilia dorsalisLowlands had higher popula- Th restriction of upland rice and weed hosts to the rainy tions Katanyukul and Chandaratat (120) monitored hopper season the small area planted to upland rice and the hopper numbers in upland and lowland fields of northern Thaiiand vectors having narrow host ranges all work against virus and found similar results A comparative study of plant- diseases in upland rice hoppers in Fiji showed more brown planthoppe BPH) and In Latin America Sogctdhs o(rzicola and S cuhanus whitebacked planthopper (WBPH) in wetland rice than in transmit hoja blanca virus disease but reports on upland lowland rice and hopperburn was prevalent in lowland rice are rare except in the favored upland areas of Colombia areas with standing water (89) Peru and Venezuela (35 42) S rricolaocurs in Brazil

Hopperburn in upland ricefields in Asia is rare (84) In on upland rice but hojaI blanca has not been reported (67) Batangas Philippines isolated fields of upland rice become S orzicola has a wide host ramgc (10) similar to WBPI- in hopperburned infrequently from WBPH A report from Fiji Asia WBPH however is not known to transmit any virus cites hopperburn from BPH and WBPH (250) In these two disease situations upland rice was grown on highly fertic soil and In Africa pale yellow mosaic virus is endemic to swampy N fertilizer was applied Hoppers respond to better nutrition lowland areas where rice ratoons year-round (19) and is Normally upland rice is grown on poor soil and receives no mechanically vectored mainly by hispid beetles A minor fertilizer vector a flea beetle Chaeiocnema spp is an upland rice

In Asia upland rice ras are normally not far from pest and upland ricetields near endemic areas occasionally lowland ricefields-- a source of hoppers Otherwise hopper contract the virus species with greater polyphagy than BPH or Nelhotettiv Spittle bugs Cercopid nymphs producc a protective virescens would be dominant frothy covering that looks like saliva In Latin America

In Africa leathoppers and planthoppers do not become spittle bugs damage upland rice In parts of Brazil where abundant on upland rice (6 II) probably because lowland upland ricefields are surrounded by pastures adults rice is not widely rlanted nearby The only reported migrating from grasses can damage rice severely and cause hopperburn as from Nilapanrata maeander in breeder complete hopperburn in a young rice crop within a week of plots receiving high N rates in Nigeria (98) Hoppers colonization (194 195) Dispersal to rice occurs even if however are a greter problem in Latin America where pastures are verdant and spittle bugs prefer pasture grasses large grassland areas breed polyphagous species (Gra- to rice (168) Nymphs are rarely abundant enough to phocephala spp Hortensia spp lEvitianus ohscurunervis damage rice Regular rainfall favors their development Balclutha spp and Draeculacephala spp) vhich then (233) disperse to cause hopperburn on seedling rice (68) Rosetto et al (195) reported Graphocep7hala spp and Hortensia spp Seed pests to be equally abundant on upland and lowland rice Alydid and pentatomid bugs are seed pests of upland rice

The leafliopper and planthopper epidemics in Asia that worldwide In Asia Leptocorisa sp prevail Their habits caused severe losses from hopperburn and vijus dicases make them particularly suitable for upland rice 1)thev feed over the past several decades occurred principally on on many grasses (and therefore can survive the earl y wet lowland rice Except in a few isolated cases (160) upland season before rice sets its grain) 2)adults are long-ied and rice has been spared from tungro grassy stunt and ragged mobile allowing them to find isolated plantings 0 rice stunt virus diseases vectorcd by green lcafhopper (GLH) (besides seed bugs only birds and rats appear to have shyand BPI] (123) Virus diseases have occurred in upland rice capability) and 3) adults aestivate in forested areas or where lowland rice was nearby Rice dwarf disease however sugarcane fields during the dry sca son when neither rice nor was first recorded on upland rice in the Philippines (187) grassy weeds are present (I199) Rice seed bugs concentrate Virus diseases can be perpetuated only on livir plants or on small-scale upland riceficlds because they can actively insect vectors and no living rice - plans tooned or search them out The prevalence of groves of trees characshyvolunteer - grows year-round in upland arcas excert in a teristic of upland environments ensures nearby aostivation few with 12 mo of rainfall sites keeping the seed bugs close by

The smaller brown planthopper Laodcelphax striatellu Leptocorisae are uncommon in lowland rice plains occurs in upland rice in Japan (18) It spreads black- because 1) their populations are diluted in a sea of rice streaked dwarf and stripe viruses between winter wheat and planted more or less at the same time (they ca n only feed on barley to lowland rice Early planting of lowland rice rice during the milk to hard dough stages) 2) they lack nurseries spreads these viruses but upland ice has remained aestivation sites and 3) weedy fallow areas are limited uninfected The lowands of Asia are dominated by L oratorius

16 IRPS No 123 January 1987

which lays its eggs hign on tile foliage (199) L acuta and L solonlonlesis oviposit at ground level the former on litter and the latter loose on the ground These egg laying habis explain theirtenvironmental prefcrencecs(214) In Japan the postdiapause adults of L chinensis migrate from moun-tainous areas to upland ricefields during floweiing (1 14)

Pentatornid bugs prevail in Latin America (8182 195) and in Africa pentatomid and coreid bugs arc equally cited (6 82) l)amage is often characterstic of the species (5)rcoris ventralis and Mnida spp are the common

pentatomid seed bugs of Philippine upland rice but they prefer lEchinochva spp grass seed to rice Vearaviritda occurs worldwide and hoften more abundant in the uplands(54) We saw few reports of seed bugs on upland rice but see(d bugs apparently opelrate cqually between by land and upland rice Good dispersal is necessary ior species that call feed for only 2-3 xon a crop and must first find those crops at ripening stages

Other pct species feed n flowers In Mato Grosso Brazil (194) and Santa Cruz Bolivia the panicle weevil Aeo-hariuia amllitarsis attacks rice during flowering (68 233)The larva e rice sten horers adutlts feed at the bases of spikelets caIusing empty grains Also inLatin America thecydnid -lkiituttatralusisa seed bug as anad tilt and nymph (194)

A number of beetles ecd on poilen - coccinellids lialrotia Aulac lhora and Monohpl-a (55) - but becatise riCe is self-pollinating and fcrtiliation occurs before pikelets open they pose no threat to yield

IST ECOLOGY

We now look at characteiistics of the life cycles of upland rice insect pests to lea rn hew each plays an adaptive role in upland rice ecology

Life history strategies Orzasativadoes not tolerate drodgh well so upland rice is highly seasonal normally grown in the wettest months of the year Usually rice is present for less than half of the year The rice-fiee fallow poses serious problems of survival to upland rice insects which have evolved att least four mechanisms to ovrcolne tle cvcl ic-l lack of a hostI) polyphagy 2) longevity 3) dormancy and 4) vagility (dispersal)

Poldt)agy lolyphagy is defined as having hosts of at least two botanical families oligophagy of more than one gents and ionophagy of only one ventus (40) - inour case Orvza Most wild rices however are illi tiC and theieiirc are normally far removed from upland rice hahitat othewise more would be alternate hosts for uplanJ iice insects Thte highly mornophagous rice pests shySciriol)haita hiceritlas Nilal)arvaiahiens Iand Nealo--tettiyvircsctws in Asia and Aalirpha secparatella and 1ioj)sis lngicornis inAfrica - occur on upland rice but

are more abundant in lowland culture They can specialize in rice because they are highly vagile and can attack all growth stages By specializing in rice monophagous rice pests can outcompete related but polyphagous speciesIntensive andI extensive lowland rice culture has favored monophagous species Riqwla alhinella (Cramer) the onlymonophagous species in Latin America is rarely reported on upland ice (I1)

Because of its limited temporal and spatial existence upland rice has not favored the evolution of specialized species Clhang(44)shows evidence that upland rice has only recently been cultured by man from the lowlands thus pests have had little time to fully adapt to the crop All but the lowland-adapted insect pest species have alternate planthosts to rice Root pests with limited mobility are highly polyphagous Pests that specialize in one growth stage eg secdling maggots seed bugs are oligophagous or polyphagous

Most alternat hosts of rice pests are annual grasseswhich also are seasonal Upla_rice is also a host but not the preferred one of the most highly polypliagous spcciessoil pests grass hoppers armvwarrs and Vearaviirielda

The d ry season poses igreat obstacle to ani upland riceinsects survival and only sonic perennial plant hosts are suitable food during the off-seasoin Plolyphagy allows upland rice insects to suLrx-ixe during thC rainy season when plants- both annual and perennial -areactivelygrowing and therefore more nutritious

Loiugevitr The ability of aiinsect to live 2-3 mo without undergoing dormancy helps species sich as root-feedingwhite grbs (as larvae) and secd-feeding bugs (as adtilt) to survive unfavorable periods White grubs arc not highly fecund and their life strategy is to improve survivorship of limited progeny Their subterranean habitat hides them from manv natural enemies lBtut root tissue is not highly nutritious so they ltilst eat great quantities and need a largeinsect biomass to digest it Other soil-inhabiting rootshyfeeding insects have short life cycles but are more fecund

Seed pests need greater longcvi vas adults to locate ahost at early grain dcclopment arnd give sedentary offspring time to develop Few groups of upland rice insects live longin active development Polyphagy and longevity only increase their ability to survive during the rainy seasonOther mechanisms are needed to survive the dry season

orantc In the tropics the nonrice season is the dry season while intemperate areas tile off-+season is winter Many upland rice insects undergo dormancy inlsummer (aestivation) or winter (hibernation) Dormancy is simplyinactivity during winters with prolonged temperatures below 10 C insects find shelter arnd cease movement Activity is resu med with warmer t empcera tu res A deep state of irnactivity termed diapauise occurs in sole species that prepare physiologically (havxing low metabolism and convcrting gonad s to fatty fuel reserves) for the unfavorable season (148) They enter diapaUse inanticipation of unshy

IRPS No 123 January 1987 17

favorable weather and resume activity after favorable Armyworms skippers and locusts actively travel hundreds weather begins usually in response to photoperiod or of kilometers in air fronts such as intertropical convergencethrough a delayed response to a stimulus such as heavy zones to descend onto upland rice fields far from their rainfall (237) Not all individuals of a population have the breeding grounds (31) Several generations can build ip on same threshold for environmental responses for entering or alternate hosts in tile grass family before rice is plantedterminating diapause -- this is seen as a mechanism to These large insects are active fliers Smaller insects such as prevent total mortality if a false cue appears such as a thrips aphids and planthoppers passively migrate long sudden warm spell in winter or rain in the middle of the dry distances by the wind season (154) Otherspecies ean readily travel tens of kilometers - stem

Probably more upland rice insects undL go dormancy borers leaffolders leafhoppes cutworms spittle bugs leaf than we realize Candidates are Atherigona spp root beetles seed bugs ants and termites aphids root weevils crickets leaf beetles thrips and seed The least dispersive species are white grabs mealybugSbugs Insects known to entervarious states of aestivation are seedling maggots crickets weevils and gall midgesleptocorisaspp adults (199) Maliarphascparatellalarvae Each species has evolved a unique set of attributes to (9) Scirpophaga innotata larvae (117) Chilo partelhtsv enable it to survive and adapt to the changing uplandlarvae (21) ScirpopThagainccetulas (179) white grub larvae environments Several evolutionary avenues lead to fitness (134) Patangasuccincia adults (6 82) and Schistocerca (Table 2) but there is no reason to believe one set of gregariaeggs (82) attributes is better than another (224)

Insects known to enter hibernation are LodeIphaDstriatellus nymphs (124) Chtio Tvlpressalis larivae (125) DroughtSesaia inferens larvae (146) but not S calavisis(2) Well-drained rainfed upland soils are subject to drought ifIDiaraea saccharalis larvae (75) Scirpophaga- tcerlttas rains do not fall within 2-3 wk Prolonged drought followedlarae(5 lacchrli s lae (75)saSlOhagasshoperseggs (82)and felo pidas athias pOpae (232) by favorable rains stimulates armyworm and locust outshybreaks but mealybtigs root aphids and thrips become

Lcpmotorisa oratorisand L acuta adults aggregate on numerous onlyafter dry spells for two reasons First heavy trees sugarcane or other shady moist sites L paluwanmsis rain normally kills soft-bodied foliage-feeding thrips and confined to Sulu and Palawan Philippines lives on the mealybugs and soil-inhabiting root aphids and root mealyshygrass Brachiariamutica (Forsk) Stapf in the absence of bugs Drought eliminates this source of mortality Second rice The adults can live for 5 mo When aestivating their the rice plant responds to drought by breaking down rate of metabolism drops and they do not feed They will proteins into soluble N compounds which enter the phloem however puncture plant tissues of their aestivation host in and are taken up by these sap-feeding insects (245) Greater search of water This behavior has led to errors of the host nutrition therefore leads to greater survivorship and range of rice bugs (I) Aestivation quickly terminates with fecundity rain probably meaning aestivation is not deep Small rice area

Some stem borers aestivate as mature larvae in rice Except in the highly mechanized large landholdings of stubble or in straw stacked as livestock feed However Latin America upland ricefieldstypically are patchy within moriality increases proportionally with the duration of the a highly diverse flora The small fields result in higher ratios dry season even if the stubble remains undisturbed If the of perimeter to area than with the typically larger lowland stubble is plowed for a crop following rice few stem borers ricefields Upland ricefields themselves may be intercroppedsurvive Rainfall terminates dormancy but it normally with cereals legumes or root crops Mixed intercroppingtakes several weeks for moths to develop and emerge with a wide variety of species also occ in tribal slash-andshy

Last-instar white grub larvae tunnel 1-2 m deep in the soil burn agriculture (50 74) to construct pupal chambers in which they aestivate The Upland ricefields tend to be small because of the labor larvae are very sensitive to dryness so pupal chambers are needed to clear land in slash-and-burn areas and to till land sealed to coierve body moisture Aestivation terminates to minimize weeds Rice requires more tillage than does with the first heavy rains but as in stem borer 3-4 wk pass maize because rice competes less well with weeds Uplandbefore adults develop rice is more a subsistence than a cash crop and small areas

Insects may respond differently in each region to cold (I ha) can feed a family temperature or drought Biotypes or loal populations may Small fields favor some insect pests such as some ant evolve (56) Local populations of Scirpophagaincertulasor species that can forage from more permanent field borders Patangasuccincta may have become adapted to prevailing Trees provide food and mating sites for white grubs (134) as conditions well as aestivation sites for LetTocorisa spp (199) Field

iagiliti The ability to disperse combined with high border grasses plus tIhose within a ricefield provide alternate fecundity orshort life cycles enable some upland rice pests to food for all but the highly monophagous upland rice better exploit the temporarily favorable upland habitat colonizers and a habitat for natural enemies

0 z

Table 2 Life history patterns of upland rice insects or insect groups

Host range Longevity Dormancy Vagility FertilityMonophagy Oligophagy Polyphagy Low Medium High Yes No Low Dispersive Migratory Low Medium High(Oryza spp) (grasses sedges) (angiosperms) (lt3 wk) (gt2 mo) (lt1 kin) (lt50 eggs) (gt500 eggs)

Ants xTermites x x x

Mole crickets x x x xx x xField crickets x xx

White grubs x x x xx xRoot aphids x x xxMealybugs x

x x x xxSeedling maggots x

x x x x xLeaf beetles

x xx xArmyworms x x xx x

Thrips x x xx xChilo spp x x xx xMaliarpha x x x x

xLocusts x x Xx x xGall midges x x xxiVephotettL iirescens x x xx x x xLeprocorisaspp x

x xLeaftolders x x x

Sogatellafurcifera x

xx x x x x x x

00

IRPS No 123 January 1Q87 19

All this appeals to favor insect pest buildup but the notwithstanding the release of stored nutrients in response contrary is true Loevinsohn (138) showed that insect pest to drought that temporarily benefits philoem feeders populations respond exponentially with the proportion of land devoted to rice up to about 75 of the area then the M POR ANCE I I P1A 1) RWI P1St S

rate of response declines In most upland rico areas tie rice crop occupies less than 50i of the area- therefore the Visits to upland ricefields normally reveal 1kw insect pests potential for population buildup is low This relationship Loevinsohn (138) and Locvinsohn ct al (139) compared may not hold for seed pests that can locate small ricefields insect abundance using annual light trap atchcs in upland

Another outcome of the highly diverse flora and phical rainfed lowland and irrigated lowland sites in the Philipshyenvironment of upland rice isthateach upland ricearca has pines Although only one upland site was studied data a unique composition of insect pests showed equal or fewer insects at the upland site than at the

For example in Tanauan Batangas Philippines the rainfcd lowland sites and fewer than at the irrigated sites main upland rice pests are rice leaffolder(RLF) armvworm (Table 3) M Lltiple regression analysis showed that the key Aythntnaseparata WBPH ants Soli psis gentitataand factors explaining insect abundance among a set of cropping white grub Leucopholis irrorata Sixty kilometers away in intensity variables were the number of rice crops grown per Real Quezon the main pests are seedling maggot Athe- year followed by area devoted to rice Table 3 shows that rigona orae flea beetle (Thacitcmnra hasalis thrips the upland rice site had only I rice crop perym rand that rice Stenchaetothrips hiurnmis and rice bug Leptocorisa was planted in only 20Vi of the area Fertiliter and oratorius In a third Philippine upland rice site in Claveria insecticide use had little bea ring on the trend The armters at Misamis Oriental Mindanao the main pests are ants the upland site applied 60 kg N ha higher than at most Solenopsis gemtinata seedling maggot Alieriona orvzae irrigated sites white grubs Holotrichia in)iUaiuoan t and Lucopholiv Insect occurrence at the upland site was omparable to irrorata stem borers Sesamia ifiArelr (hio auriciliusC that at the Cagayan rainfed lowland site parlicularly in 1982stpessallis Scirpophaga incertulas S innoiata and when drought prevented many farmers fron planting(only

4c ona chrrsurapella root aphids Tetraneura niriah- 40(of the rice area eventually was planted that year) M ore dunrinalisand Rhupaosiphunm riahbdminalis lcaffolders favorable weather occurred in 1980 and 1981 which Marasniapataltis and Chaphalocrocis Medinalis army- supported greater rice areas and consequently more insects worm MIthinna separata and rice bug Leltucorisa The most intensive site for insect pests was South Cotabato oratoriusThe great diversity of insect problems is typical Mindanao where farmers plant 23-25 rice crops per year fer upland rice and makes control efforts more difficult with irrigation The Zaragoa Nutva Ecija site also had particularly regarding breeding for insect resistance high pest abundance and at this site the fields were highly

asynchronous Asynchrony was the third most important Low yield potential Iactor in explaining rice insect abundance Most upland rice is grown on low-nutrient and mineral- Rice cropping intensity in upland areas is bound to toxic soils which when combined with eiratic rainfall remain low Only one rainid upland crop is possible per make upland rice a highly riskycrop for the farmerto invest year (Farmers in Claveria Misamis Oriental Mindanao in costly soil amendments and land management Often Philippines tried asecond upland ricecrop i 1984a ycarof upland rice isgrown far from markets making inputs even favorable rains but no harvest was possible b-cause of more expensive Farmers growing a subsistence crop hosi- drought blast and birds) Rice area isaso bound to remain tate to invest in costly inputs to raise yields low because of the high labor and power rcquirelmenlts to

Even having varieties that could double existing yield prepare the land (because of weeds)and because upland rice potential under present management levels would probably wvill not compete with lowland rice as a cash crop Some not prompt many farmers to use purchased insecticide upland rice sites with prolonged rainy seasons howeverare unless tile risk of crop failure were reduced such as it was planted asynchronously such as the Clacra site with irrigation systems in lowland rice The seasonal abundance of Upland rice insect pests

The Tanauan Batangas site is atypical for upland rice attracted toa light trap wasgrapied for crop year 1980-81 in The soils and rainfall pattern are favorable and because of Tanauan FBatangas (Fig I 2) W13P11 was he most its nearness to Manila farmers have cash resources ftm common rice hopper and its population peaked at the sales of vegetables to purchase tertilier They applied an midgrowth stage as it does in lowland rice D-Vac suction average of 60 kg Nha after particle initiation and obtained samples showed an earlier peak than that from a light trap 25-35 tha yields The high fertility isone reason why RLF indicating emigration W13PI usually emigrate after the armyworms and WBPH developed into large numbers vegetative stage Populations in 1980 reached levels that Farmers however used no insecticide on upland rice caused patches of slight yellowing in some fields WBPH although they did on vegetables probably cause hopperburn in Batangas They immigrate

The poor growing conditions fOr rice generally mean year-round as shown by low levels in the light trap BPH poor growth and low fecundity for insect pests fIeding on it however were scarce in the light trap and not collected with

Table 3 Abundance of rice insect pests and factors in rice cropping intensity of Philippine upland rainfed lowland and irrigated lowland locations where light traps were operated daily for atleast I yr -I

0Rainfed Rainfed lowland Irrigated lowlandupland

Pangasinan Cagayan lloilo Santa Maria Nueva Ecija South Cotabato Tanauan Caaringayan Caaringayan Bangag Bangag Bangag Buray

LagunaCale

Batangas Manaoag 1980-81 Santa Motiica 1982-83 Cabanatuan Zaragoza Avancena NaniamanSolana 1981-82 1982-83 Oton

1980-81 1979-80 Oton 1981-82 1981-82 Koronadal Koronadal1980-81 1979-80 1979-80 1983 1983

Insect (kerosene light trapNephotcttixcatch nolyr)virescens 650 1240 500 1800 330 210 3400 1600 160N nigropictus 210 - 360 18400 5500 2900- 140 90 20 - shy 1700 -Reciliadorsalis 710 - 3200 1700180 890 630 140 170 2400 800Nilaparratalugens 130 690 - - 4200 4600300 520 750 180 60 3300 1600 740 310Sogatellafiircifera 1700 31000 13000 8200310 210 3300 610 270 980 1600 6300 -Cyrrorhinus - 12000 6700600 540 3200 290 160 330 120 410Scirpophaga spp 50 410 620 - - 21000 12000150 180 220 270 30 1000Cnaphalocrocismedinalis 70 70 540 610 270 11000 6100200 140 100 6 60 250Chilo suppressalis 0 60 60 1900 300 200- - 30 20 0 6 3 160 -C auricilius - 0 00 - - 20 3 60 20 3 0Sesania inferens 0 - 80 60 3 80 10 51 -Cropping intensity _ -

Rice crops (noyr) 10 10 12 10 10 10 13Rice area (7c) 17 19 20 20 25 2320 85 85 60 70 40 85 85 90Fertilizer (kg Nha) 60 20 85 80 70 7020 0 0 0 30 30Insecticide (no of 40 60 40 300 04 04 0 03 300 13 13 30 50 40 33 33applicationscrop)

__________

IRPS No 123 January 1987 21

Light trapped Insects Drytnd __ 2(notrap-nght) Suctioned Insects (nom )

50 -)o 9 jO40- SOW ffurciWO Light trp

D-Vc suction

20 to 4

20

Egg porasitizahion (M) 75 60

300 SOOampReOftwciffr

45 30shy

15

-insects (notrop-night) 4 Nileporvata Iuen5

Egg parastization()

Light trapped insects (no trop-nght) Suchoned insects (no m 2 )

6 8 Cytorh~nuls hvidipennii 0

4 05 2

50 0 Insects (noltrop-night)40-ui Ipanh pe g

Moohase 1 Seasonal abundance measured by light trap

Rainfall(etod) and D-Vac suction sampler of whitebacked

to ~ hopper Nilaparvata lugens and lea ffolder60)-t Cnaphalocrocis mnedinalis andplthpe g20 L I II I I I parasites and mtrid predator Ortorhjntus livi-Apr May Jun Jul Aug Sep Oct Nov Dec JaIn Feb Mar dipennis set against cropping pattern daily1 9 80o 19 8 1rainfall and moon phase Tanauan Batangas

1980-81

the D- Vac Cyrtorhinus build up on the crop coterminous are collected in light traps during full moon (109)with WBPH buildup Cyrtorhinus disperses during the Because upland rice shares most of the lowland rice pestsrainy season RLF not readily collected in light traps was and those pest species are generally less impotant to uplandprevalent only during the rice crop rice than lowland rice there is a general belief that insect

Nephotettix virescens was more abundant than Nnigro- pests are less important onpicLus in the light trap collections and on the crop GLH upland than lowland rice Arcview of limited data on upland rice shows an averag of population peaked at the reproductive stage as did zigzag 10-21 yield loss from insect pests on a par with that ofleafhopper (ZLH) Crtorhinus tracked the populations of lowland uee determined by the same methodology (132) planthoppers and leafhoppers Both GLH species were (Table 4) McGuire and Crandall (144) made a similarcollected year-round in the light trap indicating immigra- estimate for Central America Cramer (51) howevertion from nearby irrigated rice areas Possibly because of the estimated 35 yieldlow populations loss in South America based on ainsect number did not correlate with similar pest complex - stem borers and seed bugs shymoon phase although reports indicate that more hoppers attacking irrigated rice in North America This estimate did

22 IRPS No 123 January 1987

Light trapped insects D~~nd (no trap-night )

St plht Suctined insects (nom 2

)

D-Voc suction 1520

1o

0 Lih trap 5 ~105 o5

00 Eggparositizotion()5040 NephOtgtha virneens

30 20

I010 A-Insects (notrop -night)

4 Nephatelh nigropccent i

2 o I vV^J - -- ^ VA____ f _

Light trapped insects (no trap-night) Suctioned insects (no rnE

10 -1208 ReCdO doRsol

6 15

4 10 052 iwxALAiamp 0

Light trapped insects (notrap-night) 2Suc oned nsects (nom )

8 CyrtorhmnusI6o nn]10 i0075

4 05

2 0 0

025

2 Seasonal abundance measured by light trap Roinfall (mmd) Moon phase and D-Vac suction sampler of green leafhoppersNephotettix virescens and N ngropicrus and I zigzag leafhopper Recilia dorsalis their egg 60

parasites and micid predator Cyrtorhinus livi-dipennis set against cropping pattern dailyrainfall and moon phase Tanauan Batangas1980-81

40

5 Lf Apr May Jun

1980 Jul Aug Sp Oct Nov I

-aMar

1981

not account for soil pests and was not based on field data in the absence of pests Carbofuran granules are phytotonicAkinsola (I1) cited mainly data from irrigated rice areas in to rice (239) The prevailing hypothesis is that carbofuranAfrica but estimated yield losses of upland rice between 15 prevents soil bacteria from consuming fertilizer The phytoshyand 30 from insects However losses in Thailand over a tonic effect is through greater availability of fertilizer to the4-yr period ranged from I to 13 averaging 5(106) plant Denitrifying bacteria are known to be more active inYield loss data however should be interpreted with aerobic than anaerobic soils therefore it may not be merecaution (184) The Philippine data were all derived from coincidence that the highest recorded yield losses (22-69)trials on farmers fields under farmers normal agronomic occurred in those trials with 12-30 kg basal Nha On themanagement practices except for Bukidnon and Capiz other hand contro with carbofuran granules may not be aswhere researchers used fertilizer Batangas farmers are good as a seed treatment against seed pests and perhapsatypical because ofhigh levels of fertilizer used The trials in yields would have been higher if a systemic seed treatmentBrazil were carried out at an experiment station but insecticide had been used conditions were similar to those of local farmers The 1979 and 1980 trials in Batangas had two full-

Carbofuran granules were used in every trial as a broad- protection treatments one using carbofuran granules andspectrum insecticide applied basally with the seed An the other using bendiocarb WP as a seed treatment Plotsobjection to using insecticides to measure yield losses isthat protected with bendiocarb yielded higher than those proshyinsecticidescandirectlyorindirectlystimulateplantgrowth tected with carbofuran granules The other trials in the

Table 4 Yield losses to insect pests and yield responses to insecticide in upland rice in the Philippines Brazil and Upper Volta

Yield (tha) Yield loss (5) Location Year Cultivar Protected

with Protected

without Check Total Vegetative Reproductive Ripening

Basal N (kgha) Reference

carbofuran carbofuran Protected No with carbofuran

carbofuran

Philippines Pili Camarines Sur Tanauan Batangas

Pangantucan Bukidnon

Dumarao t-apiz

BrazilGoias Minas Ge~ais

Upper Volta6

1975 1974 1974 1975 1975 1976 1977 1978

1979

1980

1979 1980 1979

1977 1978 1979

Bursiging puti Kinanda Dagge Kinanda Dagge Dagge Dagge Dagge UPL Ri5 Dagge UPL Ri5 Dagge UPL RiS UPL Ri5 UPL Ri5 UPL RiS

IAC47 IAC47 IAC47

14 33 11 36 35 31 30 27 37 26 40 30 40 42 46 35

19 21 09

--

-34 24 24 27 --

29 43 31 42 -

47 25

---

10 28 05 27 27 30 26 29 33 28 33 29 43 23 36 11

15 15 06

29 15 55 25 23

3 13 -7 11

-7 18

3 -7 45 22 69

24 28 35

--

21 -11 -20

4 -

-3

23 7

-2 -23 56

-

-

-__0

6 --

-

-23 14

5 0 8 2

22

-

--

0

-

--18 11

3 -10

4 10

7

--7 -

14 -

-1 6

10 -5

8 9 7

-

--

0 0 0 0 0 0 0 0 0 0 0

30 30 30

12 12 12

99 99

100 100 101 101 102 102 102 102 103 103 104 104 104

66 66 66

Farako-Ba

Mean

1977 1978

IRAT10 IRATIO

38 66

--

40 43

-5 35

21

-

10

--

--

--

- 242 243

Z

tJ

-w

24 IRPS No 123 January 1987

Philippines which had two treatments offering protection In the 9 Philippine trials where yield losses were measuredwith and without carbofuran granules relied on further by growth stage 10 of the yield loss (21) occurred at thecontrol with foliar spray and not on seed dressings The vegetative stage and 517 occurred at each of the reproshycarbofuran-protected plots had an 18yield los compared ductive and ripening stages Ibis result is similar to thewith only 10 with foliar sprayS This yield difference is pattern of yield loss in lowland rice measured by successiveprobably due to differences in protection against sown-seed treatments where the crop is unprotected at each growthpests (foliar spray do not control sown-seed pests) rather stage but proticted during the other growth stagesthan phytotonic effects became basal fertilizer wa used in However the insect complexes r sponsible for vegetativeonly 2 of the 10 comparisons stage yield losses in upland rice are entirely diftfrent than in

The recorded yield losses were not related to yield lowland rice The key vegetative stage pests in lowland ricepotential (Fig 3) One might expect higheryielding crops to arc aquatic whereas those in upland rice are sown--seedhave higher percentage of yield losses but this ws not tile (sown in soil) root and seedling pests This dift rence maycase One reason is that insect damage exacerbates plant explain why agricul torists have overlooked tile ilmportanccinjury from drought stress low yielding fields from d ought of upland rice insect pests Many reports of upland rice pestsstress would show disproportionate differences between in Asia ha Ve focused on major lowland species (46 108)insect-protected and -unprotected treatments Shoot and It isalso surprising that tei vegctative stage of upland rice root pests would be more responsible for losses from would record the highest yield losses becatuseseeral studiesdrought stress than sown-scd pests A second reason have shown that upland rice can readily recover from foliageconcerns seed pests which would have a greater impact on reinoval during that stage (117 152 165 188) Upland ricelow tillering low yielding varieties such as )aggc Low poscsses a hitgh obilitV to compensate for ca rly loss oftillering varieties nnot fill in the space created by removal foliage and will even be stinulated by foliage removal toof seeds oweer 1) R i 5 is high yielding because it tillersP1 produce a higher viel than plants ithout any foliageactively which can close the canopy to provide greater weed removed (63 192 231) [ie reason for this apparentcontrol discrepancy hovce is that yield loss at the vegetative

stage does not octUr frol leat arca loss but fron pestsYield loss () removing sown seed and feeding on devcloping tillers and roots -- damage that cannot be readily conpLtnsatCd for

70 - Measuring yield losses in upland rice cnvironrnents is further complicated by a report from Brazil that greater yield losses occurred in response to better insect control60 - =-09 as Insecticide seed treatments resuled in denser plant stands

(n=21) which in turn created a selectively more favorable environshyment for blast (69) This resuIt was parlticurla rlxexacerbated50 when cirbofuran granules were used in seed furrows Aside

ffrom a greater plant stand ca rbofuran iay have auigmented the N lecl and stimulated blast If so ca rbofuran should be40 - replaced with another insecticide or the seeding or fertilizerI rates decreased

Other methods to determine the impact of upland rice30 shy pests wvcrc to interview farners or to develop single speciesbull bull correlations of population levels and yield loss Feneira (66)2 NIM)st (75(1) Golas farmers reporteddid both in Brazil

20- insect losses in ufpland rice and 251 said those lossesS avcragcd 37(1 [his result is consistent with tie yield losses 0 0 measured by field trials in the same statc(Table4) For every

I0 4 of seedlings removC d by leaf-cutting ants a I 7yield loss is predicted Similarly Ferreira (66) found that ai

0 - average of 53 thrips per panicle before panicle initiation meant 3 times more unfillh d grains

-IO0 I 1 1 I I-101 0 2 3 4 5 6 7

Yield (tho) C)N [ROt IM FI IODS 3 Correlation between upland rice yield and yield lols caused by Upland rice insect pests may be controlled by culturalinsects measured in 21 trials in the Philippines Ivory Coast andBrazil genetic biological and chemical methods

IRPS No 123 January 1987 25

Cultural control dry season plowing will have little effect on them Plowing Crop husbandry to reduce upland insect pest populations when the soil is wet from the first monsoon rains will expose can be classified into practices effective in a single field and soil insects that tunnel close to the soil surface to predatory those effective only at the community level Cultural birds chickens dogs and even man (field crickets nole controls should come first in a pest management program crickets and white grubs a re delicacies in man tra ditional They have broad stable effects as pests have little pos- diets) sibility of overcoming them through selection of biotyps The greatest degree i1 inIscct control fron tillage comes (138) On the other hand some cultu ral controls work only from plowing soon after rice ha rvest when most soil insects with high labor or power inputs and connunity-wilt lie close to the soil surface anrd stern borers are in the rice methods require coordination of iany farniters stuble Whitegrubsare mature and tillage exposesthem to

Pantin time The luxury of being able to shift the time of predators Tirnelv control will prtec the crop following pla ntring upland rice is open onl hr la rniers in areas with a rice and lowr the populationIfor succeedinig years Plowingprolonged rainifall pa ttern lI copping intensity or underthe ice stulble helps it decompose killing sten borer rnechanied land preparatiot Gcnerally planlting as soon larvae inside as the rFiinv season begins will lower populatiois of 1110st [requent tillage destroys ait nests bit not termites insect pests Insect populationsarelo aftcra dlvseoll or Tillage and clean culture howes er rcliove food for winter fallow delayed plaiting lets them build tip on termites which then mtay attack a young rice crop Scarab alIternate hosts that gi vigorotsly with the first seasonial beetles also prefer to miposit in eceitly tilled fields Noshyratiris This first flush iof weds cros lItxuttriartlv with iceCult is dibbling faors the survival oftillace Clre Such rineralied N released during the dr season aid made stils i aels tnd insects thtt pttpate or lay eggs- in the soil

available to the plants with the rains It iliso igrtmitcally Zero tilla hio eerl conserves predators ofthe GIH (37) advanttgeous for the rice ciop to tap this trlLItlaI fertility nt it mcttoi lo lessen the impact o0 soil insects and grow vigorotsly to compete with wrus and to tolerate brotdcasting seed or planting in ftltrrows is prefc ired over pest infestation dihlirig in hills Crickets are large enotlgh to eat seedlings in

Larly plantngs terd to escape seedling maggolts Upland a hill atrd tillering normnally ca iriot fill in tlhe missing space rice in Bitangas is planted in Ma and ine itli the Iirst Oftcn dibbll rd is not well covered and iseasily found by rains Fairners are highly mtotiated to plant early tocscape foraging insets ioIdents and birds drlitght and typlotns ricar harct Arn Atrgtst planting Plan uit sii Incrcatsini sced density protects tile cropwould be evcely attacked by It riim orivzo At tti againrst seed and seelling pests especially ants particularly other Philippiie tipland sites Rea l Qteot antd (laelit if the crop will germinate and ermerge quickly M isainis Oriental seedlig mnaggots arnrige ric boe t se hr trftinll itI India rice intcrclopped witi cotton or farnes have a longer growing season and plnit 2 rin or pigeon pci had lower 6IIt and WBIlII poptrlations than mote after the ortset of the rainy season liaaiangas ftrmners rice alone (211) NfailC arid Urplald ir_ 2re coilltori in the plant ca rly becia use ofa shorter raiinfall period arnd the desi re Philippines he chatngC in pest status folii iniercropping is to lliarvet before the typhoor sea son peaks ltighly locatioll-specific (133) cmianarid the net elfc be either

Early plantings cornbind with early-iatiuring varictics nil bcnelficial ortdetriential (189) For example BIatangias provide higher crp tolerartce for white grubs Aestiatior farmers intercrip Itro in their ipland ricefields A liornshyterminates with the first rains but 5-6 moare ineeded for the nrii iotd-Ahis i (I iiiacius) which norlially feeds resting larvae to develop into beetles emerge and lay eggs on taro ill feed or miarbv rice plants Intercropping and for larvae to reach the danriigii g ird-instar stage probably would not afft0 seed pests A crluanion crop (134 In lataigas this normually iseiough tiite for harvest planted ith rice and haing gleater tolerarnce for root

It Brazil howecver delayed planting is a suggested dariage or being toxic nayact isi trapcrop to root-feeditig cultural control for the lesser cornstalk borer and insect pests Acial-fei i ig pests voultId be most affected by pests favored l drought (66 195) Cropping internsity in intrer ropping Shoot aphids leaf beetles sten borers and Brail is low and delayed plraing places the rice crop in the hopplers would be likely ca ntidates to note effcts On the niost stable rainfall period reducing likelihood old rought othlt halnd the corlipalior crop stch~ sitniaie Iiiv be more

In rilniy arca s rinmirsare roi atl to plant other crops benehited than rice from initecropping ( 137) before rice and althNgh it woultl be desirabh to avoid 11echi Weeuling luring the first month after clop infestations by planting rice cuatl lariners lack the labor cstablishent will force pcts such is anyvworris which and time to do so ol a low priority crp prefer grassy cetds ontor tle vorling rice Soil iampsecs will

77age Uplild rice Stils ae light textured and Call be mvnye to the rice crop Cleani citrltirc also lay lore territes plowed duiring the d rv seasor notl only to grin time for to attack I u cetincti inrice ceds i ay he le optinal planting with lie fitst Inilsoon riiains but also to desiccate solition 13) soil insects arid weeds such is ntsedge that foim rhizomes Irtlizr Itneatsing N Iciiliier oin hlwland rice has Htowever becta use most soil insects lie below die plow layer fAtwvored Irigher populains of pliithoppeiv Imlhoppers

26 IRPS No 123 January 1987

and leaffolde rs However because of the need to temper fertilizer use on upland rice o prevent blast fIrtilizer rates will be lower than is comonia r for lowlad rice causing relatixely less hopper and IaIfTfoide r population buildup However inupland rice areas without a history of blast such Is Batangas farmers apply high rates of N and conscqtrently the crop has high infcstation levels of WBPI]and RIF The fhrmrcrs leec insect buildup by splitting the fertili11r in0to two to three applicatiois bit do not applV fertilizer at the vegetati estage because this prodrces a tall crop which illeasily lodge during typhoons Greater crop fertility leads to greater insect survival (better nutrition) larger insects (to cat no re foliage) arid greater fec titd t (to lav nmre eggs) (128 145 171) Not all insect species respondequally hovr Increasing rates (if N l anrid K increased Dialraa s-acchralis and (iaeiurema flea beetle but reduced thrips (66) Zn rcdLuced deadheart density from the lesser cornIstalk borer

Fnldingi Although most Upland rice areas do nIot hawxcaccess to wa ter floodiig the fields is rRrrnriierdcd in areas of Brazil or root bug and white gib control ( 195)

Crop rotatioti Tillage before planting a crop after rc harvest will control stern boreis and other pests renain n the stubble arid will unearth soil pests Planting a non-gratriiaceous crop after rice is recommncrded for termite control (66) Crop rotation is less nttdcd in upland rice culture than in lowlands because atcrop-free period is assured even after harvest of crops following rice

t1ulchilt iryl plastic irli irig designed for weedcoitrol ii upland rice inJapan resulted in ntr highanri liy infestation of (ihilo sulytresvais (93 9-1 95) NIulched Upland rice grew more luxuriantlv and attracted more (Avipositing mothis than u lched rice Also the rice plant and stein borers imatoured 10rC quickly and a partialgeneratiori af vouing ista iNoverwintcred in the stubbleLotili--malitrt varieti Quickly maturing crops reduce the tmbCr of pCst generations that Calt build up Rapid crop establishment olan ca rl v-natu ring vaitvty if)pa 11icLa r will lessen white gib and stem borer damiage White grub larvae intle last iistar are highly destIruCtive so aii early-iiatri ng variety will escape serious root loss Stem borers build 1ip slowlyx in rice to ecormic abundant in a late-Maturing crop anrd caiuse whi tlieads Numibers of somespecies increase exponeitially with each succeeding genera-ti so a xarietyexhat atures I trowh earier thian another will hac less insect damage Early-naturing vaiicties however will not lessen seed pest daiage either ii the soil or oina standing crop

Strwhoi hAIJttam Inrscts tcd irig on thlie aerial portions of rice plats disperse from carly to late plantedfields AS iiiibCirs increase exponentially with eachinsect

gtscnrattm e eiglbrig f rmc~s1(uil upp wh rc generation pests build tIP where neighboring fihrTC e9r1sstagger their plantings II Batangas farmers plant uplandrice within I iro use 120-d s-arieties and plant at the start of the rainy season Ini Misarnis Oriental laricrs stagger their

plantings up to 3 mo apart and use a set of atieties that mature in 4-7 mo Insect pest danage particularly from stem borers is greater in Misarmis Oriental Also seedling maggots damage rice there because rainfall favors frequent planting of maic In Batangas rice escapes seedling maggots bCca usC of distinct drs season and synchron0us early planting with the onset of ains

Therefore combinations of cultural practices ea rlyplanting synchronous planing crop rotation and earlyshymaturing vaneties protect the rice crop against most insect pests

Ilmant resistance Beca tse upland rice has few specialized insect pests and many insect types attack the crop worldwide regional breeding and strong internatiornal cooperation are required

Sown-sced pests arc economically controlled by insectishycide seed treatment and should the relore receive lowxpriori ty inbreeding objectives

Root pests are normally cost ly to control with insecticides but finding resista t sonres mayvbe di fficult Tanaka (229) for exa -ple fi led to find varieties resistant to root aphids Selecting for large root bioniass perhaps should be the strategy for pests sLuch as Whlite grubs that renliove roots Fainter(174) howcvcr cites varietiesof crops tolerant ofor resistant t(i root pests Some sorghun varieties are high in cyanide so rice sorghun might incorporate broadshyspcctruri chemical resistance

Seedling vigor and drought tolerance are high prioritybreeding objectives so tupland rices should have large r root systernsand be able to tolerate ligherlevels of root loss Sonie root loss inolder plants is beneficial if it stimulates new root developnrut and therefore crihances intake of soil nutrients (53) The rice seedling inaggot rtiLotia spp may be

controlled thirough resistant aittics A hi rgc breeding effort on sorghniu has had fair success inideveloping arreties esrstait to Asoccata Rodani (251) A field trial comparing 10 upland ricc varieties showed differences aniong varieties va ring from II to 4(- damaged tilleis (215) Hovver resistarce to 1Iill-0r gtia species may rot cross ox-rti otier

Shiraki (210) reported that upland i-ice stens wcrc harder than lows-land rice stemsperhaps becaIusc (f a higher silicoricontent and thus were more resistant to yellow stern borer hc tortia Iow tillering (f uplaid iice xarneties prodtres a

higher percentage of infested tillers than would occor in higher tillering lowland iice under a siiilar egg density The corollary therefore is to select for high tilleriig upland rices

Stemii bor amcstheh main targes dobrecders inAfrica (571 204 )but nticl oifth e sc reen irig is dotie unde r lo wland co d t osrm s wih A h r~ o l pe i s n acorditions (11 96 218) As with lthrinrnwspecies mayhave to be dealt with independently however initial results show cross resistance between African and Asian species(92)

In Brazil screening Ior insect resistance focttses oil lou I pest problems Varieties tolerant of lasmolwalpus ligi-sellus and resistant to )iatraea saccharalis have been identified (70 153) Current work also emphasies 7iraca limativetntris and Thmialspis (=Iosis) flavouicta (66) These insect pests Ilt onlV citise IIigh yield losses tihey also ire difficult to control chemically ofr by other ilealls

ManV upland rice vaieties aic early maturing and are resistant to pests such as gall midge (I Xl ) Early mIturity is often a highly desirable trait to esuilpe pest buildup but it is not true resista nce bca usc ilthose va reties are plinted late they will be itsdamaged as ain early planted susc ible variety

P~erhaps tile cosiiiopolitall of all upland rice pestsimost are the sced pests (ripenling stagc) which pose a particular problem beta use scuds re ised IsIO for hitnit4ans Ayltoxic substatice introd tcld geticticllyilmay also be toxic to nal iniless it caln he placed old in thfe IdhuffSelectmi fo hairiness or lotng wns kill mIlct sthilrohleiiis dutilig threshing pa rtictfarl maitial threshinfl [hireshers ill complain of itcfimin atid itteascd (fust levels

Attempts itIRRI to fitid tcistatncc to the rice bug plorisa have ilotbeCnl prmitisilg lIIetIattonlids iorc

through the lenilna or palca hillhairs sedstVIor seeds wilh awns that deter bugs promote complaints from threshers Seed bugs liptocorisa and Stcnocoris enter the seed through the opening between tile lemna and palC but varieties with narrow openings are diifficult to mill

Breeding priorities should concentrate ott inotCI gruLtLIps that are most difficult to c nttrol by other meains Steil) horers appear to be the most widespread group that sigiiificantly dalag s rice aind is hard to manage by other Ilnca us

Biological control Very fe stutdic of mitatt~l eneties of rice pests in upland envirollnletits wee in tile wefouiid literature Therefore have not compiled a list of species Ve present a list of natural enemies iOtid in upland ricefields tuic latialln Batangas for some conitno iOldiar pest species ([itble 5)

Muich ofl thc lollowinle itfor ittiol cotlliCs rol Stutdies in the lowlatnds or roi crops otithcr thlin tice

Qu(ailtr o 11ll t11t( Nat trrtf lii hcoeits ight Ile used agaitist ile social insects tilt and termites Sollie predator specice live ill theirf nests n ilnic theirnllt ip-pearanIlCe aMRidbehavior IJtrouMdCing a mote aggressise ant species to displace pest ant codonies is a possibility (169) Patihogens as potential natural encnies have reccived little atteition (I13) Natural ncnuics asa group however hac not been toIl iporta it in Iegulating aintshown be and ternite numbers

The nocturnal behavior aid subteitmien habitat of lole crickets and field crickets protect them from many ntllral enemies There are only limited records of natural cnelics for these groups of pests some pathogens (87) but mainly

IRPS No 123 JattuIny I 271987

parasites - scelionids on eggsand sphecids on ilymphs and adults Sphecid wasps Larra carutmaria (SImitI hI L huzwuensis Rohwer and lsaltiuhta Willi ams speciali in mole crickets pa ralyzing them nd dragging the ninto thcir nests as food lot their voting ()tier sphCcid species Lins aurulenoa (Iabricius) lot maiac (Ashmead ) 1 lhshylssellaus (Smith) and Ml lal)oriosu (Snithf specialie in field clickets

White grUbsO the other hand hae Icti scriuns enuigh to have been extensively studied for cotttroi 1W nati al enemies( 112 140 141) Scoliid parasites C(ajsoMcispp have beetn intrtduced itto tilePhilippites for control of Ielxl ]udisiro la The rcsearchcts w Intttrdced themle during ill era claimed that the white gttionthreak as controlled (142) howevcr it isnormal for pest epidemics to subside and there was no diect evidence to indictte that tfe parasite was responsible A tmore promisitg titctllod might he to identify iso ltcttass prt~f tcartd apply hiactelaln fingal pathogens to tile soil Ihe lalattese beetle in North America has hcci coitrolled hv tite conmnleciall aailable hactcrial prcpa ration itacihlpu pav ofr milksdiscas (71) Spote prtyefott the soil itloted ilrtier reitiain actic fr Veas

Root aphid sardtnctly bugs readilslic tilib to predatots and parlsites once their tetndine ants are controlled The larvac and adtilts of a coccinellid beetle 1Scynms sp prey on llriadlouais in slash and burn upland rice Baits treated with insecticide may be used to control tending ants by killing their ourig (244)

Root-tkcditig blugs false wireoris wireorms and root wccils aic r ofsuch little ifilpoltatice that thenomttll role of liatural Cticniics hias tiot beei asscssed A possibility hoevrl tiglt he tit ItaIIitic niaCIMtodcs lintt1g11en Itese soil-itlhlfbitine mCtitodcs W5uhtfhave to be imiass Produiced

large miltibcr of niatltrll cnemics ircknown ft CltIoriS IC IpblCeii is thtt clttvortnlls ofonlie the crop soott alter land ptel)tritiOti a1tithe bcgit iniig of the raittv

Citsoln hfien tiatllral leniiiy populations aric lo Colltloil WOtuld iii ole mass prodtticing andirclcaslng key spccies

le egg stage is the fpirt (olthe Cedlitgt Ialot life cycle tI hich to f)ctis ftor tittlril elctit cottntrol hplraites o1 pleditors ttsit elleggs highly expoIsYCl I Sgg pailsites or predators stolld probahly e ittass andtbe produced released Littlh i toruialioll is a ailable ott what species to trV o ho to Iniss podLice thiCIi A etiloplhid wasp Icmihlaol sp effctiicly regushy

lates the leatf lcloalapolm2a spioanoiiitr heat andc rice antid should be collsctvcd

leaf beetles whCllose e am] eggs ate laid in the grutindhirO ould be motC difficutlt to control by niatulral enelV Mnll ipuhltiOn But species sUici IS c_gsthispid beetles ith

and larVle oilleascs ate vtilcratble to iparaites and pledators

Natural enemisc of tlirips otl rice are little known

28 IRPS No 123 January 1987

Table S Parasites recorded from upland rice insect pests in Tanauan Batangas Philippines 1977-84

Order Family Species Host stage

I lymttenoptera Braconidae Leaffolders Cnaphalocrocis Marasinia spy

Cardiochilesphilipplinensis Ashmead a Larva

Cotesia angustillasis(Gahan)Cotesia or cypris (Nixon)

Larva Larva

Cotesia or taeniaticornis(Nixon) Larva Cilonusinotakatae Munakata Chelonits SppAlacrocetrts philippinensis (Ashtmead)

Egg to larva Egg to larva Larva

Iclineumonidac Orgilts spp Isclutojppa hireator(Fabricius)

Larva Larva

Itoplectis naraigav(Ashmcad) Larva Tenelhtcha philippiensis (Ashmead) Larva Tenelchha stangli (Ashmead) a

irichomma cnaphalocrosis Uchida Larva Larva

Bethylidac Elasmidac

Goniozms or trianguli r Kieffer lasmtusalhopictus Crawford

Larva Larva

Diptera

Encyrtidac Chalcididac

Tachinidac

Elasmtus spp Copidosomopsisnacohiae (Fidy) Brach nteria evcarinata Gahan Biachyieria lasus (Walker) Ztgohothria ciliata (Wulp)

Larva Egg to larva Pupa Pupa Larva

Argyrophylx nigrotihialis(Blaranov) Larva

l-ymenoptera Braconidac Yellow stern borer Scirpopiaga incertulas

Braco chinensisSzeplegeti Larva CIthelotis spp Chelonus niunakatae Munakata Cotesi flaripes Cameron Stenoracon nicerilli (Bingham) a

Larva Egg to larva Larva Larva

Ichneumonidac Tropobracon schoenobii (Viereck) Amatroinorpiaaccepta metathoracica(Ashmiead)Frihorus sinicus (I lolmgren)

Larva Larva Larva

Ischnojoppaluteator (Fabricius) Larva Isotima nr dammeratiti Rohwer Larva Tenelucha philippinensis (Ashmead) a

Larva Tenehicha stangli (Ashmcad)Trichomma enaphalocrosis Uchida

Larva Lara

Chalcididac Eulophidac Trichogranmmatidae

Xanthopimnpla stemmator (Thunberg) Charops hrachYpterumn Gupta and Malteswary Brachtm eria sppTetrasticttus schoenohii Ferriere Trichorigranm chihunis Ishi

Larva to pupa Larva Pupa Egg Egg

Diptera

Scelionidac Pteromalidae Tachinidae

Tt ichograintnta aponiicuitt (Ashniead) Telenomits rowani (Gahan) a

Trichoinalopsis apattelctena (Crawford) Jeirhaea spp

Egg Fgg Egg pupa Larva

Z4gobothria ciliata (Wulp) Larva

Ilymenoptera lecneumonirlac Gol fringed borer (hilo auricilius

Trichomm enoptalocrosisUchida Larva

Ieioenoptera Braconidae

Aantholimlh steiniator (Thtinberg) a

Pink stet Ioret Sesamia inferens StoInacoil nicrilhi (Bingh am)

Loarva to pupa

larva

Diptera Ilyinenoptera

Armvtnyorms Mythimna separata Spodoptera mauritia acronyetoidesTachinidae Zvgohothriaatropiora( Robinea i -Desvoid y)Eulophidae It-phctrtiscitapadoe (Ashimead)

Larva Larva

(halcididae Braconidac

Brachpin ria spip (oesia spp

Larva larva

IHyntenoptera I)iptera

Rice skipper Ielop ida (halcididae Tachinidae

math ias aidgreen iortedci ciliarNIelan itis eda Brat lhincria sp iii matuqitatt Cameron Irgyrolhirlaxi trotlhiais(marumtv)a1

ismtne Larva Larva

Diptera Ilymenoptera

Tachnidae lelineutionidae

fuown se tilo per Mois frugalisIrgtrophrlax 1igrothiahiv 3 a rat ov) anthopimpa pmnctata (Iabricius)

Larva Larva to pupa

aMost dominant

IRPS No 123 January 1987 29

Normally predatory thrips prey on herbivorous thrips strepsiptera on both Spiders dominate tilenymphaladultNatural enemies of stem bugs --Scolinoplara and Tibraa predators Spittle bugs move to rice from pasture grasses - are in order of importance egg par~isites and predators and biocontrol would be successful only ifcarried out before fungal pathogens and nymphal adult parasites Egg tiley immigrate into ricefields parasites could either be introduced fron other areas or Biological control tactics for hemipterous seed bugs mass produced and released ould follow those outlined for stein bugs

Armyworms are normally lwld in hcck by the activities Coml)arison of environnewtns Natural enemies are of egg and larval parasites When these natural enemies fail perhaps faced with even greaterconstraints than insect pestsusually because of drought armyworns become epidemic to survive ind reproduce in upland environments TheyIt may be worthwhile therefore to release parasites during have adapted ani survived by having wide host ranges the the rainy months following a drought Virus diseases of ability to aestivate or hibernate over unfavorable seasonslarvae would beanotheravnue to exploreforarnyworms and the ability to disperse or by other mechanisms

Parasites have ben traditionallyconsidered for biolegical Comparing the most dominant and therefore the most control attempts against stein borers usually by rekasing adapted species of natural enemies collected in various exotic species( 166) This approach has not met with success environments in the Philippines we can understand how and new avenues should be explored The role of predators different rice environments are arid pathogens is little inderstood and deserves greater Anagqrms olptahiisa mymarid planthopper egg parasiteattention Egg parasitism rates are normally reasonably is cquallyadapted to all Philippine rice environments (Tablehigh but should be supplemented with effective egg 6) It is selective for planthoppers and even parasitizes eggspredators Orthopterans are potential egg predators of the n-iize planthopper Peregrinitsmaidis itt upland areas Metioche vittaticollis (Sta ) Anavipha nifiwnnis (Saus- Alavohis also parasitizes maize planthopper eggs but is sure) and Conocephahs hongipnni (de i-laan) feed on most prevalent on BPH in the irrigated wetlands (onashystem borer eggs etioche and Anaxil)ha specialize in eggs tocerus spp like A optalhilis on BPH and WBPI Iattacks with no hair such as Chilo spp The omnivorous (ono- the eggs of GLH in ell rie environments It is also found cephahsspecializes in eggs cov red with hairs it is a proven parasitizing the eggs ef the white rice licahopper (oimna egg predator of Scirlouphaga spp but has the discouraging spetra Oirosita naias a trichogrammatid egg parasitehabit of eating rice guin (65) attacks BIPH and WBPH in the wetlands and has no other

Grasshoppers have egg (scelionid) and nymphalad ult host A related species 0 aesopi specializes in G1-H in (nemestrinid tachinid and sarcophagid) v irasites Among wetland environments Dryinid strepsipt-an and pipunshypathogens protozoans have been recorded most frequently culid nymphal and adult parasites of planthoppers and Control of locust species by natural enemies could con- leafhoppers also specialize either in hopper species or centrate on their habitual breeding areas environments Among the dryinids Pseuhdgonatoipsflavi-

Leaffolders have rich complexes of natural enemies some feimur prefers the lowlands and BPH while P nuch has no adapted to upland rice (22 27) ranging from egg predators environmental prefirences but is adapted only to WBPH (gryllids coccinellids) to larval parasites (braconids ichncu- The main d ryinid species on GLH aptogonatop)usspp is monids) to larval prcdators (ants and carabids) to larval most dominant in rainfed lowlands and also attacks other pathogeiis (vira and fungal diseases) Efforts to augment leafhoppers within Amnrasca Cicadulinaand Bachaltuathese natural enemies should focus on the egg predators Strepsiptera are more dominant on plarithoppers than (rearing releasing)and viral or fungal pathogens (culturing lealhoppers The Elenchus species in the Philippines attack spraying) all species of planthoppers within the genera Sogawilla

Parasitization of the large conspicuous larvae of Sogatodels NilaparsataHarualia and Opiconsiva Elenshyskippers green horned caterpillars and other polyphagous cusrvasuntaisuiis adapted to the major rice environments Lepidoptera by tachinids and chalcids is normally low ELg but prefers BPH in rainfed arid WBPIH in wetland environshyparasites and predators are perhaps the key to their control ments Ilalictoph(utsmtnroeiand HVpectrus attack G LH

Gall midges are parasitized by pterormalids and platy- and COfimravcctra respectivelygasterids that aestivate within their larval host between Pipunculids are mainly parasitic oii lealhoppers Each seasons Shoot aphids are preyed upon by coccinellids species has a unique environmental preference Tomosshysyrphids and chrysopids varireIa oraetoraprefers Ne7 otetltix nigropictrs and

Planthoppers arid leafhoppers have egg parasites and Pilunwulhtsjavanensisattacks Dehoceplhahls spp which predators as well as nymphialad nlt parasitesand predators are more abundant in upland than irrigated lowland their rich complex of natural enemies includes fungal environments T suhvirescensattacks Nephotettix virescens pathogens The principal egg parasites are myrnarids and and prefers the lowlands Pilpuiulus nttillattls is an trichogramniatids Egg predators are norinally mirid plant upland rice parasite arid attacks only Nephotetix speciesbugs that suck out the yolk Nymphal ad nIt parasites are More than 17 recorded species of leaffolder parasites in dryinids on planthoppers pipunculids on lealhoppers and the Philippines attack species of C(naplwocrocis and

30 IRPS No 123 January 1987

Table 6 Dominant natural enemies of rice pests in upland rainfed lowland and irrigated lowland rice environments of the Philippines

Natural enemy

Host stage Species

Brown planthop-Parasite Egg

MymaridaeAnagrus optabilis(Perkins) Anagrusflaveolus Waterhouse

TrichogrammatidacOligosita naiasGirault

NymphAdult Dryinidac

Pseudogotatopuslaiifemttr Esaki et Hashimoto Elenchidae

Elenchus yasurnatsui Kifune and Hirashima

Environment

Upand Rainfed lowland Irrigated lowland

Ailaparata lugens

x x X x

x

x x

x X X Whitebacked planthoppers Sogatella furcifera and Sogatodes pusanusEgg

MymaridaeAnagrus optabilis(Perkins)

TrichogrammatidacOl~gositanaias Girault

NymphAdult Dryinidae

Pseudogonatopusnudus Perkins Elenchidae

Elenchus yasutatsui Kifune et Hirashima

Parasite Egg

MymaridaeGonatocerus spp

TrichogrammatidaeOligositaaesopi Girault

NymphAdult Pipunculidac

Tomosvaryella subviresccns Loew Tomosvaryella oryzaetora (Koizumi) Pipunclusmutillatus (de Meijere)

DryinidaeIlaplogonatopusspp

Egg Copidosomopsisnacoleiae (Eady)

Larva BraLonidae

Cardiochiles philipphnensisAshnead lchneumonidac

Trichiomnia cnaphalocrosisUchida Temelucha stangli (Ashmead)

Pupa Chalcididae

BrachymeriaexcarinataGahan Brachymerialasus (Walker) Brachymeria nr tachardiaegroup

Parasite Egg

Scelionidae Telenomus rowani (Gahan)

EulophidacTettasticusschoenobhi Ferriere

Larva Braconidae

Stenobraconnicevillei (Bingham) Iclhncurnonidac

Temelucha phiiippinensis(Ashinead)Temelucha stangli (Ashmead)

x x x

x x

X x X

x x Green leafhopper Nephotettix virescens

X x x

x x

X X X X

X

X Rice leaffolders CnaphalocrocisMarasmia

x X x

X

X X

x x x x x X x x x

Yellow stem borer Scirpophaga incertulas

X X x

x

x

x X XAinauromorphaaccepta metathoracica(Ashmead) x

continued on next page

IRPS No 123 January 1987 31

Table 6 cntinued

Natural enemy Environment

Host stage Species Upland Rainfed lowland Irrigated lowland

Larva-pupa Gold fringed borer Chilo attriciiuslchneumonidae

Xanthopimpla stemmator (Thunberg) x aPupa x

Pteromalidae Trichoinalopsis apanteloctena (Crawford) ax

Pink stem borerb Sesamnia inferensParasite

Larva Braconidae

Stenobracon nicevillei (Bingham) x b b Armyworms Mvthinna SpodopteraLarva

Tachinidac Zygobothriaarroptvora (Robineau-Desvoidy) x x

Braconidac Cotesia spp

X Skipper Pelopidas mathias and green horned caterpillar Melanitis leda ismeneF-gg

TrichogramrnatidacTrichogrannaspp x

Scelionidae Undetermined Scelionid

x Larva

Tachinidae Argyroph lax nitrotibialis (Baranov) X x

Pteromalidae Trichomnalopsis apanteloctena (Crawford)

X Predator

CoenagrionidaeAgriocnenis femina femina Brauer x xIschnura senegalensis tambur

Miridae CrtorhinsliidipennisReuter x x

Nabidac Stcnonabis tagalicus (Stal) x

VeliidaeAlicrovelia douglasi atrolineata (Bergroth) x

Coccinellidae Micraspis crocea (Mulsant) X x x

Vespidae Eumenes campaniformis (Fabricius) x

Formicidac Diaecamna spp xOdontoponera transversa (F Smith) x

Carabidae Chaenius spp OphioneaW- aJasciataSchmidt-Goebel

x

Ophionea ishii ishii Habu x x X

Gryllidav Anaxipta longipennis (Saussur-) x xMetioche ittaticollis (Sial) x x x

Tettigoniidae Conocephalus longipennis (de Haan) x x x

aNo rearing made on rainfed lowland bNo rearing made on rainfed lowland or Irrigated lowland

MrasmiaAmong the Laval parasites the braconid Cardio- lowlands and parasitizes oriental maize borer OsIriniachilesphilijpphensis is most adapted to an upland environ- furnacalis (Guenee) larvae The leaffolder pupal parasitesment It also parasitizes Hydelepta indicata a common not only occur equally in all environments but also haveleaffolder of legumes The ichneumonids Trichomma cna- wvide host ranges- parasitizing species of OsIrinia Marucaphalocrosis and Temehcha stangli occur more in the Hydelepta and Honiona - in maize and legumeslowlands T stangli is most adapted to irrigated lowlands The principal egg parasite of yeltow stem borer (YSB)and also parasitizes Chilo and Scirpophaga stem borer Telenomus rowani is widely adapted to all environmentslarvae T cnaphalocrosis is most prevalent in rainfed and has an unusual alternate host -Tabanus eggs (25)

32 IRPS No 123 January 1987

Tetrastictu schoenobii specializes only in YSB and occurs mainly in irrigated lowlands -- further proof that YSB is most adapted to the wetter environments

Larval parasites of YSB are highly environment-specific The braconid Stenohracon nicevillei occurs only in upland rice but also attacks the pink stem borer (PSB) Tieinchwa phiiiplPwnsis an ichneumonid occurs in upland rice and also parasitizes the larvae of (7hiu stqpresaisinatro-morpha accepta melaihoracica like 7 stangi is mnost adapted to the irrigated lowlands

The dark-headed stem borer(l)HSB) (ih pochit3sus was not collected in all rice environments and the gold fringed borer Chilo auricilius nd PSB were collected only in upland rice Xaantiholiplastenrnaloran ichneumonid larval parasite appears to be widely adapted whereas TrihomaslopisapanitI)cteiiaa pteromalid pupal parasite occurs mostly in wetlands and also parasitizes larvae of Chilo suppressais and Pelhuidas mathias

Armyworms Myit hinin and Spodujipera have two main larval parasites - a tachinid Zrgohothriaatropivora in rainfed environments and a braconid Cotesia spp in irrigated lowlands

The butterfly pests Pelopidas and 4elanitishave tricho-grammatid egg parasites found only in irrigated lowland environments

Two larval parasites show environmental preferences --Argrroph av nigrotihialis a tachinid in rainfed habitats and Trichonialopsisapantelkcnela in irrigated ricefields A limgrotibialisalsoattacks the sweet potato hornwormAgrius convolvtdi and T apanteloctena parasitizes DH S1B

Predators as a group show more distinct environmental preferences The aquatic damselflies Coenagrionidae are most abundant in the lowlands Cvriorhinus is also most adapted to the lowlands The nabid Stenonabis tagalicus occurs only in the uplands and isalso prevalent on legumes The coccinellid lady beetles wasps and ants are more prevalent in upland areas AlicrasIpis cit-ea larvae prey ona variety of aphids attacking legumes and maize Etunienes campanonia vespid mud wasp makes nests in trees and is therefore most adapted to the more botanically diverse uplands Soil-dwelling ants cannot tolerate flooding

Upland environments are habitats to arboreal carabid beetles Three species of Chlaeniusbeetles prey on leaffolder larvae and are perhaps more important than parasites (27)

The gryllids and tettigoniids are widely adapted egg predators not only on rice but also on maize and legumes They feed on eggs of most insects that are laid on leaves

The spidercommunity of upland rice environments isrich in species At one site (Batangas Philippines) 31 species have been recorded (23) Of the 176 spider species recorded in Philippine ricefields about one-half (82) occur in upland rice The spiderspecies of upland rice environments overlap more with those of rainfed lowland than with those of irrigated lowland rice (24) The same study showed that of the three environments irrigated rice has (ie greatest spider species diversity foiiowed by rainfed lowland and upland

However of the 10 most prevalent ricefield spiders 9 were abundant in upland rice showing wide adaptation All three spider guilds -- orb-web space-web and huting spiders shywere prevalent in upland ricefields There were however differences in environmental prelerences for some spider species

Of the orb-wcavers Tetragniathamanlihulatawas partishycila rlv abundant 7 japonica was low and Ltucatge dehcorata was absent in upland rice

As in other environments Aiypeiwnaifirtosana was the most dominant space-web spider however its relative numbers were lowerthan for lowland sites Among hunting spiders Ox|opes javanus and Lcosa leatcost fgla were more abundant and Lvcosa lpseudoanmnlata was less abundant in upland compared with lowland rice envishyroniments

Extensive rearing of rice insect pests in upland rainfed lowland and irrigated sites over an 8-yr period in the Philippines provided insight into the effectiveness of parasites as natural enemies (Table 7 8) Most of toe data are from large sample sizes taken over at least one crop which overcomes the pitfalls expressed by Van Driesche (238)

VSB egg parasitization was surprisingly similar across all environments Larval parasitization was highly dynamic among sites and even years within the same site but showed somewhat lowerlevels in uplandsthan in irrigated wetlands This would have been predicted for egg parasites as well since YSB is adapted to the lowlands and so must be its parasites This isevidence that Telenomus rowani is as good at dispersing as its main hosts Trowani clings onto the anal tufts of female YSB and parasitizes eggs as they are laid This phoresy may explain its wide adaptation The more specialized lowland YSB larval parasites therefore appear more effective than their upland counterparts with more alternate hosts

Low collection levels of PSB larvae especially in the wetlands make comparisons between environments difficult Low levels of larval parasifization were recorded in both rainfed environments Except in the Cagayan rainfed wetland site in 1980 low levels ef larval parasitization were evident at irrigated and rainfed sitesalike No differences by environment were evident Higher levels of parasitization occurred in Sarawak during a 1967 outbreak also from tachinids Rothschild (197) indicated that higher parasishytization occurred in upland rice areas

Skipper and green horned caterpillar larval parasitization rates were generally low and showed no environmental effect htighest activity occurred in Cagayan in 1980 (48 parasitization) Although larval parasitization levels of RLF were variable year to year activity appeared greater in the irrigated lowlands than in the uplands Again the highest levels were recorded in Cagayan in 1980 Cagayan has extensive grasslands and fallow around the rice areas and a relatively long rainfall period Rainfall in 1930 was good for crop growth and apparently for increase of larval parasites

IRPS No 123 January 1987 33

Table 7 Parasitization rates of rice planthoppers and green leafhopper in upland rainfed lowland and irrigated lowland environments Philipshypines 1976-840

Environment and site Year Egg parasitization () Nymphaladult parasitization ()

BPH W13P GLH BPII WBPH GLIH

UplandBatangas

Rainfed lowland

1976 1979 1980 1982

15 17 12

2(47)8(39)

4(51)

9(1800)14 (212)

12(304)

8 (101)6 (127)

9(116)

Iloilo

Cagayan

1976 1977 1978 1981 1980 1981 1982

0 (69)

11

44 48 40

-

-

46 55 44

0 (471)

11

52 56 49

0 (200)9 (214)

10 (1259)8(235)6 (900) 3(800)

-7 (462)

11 (270)3 (313)5 (900) 1 (800)

4 (600)10 (725)10 (362)12(870)5 (900) 3 (800)

Pangasinan

Irrigatedlowland

1984 1976 1978 1979 1982

13 (436) 13(379) 14 (457)

6(14)7 (197)

17(198) 11 (486) 18 (275)

3 (81)10 (274)3 (116)

16(871) 19(88)

28(211)9 (764)5 (647)7 (1204) 8(305)

Laguna 1977 11 7 21 1978 31 1979 14 4 4 28 1980 7 8 20 1981 1983

56 7(278)

26 16 (1650)

16 27(1921)

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R I Apostol unpublished 1lo1lo and Pangasinan 136 Cagayan BCanapi unpublished Liuna 175 I C Pantua unpublished 118 234 43 Carino and Shepard unpublished BPH = brown planthopper Nila parvara htgens W131Pi =whitebacked planthopper Sogatelia furcifera GLH = green leafhopper Nephotettrix virescens Figiires in parentheses are sample sizes

Tauher et al (230) reported that internal external and genetic factors influence the seasonal activity of parasitoids

Hopper egg parasites but not nymphaladult parasites were also parlicularly abundant at the Cagayan site possibly the egg parasites have more alternative hopper prey in the weedy areas than nymphaladult paisites

The seasonal dynamics of hopper egg parasites can be seen for upland rice in Batangas(Fig I 2) and for rainfed lowland lice in Pangasinan( 135)and Cagayan( 105) Levels of BPHWBPI-L and GLH egg parasitization were corn-parable in upland and irrigated lowland environments Except for some high rates of paraMsitization early in the season the parasitization rate tended to be steady in all envilonments However hopper nymphaladult parasi-tizution levels were more similar in the rainfed environments and lower than in irrigated rice

Overall parasites do not appear highly effective by themselves against upland or even lowland pests Among natural enemies parasites have attracted the most attention because they can be more readily assesscd

Predators and pathogens lend themselves better to management practices Predators the most important group of natural enemies in riccfields are difficult to quantify On the other hand pathogens appear less im-portant but can be readily cultured and disseminated

The natural enemy commulnity of upland rice is rich in species and differs significantly from that of lowland rice

These beneficial organisms must be conserved by applying insecticides judiciously particularly sprays Seed or soil placement of chemicals minimizes exposure to natural enemies The strategy to derive the greatest benefit from natural enemies isto allow the greatest number of beneficial species to thrive Then perhaps at least one species will be effective against each pest at any tine overcoming the variable abundance of each species within and between years

Programs to introduce exotic species or mass produce indigenous ones are ambitious and expensive because they require trained people on a sustained not ad hoc basis

Chemical control Insecticides offer rapid and efficient control of upland insect pests (190) but-should be used only after other control measures have been considered Insecticides are rarely used on upland rice because of cost (32) Upland rice yields are normally too low to justify the expenditure Also most upland rice is grown as a cash crop Spraying upland rice is more difficult than lowland rice because water is less accessible Government subsidy programs to provide pestishycides to tribal peoples for upland rice production have created problems of toxicity to humans because isolated people have had no experience to be able to handle pesticides safely

However ins-eticide use has beenjustified when a crop is

0Z

Table 8 Parasitization rates of lepidopterous pests in upland rainfed lowland and irrigated lowland environments Philippines 1976-84a

Parasitization (5)

Environment and site Year YSB GFB PSB Earcutting Swarming Skipper Greenborned RLF Brown larva larva caterpillar caterpillar caterpillar caterpillar larva semilooperEgg Larva larva larva larva larva larva

UplandBatangas 1977 - 0 (57) - 0 (63) - 0 (71) - 0 (106) 11(142) 2 (101)1978 43 (4697) 8(451) 5 (38) 3 (40) 6 (61) 2(58) 15 (57) 13 (129) 13 (1286) 4 (30)1979 59(4135) 12(320) 13 (11) 5(28) 9(45) 6(46) 7(23) 8(160) 9(1152) 8(26)

Rainfed lowland Iloilo 1976 - 0 (10) 0(10) 0 (7) 0 (19) 2 13) 15 (180) 5 (76)1978 0 (54) 0 (3) 0 (17) 9 (64) 4 (78) 3 (73) 0 (121) 0 (191)1979 64 (2350) 16 (84) 6 (17) 0(6) 18 (68) 25 (104)

1981 56 (1322) 18(304) 0(5) 12(51) 18(138)Pangasinan 1976 60 (7 7 )b 62(82) 0(111) 5 (39)

1978 58 (53)b 5(51) Cagayan 1979 59(713) 8 (101) 6 (34) 0(7)

1980 0(11) 4 (28) 15 (286)

62 ( 48(177) 61(140)1981 0(5) 12 (2) 13(45) 14 (92)

Irrigatedlowland Laguna 1974 62 8

1974-75 50 41 1975 43 72 1978 96 (623) 21(24) 33 (60)1979 83(2017) 18(110) 8(62) 11 (89) 22(400)1980 78 (1625) 20(75) 0 (16) 4 (72)1983 60

9 (80) 7 (270) 30

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R F Apostol unpublished Iloilo and Pangasnan 136 Cagayan B Canapi unpublished Laguna 175 P CPantua unpublished 118 234 143 Carino and Shepard unpublished YSB = yellow stem borer ScirpophagaincertulasGFB = gold fringed borer Chilo auricilius PSB = pink stem borerSesamia inferens earcutting caterpillar Alythimna separata swarming caterpillar Spcdoptera mauritia acronyctoides skipper Pelopidas mathias reen horned caterpillar Melanitls ledaismene RLF = rice leaffoldersCnaphalocrocismedinalis and Marasmiaspp brown semilooper Mocisfrugalis Figures in parentheses are sample sizes Egg masses

IRPS No 123 January 1987 35

Table 9 Effect of seeding rate and insecticide on plant density and Yield losses from insect pests in upland rice often areyield of Dagge upland rice Batangas Philippines 1976a comparable to those of lowland rainfed or irrigated rice

Seeding rate Insecticide Dosage Plant density Yield (0-Y30o) but usually from different pests The principal (kg aiha) seed treatment (kg aiha) (noIra-row) (tha) groups are soil pests feeding on sown seed and rootsat 14 DE followed by seedling pests These groups have largely gone

50 None - 35 a 20 a unnoticed leading researchers to conclude that stem borers50 Carbofuran F 10 38 ab 23 a and seed bugs are the most important pests (17) Yield loss 10 beCabofuranF100 None - 4551 c 23 a trials that pinpoint major pests need to includc seed

100 Dieldrin WP 10 52 cd 22 a treatments Granules applied in the furrow may not give150 None - 59 d 22 a adequate protection from many seed pests

aAv of 8 fields DE = days after crop emergence In a column means The floristic and pedological diversity of upland rice followed by a common letter are not significantly different at the environments is matched by the diversity of arthropods5level by D)MRT F = flowable Wi = wettable powder both pests and their natural enemies Irrigation has homoshy

genized lowland rice environments Flooded soils tend tothreatened by epidemics of locusts or armyworms New have the same properties and to eliminate soil pests The technology stressing efficiency has minimized insecticide spatial and temporal dominance of cultivated rice has cost benefited specialist pests

Seed treatment is normally inexpensive and can be Upland rice has no specialist arthropod species The economicallyjustified in -nanycases(66) Treating seed with monophagous pests on upland rice prefer the lowlands systemic insecticide (216) but not chlorinated hydrocarbons where rice evolved Rice in the uplands exists only because (247) is most effective in controlling seedling maggots of man and even though lowland rice predates upland riceFoliar sprays have to be repeated to obtain control and specialist species evolved Wheat maize and sorghum do often are washed off by frequent early season rains not have monophagous pests leading us to conclude that Granular insecticides are effective but required dosages are there is no evolutionary advantage for inssects to have too expensive to justify except under heavy infestation narrow host ranges in upland graminaceae(247) When directed against first-instar larvae banding Pests have adapted to the uplands not only by having granules in seed furrows (177) can be effective against white wider plant host ranges but also by having long life cyclesgrubs at low dosages (025 kg aii ha) (134) Seed treatments undergoing dormancy or being dispersive Each pest has a are not promising for white grub control (248) unique life history strategy suited to highly unpredictable

Seed treatment is also effective ainst ants and may be environments In agivenyear someinsect specics an better cheaper than increasing the seeding rate In Batangas exploit upland rice as a food source but no pest is a senious treating 50 kg seed ha was as effective as increasing the one every year on upland rice seeding rate to 100 kgha and using untreated seeds (Table The low yielding potential of upland rice means less 9) In Brazil seed treatments control termites and other soil quantity and lower quality of food for pests to exploit but insects including the lessercornstalk borer(66) Domiciano also less incentive for farmers to undertake control (58) however found seed treatments inconsistent in their The most effective control would be early and synchroshyeffect against high populations of lesser corn stalk borer nous planting of early-maturing upland rice varieties to

Baiting can be an inexpensive way of controlling ants prevent insect pest buildup Breeding for insect resistance seedling maggot flies mole crickets and field crickets Baits should initially focus on stem borers because they are verymade of locally available material and impregnated with difficult to control by other means Biocontrol efforts insecticide can be sparingly distributed taking adantage of should first stress conservation of natural enemies preshythe pests ability to disperse and encounter bait sites dators and pathogens rather than parasites Efficient

The new low-volume hand-held controlled-droplet chemical control includes seed treatment or baiting for seed sprayers may offer an advantage over high-volume knap- seedling and some root pests and using controlled dropletsack sprayes for controlling foliar pests Upland rice sprayers for spot treatments against foliar pests and seed canopies are more open than those in lowland rice and bugs when economic thresholds are reached better droplet penetration should result Controlled droplet applicators would be ideal for seedbugs and defoliators

CONCLUSION

Upland rice is attacked by a wider array of insect pests than is lowland rice mainly beca use of the addition of soil pests Generally population levels of lowland rice pests are lower in upland than lowland rice leading many people to conclude that insect pests are not important on upland rice

36 IRPS No 123 January 1987

REFERENCES CITED

1Abraham C C and K S R Mony 1978 Occurrence ofLeptocorisa acuta Fabr (Coreidae Hemiptera) as a pest of nutmeg trees J Bombay Nat ttistSoc 74553

2 Agyen-Sampong M 1975 HlierogrlphusdiaganensisKrauss (Orthop-tera Acrididae) a new pest of rice in Northern Ghana Ghana J Agric Sci 8249-253

3 Agyen-Samnpong M 1976 Varietal resistance and other inle-ratcd pest control components as factors of rice varietal improvement in West Africa Prcsented at the Second varietal improvement serinar 14-18 Sep 1976 Bouak Ivory Coast West Africa Rice Development Association Monrovia Liberia 18 p

4 Agyen-Sampong M 1977 Outbreak of Ihccrorrdnisorae in rice seedlings near Rokupr Sierra Leone lit Rice Res Newsl 2(6)16-17

5 Agyen-Sampong M 1980 Dirty panicles and rice yield reduction caused by hugs IntRice Res Newsl 5(l)11-12

6 Agyen-Sampong M 1982 The major pest problems of irrigated upland and mangrove swanip rice ecosystems ir the humid tropical Guinea savanna and Salel climate zones Pages 318-328 in Pro-ceedings training course cn concepts techniques and application of irtegrated pest management in rice inWest Africa 10-20 Jan 1982 Fendall Liberia West Africa Rice l)evelopioent Association Monrovia Liberia 505 p

7 Akibo-Betts V I and S A Raymundo 1978 Aphids as rice pests in Sierra Leone lit Rice Res Newsl 3(6)15-16

8 Akinsola F A 1975 Present status of different rice stemborers in parts of Nigeria iicL Entomo Newsl 328

9 Akinsola FA 1979 Ihe biology and ecology of rice stemborers in Nigeria Ph 1) thesis University ol lbadan Nigeria 217 p

10 Akinsola E A 1979 Guidelines of integrated management of insect pests of upland rice in West Africa Presented at the Seminar on integrated management of rice diseases and insect pests at Bobo-Dioulasso Upper Volta 17-22 Sep 1979 West Africa Rice Develop-ment Association Monrovia Liberia

II Akinsola IA 1984 Insect pests of upland rice in Africa Pages 301-305 in An overview of upland rice research Proceedings of the 1982 lIouak Iwsrv Coast upland rice workshop International Rice Research Institute 13O Box 933 Manila Philippines

12 Alam M S V TJohn and Kaung Zan 1985 Insect pests and diseases of rice in Africa Pages 67-82 in Rice improvement in Eastern Central and Southern Africa proceedings of the international rice workshop at Iusaka Zambia 9-19 April 1984

13 Alticri M A A van Schoonhovcn and J D Doll 1977 The ecological role of weeds in insect pest management systems a review illustiated with bean (Phaseohsvulgaris L)cropping systems PANS (Pest Artie News Summ) 23195-205

14 Anmante F 1967 The leaf-cutting ant Atia capiguaraa pest of pastures [in Portuguese] Biol6gico (Sao Paulo) 33113-120 (Rev Appl Entomol A56207)

15Andrade A C and D Puzzi 1951 Preliminary results of experi-merits on the control of Sculptcoris iastantus on sugarcane [in Portuguese] Bioulgico (Sao Paulo) 1744-49 (Rev Appl Entomol A41421422)

16 Appert J 1970 Maliarplia separaelha (rice stem borer) new observations of entomological problems of rice in Madagascar [in French] Agron Trop (Paris) 25329-334

17 Ariadeau Mand Z Harahap 1986 Relevant upland rice breeding objectives Pages 189-197 in Progress in upland rice research Proceedings of the 1985 Jakarta conference International Rice Research Institute 1O Box 933 Manila Philippines

18Asar S and Kisimoto 1975 Seasonal occurrence of the smaller brown planthopper in upland rice [in Japanese] Proc Kanto-Tosan Plant Prot Soc 2228

19 Bakker W 1974 Characterization and ecological aspects of rice yellow mottle virus in Kenya Meded Landbouwhogesh Wage-ningen 152 p

20 Balut F F 1970 White grubs attacking rice Oryza ativa L [in Portuguese] Biol6gico (So Paulo) 36321-322

21 Banerjee S N and L M Pramanik 1967 The lepidopterous stalk borers of rice and their life cycles in the tropics pages 103-124 in The major insect pests ofthe rice plant Proceedings ofa symposium at the International Rice Research Institute September 1964 The Johns Hopkins Press Baltimore Maryland USA

22 Barrion A T and J A Litsinger 1980 Ants a natural enemy of cliaiarac in dryland rice IntRice Res Newsl

5(4)22-23 23 Barrion A T and J A litsinger 1981 The spider fauna of

Philippine rice agroecosystem 1Dryland Philipp Entomnol 5139-166

24 Barrion A T and 1 A Iitsinger 1984 The spider fauna of Philippine rice agroccosystcms II Welland Philipp Eiitomol 611-37

25 Barrion A T1 and J A Litsinger 1984 lihanus (Diptera Tabanidae) eggs an alternative host of rice stei borer (SI3) egg parasite Telenoonis digns (Iliymenoptcra Scelionidae) lit Rice Res Newvsl 9(6)19

26 Barrion A T and J A Litsinger 1985 Identilication of rice leaffolders by wing markings Int Rice Res Newsl 10( 1)24

27 Barrion A T and J A Litsinger 1985 Chctiosspp (Coleoptera Carabidae) a leaffolder ILF)predator lit Rice Res Newsl 10(1)21

28 Bertels A 1970 Insect pests of rice and theircontrol [in Portuguese] Pelotas Brazil Instituto de Expcrimentacao Agropccuararias do Sul Circular 43 24 p

29 Bonzi S M 1982 Ctilo (flisilineus J de Joannis (Lepidoptera Pyralidae) stem borer of irrigated cereals in Upper Volta LioFrench] Agron Trop Riu Rizic Cult Vivrieres Trop 37207-209

30 Bourke T V T L Fenner J N LStihick G Baker EHassan D F OSullivan and C S li 1977 Insect pest survey for the year ending 30th June 1969 Papta New Guinea )epartment of Agrishyculture Stock and Fisheries Port Moresby 57 p

31 Bowden 1 1973 Migration of pests in the tropics Meded Rijksfac Landbouwwet Gent 38785-796

32 Breniere J 1973 The main rice parasites in West Africa and their control Pages 27-42 in Proceedings seminar on plant protection for the rice crop West Africa Rice Development Association Monrovia Liberia

33 Breniere J 1976 The principal insect pests of rice in West Africa and their control West Africa Rice Development Association Monrovia Liberia 52 p

34 Brown ES 1962 The African arinyworn Spoiolptera eXI)Ina (Walker) (Lepidoptera Noctuidae) a review of the literature Cornshymonwealth Institute of Entomology London UK57 p

35 Brown F B 1968 Upland rice in Latin America FAO-International Rice Commission 12th Session of the Working Party on Rice Production and Protection 2-14 Sept Peradeniya Sri lanka Food and Agriculture Organization Rome Italy 6 p

36 Buyckx F J 1 1962 Disease and insect pests of crops inCongo Rwanda and Burundi [iil French] Publications dc LInstilut National pour lestude Agrononiquedu Congo M Weissenbruck S A Brussels Belgium 708 p

37 Cariiio F 0 1981 Role of natural enentics inpopulation suppression and pest management of green rice lealhoppers Ph D thesis University ofthe Philippines at Los Bafitos PhilippinesLaguna 136 -

38 Carpente A J 1983 The development of rice cultivation and extension of vegetable productin in Zanzibar Final Report Vol II1 October 1983 Food and Agricultme Organization Ronte Italy

39 Carroll C P and S1 Risch 1983 Tropical annual cropping systems ant ecology Environ Manage 751-57

4(0 Cates R G 198 1 Host plant predictability and the feeding patterns of monophagous oligophagous and polyphagous insect herbivores Oecologia (Berl) 48319-326

IRPS No 123 January 1987 37

41 Cendcafia S M and F B Caora 1967 Insect pests of rice in the 62 Ene 1 C 1981 I)iiorldhol)ierusihilis Slater (licteroplera Philippines Pages 591-616 in The najor insect pests ofthe rice plant lygacidue Illissinac) a new rice pest front Nigeria lit Rice Res Proceedings of a symposium at the International Rice Research Newsl N3)15 Institute September 1964 The Johns Hopkins Press Baltimore 63 Fugenio T S 1965 rhe effect oi e-vCgetation on rice production Maryland USA 729 p and the possibility of overcoming it by leaf pruning MS thesis

42 Centro Ihternacional de Agricklittira Tropical 1984 Upland rice in University ol the Philippines at Los Baijos ILaguna Ihilippins 69 p Iatin Anerica Pages 93-119 hi An os ervies of uplaind rice research 04 Farrow R A 1975 The Alrican nmigratory locust ii is main Proceedings of the 1982 Boinakamp Ivory Coast upland rice workshop outbreak area of the middle Niger qlinlitatise studies of solitary International Rice Research Institute P0 Blox 933 Manila populations in relation to environmental factors Locusla No II Philippines 198 p

43 Chandra G 1979 Taxonomy ad hn riononlics ofthe imect parasites of 65 Fennal R G 1943 Food crop investigation Windward and rice Icalloppers and planthoppers in the Pihilippines and their Ieeward Islands First report Report onl work dunring the periodiiiportairce iii natural biol)gical cointrol Philipp Fntoltol 4119-139 April 1942-April 1943 Imperial College of Tropical Agriculture

44 Chang T T 1976 The originii colut ion enlIivation dissemination Trinidad 16 p (Rev of Appl Fintornol A3222-25) and diersificition of Asian and African rices -uphytica 25425-44 1 66 Ferreira F 1983 Integrated pest ritaageiret of upland rice [ill

45 Chatierjee P B 1979 Rice leaffolder attacks in India lit Rice Res Portuguese English summary] Pages 323-341 in Upland riceculttre Newsl 4(3)21 factors affecting production E Ferreira T Yamada and F-

46 Cheng Cbien-pan 19(4 Upland rice in Taiwan International Rice Malmolta eds Institutoda Potassae Foslato Institnto Internacional Cnmmission Wiorking Party oit Rice Production ard Protection I()th da Potassa Piracicaha Sao Paulo lraid Meeting 3-10 March 1964 Manila Philippines Food and Agricul- 67 Ferreira E arid F dl Silva Martins 1979 Occurrence of ture Organiaion Rome Italy 5 p Sogatohs orvziola (Muir) inupland rice in Goias Ilrail lit Rice

47 Chcrrett 1 M and 1) 1Peregrine 1976 A review of the status of Rcs Ncwssl 4(4)18 leaf-cutting ants and their control Ann Appl Biol 84124-128 68 Ferreira E and 1F da Silva Martiris 1984 Insect pestof rice ill

48 Chipeta F M 1974 Pests of rice in Malawi Pages 90-95 in Brazil and their control Jill Portugucsej FMIlRAtPA-CNPAF Proceedings Sith eastern Africa cereals research conference 10-15 Goiania Brmiil 67 p March 1974 DRB Mialda ed Ministry of Agriculture lilongwe 69 Ferreira FI F ca Silha Miartins S S Nero and 1 It P de Malawi Carvallio 1982 M tltiplc regression analysis todeterimine the effect of

49 Chu Yau-i 1979 The d atiaging period of the rice seedling fly insect populatiois ott upland rice yield [inIPortuguese Pesqui (Atlterigotiavxigu Stcin) (NItiscide Diptera) oil cori) at Fast Java Agropecu Bras 17671-675 Indonesia Plant Prot Bill 21403413 70 Ferreira F 1F da Silva Martins and F 1PZimmermann 1979

50 Conklin H C 1957 IHaninoo agriculture a report on aln integral Rice varietal resistance to tle lesser corn stalk borer [in Port uguese] systent of shi fti rig cult ivationi in the Pltilippincs Food and Agriculture Pesqui Agropccu Biras 14317-321 Organization Rome Italy 209 p 71 Fleming W F 1968 Biological control of lie hapanese beetle U S

51 Cramer H II 1967 Plant proection and world crop production Dep Agric RLs Sers [ech Bull 1383 78 p Pilaniensclhutz-Nachr 201-524 72 Food and Agriculture Organiation of the United Nations 1982

52 l)ani R C and Majunidar 1978 Prelininary observations on tie Country report surmimary Pages 27-37 in Report on the first incidence of root aphids on differeit rice cultnrcs Sci Cult 4488-89 FAOIUN)IPhail and regional training coiuirse in iniproved ctltural

53 Davidson R I 1979 lffects of root feeding on foliage yield Pages practices for upland rice 5-23 Oct 1981 Northern Region Agrculturd117-120 in T K Ci osb and R B Pottinger eds Proceedings of the Developririt Center Chiang Mai Food and Agricuilture Organiiashy2nd Australian conlerence on grassland invertebrate ecology Govt tion Rome Italy 37 n Printer Wellington New Zealand 73 Franco do Ariaral S 195 1 Residual action of insecticides in turfy

54 l)ean G 1W 1976 Rice insect pests inlaos Int Rice Res Newsl soil--the results of a field experiment [in Portuguese] Biol6gico (Sao 1(2)15 Paulo) 17209-214 (Rev Appl Entomol A41423)

55 I)ean G J W 1978 Insect pests of rice in Laos PANS 24(3)280-289 74 Freeman ) 1970 Report onithe Iban london School of -coroiics 56 )ingle H 1984 Behavior genes and lifc histories conplex adapta- Monographs on Social Anthropology No 41 Athlane Press

tions to uncertain environments Pages 169-194 in A new ecology London UK 317 p novel approaches to interactive systems P W Price C N Slobod- 75 Fuchs T W1 A Harding and 1W Smithlr 1979 Induction and chikoff and W S Gaud cdsl ohn Wiley New York USA termination oldiapause in the sugarcane borer Ann Flitonlol Soc

57 Dobelmann-JP and M Falais 1968 Upland rice cultivation ill tie Am 72271-274 North-western part of Madagascar in 1966-1967 [in French English 76 (hbriel B1) 1975 A review tf the major insect pests of some tpland summary] Trop Agron 2313-21 crops in tlc Philippines Pages 86-111 in Ad Htoc panel of experts on

58 Doniiciano N L 1979 Chemical control of tie lesser corn stalk pest disease arnd weed problems in some rainfed crops Bangkok (lolsonno~paloIgnowslhts) (Zeller 1848) (I-epid(ptera Ihycitid ae) Thailand Food aid Agriculture Organization Roire Italy

on rice in Ponta Gressa Parara 1977-1978 [in Portuguese] Pages 77 Garg D K and N K Shal 1983 White grub outbreak on rainfed 291-303 in Anais 1 Reuniao de tecnicos cmiirizicultura do estado de dryland rice in Uttar Pradeslh Int Rice Res Ncwsl 81(4)16-17 Sao Paulo-Campinas Cati Brazil 78 Garg I) K and IP Tandon 1982 Major insect pests of rice on hilly

59 Dunsmore 1 R 1970 Investigations on tlie varieties pests and tracts of Uttar Pradesh India lilt Rice Res Newsl 7(11)11-12diseases of upland rice (hill padi) in Sarawak Malaysia Int Rice 79 Garrity 1) P 1983 Upland rice research an international bibliograpy Comm Newsl 1929-35 1975-1982 Iternational Rice Research hnstitnte PO Box 933

60 Elias 1R1965 The new decline of rice seedlings is not caused by Manila Philippines 121 p parasites [in Portuguese] Biol6gico (Sao Paulo) 313-6 80 Ghauri M S K 1982 A new subspecies of Iinorh)rone bull

61 Empresa Brasiliera de Pesquisa Agropecuaria 1984 Upland rice itt cornlus Slater (Hemiptera Iygaeidae) front Papua New Guinea on Brazil Pages 121-134 it Av overview of upland rice research rice and carpet grass Ihll lEntoniol Res 72133-137 Proceedings ofthe 1982 Blouak Ivory Coast Upland rice workshop 81 Gonzales F 0 PArregoces R I ihernandef and OT Parada International Rice Research Institute PO Box 933 Manila 1983 Insect and imite pests of rice arid their control in Latin America Philippines [in Spanish] la Federacion Nacional de Arrocerts (Fedearroz)

Bogota Colombia 60 n

38 IRPS No 123 January 1987

82 Grist D H and R J A W Lever 1969 Pests of rice Longmans London UK520 p

83 Hale P R 1979 Two subterranean pests of upland rice in Papua New Guirnca IntRice Res Newsl 4(2)16-17

84 Hale P R and J T Hale 1975 Insect pests of rice in the New Guinea Islands Rice Entomol Newsl 34

85 Harris W V 1969 Termites as pests of crops and trees Common-wealth Institute of Entomology London UK 41 p

86 Hattori 1 1968 A method to distinguish between grass leaf roller (Ctaplha rocis medinalisGuenee) and the rice leaf roller (Susurtia exigua Butler) [in Japanese] Plant Prot (Japan) 23167-170

87 Henry I E and 1A Onsager 1984 Experimental control of the mormon cricket Iahrosshnph by NJsena locustae(Aficrospora Microsporidae) a protozoan parasite of grasshoppers (Ort Acri-didae) Entomophaga 27197-201

88 Hidaka 1 P Vungsilabulr and SKadkao 1974 Studies on ecology and control of the rice gall midge in Thailand Tech Bull 6 TropicalAgricultural Research Center Tokyo Japan 113 p

89 Hincklcy A 1) 1963 Ecology and control of rice planthoppers in Fiji Bull Entomrol Res 53467 481

90 Hirao J 1982 Ecology and chtInical control of the rice leaffolder Jpn Pestic Inf 4114-17

91 Ho DT 1984 Sten borer incifience in rice ecosystems in KenyaEast Africa Pages 307-320 in A overview of upland rice research Proceedings of the 1982 Bouac Ivory Coast upland rice workshop International Rice Research Institute PO Box 933 Manila Philippines

92 Ho )T J 1Njokah and J GKibuka 1983 Studies on rice sten borers in Kenya with emphasis on ifaliarpiaipiraellRag Insect Sci Appl 465-73

93 Ichihara I T Ito A NIatsuo and M Sawada 1971 Chemical control of the rice striped stem borer in upland rice grown in plastic mulch [iii Japanese] Proc Kanto-Tosan Plant Prot Soc 1882

94 IclhiharaIand A Matsuo 1969 Damage caused by rice striped stett borer to upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1686-87

95 Ichhara I atid A MIatsun11970 Conside ration oft lie extraordinary prevalence of rice striped stem borer in upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1778-79

96 Institut de Recherches Agrolnliqes Iropicale et des Cultures Vivrieres 1978 Insect resistance Pages 12-13 in Upland rice 1978 atnual report Bouak Ivory Coast 27 p

97 International Institute for Tropical Agriculture 1983 Aphids itt rice roots I ITA Res Briefs 4(1-3)4-5

98 International Institite fi Tropical Agriculture 1983 Hopperburn in rice caused by brown plathopper IITA Res Briefs 4(l-3)6-7

99 International Rice Research Institute 1975 Annual report for 1974 PO Box 933 Manila Philippines p 216

100 International Rice Restarch Institute 1976 Annual repiort for 1975 PO Box 933 Manila Philippines p 228

101 International Rice Research Institute 1979 Cropping systems program Component technology development and evaluation Page 404 i Annual report for 1978 11O Box 933 Manila Philippines

102 International Rice Research Institute Department of Entomology 1981) Insecticid evaluation 1979 Tables 73 74 PO Box 933Manila Philipes

103 International Rice Research Institute Department of Entonmology 1981 Insecticide evalomtion 1980 Table 81 POBox 933 Manila Philippines

104 International Rice Research Institute 1981 Report of a meeting of Asian rice-based cropping systems entomologists Eleventh cropping systems working group meeting 18-22 May 1981 PO Box 933 Manila Philippines p 132 133 144

105 International Rice Research Institute 1984 Cropping systems program Pages 368-373 in Annual report for 1983 PO Box 933 Manila Philippines

106 International Rice Research Institutc 1984 Upland rice in Asia Pages 45-68 in An overview of upland rice research Proceedings of the 1982 Bouakamp Ivory Coast upland rice workiiop loternational Rice Research Institute 11O Box 933 Manila Philippines

107 Irshad M 1977 Grasshoppers in rice in Pakistan lit Rice Res Newsl 2(3)8-9

108 Ito R and TOno 1964 Upland lice its culture and varieties in Japan lIternational Rice Cormmission loth Meeting of the Working Party on Rice Production and Protection 3-10 March 1964 Manila Philippines Food and Agriculture Organization Rome Italy 12 p

109 Jeffrey M I and V A Dyck 1983 Effect of moonlight on lightshytrap catches of brown planthopper lit Rice Res Newsl 8(6) 13

110 Jennings P R and T Alicia Pineda 1983 Management ofthe rice plnthoppers Sogatodsorivi(coIl in Latin America Presented at the meeting on stable plant resistance FAO and lIndocsian I)inectorate of Food Crops Protection Bali lndonesia 29 Novcniber-2 l)ccemshyber 9 p

IllJimenez Gomez J A 1979 Rice insect pests i lla ios ()riCntales lil Spanish] Pages 99-103 in Rice ciltivation Instituto (olotibiatto Agropecuario Compndio No 29

112 Jongeleen F JJ Soesanto and A Mockidjo 1979 Studies oti pathogens of two species of Scarabacidac froinIndonesia Z Angew Enlomnol 87412-417

113 Jouvenaz 1) PC S lofgren and W A Baiks 1981 Biological control of imported fire antsa review of cirrent knowledge Bull Entomol Soc Am 27203-208

114 Kainoh Y K Shiniiu S MarUani Y Taiaki 1980 lost finding behavior of tie rice bu lplotori+JchinvittjI)allas (liciniptera Corcidac) with special reference to diet pattern ol aggregation and feeding on the rice plant Appl Entomol Zool 15225-233

115 Kakkar K 0 P Sharma and P I Sharma 1977 Note on Ihteronyrchus liodresRedt (Coleopicra I)ynastidae) as asporadic pest of paddy and maie inHimachal Pradesh Corr Agric 156-57

116 Kalode M131 79 Major insect pest problems and their control in upland rice (Unpublishcd mineogriph CWI oJNaaiat lni Chandaratat (198 1)original not seen)

117 Kalshoven L i 98 I The pests of crops in I ldoniesia (translated by P A vatn der Laan) 1 T Ichtiar B3aru-vati Iloeve lakarta Indonesia 701 p

118 Kamal N Q 1981 Suppression of whitebacked planto ner Sogatella fircaifra (Horvath) aLId rice lealfolder (nhal -cis licdb lis (Guenee) populations by natural cnclmie hlI) tiesis Grcgorio Araneta University Foundation Victoncta Park Metto Manila Philippines 179 Ip

119 Karim A N M Reaul 1975 A species ofliealy bug Trioni i70s ceres new oii rice in the Philippines Rice Entoniol Newsl 247

120 Katanyukul Wand C Chandaratat 1981 Insect pestsof uplaid rice and their control Report firstFAOUNlFi Tliailand training course on improved cultural practices for tplad rice Chiang Mai Thailand 5-23 Oct 1981 Food and Agriculture Organiation Rome Italy 22 p

121 Kern F 1955 lhletwioa(I)uninemts) bhhit Biturm(Coleoptera) a new pest of sugarcane inVenezuela [inSpanish with Etnglish sunmary] Agron Trop (Maracay) 527-29 (Rev Appl Eitoniol A4572)

122 Khan K M and S Ghai 1974 White grubs ald their control in India Pesticides (1Bombay) 819-25

123 Kiritani K 1975 Integrated pest management of rice pests Pages 66-70 itt Proceedings platning workshop ott cooperative hIeld research in pest managementP SMotooka ed East-West Center Honololu Hawaii USA

124 Kisimoto R 1967 Ecology of insect vectors forecasting and chemical control Pages 243-255 in The virus diseases of tle rice plant Proceedings of a symposium atthe International Rice Research Institute April 1967 The Johns Hopkins Press altimore Maryshyland USA

125 Kisimoto R and V A Dyck 1976 Climate and rice insects Pages 367-391 in Climate and rice International Rice Research Institute PO Box 933 Manila Philippines

126 Kureha S S Kobayashi SAsari and T Ishihara 1977 Life history and control of the mole cricket Grrllotalpa africana Palisot de Beauvois [in Japanese English translation 7520 1RRI library] Proc Kanto-Tosan Plant Prot Soc 21192-197

127 Kushwaha K S 1981 The black beetle as a pest of rice in Haryana India Int Rice Res Newsl 6(4)14

128 Lawani S M 1982 A review of tile effects of various agronomic practices on cereal stem borers Trop Pest Manage 28266-276

129 Leuck D B 1966 Biology of the lesser cornstalk borer in South Georgia J Econ Entomol 59 797-801

130 lever R J A W 1955 Insect pests of the rice plant in Malaya Int Rice Comm Newsl 1612-21

131 Li Ching-Sing and Shin-Foon Chiu 1951 The study ofthe rice gall midge Iachydiplosis orvza Wood Mason [in Chinese with English summary] 1Taiwan Agric Res 21-13

132 Iitsinger J A 1984 Assesament of the need-based application for rice Proceedings TechnologyTransfer Workshop 15-16 March 1984 International Rice Research Institute 1PO Box 933 Manila Philippines 6 p

133 Litsinger J A 1984 Environmental diversity and insect pest abundance with reference to the Pacific Pages 285-307 in Proceedings sub-regional training course on methods of controlling diseases insects and other pests of plants in the South Pacific 4-20 Oct 1982 Vaini Kingdom of Tonga Nukualofa

134 Litsinger J A RF Apostol and M BObusan 1983 White grub Leucopholis irrorata (Coleoptera Scarabaeidae) pest status popula-tion dynamics and chemical conllol in arice-maize cropping pattern in the Philippines J Econ Entoinol 761133-1138

135 Litsinger J A C de la Cruz F Raymundo A T Barrion M D Lumaban J P Bandong M S Venugopal F Paragna and E Balete 1982 Insect pests and insecticide reponse of dry-seeded bunded rice Pages 179-193 in Report of workshop on cropping systems research in Asia International Rice Research Institute Los Bafos Philippine

136 Litsinger J A M 1) Luinaban J P Bandong A T Barrion W Katanyukul and N Panda 1978 Insect pest management in rainfcd lowland rice agroecosystms of tropical Asia International Rice Research Conference 17-21 April 1978 International Rice Research Institute Los Bafios Philippines

137 Litsinger J A and K Moody 1976 Integrated pest management in multiple cropping systems Pages 293-317 in R I Papendick P A Sanchez and G B Triplett eds Multiple cropping American Society of Agronomy Madison Wisconsin USA

138 Loevinsohn M E 1984 The ecology and control of rice pests in relation to the intensity and synchrony of cultivation Ph D thesis University of London 36) p

139 Loevinsohn M E J A Litsinger N Panda M A Austria R F Apostol C Andrion and J Bandong 1982 A basis for predicting rice insect pest abundance in cropping systems Paper presented at the 1982 International rice research conference on rice and cropping systems 19-23 April 1982 International Rice Research Institute Los Bafios Iaguna Philippines 25 p

140 Lopez A W 1930 Repot of the Entomology Department Pages 145-172 in Annual report of the Philippine Sugar Association 1929-30 Manila

141 Lopez AW 1931 Entomology Department Pages 227-266 in Annual report of the director or research 1930-31 Philippine Sugar Association Manila

142 Lopez A W 1932 Entomology Deparment Pages 252-279 in Annual report of the Philippine Sugar Association 1931-32 Manila

143 Lungibill P and G G Ainslee 1917 The lessercornstalk borer US Dep Agric Agric Res Serv Tech Bull 539 27 p

IRPS No 123 January 1987 39

144 McGuire J U Jr and BS Crandall 1967 Survey of insect pests and plant diseases ofselected food crops of Mexico Central America and Panama U SDepartment of Agriculture International Agriculshytural Development Service Agricultuial Researrh Service October 1967 11 p

145 McNeill Sand T R ESouthwood 1978 The role of nitrogen in the development of insectplant relations Pages 77-9 in lliochemical aspects of plant and animal coevolution J B Ilarbornu ed Academic Press London UK

146 Maebara H 1955 Spatial distribution of lastinstiarlanaeof purplish stem borer (Sesamia iqferens Walker) hibernating in an upland rice field Bull Kagoshima Univ Fac Agric 438-41

147 Mallick E H and N G Hajra 1977 Response of rice plant to defoliation treatments at different developmental stages Indian Agric 2151-56

148 Mansingh A 1971 Physiological classification of dormancies in insects Canl Entomol 103983-10)9

149 Martell C G 1974 Catalogue of plant feeding insects of Mexico [in Spanish] Fitofilo 27(69)1-176

150 Martinez 0 J 1982 Principal rice pests in Ilanos orientalis [in Spanish] P~ages 80-94 in Curso deact nalization de conocimientos en arroz Villaniencis Instituto (olonmbiano Agropecuario

151 Martins J F da Silva F Ferrcira A S Prabhu F 1 P Zimmermann 1980 Use of pesticides for the control of the main subterranean upland rice pests esqui Agropecu Bras 1553-u

152 Martins I F da Silva F Fe eira rod BdamSilveira Pinheiro I Q2 Damage simulation caused by defoliators of upland rice [in Portuguese with English summary] Pesqui Agropecu Bras 171113-1119

153 Martins 1F da Silva N(iyuen van Tan and 1Ida Silveira Pinheiro 1981 Stem borer resistance in upland rice associated with plant morphological characters [in Portuuce Iesqui Agropecu Bras 16187-192

154 Masaki S 1980 Sumnmer diapause Ann Rev Entomiol 251-25 155 Masuzawa T 11 Suwa and F Nakasoji 1983 l)ifkleences of

oviposition preferencc and survival rate of two skipper butterflies Parnara guttata and PIchlpidai iathiais (Iepidoptera HIesperiidac) on rice plant and cogon grass New Entomol 3247-56

156 Maurya D M and C P Vaish 1984 Upland rice in India Pages 69-92 in An overview oftpland rice research Proceedings ofthe 1982 Bouakamp Ivory Coast upland rice workshop International Rice Research Institute IO Box 933 Mnila Philippines

157 Menon M G R R N Katiyai S Kuar and S S Misra 1970 Termites attacking the paddy crop Sci Cult 36294-295

158 Miyahara Y and 1 Nishikubo 1967 Behavior of young nymphs of the red rice root aphid Iholahsipiun ri liabhdontinalisSasaki in relation to movement from the leaf to the roots of upland rice [in Japanese] Pioc Assoc Plant Prot Kyushu 1363-66

159 Mo ljoa Tjien 1952 White grub of upland rice (Holotrichiahelleri Brsk) [in Indonesian with English summary] Contrib Agric Res Stn Bogor No 13134 p

160 Mochida 0 and T Suryana 1979 The occurrence of the brown planthopper and ragged stunt on lowland and upland rice ill Central Java and logyakarta in tile late of January 1979 Entomnolgy Survey Report No 15 Central Research Institute of Agriculture (CRIA) Sukamandi Indonesia I February 1979 9 p

161Monet C 1979 Varietal resistance to upland rice pests [inI French] Pages 35-39 in Annual report 1977 Institut des Savanes Department des Cultures Vivrieres Bouak6 Ivory Coast

162 Monte 0 1942 A caterpillar ofrice-fields[in Portuguese] Biol6gico (Sao Paulo) 8161-163 (Rev Appl Entomol A31414)

163 Munroe I) D 1979 Pest management strategy for hill rice in four resettlement areas in Sarawak Malaysia Pages 89-102 i1 Guidelines for integrated control of rice insect pests Food and Agriculture Organization Plant Production and Protection Paper 14 Rome Italy 115 p

40 IRPS No 123 January 1987

164 Nakasuji F M Ishii and Jen-Zon Ito 1984 Rice skippers in Taiwan and their life histories ZZrC (Taipei) Tech Bull 82 10 p

165 Navas I) 1976 Fall armyworum in rice Pages 99-106 il Ecological animal control by habitat management Proc Tall Timbers Con-ference 28 Feb-I Mar 1974 Fall Timbers Res Stn Misc Publ 6 Tallahassee Horida USA

166 Nickel 1 1 1961 Biological control of rice stem borer A feasibility study Tcch Bull 2 International Rice Rcarch Institute PO Box 933 Manila Philippines IIl p

167 Nigan P M and R A Vernia 1985 Insect pests of uplaild rice in Uttar Pradesh Int Rice Res Newsl 10(6)22

168 Nilakhe S S 1985 -cological observations on spittle bugs with emphasis on their occirence in rice Pcsqtui Agropecu Bras 20407-414

169 Nixon G EJ 195 1 lie association of ats with aphids and coccids Commonwealth Iusti lcte of Entonology Iondon UK 36 p

170 Nurdin F an Ishak Manti 1985 Study oit seedling fly of upland rice Interlational Upland Rice Moitoring Tour Report 26 lebruary-2 MarLI Sukaranti West Sumatra

171 Oka 1 N 1979 Ciltural conttol of the brown planthopper Pages 357-369 in Brown planthopper threat to rice production in Asia International Rice Research Institutc 1p0 Box 933 Manila Philippines

172 Ohni M and A Vilnaii 1975 Rice insect pests of Italy biology of Sipha grcria (Kaltenbach) and its control [in Italian] Riso (Milan) 2459-71

173 Otanes F Q atnd 1 I Sison 194 1 Pests of rice Philipp J Agric 12211-259

174 Painter R I 1951 Insect icsistance in crop plants University Press of Kansits Lawrence Kansas USA 520 p

175 Pantua P C and 1A litsinger 1981 Arthropod abundance in continuous vs biannual rice cropping systeis Int Rice Res Ncwsl 6(2)20-21

176 Pathak M ) and V A l)yck 1975 Sttudics ott insect pests of upland rice Pages 186-191 in Major research in upland rice International Rice Research Institute PO Box 933 Ilanila Philippines

177 Patil B R B M IHasabe and 1) S Ajri 198 I Chemical control of white gritb lIditriihinhorraia Fabr intfesting groundntut paddy atnd sorghum Indian 1Entomol 43232-236

178 Pawar A 1) 1974 (ollection and identification of insect pests of rice rlerminal repirt lInternational Rice Research Institute PO Box 933 Manila Philippines 72 p (tnimco)

179 IerrajU A aid A S Reedy 1965 Observations ott diapausc itt tie steti borer ohrice Rice News Teller (lai-Mar)9-13

180 Pont A C 1972 A review of the Oriental species of theri ona Rondani (I)iptera Muscidac) of econtomic importance Pages 27-102 in Control of sorglium shoot fly M G lomtani and W R Young eds Oxford atd IH1 Ptthl Co New I)clhi India 324 p

181 Prasad S C 1PChand aid M 1PSingh 1983 Incidence of rice gall midge in the upland plateau region of Hihar Oryza 20235-236

182 Rai P S 198 0 Nisaga simpex damage to rice in the hill tracts of South India Int Rice kles News[ 5(I)12

183 Rathorc Y S 1969 Gonocplhahli dcprssunt Fabr as itpredator of (tilo zonwIlhis (Swinhoc) and other moths in field cage at Patnagar Uttar Pradesh Indian 1Entotnol 3127

184 Reed W 1983 Crop losses causcd by insect pists in the developing world Proceedings 10th Internatial Congress of Plant Protection lrighton UK 174-80

185 Reedy M V 1981 Ateactogasterinhens(Walker)a new root-grub pest of tipland paddy and inaize Curr Sci 501076

186 Reveno M A 1968 Catalogue of the insect pests of economic crops in Colombia [in Spaiish] Asociacion Latino Americana de Entomo-logia (AIAE) 129-33

187 Reyes G M 1957 Rice dwarf disease in the Philippines Food and Agriculture Organiiation Plant Prot Bull 617-19

188 Rice S E A A Grigarick and M 0 Way 1982 Effect of leaf and panicle feeding byarmyworm (Lepidoptera Noctuidae) larvae on rice grain yield Environ Entomol 7593-595

189 Risch S J 1) Andow and M A Altieri 1983 Agioccosystem diversity arnd pest control data tentative conclusions and nizw research dirctions Environ Entomol 12625-629

190 Rivera C T 1956 Three organic insecticides in thecontrol of insects affecting upland rice Philipp Agric 39465-472

191 Roffrey 1 1972 Locusta outbreaks in the Philippines Acrida 1177-188

192 Rojas R 1) 1941 The effect of pruning at varying ages upon the yield of Ramairio Phipp Agric 29851-860

193 Roonwal M i 1979 Ternite life and termite control in tropical South Asia Scientilic Pnbl odhpr India 139 p

194 Rosetto C1 F da Silva Martins N C Schmidt and L EAzzini 1978 Damage caused by pasturespittle bugs 4Dvoi)lavopictaandDJ schach) on rice lBragantia Nota 6 (Matcli)25-27

195 Rosztto C1- S S Neto I- Amantc I) Iink 1Garcia-Vieira D M de SoLza N V Bansa t and A M de Olivcira 1971 Rice pests of Brazil [in Portuguese] Pages 149-238 inrFA()-International Rice Commission 2nd meeting of the Rice Committee for the Americas 6-11 l)ecember 1971 Pelotas Rio Grande do Sul Brazil

196 Rothschild G H L 1969 Obseratiois on the armyworm Spodoptera matritia acronrctoiiles Gn (Lep Noctuidae) Bull Entomol Res 59143-160

197 Rothschild G 1- L 1974 Parasites of the armworm Spolopiera mauritiaacroncitoihes in Malaysia Fntornopliaga 19293-299

198 Sachan i N S K Gangwar and 1N Katujar 1980 Field crickets damaging cages in North Eastern IHill Region Indian J Entomol 42526-527

199 Sands 1) P A 1977 The biology and ecology of Lepocorisa (Hemiptera Alydidac) in Papua New Guinea Res Bull 18 Departshyment of Primary Industry Port Moresby Papua New Guinea 104 p

200 Sarawak 1968 titomology division Pages 97-106 in Report of the research branch of the Department of Altriculture 1966 Kuching (Rev AIppl Eintomol A56603)

201 Satpathy JM M S Das and K Naik 1977 Effect of multiple and mixed cropping on the incidence of scine important pestsI Entomol Res 178-85

202 Sauer IHF G 1939 Notes Ott Elasmpnlpualtsligniosellths Zeller (Lep Pyr) a serious pest ofcereals in Sao Paolo [in Portuguese] Arq Inst Biol 10199-206 (Rev Appl Entotnol A28378)

203 Satinders 1L A 1 SKing and C L Vargas S 1983 Crop pests of Central Anerica with t bibliography [in Spanish] Centro Agroshynotnico Tropical de Investigacion y Ensenanza Turrialba Costa Rica (CATIE) Tec Bol 9 90 p

204 Sauphanor B 1985 Some factors of upland rice tolerttce to stem borers in West Africa Inseu Sci Appl 6429434

205 Sen P atnd S Chakravorty 1976 Stentflics in Haritighata Nadia West Bengal India [it Rice Res Newsl 1(12) 15

206 Senapai B and 1M Satpathy 1983 The seedling fly Atherigona orrzae Malloch is a pest of rice in Orissa (India) J Entomol Res 777-79

207 Shhjahan M S Alatn M Ah nad and BN Islam 1983 A new scarabaeid beetle pest of dryland rice in Bangladesh Int Rice Res Ncwsl 8(I)16-17

208 Sharma G L S C Modgol and R C Gautam 1979 Effect of depth of submergence ott incidence of bacterial blight and white grub infestation in transplanted rice Int Rice Res Newsl 4(6)12-13

209 Sharma S 1985 Scarabaeid beetle outbreak itt upland rice Int Rice Res Newsl 1(1(l)24

210 Shiraki T 1917 The paddy borer Schoenobhits incertulas Walker [achaku Agric Exp Stn Formosa 256 p

211 Silva A de Brito and 11P Magalhaes 1981 Insect pests of rice [in Portuguese] Circular Tecnica No 22 EMBRAPA Centro de Pesquisa t ecuaria de Tropico Unido (CPAJU) Belem Brazil 14 p

212 Singh S A 1979 Emergence of rice leaf butterfly as a rice pest in Manipur India int Rice Res Newsl 4(4)16-17

213 SinghS A 1983 Changing trends of insect pestsof rice in Manipur Int Rice Res Newsl 8(f)12

214 Siwi S S and P H van Doesburg 1984 Lenovorisa Latreille in Indonesia (Heteroptera Coreidae Alydinae) Zool Meded (Leiden) 58117-129

215 Soejitno J 1977 Reaction of some upland rice varieties toseedling fly eltherikonaexigua Stein [in Indonesian] Siri Hana I Penyakit 630-35 Lembga I osat Fenclilian Pertanian Bogor Indonesia

216 Socnardi 1967 Insect pests of rice in Indonesia Pages675-683 in The major inse pests of thu rice plant Proceedings of a symposium at the Internatital Rice Rec Wi itt S pternber 1964 The Johns Hopkins Press Baltimore Maryland USA

217 Soto P E A T Petcz and I W Buddenhagen 1976 Survey of insect pests and diseases of rice in different ecological zones in Nigeria Rice Entomol Ncwsl 435-46

218 Soto 11E and Z Ziddiqi 1976 Screening for resistance to African rice insect pests Varietal improsetnent sCmi n r Bouak Ivory Coast 13-17 Sept 1976 West Africa Rice l)evelopment Association Monrovia Iiberia

219 Squire F A 1972 Entomological problems in Bolivia PANS (PestArtic News Sumr) 18249-268

220 Stapely 1 II 1978 The rice leaf roller in the Solomon Islands Int Rice Res Newsl 3(l)13

221 Suenaga H 1937 On aphids attacking the roots of rice Oyo-I)oubts Zasshi 9150-152 [in Japanese] (Rev Appl Entomol A25745)

222 Suenaga H 1952 On the seasonal abundance of three subterranean aphids on upland rice Bull Kyushu Agric Exp Sti 1249-262

223 Sutynso Yas and Dandi Soekaria 1964 Beberapa hana yang ttenperang tana man padi tanah kering [in Indonesian] Rapat kerja Padi Tanah Kering 6 sd 8 Juily 1964 Kaliurang 9 p

224 Tallamy I) W and R F Denno 1983 Alternative life history patterns in risky environments in example from lace bugs Pages 129-147 in Insect life history patterns R F l)cnio and If Dingle eds Spring-Verlag 225 p

225 Tamura 1 1939 Observations on some insect pests of field crops [in Japanese] 1 Plant Prot 26414-420 (Rev Appl EntomolA27-556)

226 Tamura and Suzuki 1964 Influences of the early-transplanting practice in rice culture upon development of the rice stem borer Chilo siql)ressalisWalker Bull Ilokuriku Natil Agric Exp Stn 761-94

227 Tan J P 1924 The rice root aphid Drowhi ihirsoia Philipp Agric 13277-288

228 Tanaka 1 1957 Taxonomy and distribution of some subterranean aphids injurious to upland rice in Japan with description of a new species [in Japanese English summary] Botyu-Kagaku 22168-176

229 Tanaka T 1961 The rice root aphids their ecology and control Spec [full College of Agriculture Utsunomiya University Japan F3 p

230 Tauber M IC A Tauber R Nechols and 1Obrycki 1983 Seasonal activity of parasitoids control by external internal and genetic factors Pages 87-108 itt I)iapause and life cycle strategies in insects V K Brown and I Ilodek eds )r W Junk The Hague Netherlands

231 Taylor W F 1972 Effects of artificial defoliation (simulating pest damage) on varieties of upland rice Exp Agric 879-83

232 feotia T P S and S Natud 1966 Biotoomics of the rice skipper Parnaraniathiis Fabricius (Lepidoptera Hesperiidac) Indian J Entomol 28181-186

233 Teran F 0 1971 Rice pests of Santa Cruz [in Spanish Ministerio de Agricultura Sita Cruz Bolivia Bull 45 8 p

234 Than Itun 1976 Population dynamics of the yellow rice borer Tryporza itcertulas (Walker) and its damage to the rice plant MS thesis University of the Philippines at Ios Bafios Laguna Philippines 92 p

235 Uichanco L B 1911 Pests Pages 55-74 it Rice Philipp Agric 11-200

IRPS No 123 January 1987 41

236 van der Laan P A 195 1 Life history of the rice seedling fly Contrib Gen Agric Res Stn Boor Indonesia 118 15 p

237 van der Laan 11A 1959 Correlation between rainfall in the dry season and the occurrence of the white rice borer (Scirpolhaga innotala WIk) in lava Entomol Fxp Appl 212-20

238 Van Driesche R G 1983 Meaning of percent parasitismin studies of insect parasitism Fnviron Entomol 121 I1- 122

239 Venugopal M S and A litsingcr 1984 Fffect ofcarbofuran on rice growth Prot Ecol 7313-317

240 Vietnam Directorate of Rural Affairs 1962 Annual work progress report on crop improvement program of rice sugarcane vegetable and field crops (lor the period from July 1961 to June 1962) (11) 299 p (Rev Appl Entomol A51526-527)

241 Wan M T K 1972 Observations on rice leaf- and planthoppers in Sarawak (Malaysian Borneo) Malay Agric 148308-335

242 West Africa Rice I)esclopnient Association 1978 Annual research report Vol 2 Statistics Monrovia Liberia

243 West Africa Rice Development Association 1979 Annual research report p47 Monrovia Liberia

244 Williams ) F and CS lofgren 1981 Fli I illy l-468 anew bait toxicant for control of the red imported lire at Fla Entomol 64472477

245 Williams I) 1 1970 The mealybugs OFlomoptera Coccoidea Iscudococcidac) of sugar-cane rice and sorghum Bull Entomol Res 0109-188

246 Williams D) 1 1982 Pulinaria icervi (Signoret) and its allies ott sugar cane and other grasses Btll Entoinol Res 72111-117

247 Wirjosuiardjo S 1976 Study ofchenmical cont rot of insect pests ott upland rice Imiu Pertanian (Jogjakarta) 1323-329

248 Wirjosuhardjo S and N Wirarta 1976 Chemical control of the white grub tIolorithiatlltriin upland rice in M uneng Irobolinggo Indonesia) [in Indonesian] Imut Pertanian (Jogjakarta) 1330-332

249 Yano K T Miyake and V F Eastop 1983 The biology and economic importance of rice aphids (Henuiptera Aphid idae) a review Bull Entomol Res 73539-566

250 Yelp D and P Rhodes 19(4 lhe success of drilled dryland rice iii the Fiji Islands International Rice Commission Working Party on Rice Production and Protection Manila Philippines 3-10 March1964 Food and Agriculture Organization Rome Italy 13 p

251 Young W R 1981 Fifty-five years of research on the sorghum shootfly Insect Sci Appi 23-9

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Page 3: UPLANDRICE. INSECTl--1i M T -E!R.EOLOU t RTANCE ,AND …delphacid.s3.amazonaws.com › 6958.pdf · The significance of abiotic factors in upland rice insect ecology will be apparent

UPLAND RICE INSECT PESTS THEIR ECOLOGY IMPORTANCE AND CONTROL

Upland ricefields are nonpuddled and unbunded without the expectation of impounding water They may be located in many settings - in isolated pockets surrounded by irrigated wetland fields or along steep slopes of recently cleared forest Each enviionment produces a unique composition of insect species

Information on upland rice insect pests is limited and scattered among reports dealing with other rice cultures We found eight reports specifically treating insect problems of upland rice culture (11 59 66 79 116 120 163 176) but none worldwide in scope Most literature focuses on one species or a group of related species Table I lists upland rice insect pests that others have identified and those we have observed Many citations refer to lists of pests with rice as a recorded host or are based on insect habits

Papers concerning upland rice often fail to distinguish between the two ecologically significant aspects ---culture and environment embodied in th term upland rice It is dangerous to extrapolate results of studies of insect pests in well-drained fields where seeds of upland rice

varieties are sown into dry soil (upland rice culture) but in an otherwise lowland environment (fields surrounded by flooded rice) Insects readily fly from the lowland rice-fields to the fields planted in upland rice culture These insects therefore are not necessarily upland rice insects They breed on lowland rice and colonize upland rice We gain greater ecological significance from faunistic records and ecological studies from an upland rice environment where lowland ricefields ar- out of the effective dispersal range of most insect species (tens of kilometers away) In border areas where lowlands and uplands meet studies cannot accurately represent either upland or lowland rice (181)

IPEST (iROUPS

Upland rice insects are more influenced by physical than by biological or socioeconomic parameters I) well-drained soils (lack of prolonged flooding or soil puddling) 2) high probability of drought during crop growth 3) restricted growing season (lengthy nonrice fallow) 4) ricefields interspersed with other crops (diversified flora) and 5) low use of agrochemicals (because of low and unstable yield)

Although a dryland habitat represents an extreme hydrological condition upland rice is host to all but the most aquatic insects (11 176) Whorl maggots caseshyworms water weevils and bloodworms require ponding Many soil and scedling pests are not common in lowland ricefields Deep water rice is established in dry soil and therefore has more in common with upland and rainfed lowland rice than with irrigated rice even though water depths may later reach 1-3 inSecond to deep water rice upland rice represents the most unstable rice environment for foliar insects But upland rice is highly stable for soil insects

The significance of abiotic factors in upland rice insect ecology will be apparent in a discussion of the most prevalent groups of insects attacking upland rice worldwide

Soil -inhabiting pests Well-drained nonpuddled upland rice soils favor pests that pass at least one growth stage underground Soil pests feed on underground plant parts (sown seed or roots) develop entirely in a subterranean habitat and leave only for adult mating and dispersal flights Soil pests include ants (Formicidae) termites (Isoptera) mole crickets (Gryllotalpidae) field crickets (Gryllidae) white grubs and black beetles (Scarabacidac) root aphids (Aphididae) root-feeding mealybugs (Pseudococcidae) root-feeding bugs (Lygacidac Cydnidae) false wireshyworms (Tenebrionidac) wireworms (Elateridae) root weevils (Curculionidae) and soil-inhabiting cutworms (Noctuidac)

A subterranean environment limits mobility particularly in locating food and soil insects have adapted by 1)being long-lived either as individuals (beetles or orthopterans) as colonies olsocial insects (ants or termites) or as dependents on social insects (mealybugs and aphids) and 2) having a wide host range (all species)

Anis Species of -rrestrial ants notably the ubiquitous fire ant Solenopsis geminata and harvester ants Pheidole spp specialize in feeding on ungerminated grass seed Solenopsis readily colonizes disturbed habitats which in turn initially encourage the growth of grasses (39) Colonies of these granivorous ants have a specialized caste that processes seed for food Foraging workers bring seed to

4 IRPS No 123 January 198

Table 1Insect pests of upland rice worldwide

Order Family Species

Isoptera Rhinotermitidae Coptotermesformosanus Shiraki

Termitidae Heterotermesindicola (Wasmann) Anacanthotermesviarum (Koenig) Anacanthotermesrugifrons Mathul et Sen-Sarma Captiterpnesnitobei (Shiraki)Corniterrnesstriatus (lHagen) Procapriterines mnushae Oshima Pericapritermes nigerianus Silvestri (=socialis)Macroternes nutalensis (Haviland)Macrotermes beiiicosus Smcat hman Macroterines gilius (Hagen) Macrotermnes spp Odontoterniesformosanus (Shiraki)Procornitermesarauloi Emerson Procornitermws triacifer (Siivestri) Trinerviterniesgenzinatus Wasmann

[= ebenerianus Sjostedt]MicroceroterynessppMicroterinesspp Synternes nolestits (Burmeister)

Dermaptera Forficulidae Termites unspecifiedDiaperasticuserythrocephalusOlivier

Orthoptera Gryllotalpidae Don lineare (Eschscholtz)GryllotalpaafricanaPalisot de Baeuvois Gryllotalpaorientalis (=africanu) Burmeistera

Gryllidae

Neocurtilla (= Gryllotalpa)hexadacryla (Perty) Scapteriscusdidact)hts(Latreille) Grjllusassinilis(= bimaculatus)(Fabricius)Gryllus (=Liogryllus) bimaculatusde Geer Brachytrupes portentosus (Lichtenstein) Brachytnmpes mnembranaceus DruryPlebeiogrylhsplebejus (Saussure) Teleogryllus testaceus (Walker)Teleogryllus occipitalis (Serville) Loxoblenimus haani Saussure

Pyrgomorphidae

Velarifictorusaspersus(Walker)Velarifictonis sp Euscyrtus concinnus (de Haan)Atractomorpha burri I Bolivar

Tettigoniidae A tractomorphapsittacinapsittacina(de Haan)Conocephahslongipennis(de Haan) Conocephalusrnaculatus(Lethierry) Conoceptahissaltator Saussure Euconocephah s varius (Walker)

Tetrigidae Acrididae

Caulopsiscuspidatus (Scudder) Caulopsis oberthuriScudder Phaneroptera furcifera (Stal) Amphinofus spp Aiolopus thalassinustamulus (Fabricius) Acrida ivilleinsei Dirsch Gesomida mundata zonocera Navas Gonistabicolor (de Haan)Oxyajaponicajaponica (Thunberg)Oxya hya intricata (Stal) Oxyafitscovittata (Marscliall)Oxya velox (Fabricius)Gas frimargus niarmoratusgrandis (Saussure) Xenocatantopshunilishumilis ServilleHierogiyphus banian (Fabricius) ltieroglyphusdaganensis Krauss Hicroglyphus nigrorepletus(1 Bolivar) Hieroglyphus oilziiorosCarl Heteropternisrespondens (Walker)Patangasuccincta (Linnaeus) Stenocatantops splendens (Thunberg) Chondracrisrosea brunneri Uvarov Trigonidium cicindeloides Rambur Locusta migratoriamanilensis (Meyen)Locusta migratoriacapito Saussure

Distribution

Asia Asia Asia Asia Asia Latin America Asia Africa Africa Africa Asia Africa Asia Latin America Latin America Africa

Asia Asia Africa Latin America Asia Africa Latin America Africa Asia Latin America Latin America Asia Asia Asia Africa Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Latin America Asia Latin America Latin America Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Africa Asia Asia Asia Asia Asia Asia Asia Asia Africa

Reference

85 82 190 190 82 85 151 82 85 7 85 85 178 12 82 85 85 195 195 85

193 11 12 157 60 85 195 72 36 186 12 32 59 72 76 84 81 150 195 203 211 81 76 178 b 48 b 76 178 b b b b b 55 178 178 178 65 178 68 81 68 178 178 178 178 178 55 72 178 178 78 178 178 55 55 72 6 82 82 55 54 55 59 130 55 178 55 78 55 82

IRPS No 123 January 1987 5

Table I continued

Order Family Species Distribution Reference

Locusta migratoria migratorioidesReische Africa 33 et Fairmaire

Schistocercagregaria (Forskal) Africa 33 Schistocerca americana (Drury) Latin America 82 195 Valanga nigricornis (Burmeister) Asia 82 Orphulella inticata Scudder Latin America 68

lomoptera Coccidae Pulvinaria sorghicola De Lotto Africa 246 Pulhinariaiceryi (Guerin) Latin America 186 246

Pseudococcidae taennococcus spp Asia 120 Lhorizococcus iu Williams Asia 245 Dysmicoccus boninsis (Kuwana)C Asia Africa 245

Latin AmericaDysmicoccus brevipes (Cockcrell)c Asia Africa 117 203245

Latin AmericaDysmicoccus oryzae (Wirjati)c Asia 245 Geococcus (=Ripersia) oryzae (Kuwana)C Asia 245Brevennia (= tererococcus)rehi (Lindinger) Asia 72 245Nipaccoccus graminis (Maskell) Africa 245 Planococcoides lingnani (Ferris) Asia 245 Pseudococcus saccharicola Takahashi Asia 245 Pseudococcus spp Latin America 60 Saccharicoccus sacchari (Cockerell) Africa Asia 245

Latin America Trionymus ceres Williamsc Asia 119245j rionymus sp Asia b Mealybug unspecified Asia 83 84

Aphididae Capitophorus[= Rhopalosiphumn] prunifoliae Asia 221 (Shinji)

Rhopalosiphum maidis (Fitch) Asia 249 Rhopalosiphum padi (Linnacus) Asia Africa 12 97 249 Rhopalosiphum rufiabdominalis (Sasaki) Asia Africa 66 76 120 195203

Latin America 221229240249 Tetraneura akinire Sasaki Asia 221249Tetraneurabasui ltillcRisLambers Asia 249Terraneura nigriabdominalis (Sasaki) Asia Africa 712 30 76 227229 (=Dryopeia hirsuta)

Tetraneuraradiciola Strand Asia 249 Paracleus ciniciformnis von Heyden Asia 249

[= Forda harukawai Tanaka]Pineus hanikawai (lnouye) Asia 228 Geoica hcifitga (Zehntner) Latin America 76 195 229 249

AsiaGeoica setulosa (Passerini) Asia 249 Anoecia fuliabdominalis (Sasaki) Asia 249 Anoecia comi (Fabricius) Asia 221229249Hysteroneura sctariae (Thomas) Africa Asia 7

Latin America Sipha glyceriae (Kaltenbach) Asia 249 Root aphid unspecified Asia 72 120

Cercopidae Deois schach (Fabricius) Latin America 61 68 194 Sepullia (=Denoplux) nigropunctata Stal Africa 29 Tomaspis ( Deois) fluxtuosa (Walker) Latin America 61 68 194 Tomaspis ( Dcois) flaiopicta (Stal) Latin America 61 61 194 To naspis (Dcois) complcta (Schmidt) Latin Aincrica 68 Tomaspis ( Zulia) entreriana (Berg) Latin America 68 Tomaspis ( Mahanarva) fimbriolata(Stal) Latin America 68 194 Tomaspis ( Aeneohna) spectabilis (Distant) Latin America 219 233 Aeneolamia varia Fciinai Latin America 81 111 Aeneolamia postica (Walker) Latin America 203 Abidama producta Walker Asia 54Delphacidae Sogatella furcifera (Horvath) Asia 59241 Niapanatalugens (Stal) Asia 30 59 72 84 241 Sogatodes oryzicola (Muir) Latin America 6167150Sogatodes cubanus (Crawford) Latin America 150 Sogatodes pusanus (Distant) Asia bPeregrinus maidis (Ashmead) Latin America 186 Laodelphax striatellus (Fallen) Asia 18

Cicadellidae Recilia dorsalis (Motschulsky) Asia 59 241 Nephotettiv nigropictus (Stal) Asia 59241

6 IRPS No 123 January 1987

Table I continued

Order Family

Hemiptera Lygaeidae

Cydnidae

Alydldae

Coreidae Corimelaenidae

Pentatomidae

Thysanoptera Thripidae

Species

Nephotettix virescens (Distant) Nephotettix modulatus Melichar Cofana spectra(Distant)Graphocephala spp Hortensiasimnilis (Walker) Exitianusobscurinervis (Stal)Balclutha spp Draeculacephalaclypeata Osborn Cicadulinabipunctella (Matsumura)Cymoninus turaensis(Paiva) Ninus insigniStal Pachybrachiusnenosus Horvath Caenoblissuspilosus Barber Blissus leucoptens (Say) Blissus spp Dlmnorphoptentscornutus novaeguineae GhauriDimorphopterus similis Slater Paroniuspiratoides (Costa)Pangaeus spp Aethus indicus (Westwood)Geotomnus pygmaeus (Dallas) Stibaropus inolginus Schiodte Cyrornenusbergi FroeschnerCyrtomnenus ciliatus (Palisot de Beauvois) Cyrtonenuscrassus Walker Tominetus spp

Riptortuslinearis (Fabricius) Riptorts spp Leptocorisaacuta (Thunberg)Leptocorisabiguttata (Walker)LeptocorisapalawanensisAhmad Leptocorisaoratorius(Fabricius)Leptocorisachinensis Dallas LeprocorisasolomonensisAhmad Leptocorisaspp Stenocorissouthiwoodi Ahmrad Stenocoris claviformis Ahmad MirperustorridusWestwood aetus trigonus (Thunberg)Alkindus atratusDistant ScaptocorisdivergensFroescher Scaptocoris castaneusPertyScotinopharatarsalis (Vollenhoven) Scotinopharascotti Horvath Eysarcoris (=Stollia) ventralis (Westwood) Dolycorisindicus Stal Macrinajuvenca Burmeister Tanda antiguensis (Westwood) Tantiageifi Schout Tantia perditor (Fabricius) Pygotnenidavaripennis (Westwood) Menida sppNezara viridula (Linnaeus)

Oebaluspoecilus (Dallas) Oebalusypsilon-gnseus (de Geer)Oebalusgrisescens (Sailer) Tibracea limbativentris (Stal)

Aspavia armigera(Fabricius)Diploysfissa Erichson Acrosternum marginatum (Palisot de Beauvois) Acrosternum sppFrankliniellarodeos Pergande Breginatothrips enustos Watson Stenchaetothrips(=Baliothrips)biformis

(Bagnall) (=(Jlloethripsoryzae (Williams)HaplothripspriesnerianusBagnall

Distribution

Asia Africa Asia Latin America Latin America Latin America Latin America Latin America Asia Asia Asia Asia Asia Latin America Asia Asia Africa Asia Latin America Asia Asia Asia Latin America Latin America Latin America Latin America

Asia Africa Asia Asia Asia Asia Asia Asia Asia Africa Africa Africa Asia Latin America Latin America Latin America Asia Asia Asia Asia Africa Latin America Africa Latin America Asia Asia Asia Africa

Latin America Latin America Latin America Latin America

Africa Africa Latin America Latin America Latin America Latin America Asia

Asia

Reference

59 241 217 59 241 6166 61 66 81 150 6166 66 81150 b 178 178 30 83 42 81150 203 30 80 62 178 195 178 178 178 203 203 203 203

55 178 12 72 78 120 214 214 b 59 120178114 178 54 636 82 12 6 12 82 178 81 150 60 is 195 59 120 178 78 36 186 36 186 b 55 12 30 54 55 78 82 120 61 66 81195 61 65 66 195 211 150 195 61 66 81 150 219 233 6 12 82 6 82 186 186 61 66 68 213235

178

7 IRPS No 123 January 1987

Table I continued

Order Family Species Distribution Reference

toleoptera Scarabacidae leteroligus (=Aphonoprocts) 15elcs Billberg Africa 36 Dynastinac (=Ileteroligusmees robustui Prell)

Alissonotum pauper Burnieiser Asia 82 Alissonotuon simile Arrow Asia 209 Dscinetusdubibs (Olivie) Latin America 28 195 233 Devscinetusgagates Burmncster Latin America 20 195 Dyscinctt spp Latin America 60 Eutheola Inimilis Bturmcistr Latin America 20 28 8 1 195 203

233Eutheola identarr(Biurnciter) Latin America 121 203 Heteron)ciwv aw rsonJack Africa 82 Hcreron)chusbtuterculatus Kolbe Africa 82 tleteronichtslicas (Klug) Africa 82 fhteron)chus mosamhicus Peringuey Africa 4 6 48 82 (=Veteronychus orzae Britton)

leteronychts lioderes Redtnbacher Asia 77 82 207 ltcteronchusarator (Fabricius) Africa 82 tleterontchusplebepus (Klug) A frica 57 82Ileteron chus pseuocongonnsis Ferreire Africa 32 Ileteronvchns rogiJrons Fairmaire Africa 82 Heteron chus nrsticus niger (K lug) Africa 82 Heteronvchus spp Africa 17 Lachnosterna longipennis (Blanchard) Asia 82 Lachnosternaspp Asia 120 Ligvros fossator Burmeister Latin America 82 Ligyrus nasutus Burmeister Latin America 203 Ligyrus (=Scarabaeus)ebenus (de Geer) Latin America 20 195

Dynastinac Ligtvros huonilis Burmeister Latin America 60 82 Aaladera castanea (Arrow) Asia 82 Maladera orientalis Motschulsky) Asia 82 Malaaerajaponica Motschulsky Asia 82 Phvlognathusdicynsius (Fabricius) Asia 82

Melolonthinae Exopholis h)ypoletca (Wiedemann) Asia 117 Lerlinosterna[= lolotrichia] consanguinea Asia 122

(Blanchard) Leucopholis rorida (Fabricius) Asia 82 117 Lepidiota blanchardi Dalla Torre Asia 76 82 Leucopholis irrorata (Chevrolat) Asia 76 134 Scrica internpta Walker Asia 76 Holotrichialonipennis (Blanchard) Asia 122 Holotrichia serrata Fabricius Asia 122 Holotrichia leucoph thabna (Wiedemann) Asia 82 Holotrichia seticollis Moser Asia 77 78 Holotrichia pnindanaoana Brenske Asia b Stenoeres spp Latin America 71 195 Atoenius spp Latin America Asia 203 Ph luplg aequata (Bates) Latin America 203 Piyvilophagacaraga Saylor Latin America 203 Phyllophagachiriquina (Bates) Latin America 203 Phyllophagadas)poda (Bates) Latin America 203 Phyllophagaclegans Saylor Latin Ameica 203 Phyllophagahondura Saylor Latin America 203 Phyllophagalatipes (Bates) Latin America 203 Phyllophagamontriesi (Blanchard) Latin America 203 Phyllophaga pamisetis (Bates) Latin America 203 Ph vllophaga sanfosicola Saylor Latin America 203 Phllophagasetifera (Burmeister) Latin America 203 Phyllophaga (=tlolotrichia)helleri Breuske Asia 82 117 159 Phvllophagaspp Latin America 81 150 White grub unspecified Asia 84

Rutclinae Lagochile trigona (Ilost) Latin America 82 Anomala dinidiata var barhata Burmeister Asia 77 78 82 Anomala huteralis(Burmeister) Asia 76 Anoimala lurida (Blanchard) Asia 76 Anomala sulcatula (Burmeister) Asia 76 Anontala varians (Olivier) Asia 82 Anomala antigua (Gyllenhal) Asia 82 Anoinala palida (Fabricius) Asia 82

8 IRPS No 123 January 1987

Table 1 continued

Order Family

Tenebrionidae

Elateridae

Languriidae Chrysomelidae

Chrysomelidae

Coccinellidae Curculionidae

Lepidoptera Lymantriidae

Amatidae

Arctiidae

Limacodidae Eupterotidac Noctuidac

Species

Anomaa volita (Blanchard) Adoretus caliginosus Burmeister Adoretus comprcssus (Weber) Papuana huhneri (Fairmaire) Papuana incrmnis Prell Popillia capricollis Hlope Gonocephalu depressur (Fabricius)Gonocephahm acuiangulum (Fairmaire)Gonocephalum sinphx (Fabricius) Epitragus sallei Champion Anaedus ptnctatissimnts ChampionAeolus spp4griotes spp Agriotes mancus (Say) Conoderts spp Aeoloderma brachmana (Candeze)Wireworni unspecified Anadastus filijf rmis (Fabricius)Dicladispa viridicanca(Kraat z)Trichispasericea (Guerin)Oediopalpastervalis (Weise) Oediopalpa guerini (Italy) Chaetocnema cvlindrica (Baly)Chactocnema denticulata S tephens Chaetocnemabasalis (Baly)Aulacophora sp nr similis Olivier Monolepta caripemne Ialy Monolepta bilasciata (Ilornstedt) Monolepta signata Olivier Cerotoma airojcsciata J acoby Diabrotica adelpha Ilarold Diabrotica balteata Leconte Diabrotica graminLa ltaly Diabrotica speciosa (Germar) Diabrotica limitafa quindecinmunctata(Germar)Diabrotica melanocephala (Fabricius)Diabrotica spp Altica (= Fondia)madagascariensis(Allard)Altica spp Epitrix cuctmeris (Harris) Epitrix spp Oulenjo or pae(Kuwayama) Disonchq (= Donacia) spp Flea beetle unspecified Chnootriba (= htIilachna)sinilis assimilis Mulsant Atactogaster iduriens Walker Hypomeces squtamosus (Fabricius) Tan vinecus discoidalis GyllenhalPhaulosoinusmusculinus (= mnus) Csiki Neobaridia anplitarsis CaseyPsalis pennatula (Fabricius) Laeliasufihsa (Walker) Euproctis irgtncula (Walker)Euproctis minor (Snellen)Euproctis xanthorrhoea (Kollar)Amata sp

Diacrisia obliqua (Walker) Creatonotusgangis Linnacus Latoia (= Parasa) bicolor (Walker)Nisaga simplex Walker Sesamia botanephaga Tams amp Bowden Sesamnia nonagroides botanephagaTfums et Bowden Sesamia calamistis llampson Sesamia inferens (Walker) Spodoptera eridania (Cramer)Spodoptera mauritia acron)ctoides Guenee Spodoptera frugiperda ( ESmith)

Distribution Reference

Asia 82 Asia 82 Asia 82 Asia 82 Asia 82 Asia 78 Asia 82 117 178 Asia 82 117 Africa 82 Latin America 203 Latin America 203 Latin America 195 Latin America 195 Latin America 203 Latin America 195 Asia h Asia Latin America 60 73 117 Asia 200 Africa 36 Africa II Latin America 6166 68 Latin America 6166 68 Asia 225 Latin America 61 66 68 81

Asia 178 Asia 55 Asia 55 Asia h Asia 55 Latin America 203 Latin Amefia 203 Latin America 203 Latin America 203 Latin America 6166 68 Latin Amcrie 195 Latin Amw 195 Latin Aierwca 61 81 Africa 57 Asia 178 Latin Americ 203 Latin America 81 Asia 82 Latin America 195 Asia 120 Africa 6 11 Asia 185 Asia 120 Africa 82 Africa 82 Latin America 66 68 195 233 Asia 117 Asia 117 Asia 117 Asia 117 Asia 117 Asia A S Pradhan

unpubl dataAsia 82 Asia 82 Asia 82 Asia 182 Africa 12 Africa 6 36 Africa 6 8 11 12 36 Asia 59 82 84240 Latin America 150 Asia -0 59 196 Latin America 28 61 66 68 81 150

152195203219233

9 IRPS No 123 January 1987

Table I continued

Order Family Species Distribution Reference

Spodoptera exigua (Hubner) Spodoptera exem pta (Walker) Spodoptera ornithogalli (Guenee)

Africa Africa Latin America

32 217 32 33 36217 150

Mythimna loreyi (Duponchel) Asia 30 Mythimna separata (Walker) Mythimna (- Pseudaletia)latifascia [= adultera

Asia Latin America

178 195

(Schaus)] (Walker)Mythimna (=Pseudaletia) sequax (Fabriclus) Latin America 68 Mythimna rosellinea (Walker) Asia b Mythimna yu (Guenee) Asia b Mocisfrugalis (Fabricius) Asia 30 117 Mocis latipes (Guenee) Latin America 66 68 149 150 203

Platysenta (=Spodoptera) compta (Walker) Asia 219233A Barrion unpubl data

Agrotis ipsilon (Hufnagel) Latin America Africa

33 68 81 150 1)5 203

Agrotis spp Latin America 60 Achaeajanata (Linnaeus) EldanasaccharinaWalker

Asia Africa

117 36

Armyworm unspecified Asia 72 84 Pyralidae Cnaphalocrocis medinalis (Guenee)

Marasmia bilinealisHampson Asia Africa

30 4559 72 84233 36

Marasmia trapezalis(Guenee) Africa 32 217 Marasmia (=Sumia) exigua (Butler) Asia 220 Marasnianmralis (Walker) Asia b Marasiniapatnalis Bradley Marasiniaspp

Asia Asia

b b

Leaf roller unspecified M2liarphaseparatellaRagont

Asia Africa

72 12 57

Maliarphasp Asia b Chilo auricilius Dudgeon Asia b Chilo partellus (Swinhoe) Chilo polychrysus (Meyrick)

Africa Asia

12 92 59 72

Qhilo diffusilineus J de Joannis Africa 6 1229 Chilosuppressalis(Walker)Cibozacconius Blezzinski

Asia Africa

59 72 6 11 12

Cravubus spp Latin America 203 Acigona loftini (Dyar) Latin America 203 Acigona chrysographella(Kollar)Scirpophagaincertulas(Walker)

Asia Asia

82 59 72

Scirpophagainnotata (Walker) Asia 59 200 Scirpophaganiuella (Fabricius) Asia b Elasinopalpuslignosellus (Zeller) Latin America 28 60 61 66 81 150

Diatraeasaccharalis(Fabricius) Latin America 162195203219233 28 66 81 111 150 195

Rupela albincllaCramer Latin America 111

Gelechlidae

Stem borer unspecified Undetermined stem borer nr Maliarpha BrachniaarctraeaMayr

Asia Asia Asia

72 b 55

Brachmi spp Africa 32 Hesperiidae Parnaraguttata Bremer et Grey Asia b

Parnaranaso Fabricius Asia 55 Pelopidasagnaagna (Moore) Pelopidasmathias (Fabricius)

Asia Asia Africa

164 36 155217

Pelopidasconjuncta conjuncta (Herrich-Schaffer) Asia b Borbo fanta Evans Africa 6

Satyridac Melanitisleda ismene Cramer Asia 84213 Mycalesis asophisHew Asia 30

Diptera Cecldomyiidae Mycalesis spp Orseoliaoryzae (Wood-Masn)

Asia Asia

b 72 88 131 181

Orseoliaoryzivora Harris et Gagne Africa 61132218 Chloropidae Steleocerellus (=Mepachymens) ensifer (Thomson)

Oscinella spp Asia Asia

205 205

Gaurax sppChloropsoryzae Matsuntura (= kuwonae Aldrich)

Asia Asia

205 82

Mucldae Atherigonaoryzae Malloch Asia 30 82 117130 Atherigona exigua Stein Asia 82 117 178

10 IRPS No 123 January 1987

Table I continued Order Family

Diopsidae

Agronyzidae

lHymenoptera Formicidae

Species

Atherigona oritentalisSchiner Atherigona indica MallochDiopsis longicornis(= thoracica)Macqurt Diopsis spp Pseudonapnyza asiatica Spencer Pseudonapomyza spicata (Malloch) Agronoza orvzae MunakataAcromyren landoltibalzani(Emery) Acromyrmex landoltifracticornis(Forel)Acromyrnex heyeri Forel)Atta bisphaericaFore Atta capiguaraGoncalves Atta laeLigata (F Smi h1) Atta opacicepsBorgmeierSolenopsisgeminata (abricius)Pheidole sp Pheidolegeton diversus Jerdon Monomorium pharaonis (Linnaeus) Ants unspecified

Distribution Reference

Asia 82 Asia Africa 82205Africa 11 1248 Africa 217 Asia 213 Asia h Asia bLatin America 6168 195 Latin America 61 195 Latin America 61 68 195 Latin America 61 68 195 Latin America 14 61 68 195Latin America 61 68 195 Latin America 61 195 Asia b Asia b Asia 1) Asia b Asia 120

aG orlentalls formerly recorded in Asia as G africana thPhilippines reported in this study CSubterranean root feeders

subterranean nests Foraging ants will feed on germinatedseeds only when the supply of ungerminated seed is low However these ant specialists feed on a wide varietyplants and thus help control grassy weeds (39) This dual

of

pest benefit role extends from tending ants which protecthoneydew-excreting pest species including root aphids and mealybugs (workers repel potential predators and even selectively kill parasitized aphids) to predators of leaf- and stem-feeding rice pests such as leaffolders armywormswhite grubs and stem borers (1 17)

The upland rice environment is particularly well suited to ants because of the lack of flooding Frequent tillage perpetuates grasses and granivorous species that are highlyadapted to constructing new nests in recently tilled fieldsAnt nests however are relatively shallow and tilling with a moldboard plow can destroy them Also upland ricefields are uually small and isolated and a given area in a smallfield has more field borders than it has in a larger fieldallowing high rates of infiltration from more sedentaryspecies such as Plieidole that do not colonize tilled fields but nest in soil under shrubs and trees along field bordersPheidole workers forage into fields from borders and store seed in their nests In slash-and-burn or other no-tillage rice cultures nests would be preserved in the field and possiblylead to high rates of seed predation However granivorous species do not prefer forest habitats

Damage is characterized by reduced usually patchy plantstand (120) Seed-thieving ants greatly prefer rice but they can be selective regarding plants they collect and store Theyharvest more seed than they can consume so patches of germip ing surplus seed (such as Rottboellia) will sproutfrom their nests during the rainy season

In Latin America leaf-cutting ants (Aita spp and Acromyrmex spp) occur only in upland fields and defoliate

young rice plants (47 68) They take leaves to fungal gardensin underground nests where fungi predigest the plant material

Termites Subterranean termites of the family Termitidae which lack symbiotic protozoa to help digest plants are the most frequently mentioned termite pests of upland rice (1185 120) This family cultures fungi in special undergroundcells --- fungal combs made of half-digested plant material (117) The fungi inoculated onto the combs break down plant material into food that termits can digest The termites then consume the fungal combs These grasslandtermites build nests below the plow layer in uplandricefields The colonies attack living plants only when dead plant material is not available They will attack a droughtshystressed crop but prefer older plants having greater cellulose content They will damage newly planted crops where clean culture has removed vegetation

Infestations are worse in deep light-textured soils with low moisture content The first sign of damage is yellowingof older leaves (195) Termites feed on roots (the plantyellows then wilts and finally collapses) and germinatingseed (loss of stand) or move above ground at night to cutseedlings at ground level which they cover with soil for later consumption Soft-bodied termites are highly sensitive to desiccation They live within a self-contained system of airtight chambers and build surface tunnels linei with nudand body secretions (carton) to maintain more than 90 relative humidity Termites can be located by their tunnels

Termites apparently can withstand limited submergence (157) and can be a pest of dry lowland seedbeds (117)perhaps by having nests below perched water tables

Termites appear to do more damage in Latin America where large ricefields are plowed with tractors deprivingtermites of preferred vegetation and forcing them to live on

rice plants In Asia upland ricefields are typically small with Iprge field border areas (ecotones) where termites can forage for preferred food other than growing rice plants All grassland termites do not feed on rice In an upland area in Batangas Province Philippines ricefields are commonly infested by Hosiititernesluzonensis (Oshima) however this termite does not damage rice In the same province though Mrroternesgivus(Hagen)feeds on rice seedlings (178)

In Africa termites may damage rice even though fields are small and surrounded by perennial vegetation These termite species perhaps prefer rice over other hosts

Mole crickets Nymphs and adults are nocturnal and burrow through soil feeding on roots of a wide variety of plants or forage above ground as predators of insects Adults live 3-5 mo and are even cannibalistic Mole crickets prefer low-lyin moist upland soils with high organic matter (II 81) In the lowlands they inhabit rice bunds or nonflooded fields Adults are highly mobile and can leave a flooded field to locate a more suitable habitat Losses show as wilted plants (from root feeding) or reduced stands (120) Damage is more common near field borders where mole crickets relocate after tillage (126)

Fiehl crickets Nymphs and adults of field crickets have similar nocturnal habits and damage rice as much as mole crickets do Piles of weeds removed from fields attract them They make subterranean nests and tunnel through the soil to feed on roots Some species prefer seed to roots Others feed at the base of stems causing deadhearts

Tunnel entrances surrounded by excavated soil are easily seen Young plants are cut at ground level and stored in underground cell Field crickets tre highly polyphagous as are all soil pests (117) Sachan et al (198) reported that field crickets prefer maize to upland rice Like mole crickets field crickets cannot survive in standing water and are therefore more prevalent in upland fields

Whitegrubsandhackbeetles Scarab beetles that feed on living roots as larvae but not as adults are called white grubs or chafers (Melolonthinae Rutellinae) and those that feed as adults but not as larvae on plant crowns are called black beetles (Dynastinac) In the tropics chafers or white grubs have a I-yr life cycle starting with adults emerging from the soil about I mo after the first downpour of the rainy season (134 159) They lay eggs at the same time farmers sow upland rice The rice passes its most susceptible stage and damage is mostly avoided when the white grubs are small After several months the long-lived larvae are large enough for two or three to denude the root system of mature rice (223) This intensity of damage is rare but wilting occurs when root loss is combined with drought stress

The larvae need damp (not saturated or dry) soil to survive and pass the unfavorable dry season 05-2 m underground in aestivation in the moist soil The first heavy rains of the season (20-30 amd) stimulate the grubs to resume activity and after several weeks they develop into

IRPS No 123 January 1987 11

pupae and adults and eventually dig their way to the soil surface and fly to nearby trees to seek food and mates The adults of most white grub species feed only on trees but in some species notably Eutheolahumilis the adults defoliate and cut off stems at ground level (28) Putheola hidentata damages sugarcane fields in Venezuela (121) Floods kill white grubs (208) Grasslands can support large populations and therefore white grubs can be more abundant in newly planted upland ricefields that were previously fallow (81)

Black beetles live up to a year adults can attack a crop at any stage by burrowing into the soil (127) and feeding at the base of stems to cause whiteheads (207) Adults emerge with the early rains and are normally more abundant and do more damage on a young rice crop with small root systems (38 81)

The larvae feed only on dead organic matter in dryland fields and do not attack rice Adults arc highly mobile and although sensitive to flooding can invade a field soon after it drains (115)

Root aplhids Several genera of migratory aphids feed on the roots of upland rice (249) Populations build up more in light-textured soils with high percolation rates Damage has also been reported from dry seed heds of lowland rice (52) Tending ants -- Pheidhle Cren aster Tetramoriuw lsius Tapinoma -- are necessary for root aphids to multiply Aiits harbor aphids in their nests over winter or during unfavorable periods and relocate them on rice plants digging tunnels along the root systems to allow these softshybodied insects to penetrate the soil Most intensive studies on aphid life histories have been done in temperate environments (158 222 229) Root aphids fly to rice plants at the beginning of the rice season and pass through several generations continually developing winged forms to relocate on alternate grassy hosts until winter when they move to perennial hosts usually trees Yield loss occurs mainly through reduced tillering but infested plants become yellow and stunted and in extreme cases wilt (227) Aphid populations build up gradually so damage usually begins in the late vegetative and reproductive growth stages

Root-feeding ineal tugs More than half of the mealybug species or rice feed on roots (245) Being soft bodied mealybugs are not adapted to living underground but they survive because tending ants (the same species that tend root aphids) dig burrows for them and move them from plant to plant Well-drained soils ihelp mcalybugs survive Although six species are recorded as subterranean root pests of rice we found no report citing damage Root-feeding mealybugs are probably more prevalent on perennial grasses than on short-lived annual rice

The most common rice mealyb ig Brevennia rehi is a foliar feeder favored by lack ol rain apparently it is responsible for high yield loss in Northeast India and Bangladesh (N Panda pers comm as stated by IRRI [1061) This report needs confirmation Mealybug damage was only briefly mentioned in another report (156)

12 IRPS No 123 January 1987

Root weevils A few species of upland weevils (IHporneces Donacia Atactogaster) attack rice roots as larvae and feed on stems and leaves as adults Damage ispatchy The larvae are not adapted to submergence and are true upland soil weevils isopposed to the so-called water weevils They rarely become sufficiently abundant to cause damage Adults on rice foliage or the soil surface show that the roots are being attacked by the grubs Root weevils are highly polyphagous (185)

Root heeths Soil-inhabiting false wireworms Gono-cephalumt spp are reported in upland ricefields The adults and larvae normally feed on decaying organic matter but land preparation for upland rice removes these sources The larvae feed on seedling roots and nocturnal adults roam on the soil surface to cut off seedlings at ground level (82) This group of ground beetles is highly omnivorous adults even prey on stem borer larvae (183) True wireworms (Elate-ridae) feed on living roots but rice as a host has been reported Infrequently (117) In Latin America the highlypclyphagous larvae of Diabrotica rootworms are also recorded on rice

Root htus Upland rice seedlings can be attacked by the adults anI nlViphs of chinch bugs Blissusand Caenoblissus and bro n bug Scaptocoris which feed on the roots with theirsucking nouthparts (80 83) Three genera of burrower bugs A tms (eotonhus and Stiharopus -are common iil grasslands in the Philippines and may attack the first crop (rice or maize) planted after the dry season Chinch bugs are most abundant after a dry spell These insects are large and can kill seedlings or stunt and reduce tillering on older plants I hey prefer other hosts and rarely damage rice Adults can also feed oii developing rice grains (81)

(utworms Soil-dwelling noctuid larvae hide under-ground during the day to avoid predation by birds and bccone active at night above grour cutting off you ing plants at ground level (hence tile naiie cvtworn) arvaedrag severed plants into hurrows llie larvae have wide host ranges arid pupatc in the soillgrotis ipsilon occurs worldwide and appears to be the most common cutworm pest of upland environments but is more prevalent inlowland rice particularly in dry seedbeds

IFoliar pests Other groups of insects are adapted to upland environments because they pass one growth stage underground either as egg (in heinrietabolous species) or pupa (in holomnetabolous groups)

Rice seedlibig maggots AitJeriota flies occur in non-fhoodcd environments in Asia and Africa (180) Feeding of larvae one per iller causes deadhearts Flies as opposed to moths only secondarily evolved as pliytophagous pestsfroni a saprophagous Origin Athrnonalarvae secondarihv feed onidecaying tillers whereas larvae of stem borer moths actively Iced on living tillers to cause deadlicarts Seedling maggots will die if they cannot sever a tiller on which to feed

as it decays Plants die under heavy attack whereas lighter infestations cause stunting delayed maturity and ragged leaves (236)

Eggs are laid only on actively growing plants (49) aid damage occurs wiihin the first week after crop emergence(170) Larvae pupate more in stems than in soil Seedling maggot attack is highly seasonal (206) normally peaking 2-3 mo after the onset of the rainy season These pests possiblyaestivatc over the dry season Rice isonly one of their many graminaceous crop and weed hosts

The rice stem maggot (hlorops orzae is adapted to temperate upland and lowland rice growing regions of Asia In Japan and China upland rice grows in mountainous regions a habitat of this chloropid fly The larvae tunnel into tilestems The stem maggot is more damaging during the seedling stage larvac hibernate in grasses

File several dipterans reported as stem maggots in dryshyseeded lowland seedlings inIndia (205) could presumably attack upland rio

Leaf urinrs Icaf miners are common on wheat and maize but are rarely reported on upland rice (213) Three leaf miners are more prevalent on upland than lowland rice inthe Philippines Pvudonalomy a asicanh Vficataand lIAgromrza orzae attack at the seedling stage The larvae develop inside the tunnels they construct in the parenchyma tissue P Vpicata and Aorvzac prefer maizeEehtsine and other grasses to rice spicatacan become abundant on wheat

Leaf beetles Chrysomelid and coccinellid eaf bcetles scrape or otherwise remove leaf tissue as larvae or adults or both This damage accelerates desiccation of plants Tile larvae of sonic species are root feeders Flea beetles Epitrix and (lraetocnetna and rootworms l)iahroticaspp feed on rice foliage only as adults Their root-feeding larvae have hosts other than rice The adults and larvae of the hispa-like Oediolmlpa spp beetles of Brazil and (hmootriha (=EPfishyachra) beetle of Africa feed on upland rice leaves Oealioshypapa spp larvae are leaf miners (mootriha is also pshyvalent in lowland environments (11) Leaf beetles prefer vegetative rice (81)

The rice hispa Dicladipaarnigera is not reported on upland rice and prcers more aquatic habitats In Central Africa however I)cladispm viridicyaiea attacks uplandrice in the vegetative stage as well as lowland seedbeds 7richipasericea is reported on upland rice in West Africa and Madagascar (57)

The rice leaf beetle 1d1ea 01izae riiuech like Chioropsor|rzae is adapted to temperate regions of Ashi and is usually a pest on lowland rice but seldom oi inpland rice in the mountains (82) The adult hibernates on grasses

Armmworms Species of grassland-adapted SpoptloeraPlatysemta and A ithimna larvae create widespread epishydemics They can defoliate ricefields generally in patchesfrom early vegetative growth to harvest (41) Armywornms pupate inthe soil and highly favor upland rice (34) Adults

can disperse long distances to colonize even remote upland areas They become more abundant than their natural enemies particularly after favorable rains following a prolonged drought (196) The drought kills the armyworm and natural enemies (mainly parasites) but the armyworm can recolonize rapidly

The natural increase insoil fertility from mineralization of soil N over the drought period promotes luxuriant plant growth to foster a rapid buildup of these highly fecund species Weeding rice increases tilelikelihood of damage The parasites normally return to control the armyworms but not until after serious damage

Outbreaks on upland rice have been reported in Ghana (3) inzibar (38) Central Africa (36) lPanama (165) Brazil (28) Malaysia (196) and India (167) Defoliation may be severe often leaving only the base of stems Armyworm larvae hide during the day under litter or in soil cracks At night they ascend plants to feed Larvae pupate in the soil

Thrips Thrips frequent upland rice but damage is less than in the lowlands Thrips larvae and adults feed on leaf blades by rasping causing leaf rolling and stippling Thrips prefer vegetative stage rice and their survival increases during drought (heavy rain washes them from foliage) although drought stress usually outweighs thrips damage Drought and thrips cause leaf rolling in upland rice (235) but thrips have been overlooked perhaps becaLse of their small size Thrips are most numerous indry season irrigated rice The combination of favorable plant growth and lack of rainfall is ideal for thrips In contrast the lack of rainfall in upland rice hurts the crop and eventually the thrips whose numbers must decline

In [atin America Frankinilla rodeos attacks panicles before they emerge from the boot calusing sterile grains

Stem hug~s Black bugs are the most common group of rice stem-feeding -lemiptera in Asia The most frequently mentioned species ----Scotinophara coarctaia and Shrida - prefer aquatic habitats and therefore could not occur in upland rice as sometimes reported Upland species in the same gerurs - notably S tarsalis and S scotti -- are morphologically similar and remove sap from stems of grasses and upland rice The upland species are notably more abundant in upland rice planted near forests

In Latin America tile Tibraca ima-large pentatomid tiventris causes deadhearts and whitehcads on upland rice Both adults and nymphs remove sap from internodes of plants more than 3 wk old Tillers are killed by mechanical damage and by the entry of secondary microbial infection This damage is often confused with that of lepidopterous stem borers (233) Eggs are laid on the leaves and stems Nymphs are gregarious This species can also be found on irrigated rice Drought reduces population buildup

Stiem borers Lepidopterous and dipterous stem borers are widely recorded on upland rice but cultural practices and alternate hosts determine abtndance Except for Elasmolmll)uS Iinosellts numbers usually build up toward

IRPS No 123 January 1987 13

the end of the crop growth cycle Therefore late-maturing varieties staggered planting and lack offloodingfavorstem borer buildup Many stem borer species can lie dormant in the stUbbleafter harvest and theirsurvival is encouraged by panicle harvesting (leaving tall stems) and lack of tillage after harvest

Stem borers disperse and lay many eggs Many species are polyphagous however the monophagous MaliarpiasepashyratellaDiopsis Ivmgicornis Scirpophaga incertlas and S innotata can also be abundant on upland rice (hilo Suplressalisdevelops fasterand becomes la rger when reared on upland than on lowland rice (226)

The lesser corn stalk borer lasmopal tt Ittoselhs is semisubterranean and is perhaps the stern borer most adapted to upland rice(129) Highly polyphagous it infests maize peanut and cowpea It attacks seedling rice (one larva can kill tip to four plants before maximum tillering) (195) (hiv and Maliarphastem borers prefer older plants But Scirpophagaspp will attack young plants even in the nursery lasn opalls larvae tunnel into the stems at or below the soil level mausing dcadhcarts Larvae are not found in their turnnels inside sterns beca use vhei disturbed they retreat into cases made from soil particles bound by silk The cases areattached to the tunnel entrance (143) This behavior probably evolved as protection from natural enemies Larvae pupate in the soil The lesser corn stalk borer is more abundant during drought damage is often confused with that of drought (233) This species prefers sandy soils arid its distribution within a field may relate to soil texture (143)

Diatraeasaccharalis aside from the lesser corn stalk borer is frequently a stem borer pest in Latin America According to Teran (233) rice and maize are more preferred hosts than sugarcaine its namesake

Atcitonacrvsograpla is a less known upland rice stem borer in Asia with habits similar to Easinopall s (82) because it is not found in lowlands Acutra has grassy weed alternate hosts As a moth it is often confused with the dark-headed stern borer(hi polur(standstriped stern borer (hi sup)lpsrcais

Other common upland rice stem borers are also reported in the lowlands The dominant species in upland rice vary greatly with location In Uttar lPradesh India (78) and in Japan (123) the pink stem borer Sesamia htfirens is more prevalent hIKenya the principal rice stem borer is Chi partelus

(92) The lowland African sten borers Aialiarpha sepashyratellaDiopsis vngicornis and Sesamia calatnisnis are less abundant Oimupland than on lowland rice(217) but perhaps cause greater yield loss because plant injury is greater if combined with drought css (92) The stalk-eyed stern borer D)ivpsis is more prevalent in wet habitats (6 1I217)

In Central and West Africa Maliarpha and Sesamia calanistis are the dominant stern borers of upland rice (8 9 10 II) (lilo dihlfilineui and Chio partellus inhabit the

14 IRPS No 123 January 1987

upland savannas whereas Chilo zacconio prefers the lowlands (II) Aaliarpha is most abundant in the mangrove swamp habitats (91) coastal regions (16) or rain forest zones (II) of Africa and appears to be the homologue ofScirlpophaga incertulas The yellow stem borr can also be prevalent in upland rice (120 167) The sugarcane borer Eldanasaccharinais so far only a potential pest of uplandrice (91) despite Grist and evers (82) warning

Grasshotl)ers andI locitas Short-horned grasshoppers and locusts -- Locvta Patanrga Schistocerca - layeggs in the soil inhabit grasslands and develop into swarms to seriously damage upland rice from time to time (173) Largenumbers can destroy a field leaving only the stubble In low cmuoroer s giassnoppeirs an cut panicles (173) Farmers may not plant if locust swarms are imminent Prolonged dryweather followed by favorable rain favors development of large swarms (173) Usually breeding in di) grasslands theydisperse toattack crops including upland rice(191) Eggsaresusceptible to desiccation d uring drough t and toinundation during seasonal rains With favorable soil moisture over consecutive seasons natural enemies ca nnot prevent a largepopulation increase leading to migratory swarms (64)

Most grasshoppers on rice are adapted to uplands (107)but lowland rice grasshoppers iervgl(rlhusspp and Ora spp occur occasionally on upland rice (2 82) They lay eggson rice foliage Nymphs are scmiaquatic

The gryllid Lsctrtus coucinnus is a pest of lowland and deep water rice in Bangladesh and Thailand In the Philippines it damages upland rice grown in isolated pockets near lakes The gryllid nymphs and ad ults feed on the central portions of folrage leasing only the midrib and leaf nargins

Leal ler Pvralid moths whose larvae fold leaves to make a feeding shelter occur worldwide (217) but arc appareritly most common in Asia Leaffolde rsattacking rice in Asia are a complex of species (26) (Ctalhahcrcisinedinalis the best-known rice Ia flolde r in Asia may prefer grassy weeds over rice (86) It is a late colonizer of uplandrice U1arasmja eviua a lowland species (220) nreflers rice to grass specres (86) and colonizes upland rice during il early growth stages llarasnria patnalis is the most corn-rnonly encountered species in upland rice in the Philippines Perennial grasses are their alternate hosts and they can survive year-round in upland rice environments The leaf-folders 11arasmia ruralisand Alarasmia spp are least aburndant Dormancy is unknown in leaffolders Cnaplha-ocrocis medhnalis however is known as a migrant (90)Leaffolders remove photosynthetic tissue attack during the flag leaf stage is particularly injurious Popurlations are normally held in check by natural enemies but leaffolders become parlicularly abundant in conditions of high plantfertility and shade

Butterflies Skippers PelopidasInathiasand Borhofanta are more prevalent in African lowland than in uplandhabitats (6) but extensive studies in Japan and China

showed that Pelopidas mathias prefers an upland habitat (155) Parnaragutata seasonally migrates between lowland and upland areas and is equally abundant in both environshyments (164)

The green horned caterpillar Melanitis leda ismene is more abundant in lowlands (212) but our experience in the Philippines suggests that Pelopidas niathiasand 11elanifis leda ismene are often more abundant in uplands than in lowlands Bamboo iaunbuiva spp annually sustains P mathiaspopulations in the Philippine uplands Adults feed on nectar from flowers and migrate to upland rice during the late vegetative stage

iPi erlled h i-1 1d dill L tIC i 7-0 l(become numerous probably because of high egg predationEgg and larval parasitization are often low Butterflies can readily disperse long distances to seek renlote uplandricefields However they have low biotic potential and even it predation rates are low the rarely become abundant Upland environments with their diverse microhabitats can provide more favorable sites than lowland rice plains

tutterfly pests of rice have broad host ranges and become dormant Juring urnfavorable times of the year

IolhagrotslaidopwraThe larvae of many polyshyphagous moths are reported to defoliate upland rice tussock moths lsaispelnnatula Iawlia sUifita luproctisirgtwltaE mnorand P vathorrhoeaslug caterpillar

litotiahicolorhairy caterpillars Nisaga sinp vandA mata spp and wooly bears l)iacrisiaohliqtia and Crvatonotts gangis Many of these polyphgous larvac migrate and are transients in rice including HelhWhis arm iera and 1t zea highly pol phagous species (82)

Gall nidges The Asian rice gall midge Orseoliaorrzae (88 131) and its African counterpart 0 ortzivora(6 11 12 32217) prelerlowland to upland rice Gall midges in rainfed areas are prevalent only during tle rainy season and dispersepoorly Their alternate hosts (wild rices LeersiaIschaetnuan and Pasahmt)are mostlyaquatic grasses and are less likelyto be near upland rice areas In an area of mixed uplandfloating and rainfed lowland rice cultures in northern Thailand upland rice which is ilanted a month earlier was attacked firstbut subsequcent populations were much lower than in rainfed lowla id or floating rice Low numbers of gallmidges passed the dry season orr perennial weeds such as Paspahim (88)

Smot (hid(s Reports of aphids attacking upland rice plants are few (249) Without maize the maize leaf aphidRho alosihhunm maidis attacks upland rice in the Philipshypines The plum peach aphid IIrsteroneuraselariae was reported in Sierra Leerne toinfest leavesand tnriperied grain(7) Siphaglrctriaecaused transverse linrea r necrotic striae iir rice leaves in Italy (172) I-lea v rains held populations in check Rice is not a preferred host for shoot aphids

Leafhoper andI )anthol)pers Cicadellids and delshyphacidsdispersecasilyand readilycolonie upland ricefrom nearby lowland areas A study in Sarawak (241) showed

IRPS No 123 January 1987 15

similar species in upland and lowland rice Nephoiettix The principal vectors of rice virus diseases generally have species dominated The most prevalent species accounting narrow host ranges and alternate hosts in the grass family for 90-95 of the sweep net samples wre N virescens N are mostly annuals dying in the dr3 season Some virus umropictus Nilaparvia hgens Soatellafiircifera Cfana vectors such as BPH can disperse lontg distances but few do

sectraand Recilia dorsalisLowlands had higher popula- Th restriction of upland rice and weed hosts to the rainy tions Katanyukul and Chandaratat (120) monitored hopper season the small area planted to upland rice and the hopper numbers in upland and lowland fields of northern Thaiiand vectors having narrow host ranges all work against virus and found similar results A comparative study of plant- diseases in upland rice hoppers in Fiji showed more brown planthoppe BPH) and In Latin America Sogctdhs o(rzicola and S cuhanus whitebacked planthopper (WBPH) in wetland rice than in transmit hoja blanca virus disease but reports on upland lowland rice and hopperburn was prevalent in lowland rice are rare except in the favored upland areas of Colombia areas with standing water (89) Peru and Venezuela (35 42) S rricolaocurs in Brazil

Hopperburn in upland ricefields in Asia is rare (84) In on upland rice but hojaI blanca has not been reported (67) Batangas Philippines isolated fields of upland rice become S orzicola has a wide host ramgc (10) similar to WBPI- in hopperburned infrequently from WBPH A report from Fiji Asia WBPH however is not known to transmit any virus cites hopperburn from BPH and WBPH (250) In these two disease situations upland rice was grown on highly fertic soil and In Africa pale yellow mosaic virus is endemic to swampy N fertilizer was applied Hoppers respond to better nutrition lowland areas where rice ratoons year-round (19) and is Normally upland rice is grown on poor soil and receives no mechanically vectored mainly by hispid beetles A minor fertilizer vector a flea beetle Chaeiocnema spp is an upland rice

In Asia upland rice ras are normally not far from pest and upland ricetields near endemic areas occasionally lowland ricefields-- a source of hoppers Otherwise hopper contract the virus species with greater polyphagy than BPH or Nelhotettiv Spittle bugs Cercopid nymphs producc a protective virescens would be dominant frothy covering that looks like saliva In Latin America

In Africa leathoppers and planthoppers do not become spittle bugs damage upland rice In parts of Brazil where abundant on upland rice (6 II) probably because lowland upland ricefields are surrounded by pastures adults rice is not widely rlanted nearby The only reported migrating from grasses can damage rice severely and cause hopperburn as from Nilapanrata maeander in breeder complete hopperburn in a young rice crop within a week of plots receiving high N rates in Nigeria (98) Hoppers colonization (194 195) Dispersal to rice occurs even if however are a greter problem in Latin America where pastures are verdant and spittle bugs prefer pasture grasses large grassland areas breed polyphagous species (Gra- to rice (168) Nymphs are rarely abundant enough to phocephala spp Hortensia spp lEvitianus ohscurunervis damage rice Regular rainfall favors their development Balclutha spp and Draeculacephala spp) vhich then (233) disperse to cause hopperburn on seedling rice (68) Rosetto et al (195) reported Graphocep7hala spp and Hortensia spp Seed pests to be equally abundant on upland and lowland rice Alydid and pentatomid bugs are seed pests of upland rice

The leafliopper and planthopper epidemics in Asia that worldwide In Asia Leptocorisa sp prevail Their habits caused severe losses from hopperburn and vijus dicases make them particularly suitable for upland rice 1)thev feed over the past several decades occurred principally on on many grasses (and therefore can survive the earl y wet lowland rice Except in a few isolated cases (160) upland season before rice sets its grain) 2)adults are long-ied and rice has been spared from tungro grassy stunt and ragged mobile allowing them to find isolated plantings 0 rice stunt virus diseases vectorcd by green lcafhopper (GLH) (besides seed bugs only birds and rats appear to have shyand BPI] (123) Virus diseases have occurred in upland rice capability) and 3) adults aestivate in forested areas or where lowland rice was nearby Rice dwarf disease however sugarcane fields during the dry sca son when neither rice nor was first recorded on upland rice in the Philippines (187) grassy weeds are present (I199) Rice seed bugs concentrate Virus diseases can be perpetuated only on livir plants or on small-scale upland riceficlds because they can actively insect vectors and no living rice - plans tooned or search them out The prevalence of groves of trees characshyvolunteer - grows year-round in upland arcas excert in a teristic of upland environments ensures nearby aostivation few with 12 mo of rainfall sites keeping the seed bugs close by

The smaller brown planthopper Laodcelphax striatellu Leptocorisae are uncommon in lowland rice plains occurs in upland rice in Japan (18) It spreads black- because 1) their populations are diluted in a sea of rice streaked dwarf and stripe viruses between winter wheat and planted more or less at the same time (they ca n only feed on barley to lowland rice Early planting of lowland rice rice during the milk to hard dough stages) 2) they lack nurseries spreads these viruses but upland ice has remained aestivation sites and 3) weedy fallow areas are limited uninfected The lowands of Asia are dominated by L oratorius

16 IRPS No 123 January 1987

which lays its eggs hign on tile foliage (199) L acuta and L solonlonlesis oviposit at ground level the former on litter and the latter loose on the ground These egg laying habis explain theirtenvironmental prefcrencecs(214) In Japan the postdiapause adults of L chinensis migrate from moun-tainous areas to upland ricefields during floweiing (1 14)

Pentatornid bugs prevail in Latin America (8182 195) and in Africa pentatomid and coreid bugs arc equally cited (6 82) l)amage is often characterstic of the species (5)rcoris ventralis and Mnida spp are the common

pentatomid seed bugs of Philippine upland rice but they prefer lEchinochva spp grass seed to rice Vearaviritda occurs worldwide and hoften more abundant in the uplands(54) We saw few reports of seed bugs on upland rice but see(d bugs apparently opelrate cqually between by land and upland rice Good dispersal is necessary ior species that call feed for only 2-3 xon a crop and must first find those crops at ripening stages

Other pct species feed n flowers In Mato Grosso Brazil (194) and Santa Cruz Bolivia the panicle weevil Aeo-hariuia amllitarsis attacks rice during flowering (68 233)The larva e rice sten horers adutlts feed at the bases of spikelets caIusing empty grains Also inLatin America thecydnid -lkiituttatralusisa seed bug as anad tilt and nymph (194)

A number of beetles ecd on poilen - coccinellids lialrotia Aulac lhora and Monohpl-a (55) - but becatise riCe is self-pollinating and fcrtiliation occurs before pikelets open they pose no threat to yield

IST ECOLOGY

We now look at characteiistics of the life cycles of upland rice insect pests to lea rn hew each plays an adaptive role in upland rice ecology

Life history strategies Orzasativadoes not tolerate drodgh well so upland rice is highly seasonal normally grown in the wettest months of the year Usually rice is present for less than half of the year The rice-fiee fallow poses serious problems of survival to upland rice insects which have evolved att least four mechanisms to ovrcolne tle cvcl ic-l lack of a hostI) polyphagy 2) longevity 3) dormancy and 4) vagility (dispersal)

Poldt)agy lolyphagy is defined as having hosts of at least two botanical families oligophagy of more than one gents and ionophagy of only one ventus (40) - inour case Orvza Most wild rices however are illi tiC and theieiirc are normally far removed from upland rice hahitat othewise more would be alternate hosts for uplanJ iice insects Thte highly mornophagous rice pests shySciriol)haita hiceritlas Nilal)arvaiahiens Iand Nealo--tettiyvircsctws in Asia and Aalirpha secparatella and 1ioj)sis lngicornis inAfrica - occur on upland rice but

are more abundant in lowland culture They can specialize in rice because they are highly vagile and can attack all growth stages By specializing in rice monophagous rice pests can outcompete related but polyphagous speciesIntensive andI extensive lowland rice culture has favored monophagous species Riqwla alhinella (Cramer) the onlymonophagous species in Latin America is rarely reported on upland ice (I1)

Because of its limited temporal and spatial existence upland rice has not favored the evolution of specialized species Clhang(44)shows evidence that upland rice has only recently been cultured by man from the lowlands thus pests have had little time to fully adapt to the crop All but the lowland-adapted insect pest species have alternate planthosts to rice Root pests with limited mobility are highly polyphagous Pests that specialize in one growth stage eg secdling maggots seed bugs are oligophagous or polyphagous

Most alternat hosts of rice pests are annual grasseswhich also are seasonal Upla_rice is also a host but not the preferred one of the most highly polypliagous spcciessoil pests grass hoppers armvwarrs and Vearaviirielda

The d ry season poses igreat obstacle to ani upland riceinsects survival and only sonic perennial plant hosts are suitable food during the off-seasoin Plolyphagy allows upland rice insects to suLrx-ixe during thC rainy season when plants- both annual and perennial -areactivelygrowing and therefore more nutritious

Loiugevitr The ability of aiinsect to live 2-3 mo without undergoing dormancy helps species sich as root-feedingwhite grbs (as larvae) and secd-feeding bugs (as adtilt) to survive unfavorable periods White grubs arc not highly fecund and their life strategy is to improve survivorship of limited progeny Their subterranean habitat hides them from manv natural enemies lBtut root tissue is not highly nutritious so they ltilst eat great quantities and need a largeinsect biomass to digest it Other soil-inhabiting rootshyfeeding insects have short life cycles but are more fecund

Seed pests need greater longcvi vas adults to locate ahost at early grain dcclopment arnd give sedentary offspring time to develop Few groups of upland rice insects live longin active development Polyphagy and longevity only increase their ability to survive during the rainy seasonOther mechanisms are needed to survive the dry season

orantc In the tropics the nonrice season is the dry season while intemperate areas tile off-+season is winter Many upland rice insects undergo dormancy inlsummer (aestivation) or winter (hibernation) Dormancy is simplyinactivity during winters with prolonged temperatures below 10 C insects find shelter arnd cease movement Activity is resu med with warmer t empcera tu res A deep state of irnactivity termed diapauise occurs in sole species that prepare physiologically (havxing low metabolism and convcrting gonad s to fatty fuel reserves) for the unfavorable season (148) They enter diapaUse inanticipation of unshy

IRPS No 123 January 1987 17

favorable weather and resume activity after favorable Armyworms skippers and locusts actively travel hundreds weather begins usually in response to photoperiod or of kilometers in air fronts such as intertropical convergencethrough a delayed response to a stimulus such as heavy zones to descend onto upland rice fields far from their rainfall (237) Not all individuals of a population have the breeding grounds (31) Several generations can build ip on same threshold for environmental responses for entering or alternate hosts in tile grass family before rice is plantedterminating diapause -- this is seen as a mechanism to These large insects are active fliers Smaller insects such as prevent total mortality if a false cue appears such as a thrips aphids and planthoppers passively migrate long sudden warm spell in winter or rain in the middle of the dry distances by the wind season (154) Otherspecies ean readily travel tens of kilometers - stem

Probably more upland rice insects undL go dormancy borers leaffolders leafhoppes cutworms spittle bugs leaf than we realize Candidates are Atherigona spp root beetles seed bugs ants and termites aphids root weevils crickets leaf beetles thrips and seed The least dispersive species are white grabs mealybugSbugs Insects known to entervarious states of aestivation are seedling maggots crickets weevils and gall midgesleptocorisaspp adults (199) Maliarphascparatellalarvae Each species has evolved a unique set of attributes to (9) Scirpophaga innotata larvae (117) Chilo partelhtsv enable it to survive and adapt to the changing uplandlarvae (21) ScirpopThagainccetulas (179) white grub larvae environments Several evolutionary avenues lead to fitness (134) Patangasuccincia adults (6 82) and Schistocerca (Table 2) but there is no reason to believe one set of gregariaeggs (82) attributes is better than another (224)

Insects known to enter hibernation are LodeIphaDstriatellus nymphs (124) Chtio Tvlpressalis larivae (125) DroughtSesaia inferens larvae (146) but not S calavisis(2) Well-drained rainfed upland soils are subject to drought ifIDiaraea saccharalis larvae (75) Scirpophaga- tcerlttas rains do not fall within 2-3 wk Prolonged drought followedlarae(5 lacchrli s lae (75)saSlOhagasshoperseggs (82)and felo pidas athias pOpae (232) by favorable rains stimulates armyworm and locust outshybreaks but mealybtigs root aphids and thrips become

Lcpmotorisa oratorisand L acuta adults aggregate on numerous onlyafter dry spells for two reasons First heavy trees sugarcane or other shady moist sites L paluwanmsis rain normally kills soft-bodied foliage-feeding thrips and confined to Sulu and Palawan Philippines lives on the mealybugs and soil-inhabiting root aphids and root mealyshygrass Brachiariamutica (Forsk) Stapf in the absence of bugs Drought eliminates this source of mortality Second rice The adults can live for 5 mo When aestivating their the rice plant responds to drought by breaking down rate of metabolism drops and they do not feed They will proteins into soluble N compounds which enter the phloem however puncture plant tissues of their aestivation host in and are taken up by these sap-feeding insects (245) Greater search of water This behavior has led to errors of the host nutrition therefore leads to greater survivorship and range of rice bugs (I) Aestivation quickly terminates with fecundity rain probably meaning aestivation is not deep Small rice area

Some stem borers aestivate as mature larvae in rice Except in the highly mechanized large landholdings of stubble or in straw stacked as livestock feed However Latin America upland ricefieldstypically are patchy within moriality increases proportionally with the duration of the a highly diverse flora The small fields result in higher ratios dry season even if the stubble remains undisturbed If the of perimeter to area than with the typically larger lowland stubble is plowed for a crop following rice few stem borers ricefields Upland ricefields themselves may be intercroppedsurvive Rainfall terminates dormancy but it normally with cereals legumes or root crops Mixed intercroppingtakes several weeks for moths to develop and emerge with a wide variety of species also occ in tribal slash-andshy

Last-instar white grub larvae tunnel 1-2 m deep in the soil burn agriculture (50 74) to construct pupal chambers in which they aestivate The Upland ricefields tend to be small because of the labor larvae are very sensitive to dryness so pupal chambers are needed to clear land in slash-and-burn areas and to till land sealed to coierve body moisture Aestivation terminates to minimize weeds Rice requires more tillage than does with the first heavy rains but as in stem borer 3-4 wk pass maize because rice competes less well with weeds Uplandbefore adults develop rice is more a subsistence than a cash crop and small areas

Insects may respond differently in each region to cold (I ha) can feed a family temperature or drought Biotypes or loal populations may Small fields favor some insect pests such as some ant evolve (56) Local populations of Scirpophagaincertulasor species that can forage from more permanent field borders Patangasuccincta may have become adapted to prevailing Trees provide food and mating sites for white grubs (134) as conditions well as aestivation sites for LetTocorisa spp (199) Field

iagiliti The ability to disperse combined with high border grasses plus tIhose within a ricefield provide alternate fecundity orshort life cycles enable some upland rice pests to food for all but the highly monophagous upland rice better exploit the temporarily favorable upland habitat colonizers and a habitat for natural enemies

0 z

Table 2 Life history patterns of upland rice insects or insect groups

Host range Longevity Dormancy Vagility FertilityMonophagy Oligophagy Polyphagy Low Medium High Yes No Low Dispersive Migratory Low Medium High(Oryza spp) (grasses sedges) (angiosperms) (lt3 wk) (gt2 mo) (lt1 kin) (lt50 eggs) (gt500 eggs)

Ants xTermites x x x

Mole crickets x x x xx x xField crickets x xx

White grubs x x x xx xRoot aphids x x xxMealybugs x

x x x xxSeedling maggots x

x x x x xLeaf beetles

x xx xArmyworms x x xx x

Thrips x x xx xChilo spp x x xx xMaliarpha x x x x

xLocusts x x Xx x xGall midges x x xxiVephotettL iirescens x x xx x x xLeprocorisaspp x

x xLeaftolders x x x

Sogatellafurcifera x

xx x x x x x x

00

IRPS No 123 January 1Q87 19

All this appeals to favor insect pest buildup but the notwithstanding the release of stored nutrients in response contrary is true Loevinsohn (138) showed that insect pest to drought that temporarily benefits philoem feeders populations respond exponentially with the proportion of land devoted to rice up to about 75 of the area then the M POR ANCE I I P1A 1) RWI P1St S

rate of response declines In most upland rico areas tie rice crop occupies less than 50i of the area- therefore the Visits to upland ricefields normally reveal 1kw insect pests potential for population buildup is low This relationship Loevinsohn (138) and Locvinsohn ct al (139) compared may not hold for seed pests that can locate small ricefields insect abundance using annual light trap atchcs in upland

Another outcome of the highly diverse flora and phical rainfed lowland and irrigated lowland sites in the Philipshyenvironment of upland rice isthateach upland ricearca has pines Although only one upland site was studied data a unique composition of insect pests showed equal or fewer insects at the upland site than at the

For example in Tanauan Batangas Philippines the rainfcd lowland sites and fewer than at the irrigated sites main upland rice pests are rice leaffolder(RLF) armvworm (Table 3) M Lltiple regression analysis showed that the key Aythntnaseparata WBPH ants Soli psis gentitataand factors explaining insect abundance among a set of cropping white grub Leucopholis irrorata Sixty kilometers away in intensity variables were the number of rice crops grown per Real Quezon the main pests are seedling maggot Athe- year followed by area devoted to rice Table 3 shows that rigona orae flea beetle (Thacitcmnra hasalis thrips the upland rice site had only I rice crop perym rand that rice Stenchaetothrips hiurnmis and rice bug Leptocorisa was planted in only 20Vi of the area Fertiliter and oratorius In a third Philippine upland rice site in Claveria insecticide use had little bea ring on the trend The armters at Misamis Oriental Mindanao the main pests are ants the upland site applied 60 kg N ha higher than at most Solenopsis gemtinata seedling maggot Alieriona orvzae irrigated sites white grubs Holotrichia in)iUaiuoan t and Lucopholiv Insect occurrence at the upland site was omparable to irrorata stem borers Sesamia ifiArelr (hio auriciliusC that at the Cagayan rainfed lowland site parlicularly in 1982stpessallis Scirpophaga incertulas S innoiata and when drought prevented many farmers fron planting(only

4c ona chrrsurapella root aphids Tetraneura niriah- 40(of the rice area eventually was planted that year) M ore dunrinalisand Rhupaosiphunm riahbdminalis lcaffolders favorable weather occurred in 1980 and 1981 which Marasniapataltis and Chaphalocrocis Medinalis army- supported greater rice areas and consequently more insects worm MIthinna separata and rice bug Leltucorisa The most intensive site for insect pests was South Cotabato oratoriusThe great diversity of insect problems is typical Mindanao where farmers plant 23-25 rice crops per year fer upland rice and makes control efforts more difficult with irrigation The Zaragoa Nutva Ecija site also had particularly regarding breeding for insect resistance high pest abundance and at this site the fields were highly

asynchronous Asynchrony was the third most important Low yield potential Iactor in explaining rice insect abundance Most upland rice is grown on low-nutrient and mineral- Rice cropping intensity in upland areas is bound to toxic soils which when combined with eiratic rainfall remain low Only one rainid upland crop is possible per make upland rice a highly riskycrop for the farmerto invest year (Farmers in Claveria Misamis Oriental Mindanao in costly soil amendments and land management Often Philippines tried asecond upland ricecrop i 1984a ycarof upland rice isgrown far from markets making inputs even favorable rains but no harvest was possible b-cause of more expensive Farmers growing a subsistence crop hosi- drought blast and birds) Rice area isaso bound to remain tate to invest in costly inputs to raise yields low because of the high labor and power rcquirelmenlts to

Even having varieties that could double existing yield prepare the land (because of weeds)and because upland rice potential under present management levels would probably wvill not compete with lowland rice as a cash crop Some not prompt many farmers to use purchased insecticide upland rice sites with prolonged rainy seasons howeverare unless tile risk of crop failure were reduced such as it was planted asynchronously such as the Clacra site with irrigation systems in lowland rice The seasonal abundance of Upland rice insect pests

The Tanauan Batangas site is atypical for upland rice attracted toa light trap wasgrapied for crop year 1980-81 in The soils and rainfall pattern are favorable and because of Tanauan FBatangas (Fig I 2) W13P11 was he most its nearness to Manila farmers have cash resources ftm common rice hopper and its population peaked at the sales of vegetables to purchase tertilier They applied an midgrowth stage as it does in lowland rice D-Vac suction average of 60 kg Nha after particle initiation and obtained samples showed an earlier peak than that from a light trap 25-35 tha yields The high fertility isone reason why RLF indicating emigration W13PI usually emigrate after the armyworms and WBPH developed into large numbers vegetative stage Populations in 1980 reached levels that Farmers however used no insecticide on upland rice caused patches of slight yellowing in some fields WBPH although they did on vegetables probably cause hopperburn in Batangas They immigrate

The poor growing conditions fOr rice generally mean year-round as shown by low levels in the light trap BPH poor growth and low fecundity for insect pests fIeding on it however were scarce in the light trap and not collected with

Table 3 Abundance of rice insect pests and factors in rice cropping intensity of Philippine upland rainfed lowland and irrigated lowland locations where light traps were operated daily for atleast I yr -I

0Rainfed Rainfed lowland Irrigated lowlandupland

Pangasinan Cagayan lloilo Santa Maria Nueva Ecija South Cotabato Tanauan Caaringayan Caaringayan Bangag Bangag Bangag Buray

LagunaCale

Batangas Manaoag 1980-81 Santa Motiica 1982-83 Cabanatuan Zaragoza Avancena NaniamanSolana 1981-82 1982-83 Oton

1980-81 1979-80 Oton 1981-82 1981-82 Koronadal Koronadal1980-81 1979-80 1979-80 1983 1983

Insect (kerosene light trapNephotcttixcatch nolyr)virescens 650 1240 500 1800 330 210 3400 1600 160N nigropictus 210 - 360 18400 5500 2900- 140 90 20 - shy 1700 -Reciliadorsalis 710 - 3200 1700180 890 630 140 170 2400 800Nilaparratalugens 130 690 - - 4200 4600300 520 750 180 60 3300 1600 740 310Sogatellafiircifera 1700 31000 13000 8200310 210 3300 610 270 980 1600 6300 -Cyrrorhinus - 12000 6700600 540 3200 290 160 330 120 410Scirpophaga spp 50 410 620 - - 21000 12000150 180 220 270 30 1000Cnaphalocrocismedinalis 70 70 540 610 270 11000 6100200 140 100 6 60 250Chilo suppressalis 0 60 60 1900 300 200- - 30 20 0 6 3 160 -C auricilius - 0 00 - - 20 3 60 20 3 0Sesania inferens 0 - 80 60 3 80 10 51 -Cropping intensity _ -

Rice crops (noyr) 10 10 12 10 10 10 13Rice area (7c) 17 19 20 20 25 2320 85 85 60 70 40 85 85 90Fertilizer (kg Nha) 60 20 85 80 70 7020 0 0 0 30 30Insecticide (no of 40 60 40 300 04 04 0 03 300 13 13 30 50 40 33 33applicationscrop)

__________

IRPS No 123 January 1987 21

Light trapped Insects Drytnd __ 2(notrap-nght) Suctioned Insects (nom )

50 -)o 9 jO40- SOW ffurciWO Light trp

D-Vc suction

20 to 4

20

Egg porasitizahion (M) 75 60

300 SOOampReOftwciffr

45 30shy

15

-insects (notrop-night) 4 Nileporvata Iuen5

Egg parastization()

Light trapped insects (no trop-nght) Suchoned insects (no m 2 )

6 8 Cytorh~nuls hvidipennii 0

4 05 2

50 0 Insects (noltrop-night)40-ui Ipanh pe g

Moohase 1 Seasonal abundance measured by light trap

Rainfall(etod) and D-Vac suction sampler of whitebacked

to ~ hopper Nilaparvata lugens and lea ffolder60)-t Cnaphalocrocis mnedinalis andplthpe g20 L I II I I I parasites and mtrid predator Ortorhjntus livi-Apr May Jun Jul Aug Sep Oct Nov Dec JaIn Feb Mar dipennis set against cropping pattern daily1 9 80o 19 8 1rainfall and moon phase Tanauan Batangas

1980-81

the D- Vac Cyrtorhinus build up on the crop coterminous are collected in light traps during full moon (109)with WBPH buildup Cyrtorhinus disperses during the Because upland rice shares most of the lowland rice pestsrainy season RLF not readily collected in light traps was and those pest species are generally less impotant to uplandprevalent only during the rice crop rice than lowland rice there is a general belief that insect

Nephotettix virescens was more abundant than Nnigro- pests are less important onpicLus in the light trap collections and on the crop GLH upland than lowland rice Arcview of limited data on upland rice shows an averag of population peaked at the reproductive stage as did zigzag 10-21 yield loss from insect pests on a par with that ofleafhopper (ZLH) Crtorhinus tracked the populations of lowland uee determined by the same methodology (132) planthoppers and leafhoppers Both GLH species were (Table 4) McGuire and Crandall (144) made a similarcollected year-round in the light trap indicating immigra- estimate for Central America Cramer (51) howevertion from nearby irrigated rice areas Possibly because of the estimated 35 yieldlow populations loss in South America based on ainsect number did not correlate with similar pest complex - stem borers and seed bugs shymoon phase although reports indicate that more hoppers attacking irrigated rice in North America This estimate did

22 IRPS No 123 January 1987

Light trapped insects D~~nd (no trap-night )

St plht Suctined insects (nom 2

)

D-Voc suction 1520

1o

0 Lih trap 5 ~105 o5

00 Eggparositizotion()5040 NephOtgtha virneens

30 20

I010 A-Insects (notrop -night)

4 Nephatelh nigropccent i

2 o I vV^J - -- ^ VA____ f _

Light trapped insects (no trap-night) Suctioned insects (no rnE

10 -1208 ReCdO doRsol

6 15

4 10 052 iwxALAiamp 0

Light trapped insects (notrap-night) 2Suc oned nsects (nom )

8 CyrtorhmnusI6o nn]10 i0075

4 05

2 0 0

025

2 Seasonal abundance measured by light trap Roinfall (mmd) Moon phase and D-Vac suction sampler of green leafhoppersNephotettix virescens and N ngropicrus and I zigzag leafhopper Recilia dorsalis their egg 60

parasites and micid predator Cyrtorhinus livi-dipennis set against cropping pattern dailyrainfall and moon phase Tanauan Batangas1980-81

40

5 Lf Apr May Jun

1980 Jul Aug Sp Oct Nov I

-aMar

1981

not account for soil pests and was not based on field data in the absence of pests Carbofuran granules are phytotonicAkinsola (I1) cited mainly data from irrigated rice areas in to rice (239) The prevailing hypothesis is that carbofuranAfrica but estimated yield losses of upland rice between 15 prevents soil bacteria from consuming fertilizer The phytoshyand 30 from insects However losses in Thailand over a tonic effect is through greater availability of fertilizer to the4-yr period ranged from I to 13 averaging 5(106) plant Denitrifying bacteria are known to be more active inYield loss data however should be interpreted with aerobic than anaerobic soils therefore it may not be merecaution (184) The Philippine data were all derived from coincidence that the highest recorded yield losses (22-69)trials on farmers fields under farmers normal agronomic occurred in those trials with 12-30 kg basal Nha On themanagement practices except for Bukidnon and Capiz other hand contro with carbofuran granules may not be aswhere researchers used fertilizer Batangas farmers are good as a seed treatment against seed pests and perhapsatypical because ofhigh levels of fertilizer used The trials in yields would have been higher if a systemic seed treatmentBrazil were carried out at an experiment station but insecticide had been used conditions were similar to those of local farmers The 1979 and 1980 trials in Batangas had two full-

Carbofuran granules were used in every trial as a broad- protection treatments one using carbofuran granules andspectrum insecticide applied basally with the seed An the other using bendiocarb WP as a seed treatment Plotsobjection to using insecticides to measure yield losses isthat protected with bendiocarb yielded higher than those proshyinsecticidescandirectlyorindirectlystimulateplantgrowth tected with carbofuran granules The other trials in the

Table 4 Yield losses to insect pests and yield responses to insecticide in upland rice in the Philippines Brazil and Upper Volta

Yield (tha) Yield loss (5) Location Year Cultivar Protected

with Protected

without Check Total Vegetative Reproductive Ripening

Basal N (kgha) Reference

carbofuran carbofuran Protected No with carbofuran

carbofuran

Philippines Pili Camarines Sur Tanauan Batangas

Pangantucan Bukidnon

Dumarao t-apiz

BrazilGoias Minas Ge~ais

Upper Volta6

1975 1974 1974 1975 1975 1976 1977 1978

1979

1980

1979 1980 1979

1977 1978 1979

Bursiging puti Kinanda Dagge Kinanda Dagge Dagge Dagge Dagge UPL Ri5 Dagge UPL Ri5 Dagge UPL RiS UPL Ri5 UPL Ri5 UPL RiS

IAC47 IAC47 IAC47

14 33 11 36 35 31 30 27 37 26 40 30 40 42 46 35

19 21 09

--

-34 24 24 27 --

29 43 31 42 -

47 25

---

10 28 05 27 27 30 26 29 33 28 33 29 43 23 36 11

15 15 06

29 15 55 25 23

3 13 -7 11

-7 18

3 -7 45 22 69

24 28 35

--

21 -11 -20

4 -

-3

23 7

-2 -23 56

-

-

-__0

6 --

-

-23 14

5 0 8 2

22

-

--

0

-

--18 11

3 -10

4 10

7

--7 -

14 -

-1 6

10 -5

8 9 7

-

--

0 0 0 0 0 0 0 0 0 0 0

30 30 30

12 12 12

99 99

100 100 101 101 102 102 102 102 103 103 104 104 104

66 66 66

Farako-Ba

Mean

1977 1978

IRAT10 IRATIO

38 66

--

40 43

-5 35

21

-

10

--

--

--

- 242 243

Z

tJ

-w

24 IRPS No 123 January 1987

Philippines which had two treatments offering protection In the 9 Philippine trials where yield losses were measuredwith and without carbofuran granules relied on further by growth stage 10 of the yield loss (21) occurred at thecontrol with foliar spray and not on seed dressings The vegetative stage and 517 occurred at each of the reproshycarbofuran-protected plots had an 18yield los compared ductive and ripening stages Ibis result is similar to thewith only 10 with foliar sprayS This yield difference is pattern of yield loss in lowland rice measured by successiveprobably due to differences in protection against sown-seed treatments where the crop is unprotected at each growthpests (foliar spray do not control sown-seed pests) rather stage but proticted during the other growth stagesthan phytotonic effects became basal fertilizer wa used in However the insect complexes r sponsible for vegetativeonly 2 of the 10 comparisons stage yield losses in upland rice are entirely diftfrent than in

The recorded yield losses were not related to yield lowland rice The key vegetative stage pests in lowland ricepotential (Fig 3) One might expect higheryielding crops to arc aquatic whereas those in upland rice are sown--seedhave higher percentage of yield losses but this ws not tile (sown in soil) root and seedling pests This dift rence maycase One reason is that insect damage exacerbates plant explain why agricul torists have overlooked tile ilmportanccinjury from drought stress low yielding fields from d ought of upland rice insect pests Many reports of upland rice pestsstress would show disproportionate differences between in Asia ha Ve focused on major lowland species (46 108)insect-protected and -unprotected treatments Shoot and It isalso surprising that tei vegctative stage of upland rice root pests would be more responsible for losses from would record the highest yield losses becatuseseeral studiesdrought stress than sown-scd pests A second reason have shown that upland rice can readily recover from foliageconcerns seed pests which would have a greater impact on reinoval during that stage (117 152 165 188) Upland ricelow tillering low yielding varieties such as )aggc Low poscsses a hitgh obilitV to compensate for ca rly loss oftillering varieties nnot fill in the space created by removal foliage and will even be stinulated by foliage removal toof seeds oweer 1) R i 5 is high yielding because it tillersP1 produce a higher viel than plants ithout any foliageactively which can close the canopy to provide greater weed removed (63 192 231) [ie reason for this apparentcontrol discrepancy hovce is that yield loss at the vegetative

stage does not octUr frol leat arca loss but fron pestsYield loss () removing sown seed and feeding on devcloping tillers and roots -- damage that cannot be readily conpLtnsatCd for

70 - Measuring yield losses in upland rice cnvironrnents is further complicated by a report from Brazil that greater yield losses occurred in response to better insect control60 - =-09 as Insecticide seed treatments resuled in denser plant stands

(n=21) which in turn created a selectively more favorable environshyment for blast (69) This resuIt was parlticurla rlxexacerbated50 when cirbofuran granules were used in seed furrows Aside

ffrom a greater plant stand ca rbofuran iay have auigmented the N lecl and stimulated blast If so ca rbofuran should be40 - replaced with another insecticide or the seeding or fertilizerI rates decreased

Other methods to determine the impact of upland rice30 shy pests wvcrc to interview farners or to develop single speciesbull bull correlations of population levels and yield loss Feneira (66)2 NIM)st (75(1) Golas farmers reporteddid both in Brazil

20- insect losses in ufpland rice and 251 said those lossesS avcragcd 37(1 [his result is consistent with tie yield losses 0 0 measured by field trials in the same statc(Table4) For every

I0 4 of seedlings removC d by leaf-cutting ants a I 7yield loss is predicted Similarly Ferreira (66) found that ai

0 - average of 53 thrips per panicle before panicle initiation meant 3 times more unfillh d grains

-IO0 I 1 1 I I-101 0 2 3 4 5 6 7

Yield (tho) C)N [ROt IM FI IODS 3 Correlation between upland rice yield and yield lols caused by Upland rice insect pests may be controlled by culturalinsects measured in 21 trials in the Philippines Ivory Coast andBrazil genetic biological and chemical methods

IRPS No 123 January 1987 25

Cultural control dry season plowing will have little effect on them Plowing Crop husbandry to reduce upland insect pest populations when the soil is wet from the first monsoon rains will expose can be classified into practices effective in a single field and soil insects that tunnel close to the soil surface to predatory those effective only at the community level Cultural birds chickens dogs and even man (field crickets nole controls should come first in a pest management program crickets and white grubs a re delicacies in man tra ditional They have broad stable effects as pests have little pos- diets) sibility of overcoming them through selection of biotyps The greatest degree i1 inIscct control fron tillage comes (138) On the other hand some cultu ral controls work only from plowing soon after rice ha rvest when most soil insects with high labor or power inputs and connunity-wilt lie close to the soil surface anrd stern borers are in the rice methods require coordination of iany farniters stuble Whitegrubsare mature and tillage exposesthem to

Pantin time The luxury of being able to shift the time of predators Tirnelv control will prtec the crop following pla ntring upland rice is open onl hr la rniers in areas with a rice and lowr the populationIfor succeedinig years Plowingprolonged rainifall pa ttern lI copping intensity or underthe ice stulble helps it decompose killing sten borer rnechanied land preparatiot Gcnerally planlting as soon larvae inside as the rFiinv season begins will lower populatiois of 1110st [requent tillage destroys ait nests bit not termites insect pests Insect populationsarelo aftcra dlvseoll or Tillage and clean culture howes er rcliove food for winter fallow delayed plaiting lets them build tip on termites which then mtay attack a young rice crop Scarab alIternate hosts that gi vigorotsly with the first seasonial beetles also prefer to miposit in eceitly tilled fields Noshyratiris This first flush iof weds cros lItxuttriartlv with iceCult is dibbling faors the survival oftillace Clre Such rineralied N released during the dr season aid made stils i aels tnd insects thtt pttpate or lay eggs- in the soil

available to the plants with the rains It iliso igrtmitcally Zero tilla hio eerl conserves predators ofthe GIH (37) advanttgeous for the rice ciop to tap this trlLItlaI fertility nt it mcttoi lo lessen the impact o0 soil insects and grow vigorotsly to compete with wrus and to tolerate brotdcasting seed or planting in ftltrrows is prefc ired over pest infestation dihlirig in hills Crickets are large enotlgh to eat seedlings in

Larly plantngs terd to escape seedling maggolts Upland a hill atrd tillering normnally ca iriot fill in tlhe missing space rice in Bitangas is planted in Ma and ine itli the Iirst Oftcn dibbll rd is not well covered and iseasily found by rains Fairners are highly mtotiated to plant early tocscape foraging insets ioIdents and birds drlitght and typlotns ricar harct Arn Atrgtst planting Plan uit sii Incrcatsini sced density protects tile cropwould be evcely attacked by It riim orivzo At tti againrst seed and seelling pests especially ants particularly other Philippiie tipland sites Rea l Qteot antd (laelit if the crop will germinate and ermerge quickly M isainis Oriental seedlig mnaggots arnrige ric boe t se hr trftinll itI India rice intcrclopped witi cotton or farnes have a longer growing season and plnit 2 rin or pigeon pci had lower 6IIt and WBIlII poptrlations than mote after the ortset of the rainy season liaaiangas ftrmners rice alone (211) NfailC arid Urplald ir_ 2re coilltori in the plant ca rly becia use ofa shorter raiinfall period arnd the desi re Philippines he chatngC in pest status folii iniercropping is to lliarvet before the typhoor sea son peaks ltighly locatioll-specific (133) cmianarid the net elfc be either

Early plantings cornbind with early-iatiuring varictics nil bcnelficial ortdetriential (189) For example BIatangias provide higher crp tolerartce for white grubs Aestiatior farmers intercrip Itro in their ipland ricefields A liornshyterminates with the first rains but 5-6 moare ineeded for the nrii iotd-Ahis i (I iiiacius) which norlially feeds resting larvae to develop into beetles emerge and lay eggs on taro ill feed or miarbv rice plants Intercropping and for larvae to reach the danriigii g ird-instar stage probably would not afft0 seed pests A crluanion crop (134 In lataigas this normually iseiough tiite for harvest planted ith rice and haing gleater tolerarnce for root

It Brazil howecver delayed planting is a suggested dariage or being toxic nayact isi trapcrop to root-feeditig cultural control for the lesser cornstalk borer and insect pests Acial-fei i ig pests voultId be most affected by pests favored l drought (66 195) Cropping internsity in intrer ropping Shoot aphids leaf beetles sten borers and Brail is low and delayed plraing places the rice crop in the hopplers would be likely ca ntidates to note effcts On the niost stable rainfall period reducing likelihood old rought othlt halnd the corlipalior crop stch~ sitniaie Iiiv be more

In rilniy arca s rinmirsare roi atl to plant other crops benehited than rice from initecropping ( 137) before rice and althNgh it woultl be desirabh to avoid 11echi Weeuling luring the first month after clop infestations by planting rice cuatl lariners lack the labor cstablishent will force pcts such is anyvworris which and time to do so ol a low priority crp prefer grassy cetds ontor tle vorling rice Soil iampsecs will

77age Uplild rice Stils ae light textured and Call be mvnye to the rice crop Cleani citrltirc also lay lore territes plowed duiring the d rv seasor notl only to grin time for to attack I u cetincti inrice ceds i ay he le optinal planting with lie fitst Inilsoon riiains but also to desiccate solition 13) soil insects arid weeds such is ntsedge that foim rhizomes Irtlizr Itneatsing N Iciiliier oin hlwland rice has Htowever becta use most soil insects lie below die plow layer fAtwvored Irigher populains of pliithoppeiv Imlhoppers

26 IRPS No 123 January 1987

and leaffolde rs However because of the need to temper fertilizer use on upland rice o prevent blast fIrtilizer rates will be lower than is comonia r for lowlad rice causing relatixely less hopper and IaIfTfoide r population buildup However inupland rice areas without a history of blast such Is Batangas farmers apply high rates of N and conscqtrently the crop has high infcstation levels of WBPI]and RIF The fhrmrcrs leec insect buildup by splitting the fertili11r in0to two to three applicatiois bit do not applV fertilizer at the vegetati estage because this prodrces a tall crop which illeasily lodge during typhoons Greater crop fertility leads to greater insect survival (better nutrition) larger insects (to cat no re foliage) arid greater fec titd t (to lav nmre eggs) (128 145 171) Not all insect species respondequally hovr Increasing rates (if N l anrid K increased Dialraa s-acchralis and (iaeiurema flea beetle but reduced thrips (66) Zn rcdLuced deadheart density from the lesser cornIstalk borer

Fnldingi Although most Upland rice areas do nIot hawxcaccess to wa ter floodiig the fields is rRrrnriierdcd in areas of Brazil or root bug and white gib control ( 195)

Crop rotatioti Tillage before planting a crop after rc harvest will control stern boreis and other pests renain n the stubble arid will unearth soil pests Planting a non-gratriiaceous crop after rice is recommncrded for termite control (66) Crop rotation is less nttdcd in upland rice culture than in lowlands because atcrop-free period is assured even after harvest of crops following rice

t1ulchilt iryl plastic irli irig designed for weedcoitrol ii upland rice inJapan resulted in ntr highanri liy infestation of (ihilo sulytresvais (93 9-1 95) NIulched Upland rice grew more luxuriantlv and attracted more (Avipositing mothis than u lched rice Also the rice plant and stein borers imatoured 10rC quickly and a partialgeneratiori af vouing ista iNoverwintcred in the stubbleLotili--malitrt varieti Quickly maturing crops reduce the tmbCr of pCst generations that Calt build up Rapid crop establishment olan ca rl v-natu ring vaitvty if)pa 11icLa r will lessen white gib and stem borer damiage White grub larvae intle last iistar are highly destIruCtive so aii early-iiatri ng variety will escape serious root loss Stem borers build 1ip slowlyx in rice to ecormic abundant in a late-Maturing crop anrd caiuse whi tlieads Numibers of somespecies increase exponeitially with each succeeding genera-ti so a xarietyexhat atures I trowh earier thian another will hac less insect damage Early-naturing vaiicties however will not lessen seed pest daiage either ii the soil or oina standing crop

Strwhoi hAIJttam Inrscts tcd irig on thlie aerial portions of rice plats disperse from carly to late plantedfields AS iiiibCirs increase exponentially with eachinsect

gtscnrattm e eiglbrig f rmc~s1(uil upp wh rc generation pests build tIP where neighboring fihrTC e9r1sstagger their plantings II Batangas farmers plant uplandrice within I iro use 120-d s-arieties and plant at the start of the rainy season Ini Misarnis Oriental laricrs stagger their

plantings up to 3 mo apart and use a set of atieties that mature in 4-7 mo Insect pest danage particularly from stem borers is greater in Misarmis Oriental Also seedling maggots damage rice there because rainfall favors frequent planting of maic In Batangas rice escapes seedling maggots bCca usC of distinct drs season and synchron0us early planting with the onset of ains

Therefore combinations of cultural practices ea rlyplanting synchronous planing crop rotation and earlyshymaturing vaneties protect the rice crop against most insect pests

Ilmant resistance Beca tse upland rice has few specialized insect pests and many insect types attack the crop worldwide regional breeding and strong internatiornal cooperation are required

Sown-sced pests arc economically controlled by insectishycide seed treatment and should the relore receive lowxpriori ty inbreeding objectives

Root pests are normally cost ly to control with insecticides but finding resista t sonres mayvbe di fficult Tanaka (229) for exa -ple fi led to find varieties resistant to root aphids Selecting for large root bioniass perhaps should be the strategy for pests sLuch as Whlite grubs that renliove roots Fainter(174) howcvcr cites varietiesof crops tolerant ofor resistant t(i root pests Some sorghun varieties are high in cyanide so rice sorghun might incorporate broadshyspcctruri chemical resistance

Seedling vigor and drought tolerance are high prioritybreeding objectives so tupland rices should have large r root systernsand be able to tolerate ligherlevels of root loss Sonie root loss inolder plants is beneficial if it stimulates new root developnrut and therefore crihances intake of soil nutrients (53) The rice seedling inaggot rtiLotia spp may be

controlled thirough resistant aittics A hi rgc breeding effort on sorghniu has had fair success inideveloping arreties esrstait to Asoccata Rodani (251) A field trial comparing 10 upland ricc varieties showed differences aniong varieties va ring from II to 4(- damaged tilleis (215) Hovver resistarce to 1Iill-0r gtia species may rot cross ox-rti otier

Shiraki (210) reported that upland i-ice stens wcrc harder than lows-land rice stemsperhaps becaIusc (f a higher silicoricontent and thus were more resistant to yellow stern borer hc tortia Iow tillering (f uplaid iice xarneties prodtres a

higher percentage of infested tillers than would occor in higher tillering lowland iice under a siiilar egg density The corollary therefore is to select for high tilleriig upland rices

Stemii bor amcstheh main targes dobrecders inAfrica (571 204 )but nticl oifth e sc reen irig is dotie unde r lo wland co d t osrm s wih A h r~ o l pe i s n acorditions (11 96 218) As with lthrinrnwspecies mayhave to be dealt with independently however initial results show cross resistance between African and Asian species(92)

In Brazil screening Ior insect resistance focttses oil lou I pest problems Varieties tolerant of lasmolwalpus ligi-sellus and resistant to )iatraea saccharalis have been identified (70 153) Current work also emphasies 7iraca limativetntris and Thmialspis (=Iosis) flavouicta (66) These insect pests Ilt onlV citise IIigh yield losses tihey also ire difficult to control chemically ofr by other ilealls

ManV upland rice vaieties aic early maturing and are resistant to pests such as gall midge (I Xl ) Early mIturity is often a highly desirable trait to esuilpe pest buildup but it is not true resista nce bca usc ilthose va reties are plinted late they will be itsdamaged as ain early planted susc ible variety

P~erhaps tile cosiiiopolitall of all upland rice pestsimost are the sced pests (ripenling stagc) which pose a particular problem beta use scuds re ised IsIO for hitnit4ans Ayltoxic substatice introd tcld geticticllyilmay also be toxic to nal iniless it caln he placed old in thfe IdhuffSelectmi fo hairiness or lotng wns kill mIlct sthilrohleiiis dutilig threshing pa rtictfarl maitial threshinfl [hireshers ill complain of itcfimin atid itteascd (fust levels

Attempts itIRRI to fitid tcistatncc to the rice bug plorisa have ilotbeCnl prmitisilg lIIetIattonlids iorc

through the lenilna or palca hillhairs sedstVIor seeds wilh awns that deter bugs promote complaints from threshers Seed bugs liptocorisa and Stcnocoris enter the seed through the opening between tile lemna and palC but varieties with narrow openings are diifficult to mill

Breeding priorities should concentrate ott inotCI gruLtLIps that are most difficult to c nttrol by other meains Steil) horers appear to be the most widespread group that sigiiificantly dalag s rice aind is hard to manage by other Ilnca us

Biological control Very fe stutdic of mitatt~l eneties of rice pests in upland envirollnletits wee in tile wefouiid literature Therefore have not compiled a list of species Ve present a list of natural enemies iOtid in upland ricefields tuic latialln Batangas for some conitno iOldiar pest species ([itble 5)

Muich ofl thc lollowinle itfor ittiol cotlliCs rol Stutdies in the lowlatnds or roi crops otithcr thlin tice

Qu(ailtr o 11ll t11t( Nat trrtf lii hcoeits ight Ile used agaitist ile social insects tilt and termites Sollie predator specice live ill theirf nests n ilnic theirnllt ip-pearanIlCe aMRidbehavior IJtrouMdCing a mote aggressise ant species to displace pest ant codonies is a possibility (169) Patihogens as potential natural encnies have reccived little atteition (I13) Natural ncnuics asa group however hac not been toIl iporta it in Iegulating aintshown be and ternite numbers

The nocturnal behavior aid subteitmien habitat of lole crickets and field crickets protect them from many ntllral enemies There are only limited records of natural cnelics for these groups of pests some pathogens (87) but mainly

IRPS No 123 JattuIny I 271987

parasites - scelionids on eggsand sphecids on ilymphs and adults Sphecid wasps Larra carutmaria (SImitI hI L huzwuensis Rohwer and lsaltiuhta Willi ams speciali in mole crickets pa ralyzing them nd dragging the ninto thcir nests as food lot their voting ()tier sphCcid species Lins aurulenoa (Iabricius) lot maiac (Ashmead ) 1 lhshylssellaus (Smith) and Ml lal)oriosu (Snithf specialie in field clickets

White grUbsO the other hand hae Icti scriuns enuigh to have been extensively studied for cotttroi 1W nati al enemies( 112 140 141) Scoliid parasites C(ajsoMcispp have beetn intrtduced itto tilePhilippites for control of Ielxl ]udisiro la The rcsearchcts w Intttrdced themle during ill era claimed that the white gttionthreak as controlled (142) howevcr it isnormal for pest epidemics to subside and there was no diect evidence to indictte that tfe parasite was responsible A tmore promisitg titctllod might he to identify iso ltcttass prt~f tcartd apply hiactelaln fingal pathogens to tile soil Ihe lalattese beetle in North America has hcci coitrolled hv tite conmnleciall aailable hactcrial prcpa ration itacihlpu pav ofr milksdiscas (71) Spote prtyefott the soil itloted ilrtier reitiain actic fr Veas

Root aphid sardtnctly bugs readilslic tilib to predatots and parlsites once their tetndine ants are controlled The larvac and adtilts of a coccinellid beetle 1Scynms sp prey on llriadlouais in slash and burn upland rice Baits treated with insecticide may be used to control tending ants by killing their ourig (244)

Root-tkcditig blugs false wireoris wireorms and root wccils aic r ofsuch little ifilpoltatice that thenomttll role of liatural Cticniics hias tiot beei asscssed A possibility hoevrl tiglt he tit ItaIIitic niaCIMtodcs lintt1g11en Itese soil-itlhlfbitine mCtitodcs W5uhtfhave to be imiass Produiced

large miltibcr of niatltrll cnemics ircknown ft CltIoriS IC IpblCeii is thtt clttvortnlls ofonlie the crop soott alter land ptel)tritiOti a1tithe bcgit iniig of the raittv

Citsoln hfien tiatllral leniiiy populations aric lo Colltloil WOtuld iii ole mass prodtticing andirclcaslng key spccies

le egg stage is the fpirt (olthe Cedlitgt Ialot life cycle tI hich to f)ctis ftor tittlril elctit cottntrol hplraites o1 pleditors ttsit elleggs highly expoIsYCl I Sgg pailsites or predators stolld probahly e ittass andtbe produced released Littlh i toruialioll is a ailable ott what species to trV o ho to Iniss podLice thiCIi A etiloplhid wasp Icmihlaol sp effctiicly regushy

lates the leatf lcloalapolm2a spioanoiiitr heat andc rice antid should be collsctvcd

leaf beetles whCllose e am] eggs ate laid in the grutindhirO ould be motC difficutlt to control by niatulral enelV Mnll ipuhltiOn But species sUici IS c_gsthispid beetles ith

and larVle oilleascs ate vtilcratble to iparaites and pledators

Natural enemisc of tlirips otl rice are little known

28 IRPS No 123 January 1987

Table S Parasites recorded from upland rice insect pests in Tanauan Batangas Philippines 1977-84

Order Family Species Host stage

I lymttenoptera Braconidae Leaffolders Cnaphalocrocis Marasinia spy

Cardiochilesphilipplinensis Ashmead a Larva

Cotesia angustillasis(Gahan)Cotesia or cypris (Nixon)

Larva Larva

Cotesia or taeniaticornis(Nixon) Larva Cilonusinotakatae Munakata Chelonits SppAlacrocetrts philippinensis (Ashtmead)

Egg to larva Egg to larva Larva

Iclineumonidac Orgilts spp Isclutojppa hireator(Fabricius)

Larva Larva

Itoplectis naraigav(Ashmcad) Larva Tenelhtcha philippiensis (Ashmead) Larva Tenelchha stangli (Ashmead) a

irichomma cnaphalocrosis Uchida Larva Larva

Bethylidac Elasmidac

Goniozms or trianguli r Kieffer lasmtusalhopictus Crawford

Larva Larva

Diptera

Encyrtidac Chalcididac

Tachinidac

Elasmtus spp Copidosomopsisnacohiae (Fidy) Brach nteria evcarinata Gahan Biachyieria lasus (Walker) Ztgohothria ciliata (Wulp)

Larva Egg to larva Pupa Pupa Larva

Argyrophylx nigrotihialis(Blaranov) Larva

l-ymenoptera Braconidac Yellow stern borer Scirpopiaga incertulas

Braco chinensisSzeplegeti Larva CIthelotis spp Chelonus niunakatae Munakata Cotesi flaripes Cameron Stenoracon nicerilli (Bingham) a

Larva Egg to larva Larva Larva

Ichneumonidac Tropobracon schoenobii (Viereck) Amatroinorpiaaccepta metathoracica(Ashmiead)Frihorus sinicus (I lolmgren)

Larva Larva Larva

Ischnojoppaluteator (Fabricius) Larva Isotima nr dammeratiti Rohwer Larva Tenelucha philippinensis (Ashmead) a

Larva Tenehicha stangli (Ashmcad)Trichomma enaphalocrosis Uchida

Larva Lara

Chalcididac Eulophidac Trichogranmmatidae

Xanthopimnpla stemmator (Thunberg) Charops hrachYpterumn Gupta and Malteswary Brachtm eria sppTetrasticttus schoenohii Ferriere Trichorigranm chihunis Ishi

Larva to pupa Larva Pupa Egg Egg

Diptera

Scelionidac Pteromalidae Tachinidae

Tt ichograintnta aponiicuitt (Ashniead) Telenomits rowani (Gahan) a

Trichoinalopsis apattelctena (Crawford) Jeirhaea spp

Egg Fgg Egg pupa Larva

Z4gobothria ciliata (Wulp) Larva

Ilymenoptera lecneumonirlac Gol fringed borer (hilo auricilius

Trichomm enoptalocrosisUchida Larva

Ieioenoptera Braconidae

Aantholimlh steiniator (Thtinberg) a

Pink stet Ioret Sesamia inferens StoInacoil nicrilhi (Bingh am)

Loarva to pupa

larva

Diptera Ilyinenoptera

Armvtnyorms Mythimna separata Spodoptera mauritia acronyetoidesTachinidae Zvgohothriaatropiora( Robinea i -Desvoid y)Eulophidae It-phctrtiscitapadoe (Ashimead)

Larva Larva

(halcididae Braconidac

Brachpin ria spip (oesia spp

Larva larva

IHyntenoptera I)iptera

Rice skipper Ielop ida (halcididae Tachinidae

math ias aidgreen iortedci ciliarNIelan itis eda Brat lhincria sp iii matuqitatt Cameron Irgyrolhirlaxi trotlhiais(marumtv)a1

ismtne Larva Larva

Diptera Ilymenoptera

Tachnidae lelineutionidae

fuown se tilo per Mois frugalisIrgtrophrlax 1igrothiahiv 3 a rat ov) anthopimpa pmnctata (Iabricius)

Larva Larva to pupa

aMost dominant

IRPS No 123 January 1987 29

Normally predatory thrips prey on herbivorous thrips strepsiptera on both Spiders dominate tilenymphaladultNatural enemies of stem bugs --Scolinoplara and Tibraa predators Spittle bugs move to rice from pasture grasses - are in order of importance egg par~isites and predators and biocontrol would be successful only ifcarried out before fungal pathogens and nymphal adult parasites Egg tiley immigrate into ricefields parasites could either be introduced fron other areas or Biological control tactics for hemipterous seed bugs mass produced and released ould follow those outlined for stein bugs

Armyworms are normally lwld in hcck by the activities Coml)arison of environnewtns Natural enemies are of egg and larval parasites When these natural enemies fail perhaps faced with even greaterconstraints than insect pestsusually because of drought armyworns become epidemic to survive ind reproduce in upland environments TheyIt may be worthwhile therefore to release parasites during have adapted ani survived by having wide host ranges the the rainy months following a drought Virus diseases of ability to aestivate or hibernate over unfavorable seasonslarvae would beanotheravnue to exploreforarnyworms and the ability to disperse or by other mechanisms

Parasites have ben traditionallyconsidered for biolegical Comparing the most dominant and therefore the most control attempts against stein borers usually by rekasing adapted species of natural enemies collected in various exotic species( 166) This approach has not met with success environments in the Philippines we can understand how and new avenues should be explored The role of predators different rice environments are arid pathogens is little inderstood and deserves greater Anagqrms olptahiisa mymarid planthopper egg parasiteattention Egg parasitism rates are normally reasonably is cquallyadapted to all Philippine rice environments (Tablehigh but should be supplemented with effective egg 6) It is selective for planthoppers and even parasitizes eggspredators Orthopterans are potential egg predators of the n-iize planthopper Peregrinitsmaidis itt upland areas Metioche vittaticollis (Sta ) Anavipha nifiwnnis (Saus- Alavohis also parasitizes maize planthopper eggs but is sure) and Conocephahs hongipnni (de i-laan) feed on most prevalent on BPH in the irrigated wetlands (onashystem borer eggs etioche and Anaxil)ha specialize in eggs tocerus spp like A optalhilis on BPH and WBPI Iattacks with no hair such as Chilo spp The omnivorous (ono- the eggs of GLH in ell rie environments It is also found cephahsspecializes in eggs cov red with hairs it is a proven parasitizing the eggs ef the white rice licahopper (oimna egg predator of Scirlouphaga spp but has the discouraging spetra Oirosita naias a trichogrammatid egg parasitehabit of eating rice guin (65) attacks BIPH and WBPH in the wetlands and has no other

Grasshoppers have egg (scelionid) and nymphalad ult host A related species 0 aesopi specializes in G1-H in (nemestrinid tachinid and sarcophagid) v irasites Among wetland environments Dryinid strepsipt-an and pipunshypathogens protozoans have been recorded most frequently culid nymphal and adult parasites of planthoppers and Control of locust species by natural enemies could con- leafhoppers also specialize either in hopper species or centrate on their habitual breeding areas environments Among the dryinids Pseuhdgonatoipsflavi-

Leaffolders have rich complexes of natural enemies some feimur prefers the lowlands and BPH while P nuch has no adapted to upland rice (22 27) ranging from egg predators environmental prefirences but is adapted only to WBPH (gryllids coccinellids) to larval parasites (braconids ichncu- The main d ryinid species on GLH aptogonatop)usspp is monids) to larval prcdators (ants and carabids) to larval most dominant in rainfed lowlands and also attacks other pathogeiis (vira and fungal diseases) Efforts to augment leafhoppers within Amnrasca Cicadulinaand Bachaltuathese natural enemies should focus on the egg predators Strepsiptera are more dominant on plarithoppers than (rearing releasing)and viral or fungal pathogens (culturing lealhoppers The Elenchus species in the Philippines attack spraying) all species of planthoppers within the genera Sogawilla

Parasitization of the large conspicuous larvae of Sogatodels NilaparsataHarualia and Opiconsiva Elenshyskippers green horned caterpillars and other polyphagous cusrvasuntaisuiis adapted to the major rice environments Lepidoptera by tachinids and chalcids is normally low ELg but prefers BPH in rainfed arid WBPIH in wetland environshyparasites and predators are perhaps the key to their control ments Ilalictoph(utsmtnroeiand HVpectrus attack G LH

Gall midges are parasitized by pterormalids and platy- and COfimravcctra respectivelygasterids that aestivate within their larval host between Pipunculids are mainly parasitic oii lealhoppers Each seasons Shoot aphids are preyed upon by coccinellids species has a unique environmental preference Tomosshysyrphids and chrysopids varireIa oraetoraprefers Ne7 otetltix nigropictrs and

Planthoppers arid leafhoppers have egg parasites and Pilunwulhtsjavanensisattacks Dehoceplhahls spp which predators as well as nymphialad nlt parasitesand predators are more abundant in upland than irrigated lowland their rich complex of natural enemies includes fungal environments T suhvirescensattacks Nephotettix virescens pathogens The principal egg parasites are myrnarids and and prefers the lowlands Pilpuiulus nttillattls is an trichogramniatids Egg predators are norinally mirid plant upland rice parasite arid attacks only Nephotetix speciesbugs that suck out the yolk Nymphal ad nIt parasites are More than 17 recorded species of leaffolder parasites in dryinids on planthoppers pipunculids on lealhoppers and the Philippines attack species of C(naplwocrocis and

30 IRPS No 123 January 1987

Table 6 Dominant natural enemies of rice pests in upland rainfed lowland and irrigated lowland rice environments of the Philippines

Natural enemy

Host stage Species

Brown planthop-Parasite Egg

MymaridaeAnagrus optabilis(Perkins) Anagrusflaveolus Waterhouse

TrichogrammatidacOligosita naiasGirault

NymphAdult Dryinidac

Pseudogotatopuslaiifemttr Esaki et Hashimoto Elenchidae

Elenchus yasurnatsui Kifune and Hirashima

Environment

Upand Rainfed lowland Irrigated lowland

Ailaparata lugens

x x X x

x

x x

x X X Whitebacked planthoppers Sogatella furcifera and Sogatodes pusanusEgg

MymaridaeAnagrus optabilis(Perkins)

TrichogrammatidacOl~gositanaias Girault

NymphAdult Dryinidae

Pseudogonatopusnudus Perkins Elenchidae

Elenchus yasutatsui Kifune et Hirashima

Parasite Egg

MymaridaeGonatocerus spp

TrichogrammatidaeOligositaaesopi Girault

NymphAdult Pipunculidac

Tomosvaryella subviresccns Loew Tomosvaryella oryzaetora (Koizumi) Pipunclusmutillatus (de Meijere)

DryinidaeIlaplogonatopusspp

Egg Copidosomopsisnacoleiae (Eady)

Larva BraLonidae

Cardiochiles philipphnensisAshnead lchneumonidac

Trichiomnia cnaphalocrosisUchida Temelucha stangli (Ashmead)

Pupa Chalcididae

BrachymeriaexcarinataGahan Brachymerialasus (Walker) Brachymeria nr tachardiaegroup

Parasite Egg

Scelionidae Telenomus rowani (Gahan)

EulophidacTettasticusschoenobhi Ferriere

Larva Braconidae

Stenobraconnicevillei (Bingham) Iclhncurnonidac

Temelucha phiiippinensis(Ashinead)Temelucha stangli (Ashmead)

x x x

x x

X x X

x x Green leafhopper Nephotettix virescens

X x x

x x

X X X X

X

X Rice leaffolders CnaphalocrocisMarasmia

x X x

X

X X

x x x x x X x x x

Yellow stem borer Scirpophaga incertulas

X X x

x

x

x X XAinauromorphaaccepta metathoracica(Ashmead) x

continued on next page

IRPS No 123 January 1987 31

Table 6 cntinued

Natural enemy Environment

Host stage Species Upland Rainfed lowland Irrigated lowland

Larva-pupa Gold fringed borer Chilo attriciiuslchneumonidae

Xanthopimpla stemmator (Thunberg) x aPupa x

Pteromalidae Trichoinalopsis apanteloctena (Crawford) ax

Pink stem borerb Sesamnia inferensParasite

Larva Braconidae

Stenobracon nicevillei (Bingham) x b b Armyworms Mvthinna SpodopteraLarva

Tachinidac Zygobothriaarroptvora (Robineau-Desvoidy) x x

Braconidac Cotesia spp

X Skipper Pelopidas mathias and green horned caterpillar Melanitis leda ismeneF-gg

TrichogramrnatidacTrichogrannaspp x

Scelionidae Undetermined Scelionid

x Larva

Tachinidae Argyroph lax nitrotibialis (Baranov) X x

Pteromalidae Trichomnalopsis apanteloctena (Crawford)

X Predator

CoenagrionidaeAgriocnenis femina femina Brauer x xIschnura senegalensis tambur

Miridae CrtorhinsliidipennisReuter x x

Nabidac Stcnonabis tagalicus (Stal) x

VeliidaeAlicrovelia douglasi atrolineata (Bergroth) x

Coccinellidae Micraspis crocea (Mulsant) X x x

Vespidae Eumenes campaniformis (Fabricius) x

Formicidac Diaecamna spp xOdontoponera transversa (F Smith) x

Carabidae Chaenius spp OphioneaW- aJasciataSchmidt-Goebel

x

Ophionea ishii ishii Habu x x X

Gryllidav Anaxipta longipennis (Saussur-) x xMetioche ittaticollis (Sial) x x x

Tettigoniidae Conocephalus longipennis (de Haan) x x x

aNo rearing made on rainfed lowland bNo rearing made on rainfed lowland or Irrigated lowland

MrasmiaAmong the Laval parasites the braconid Cardio- lowlands and parasitizes oriental maize borer OsIriniachilesphilijpphensis is most adapted to an upland environ- furnacalis (Guenee) larvae The leaffolder pupal parasitesment It also parasitizes Hydelepta indicata a common not only occur equally in all environments but also haveleaffolder of legumes The ichneumonids Trichomma cna- wvide host ranges- parasitizing species of OsIrinia Marucaphalocrosis and Temehcha stangli occur more in the Hydelepta and Honiona - in maize and legumeslowlands T stangli is most adapted to irrigated lowlands The principal egg parasite of yeltow stem borer (YSB)and also parasitizes Chilo and Scirpophaga stem borer Telenomus rowani is widely adapted to all environmentslarvae T cnaphalocrosis is most prevalent in rainfed and has an unusual alternate host -Tabanus eggs (25)

32 IRPS No 123 January 1987

Tetrastictu schoenobii specializes only in YSB and occurs mainly in irrigated lowlands -- further proof that YSB is most adapted to the wetter environments

Larval parasites of YSB are highly environment-specific The braconid Stenohracon nicevillei occurs only in upland rice but also attacks the pink stem borer (PSB) Tieinchwa phiiiplPwnsis an ichneumonid occurs in upland rice and also parasitizes the larvae of (7hiu stqpresaisinatro-morpha accepta melaihoracica like 7 stangi is mnost adapted to the irrigated lowlands

The dark-headed stem borer(l)HSB) (ih pochit3sus was not collected in all rice environments and the gold fringed borer Chilo auricilius nd PSB were collected only in upland rice Xaantiholiplastenrnaloran ichneumonid larval parasite appears to be widely adapted whereas TrihomaslopisapanitI)cteiiaa pteromalid pupal parasite occurs mostly in wetlands and also parasitizes larvae of Chilo suppressais and Pelhuidas mathias

Armyworms Myit hinin and Spodujipera have two main larval parasites - a tachinid Zrgohothriaatropivora in rainfed environments and a braconid Cotesia spp in irrigated lowlands

The butterfly pests Pelopidas and 4elanitishave tricho-grammatid egg parasites found only in irrigated lowland environments

Two larval parasites show environmental preferences --Argrroph av nigrotihialis a tachinid in rainfed habitats and Trichonialopsisapantelkcnela in irrigated ricefields A limgrotibialisalsoattacks the sweet potato hornwormAgrius convolvtdi and T apanteloctena parasitizes DH S1B

Predators as a group show more distinct environmental preferences The aquatic damselflies Coenagrionidae are most abundant in the lowlands Cvriorhinus is also most adapted to the lowlands The nabid Stenonabis tagalicus occurs only in the uplands and isalso prevalent on legumes The coccinellid lady beetles wasps and ants are more prevalent in upland areas AlicrasIpis cit-ea larvae prey ona variety of aphids attacking legumes and maize Etunienes campanonia vespid mud wasp makes nests in trees and is therefore most adapted to the more botanically diverse uplands Soil-dwelling ants cannot tolerate flooding

Upland environments are habitats to arboreal carabid beetles Three species of Chlaeniusbeetles prey on leaffolder larvae and are perhaps more important than parasites (27)

The gryllids and tettigoniids are widely adapted egg predators not only on rice but also on maize and legumes They feed on eggs of most insects that are laid on leaves

The spidercommunity of upland rice environments isrich in species At one site (Batangas Philippines) 31 species have been recorded (23) Of the 176 spider species recorded in Philippine ricefields about one-half (82) occur in upland rice The spiderspecies of upland rice environments overlap more with those of rainfed lowland than with those of irrigated lowland rice (24) The same study showed that of the three environments irrigated rice has (ie greatest spider species diversity foiiowed by rainfed lowland and upland

However of the 10 most prevalent ricefield spiders 9 were abundant in upland rice showing wide adaptation All three spider guilds -- orb-web space-web and huting spiders shywere prevalent in upland ricefields There were however differences in environmental prelerences for some spider species

Of the orb-wcavers Tetragniathamanlihulatawas partishycila rlv abundant 7 japonica was low and Ltucatge dehcorata was absent in upland rice

As in other environments Aiypeiwnaifirtosana was the most dominant space-web spider however its relative numbers were lowerthan for lowland sites Among hunting spiders Ox|opes javanus and Lcosa leatcost fgla were more abundant and Lvcosa lpseudoanmnlata was less abundant in upland compared with lowland rice envishyroniments

Extensive rearing of rice insect pests in upland rainfed lowland and irrigated sites over an 8-yr period in the Philippines provided insight into the effectiveness of parasites as natural enemies (Table 7 8) Most of toe data are from large sample sizes taken over at least one crop which overcomes the pitfalls expressed by Van Driesche (238)

VSB egg parasitization was surprisingly similar across all environments Larval parasitization was highly dynamic among sites and even years within the same site but showed somewhat lowerlevels in uplandsthan in irrigated wetlands This would have been predicted for egg parasites as well since YSB is adapted to the lowlands and so must be its parasites This isevidence that Telenomus rowani is as good at dispersing as its main hosts Trowani clings onto the anal tufts of female YSB and parasitizes eggs as they are laid This phoresy may explain its wide adaptation The more specialized lowland YSB larval parasites therefore appear more effective than their upland counterparts with more alternate hosts

Low collection levels of PSB larvae especially in the wetlands make comparisons between environments difficult Low levels of larval parasifization were recorded in both rainfed environments Except in the Cagayan rainfed wetland site in 1980 low levels ef larval parasitization were evident at irrigated and rainfed sitesalike No differences by environment were evident Higher levels of parasitization occurred in Sarawak during a 1967 outbreak also from tachinids Rothschild (197) indicated that higher parasishytization occurred in upland rice areas

Skipper and green horned caterpillar larval parasitization rates were generally low and showed no environmental effect htighest activity occurred in Cagayan in 1980 (48 parasitization) Although larval parasitization levels of RLF were variable year to year activity appeared greater in the irrigated lowlands than in the uplands Again the highest levels were recorded in Cagayan in 1980 Cagayan has extensive grasslands and fallow around the rice areas and a relatively long rainfall period Rainfall in 1930 was good for crop growth and apparently for increase of larval parasites

IRPS No 123 January 1987 33

Table 7 Parasitization rates of rice planthoppers and green leafhopper in upland rainfed lowland and irrigated lowland environments Philipshypines 1976-840

Environment and site Year Egg parasitization () Nymphaladult parasitization ()

BPH W13P GLH BPII WBPH GLIH

UplandBatangas

Rainfed lowland

1976 1979 1980 1982

15 17 12

2(47)8(39)

4(51)

9(1800)14 (212)

12(304)

8 (101)6 (127)

9(116)

Iloilo

Cagayan

1976 1977 1978 1981 1980 1981 1982

0 (69)

11

44 48 40

-

-

46 55 44

0 (471)

11

52 56 49

0 (200)9 (214)

10 (1259)8(235)6 (900) 3(800)

-7 (462)

11 (270)3 (313)5 (900) 1 (800)

4 (600)10 (725)10 (362)12(870)5 (900) 3 (800)

Pangasinan

Irrigatedlowland

1984 1976 1978 1979 1982

13 (436) 13(379) 14 (457)

6(14)7 (197)

17(198) 11 (486) 18 (275)

3 (81)10 (274)3 (116)

16(871) 19(88)

28(211)9 (764)5 (647)7 (1204) 8(305)

Laguna 1977 11 7 21 1978 31 1979 14 4 4 28 1980 7 8 20 1981 1983

56 7(278)

26 16 (1650)

16 27(1921)

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R I Apostol unpublished 1lo1lo and Pangasinan 136 Cagayan BCanapi unpublished Liuna 175 I C Pantua unpublished 118 234 43 Carino and Shepard unpublished BPH = brown planthopper Nila parvara htgens W131Pi =whitebacked planthopper Sogatelia furcifera GLH = green leafhopper Nephotettrix virescens Figiires in parentheses are sample sizes

Tauher et al (230) reported that internal external and genetic factors influence the seasonal activity of parasitoids

Hopper egg parasites but not nymphaladult parasites were also parlicularly abundant at the Cagayan site possibly the egg parasites have more alternative hopper prey in the weedy areas than nymphaladult paisites

The seasonal dynamics of hopper egg parasites can be seen for upland rice in Batangas(Fig I 2) and for rainfed lowland lice in Pangasinan( 135)and Cagayan( 105) Levels of BPHWBPI-L and GLH egg parasitization were corn-parable in upland and irrigated lowland environments Except for some high rates of paraMsitization early in the season the parasitization rate tended to be steady in all envilonments However hopper nymphaladult parasi-tizution levels were more similar in the rainfed environments and lower than in irrigated rice

Overall parasites do not appear highly effective by themselves against upland or even lowland pests Among natural enemies parasites have attracted the most attention because they can be more readily assesscd

Predators and pathogens lend themselves better to management practices Predators the most important group of natural enemies in riccfields are difficult to quantify On the other hand pathogens appear less im-portant but can be readily cultured and disseminated

The natural enemy commulnity of upland rice is rich in species and differs significantly from that of lowland rice

These beneficial organisms must be conserved by applying insecticides judiciously particularly sprays Seed or soil placement of chemicals minimizes exposure to natural enemies The strategy to derive the greatest benefit from natural enemies isto allow the greatest number of beneficial species to thrive Then perhaps at least one species will be effective against each pest at any tine overcoming the variable abundance of each species within and between years

Programs to introduce exotic species or mass produce indigenous ones are ambitious and expensive because they require trained people on a sustained not ad hoc basis

Chemical control Insecticides offer rapid and efficient control of upland insect pests (190) but-should be used only after other control measures have been considered Insecticides are rarely used on upland rice because of cost (32) Upland rice yields are normally too low to justify the expenditure Also most upland rice is grown as a cash crop Spraying upland rice is more difficult than lowland rice because water is less accessible Government subsidy programs to provide pestishycides to tribal peoples for upland rice production have created problems of toxicity to humans because isolated people have had no experience to be able to handle pesticides safely

However ins-eticide use has beenjustified when a crop is

0Z

Table 8 Parasitization rates of lepidopterous pests in upland rainfed lowland and irrigated lowland environments Philippines 1976-84a

Parasitization (5)

Environment and site Year YSB GFB PSB Earcutting Swarming Skipper Greenborned RLF Brown larva larva caterpillar caterpillar caterpillar caterpillar larva semilooperEgg Larva larva larva larva larva larva

UplandBatangas 1977 - 0 (57) - 0 (63) - 0 (71) - 0 (106) 11(142) 2 (101)1978 43 (4697) 8(451) 5 (38) 3 (40) 6 (61) 2(58) 15 (57) 13 (129) 13 (1286) 4 (30)1979 59(4135) 12(320) 13 (11) 5(28) 9(45) 6(46) 7(23) 8(160) 9(1152) 8(26)

Rainfed lowland Iloilo 1976 - 0 (10) 0(10) 0 (7) 0 (19) 2 13) 15 (180) 5 (76)1978 0 (54) 0 (3) 0 (17) 9 (64) 4 (78) 3 (73) 0 (121) 0 (191)1979 64 (2350) 16 (84) 6 (17) 0(6) 18 (68) 25 (104)

1981 56 (1322) 18(304) 0(5) 12(51) 18(138)Pangasinan 1976 60 (7 7 )b 62(82) 0(111) 5 (39)

1978 58 (53)b 5(51) Cagayan 1979 59(713) 8 (101) 6 (34) 0(7)

1980 0(11) 4 (28) 15 (286)

62 ( 48(177) 61(140)1981 0(5) 12 (2) 13(45) 14 (92)

Irrigatedlowland Laguna 1974 62 8

1974-75 50 41 1975 43 72 1978 96 (623) 21(24) 33 (60)1979 83(2017) 18(110) 8(62) 11 (89) 22(400)1980 78 (1625) 20(75) 0 (16) 4 (72)1983 60

9 (80) 7 (270) 30

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R F Apostol unpublished Iloilo and Pangasnan 136 Cagayan B Canapi unpublished Laguna 175 P CPantua unpublished 118 234 143 Carino and Shepard unpublished YSB = yellow stem borer ScirpophagaincertulasGFB = gold fringed borer Chilo auricilius PSB = pink stem borerSesamia inferens earcutting caterpillar Alythimna separata swarming caterpillar Spcdoptera mauritia acronyctoides skipper Pelopidas mathias reen horned caterpillar Melanitls ledaismene RLF = rice leaffoldersCnaphalocrocismedinalis and Marasmiaspp brown semilooper Mocisfrugalis Figures in parentheses are sample sizes Egg masses

IRPS No 123 January 1987 35

Table 9 Effect of seeding rate and insecticide on plant density and Yield losses from insect pests in upland rice often areyield of Dagge upland rice Batangas Philippines 1976a comparable to those of lowland rainfed or irrigated rice

Seeding rate Insecticide Dosage Plant density Yield (0-Y30o) but usually from different pests The principal (kg aiha) seed treatment (kg aiha) (noIra-row) (tha) groups are soil pests feeding on sown seed and rootsat 14 DE followed by seedling pests These groups have largely gone

50 None - 35 a 20 a unnoticed leading researchers to conclude that stem borers50 Carbofuran F 10 38 ab 23 a and seed bugs are the most important pests (17) Yield loss 10 beCabofuranF100 None - 4551 c 23 a trials that pinpoint major pests need to includc seed

100 Dieldrin WP 10 52 cd 22 a treatments Granules applied in the furrow may not give150 None - 59 d 22 a adequate protection from many seed pests

aAv of 8 fields DE = days after crop emergence In a column means The floristic and pedological diversity of upland rice followed by a common letter are not significantly different at the environments is matched by the diversity of arthropods5level by D)MRT F = flowable Wi = wettable powder both pests and their natural enemies Irrigation has homoshy

genized lowland rice environments Flooded soils tend tothreatened by epidemics of locusts or armyworms New have the same properties and to eliminate soil pests The technology stressing efficiency has minimized insecticide spatial and temporal dominance of cultivated rice has cost benefited specialist pests

Seed treatment is normally inexpensive and can be Upland rice has no specialist arthropod species The economicallyjustified in -nanycases(66) Treating seed with monophagous pests on upland rice prefer the lowlands systemic insecticide (216) but not chlorinated hydrocarbons where rice evolved Rice in the uplands exists only because (247) is most effective in controlling seedling maggots of man and even though lowland rice predates upland riceFoliar sprays have to be repeated to obtain control and specialist species evolved Wheat maize and sorghum do often are washed off by frequent early season rains not have monophagous pests leading us to conclude that Granular insecticides are effective but required dosages are there is no evolutionary advantage for inssects to have too expensive to justify except under heavy infestation narrow host ranges in upland graminaceae(247) When directed against first-instar larvae banding Pests have adapted to the uplands not only by having granules in seed furrows (177) can be effective against white wider plant host ranges but also by having long life cyclesgrubs at low dosages (025 kg aii ha) (134) Seed treatments undergoing dormancy or being dispersive Each pest has a are not promising for white grub control (248) unique life history strategy suited to highly unpredictable

Seed treatment is also effective ainst ants and may be environments In agivenyear someinsect specics an better cheaper than increasing the seeding rate In Batangas exploit upland rice as a food source but no pest is a senious treating 50 kg seed ha was as effective as increasing the one every year on upland rice seeding rate to 100 kgha and using untreated seeds (Table The low yielding potential of upland rice means less 9) In Brazil seed treatments control termites and other soil quantity and lower quality of food for pests to exploit but insects including the lessercornstalk borer(66) Domiciano also less incentive for farmers to undertake control (58) however found seed treatments inconsistent in their The most effective control would be early and synchroshyeffect against high populations of lesser corn stalk borer nous planting of early-maturing upland rice varieties to

Baiting can be an inexpensive way of controlling ants prevent insect pest buildup Breeding for insect resistance seedling maggot flies mole crickets and field crickets Baits should initially focus on stem borers because they are verymade of locally available material and impregnated with difficult to control by other means Biocontrol efforts insecticide can be sparingly distributed taking adantage of should first stress conservation of natural enemies preshythe pests ability to disperse and encounter bait sites dators and pathogens rather than parasites Efficient

The new low-volume hand-held controlled-droplet chemical control includes seed treatment or baiting for seed sprayers may offer an advantage over high-volume knap- seedling and some root pests and using controlled dropletsack sprayes for controlling foliar pests Upland rice sprayers for spot treatments against foliar pests and seed canopies are more open than those in lowland rice and bugs when economic thresholds are reached better droplet penetration should result Controlled droplet applicators would be ideal for seedbugs and defoliators

CONCLUSION

Upland rice is attacked by a wider array of insect pests than is lowland rice mainly beca use of the addition of soil pests Generally population levels of lowland rice pests are lower in upland than lowland rice leading many people to conclude that insect pests are not important on upland rice

36 IRPS No 123 January 1987

REFERENCES CITED

1Abraham C C and K S R Mony 1978 Occurrence ofLeptocorisa acuta Fabr (Coreidae Hemiptera) as a pest of nutmeg trees J Bombay Nat ttistSoc 74553

2 Agyen-Sampong M 1975 HlierogrlphusdiaganensisKrauss (Orthop-tera Acrididae) a new pest of rice in Northern Ghana Ghana J Agric Sci 8249-253

3 Agyen-Samnpong M 1976 Varietal resistance and other inle-ratcd pest control components as factors of rice varietal improvement in West Africa Prcsented at the Second varietal improvement serinar 14-18 Sep 1976 Bouak Ivory Coast West Africa Rice Development Association Monrovia Liberia 18 p

4 Agyen-Sampong M 1977 Outbreak of Ihccrorrdnisorae in rice seedlings near Rokupr Sierra Leone lit Rice Res Newsl 2(6)16-17

5 Agyen-Sampong M 1980 Dirty panicles and rice yield reduction caused by hugs IntRice Res Newsl 5(l)11-12

6 Agyen-Sampong M 1982 The major pest problems of irrigated upland and mangrove swanip rice ecosystems ir the humid tropical Guinea savanna and Salel climate zones Pages 318-328 in Pro-ceedings training course cn concepts techniques and application of irtegrated pest management in rice inWest Africa 10-20 Jan 1982 Fendall Liberia West Africa Rice l)evelopioent Association Monrovia Liberia 505 p

7 Akibo-Betts V I and S A Raymundo 1978 Aphids as rice pests in Sierra Leone lit Rice Res Newsl 3(6)15-16

8 Akinsola F A 1975 Present status of different rice stemborers in parts of Nigeria iicL Entomo Newsl 328

9 Akinsola FA 1979 Ihe biology and ecology of rice stemborers in Nigeria Ph 1) thesis University ol lbadan Nigeria 217 p

10 Akinsola E A 1979 Guidelines of integrated management of insect pests of upland rice in West Africa Presented at the Seminar on integrated management of rice diseases and insect pests at Bobo-Dioulasso Upper Volta 17-22 Sep 1979 West Africa Rice Develop-ment Association Monrovia Liberia

II Akinsola IA 1984 Insect pests of upland rice in Africa Pages 301-305 in An overview of upland rice research Proceedings of the 1982 lIouak Iwsrv Coast upland rice workshop International Rice Research Institute 13O Box 933 Manila Philippines

12 Alam M S V TJohn and Kaung Zan 1985 Insect pests and diseases of rice in Africa Pages 67-82 in Rice improvement in Eastern Central and Southern Africa proceedings of the international rice workshop at Iusaka Zambia 9-19 April 1984

13 Alticri M A A van Schoonhovcn and J D Doll 1977 The ecological role of weeds in insect pest management systems a review illustiated with bean (Phaseohsvulgaris L)cropping systems PANS (Pest Artie News Summ) 23195-205

14 Anmante F 1967 The leaf-cutting ant Atia capiguaraa pest of pastures [in Portuguese] Biol6gico (Sao Paulo) 33113-120 (Rev Appl Entomol A56207)

15Andrade A C and D Puzzi 1951 Preliminary results of experi-merits on the control of Sculptcoris iastantus on sugarcane [in Portuguese] Bioulgico (Sao Paulo) 1744-49 (Rev Appl Entomol A41421422)

16 Appert J 1970 Maliarplia separaelha (rice stem borer) new observations of entomological problems of rice in Madagascar [in French] Agron Trop (Paris) 25329-334

17 Ariadeau Mand Z Harahap 1986 Relevant upland rice breeding objectives Pages 189-197 in Progress in upland rice research Proceedings of the 1985 Jakarta conference International Rice Research Institute 1O Box 933 Manila Philippines

18Asar S and Kisimoto 1975 Seasonal occurrence of the smaller brown planthopper in upland rice [in Japanese] Proc Kanto-Tosan Plant Prot Soc 2228

19 Bakker W 1974 Characterization and ecological aspects of rice yellow mottle virus in Kenya Meded Landbouwhogesh Wage-ningen 152 p

20 Balut F F 1970 White grubs attacking rice Oryza ativa L [in Portuguese] Biol6gico (So Paulo) 36321-322

21 Banerjee S N and L M Pramanik 1967 The lepidopterous stalk borers of rice and their life cycles in the tropics pages 103-124 in The major insect pests ofthe rice plant Proceedings ofa symposium at the International Rice Research Institute September 1964 The Johns Hopkins Press Baltimore Maryland USA

22 Barrion A T and J A Litsinger 1980 Ants a natural enemy of cliaiarac in dryland rice IntRice Res Newsl

5(4)22-23 23 Barrion A T and J A litsinger 1981 The spider fauna of

Philippine rice agroecosystem 1Dryland Philipp Entomnol 5139-166

24 Barrion A T and 1 A Iitsinger 1984 The spider fauna of Philippine rice agroccosystcms II Welland Philipp Eiitomol 611-37

25 Barrion A T1 and J A Litsinger 1984 lihanus (Diptera Tabanidae) eggs an alternative host of rice stei borer (SI3) egg parasite Telenoonis digns (Iliymenoptcra Scelionidae) lit Rice Res Newvsl 9(6)19

26 Barrion A T and J A Litsinger 1985 Identilication of rice leaffolders by wing markings Int Rice Res Newsl 10( 1)24

27 Barrion A T and J A Litsinger 1985 Chctiosspp (Coleoptera Carabidae) a leaffolder ILF)predator lit Rice Res Newsl 10(1)21

28 Bertels A 1970 Insect pests of rice and theircontrol [in Portuguese] Pelotas Brazil Instituto de Expcrimentacao Agropccuararias do Sul Circular 43 24 p

29 Bonzi S M 1982 Ctilo (flisilineus J de Joannis (Lepidoptera Pyralidae) stem borer of irrigated cereals in Upper Volta LioFrench] Agron Trop Riu Rizic Cult Vivrieres Trop 37207-209

30 Bourke T V T L Fenner J N LStihick G Baker EHassan D F OSullivan and C S li 1977 Insect pest survey for the year ending 30th June 1969 Papta New Guinea )epartment of Agrishyculture Stock and Fisheries Port Moresby 57 p

31 Bowden 1 1973 Migration of pests in the tropics Meded Rijksfac Landbouwwet Gent 38785-796

32 Breniere J 1973 The main rice parasites in West Africa and their control Pages 27-42 in Proceedings seminar on plant protection for the rice crop West Africa Rice Development Association Monrovia Liberia

33 Breniere J 1976 The principal insect pests of rice in West Africa and their control West Africa Rice Development Association Monrovia Liberia 52 p

34 Brown ES 1962 The African arinyworn Spoiolptera eXI)Ina (Walker) (Lepidoptera Noctuidae) a review of the literature Cornshymonwealth Institute of Entomology London UK57 p

35 Brown F B 1968 Upland rice in Latin America FAO-International Rice Commission 12th Session of the Working Party on Rice Production and Protection 2-14 Sept Peradeniya Sri lanka Food and Agriculture Organization Rome Italy 6 p

36 Buyckx F J 1 1962 Disease and insect pests of crops inCongo Rwanda and Burundi [iil French] Publications dc LInstilut National pour lestude Agrononiquedu Congo M Weissenbruck S A Brussels Belgium 708 p

37 Cariiio F 0 1981 Role of natural enentics inpopulation suppression and pest management of green rice lealhoppers Ph D thesis University ofthe Philippines at Los Bafitos PhilippinesLaguna 136 -

38 Carpente A J 1983 The development of rice cultivation and extension of vegetable productin in Zanzibar Final Report Vol II1 October 1983 Food and Agricultme Organization Ronte Italy

39 Carroll C P and S1 Risch 1983 Tropical annual cropping systems ant ecology Environ Manage 751-57

4(0 Cates R G 198 1 Host plant predictability and the feeding patterns of monophagous oligophagous and polyphagous insect herbivores Oecologia (Berl) 48319-326

IRPS No 123 January 1987 37

41 Cendcafia S M and F B Caora 1967 Insect pests of rice in the 62 Ene 1 C 1981 I)iiorldhol)ierusihilis Slater (licteroplera Philippines Pages 591-616 in The najor insect pests ofthe rice plant lygacidue Illissinac) a new rice pest front Nigeria lit Rice Res Proceedings of a symposium at the International Rice Research Newsl N3)15 Institute September 1964 The Johns Hopkins Press Baltimore 63 Fugenio T S 1965 rhe effect oi e-vCgetation on rice production Maryland USA 729 p and the possibility of overcoming it by leaf pruning MS thesis

42 Centro Ihternacional de Agricklittira Tropical 1984 Upland rice in University ol the Philippines at Los Baijos ILaguna Ihilippins 69 p Iatin Anerica Pages 93-119 hi An os ervies of uplaind rice research 04 Farrow R A 1975 The Alrican nmigratory locust ii is main Proceedings of the 1982 Boinakamp Ivory Coast upland rice workshop outbreak area of the middle Niger qlinlitatise studies of solitary International Rice Research Institute P0 Blox 933 Manila populations in relation to environmental factors Locusla No II Philippines 198 p

43 Chandra G 1979 Taxonomy ad hn riononlics ofthe imect parasites of 65 Fennal R G 1943 Food crop investigation Windward and rice Icalloppers and planthoppers in the Pihilippines and their Ieeward Islands First report Report onl work dunring the periodiiiportairce iii natural biol)gical cointrol Philipp Fntoltol 4119-139 April 1942-April 1943 Imperial College of Tropical Agriculture

44 Chang T T 1976 The originii colut ion enlIivation dissemination Trinidad 16 p (Rev of Appl Fintornol A3222-25) and diersificition of Asian and African rices -uphytica 25425-44 1 66 Ferreira F 1983 Integrated pest ritaageiret of upland rice [ill

45 Chatierjee P B 1979 Rice leaffolder attacks in India lit Rice Res Portuguese English summary] Pages 323-341 in Upland riceculttre Newsl 4(3)21 factors affecting production E Ferreira T Yamada and F-

46 Cheng Cbien-pan 19(4 Upland rice in Taiwan International Rice Malmolta eds Institutoda Potassae Foslato Institnto Internacional Cnmmission Wiorking Party oit Rice Production ard Protection I()th da Potassa Piracicaha Sao Paulo lraid Meeting 3-10 March 1964 Manila Philippines Food and Agricul- 67 Ferreira E arid F dl Silva Martins 1979 Occurrence of ture Organiaion Rome Italy 5 p Sogatohs orvziola (Muir) inupland rice in Goias Ilrail lit Rice

47 Chcrrett 1 M and 1) 1Peregrine 1976 A review of the status of Rcs Ncwssl 4(4)18 leaf-cutting ants and their control Ann Appl Biol 84124-128 68 Ferreira E and 1F da Silva Martiris 1984 Insect pestof rice ill

48 Chipeta F M 1974 Pests of rice in Malawi Pages 90-95 in Brazil and their control Jill Portugucsej FMIlRAtPA-CNPAF Proceedings Sith eastern Africa cereals research conference 10-15 Goiania Brmiil 67 p March 1974 DRB Mialda ed Ministry of Agriculture lilongwe 69 Ferreira FI F ca Silha Miartins S S Nero and 1 It P de Malawi Carvallio 1982 M tltiplc regression analysis todeterimine the effect of

49 Chu Yau-i 1979 The d atiaging period of the rice seedling fly insect populatiois ott upland rice yield [inIPortuguese Pesqui (Atlterigotiavxigu Stcin) (NItiscide Diptera) oil cori) at Fast Java Agropecu Bras 17671-675 Indonesia Plant Prot Bill 21403413 70 Ferreira F 1F da Silva Martins and F 1PZimmermann 1979

50 Conklin H C 1957 IHaninoo agriculture a report on aln integral Rice varietal resistance to tle lesser corn stalk borer [in Port uguese] systent of shi fti rig cult ivationi in the Pltilippincs Food and Agriculture Pesqui Agropccu Biras 14317-321 Organization Rome Italy 209 p 71 Fleming W F 1968 Biological control of lie hapanese beetle U S

51 Cramer H II 1967 Plant proection and world crop production Dep Agric RLs Sers [ech Bull 1383 78 p Pilaniensclhutz-Nachr 201-524 72 Food and Agriculture Organiation of the United Nations 1982

52 l)ani R C and Majunidar 1978 Prelininary observations on tie Country report surmimary Pages 27-37 in Report on the first incidence of root aphids on differeit rice cultnrcs Sci Cult 4488-89 FAOIUN)IPhail and regional training coiuirse in iniproved ctltural

53 Davidson R I 1979 lffects of root feeding on foliage yield Pages practices for upland rice 5-23 Oct 1981 Northern Region Agrculturd117-120 in T K Ci osb and R B Pottinger eds Proceedings of the Developririt Center Chiang Mai Food and Agricuilture Organiiashy2nd Australian conlerence on grassland invertebrate ecology Govt tion Rome Italy 37 n Printer Wellington New Zealand 73 Franco do Ariaral S 195 1 Residual action of insecticides in turfy

54 l)ean G 1W 1976 Rice insect pests inlaos Int Rice Res Newsl soil--the results of a field experiment [in Portuguese] Biol6gico (Sao 1(2)15 Paulo) 17209-214 (Rev Appl Entomol A41423)

55 I)ean G J W 1978 Insect pests of rice in Laos PANS 24(3)280-289 74 Freeman ) 1970 Report onithe Iban london School of -coroiics 56 )ingle H 1984 Behavior genes and lifc histories conplex adapta- Monographs on Social Anthropology No 41 Athlane Press

tions to uncertain environments Pages 169-194 in A new ecology London UK 317 p novel approaches to interactive systems P W Price C N Slobod- 75 Fuchs T W1 A Harding and 1W Smithlr 1979 Induction and chikoff and W S Gaud cdsl ohn Wiley New York USA termination oldiapause in the sugarcane borer Ann Flitonlol Soc

57 Dobelmann-JP and M Falais 1968 Upland rice cultivation ill tie Am 72271-274 North-western part of Madagascar in 1966-1967 [in French English 76 (hbriel B1) 1975 A review tf the major insect pests of some tpland summary] Trop Agron 2313-21 crops in tlc Philippines Pages 86-111 in Ad Htoc panel of experts on

58 Doniiciano N L 1979 Chemical control of tie lesser corn stalk pest disease arnd weed problems in some rainfed crops Bangkok (lolsonno~paloIgnowslhts) (Zeller 1848) (I-epid(ptera Ihycitid ae) Thailand Food aid Agriculture Organization Roire Italy

on rice in Ponta Gressa Parara 1977-1978 [in Portuguese] Pages 77 Garg D K and N K Shal 1983 White grub outbreak on rainfed 291-303 in Anais 1 Reuniao de tecnicos cmiirizicultura do estado de dryland rice in Uttar Pradeslh Int Rice Res Ncwsl 81(4)16-17 Sao Paulo-Campinas Cati Brazil 78 Garg I) K and IP Tandon 1982 Major insect pests of rice on hilly

59 Dunsmore 1 R 1970 Investigations on tlie varieties pests and tracts of Uttar Pradesh India lilt Rice Res Newsl 7(11)11-12diseases of upland rice (hill padi) in Sarawak Malaysia Int Rice 79 Garrity 1) P 1983 Upland rice research an international bibliograpy Comm Newsl 1929-35 1975-1982 Iternational Rice Research hnstitnte PO Box 933

60 Elias 1R1965 The new decline of rice seedlings is not caused by Manila Philippines 121 p parasites [in Portuguese] Biol6gico (Sao Paulo) 313-6 80 Ghauri M S K 1982 A new subspecies of Iinorh)rone bull

61 Empresa Brasiliera de Pesquisa Agropecuaria 1984 Upland rice itt cornlus Slater (Hemiptera Iygaeidae) front Papua New Guinea on Brazil Pages 121-134 it Av overview of upland rice research rice and carpet grass Ihll lEntoniol Res 72133-137 Proceedings ofthe 1982 Blouak Ivory Coast Upland rice workshop 81 Gonzales F 0 PArregoces R I ihernandef and OT Parada International Rice Research Institute PO Box 933 Manila 1983 Insect and imite pests of rice arid their control in Latin America Philippines [in Spanish] la Federacion Nacional de Arrocerts (Fedearroz)

Bogota Colombia 60 n

38 IRPS No 123 January 1987

82 Grist D H and R J A W Lever 1969 Pests of rice Longmans London UK520 p

83 Hale P R 1979 Two subterranean pests of upland rice in Papua New Guirnca IntRice Res Newsl 4(2)16-17

84 Hale P R and J T Hale 1975 Insect pests of rice in the New Guinea Islands Rice Entomol Newsl 34

85 Harris W V 1969 Termites as pests of crops and trees Common-wealth Institute of Entomology London UK 41 p

86 Hattori 1 1968 A method to distinguish between grass leaf roller (Ctaplha rocis medinalisGuenee) and the rice leaf roller (Susurtia exigua Butler) [in Japanese] Plant Prot (Japan) 23167-170

87 Henry I E and 1A Onsager 1984 Experimental control of the mormon cricket Iahrosshnph by NJsena locustae(Aficrospora Microsporidae) a protozoan parasite of grasshoppers (Ort Acri-didae) Entomophaga 27197-201

88 Hidaka 1 P Vungsilabulr and SKadkao 1974 Studies on ecology and control of the rice gall midge in Thailand Tech Bull 6 TropicalAgricultural Research Center Tokyo Japan 113 p

89 Hincklcy A 1) 1963 Ecology and control of rice planthoppers in Fiji Bull Entomrol Res 53467 481

90 Hirao J 1982 Ecology and chtInical control of the rice leaffolder Jpn Pestic Inf 4114-17

91 Ho DT 1984 Sten borer incifience in rice ecosystems in KenyaEast Africa Pages 307-320 in A overview of upland rice research Proceedings of the 1982 Bouac Ivory Coast upland rice workshop International Rice Research Institute PO Box 933 Manila Philippines

92 Ho )T J 1Njokah and J GKibuka 1983 Studies on rice sten borers in Kenya with emphasis on ifaliarpiaipiraellRag Insect Sci Appl 465-73

93 Ichihara I T Ito A NIatsuo and M Sawada 1971 Chemical control of the rice striped stem borer in upland rice grown in plastic mulch [iii Japanese] Proc Kanto-Tosan Plant Prot Soc 1882

94 IclhiharaIand A Matsuo 1969 Damage caused by rice striped stett borer to upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1686-87

95 Ichhara I atid A MIatsun11970 Conside ration oft lie extraordinary prevalence of rice striped stem borer in upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1778-79

96 Institut de Recherches Agrolnliqes Iropicale et des Cultures Vivrieres 1978 Insect resistance Pages 12-13 in Upland rice 1978 atnual report Bouak Ivory Coast 27 p

97 International Institute for Tropical Agriculture 1983 Aphids itt rice roots I ITA Res Briefs 4(1-3)4-5

98 International Institite fi Tropical Agriculture 1983 Hopperburn in rice caused by brown plathopper IITA Res Briefs 4(l-3)6-7

99 International Rice Research Institute 1975 Annual report for 1974 PO Box 933 Manila Philippines p 216

100 International Rice Restarch Institute 1976 Annual repiort for 1975 PO Box 933 Manila Philippines p 228

101 International Rice Research Institute 1979 Cropping systems program Component technology development and evaluation Page 404 i Annual report for 1978 11O Box 933 Manila Philippines

102 International Rice Research Institute Department of Entomology 1981) Insecticid evaluation 1979 Tables 73 74 PO Box 933Manila Philipes

103 International Rice Research Institute Department of Entonmology 1981 Insecticide evalomtion 1980 Table 81 POBox 933 Manila Philippines

104 International Rice Research Institute 1981 Report of a meeting of Asian rice-based cropping systems entomologists Eleventh cropping systems working group meeting 18-22 May 1981 PO Box 933 Manila Philippines p 132 133 144

105 International Rice Research Institute 1984 Cropping systems program Pages 368-373 in Annual report for 1983 PO Box 933 Manila Philippines

106 International Rice Research Institutc 1984 Upland rice in Asia Pages 45-68 in An overview of upland rice research Proceedings of the 1982 Bouakamp Ivory Coast upland rice workiiop loternational Rice Research Institute 11O Box 933 Manila Philippines

107 Irshad M 1977 Grasshoppers in rice in Pakistan lit Rice Res Newsl 2(3)8-9

108 Ito R and TOno 1964 Upland lice its culture and varieties in Japan lIternational Rice Cormmission loth Meeting of the Working Party on Rice Production and Protection 3-10 March 1964 Manila Philippines Food and Agriculture Organization Rome Italy 12 p

109 Jeffrey M I and V A Dyck 1983 Effect of moonlight on lightshytrap catches of brown planthopper lit Rice Res Newsl 8(6) 13

110 Jennings P R and T Alicia Pineda 1983 Management ofthe rice plnthoppers Sogatodsorivi(coIl in Latin America Presented at the meeting on stable plant resistance FAO and lIndocsian I)inectorate of Food Crops Protection Bali lndonesia 29 Novcniber-2 l)ccemshyber 9 p

IllJimenez Gomez J A 1979 Rice insect pests i lla ios ()riCntales lil Spanish] Pages 99-103 in Rice ciltivation Instituto (olotibiatto Agropecuario Compndio No 29

112 Jongeleen F JJ Soesanto and A Mockidjo 1979 Studies oti pathogens of two species of Scarabacidac froinIndonesia Z Angew Enlomnol 87412-417

113 Jouvenaz 1) PC S lofgren and W A Baiks 1981 Biological control of imported fire antsa review of cirrent knowledge Bull Entomol Soc Am 27203-208

114 Kainoh Y K Shiniiu S MarUani Y Taiaki 1980 lost finding behavior of tie rice bu lplotori+JchinvittjI)allas (liciniptera Corcidac) with special reference to diet pattern ol aggregation and feeding on the rice plant Appl Entomol Zool 15225-233

115 Kakkar K 0 P Sharma and P I Sharma 1977 Note on Ihteronyrchus liodresRedt (Coleopicra I)ynastidae) as asporadic pest of paddy and maie inHimachal Pradesh Corr Agric 156-57

116 Kalode M131 79 Major insect pest problems and their control in upland rice (Unpublishcd mineogriph CWI oJNaaiat lni Chandaratat (198 1)original not seen)

117 Kalshoven L i 98 I The pests of crops in I ldoniesia (translated by P A vatn der Laan) 1 T Ichtiar B3aru-vati Iloeve lakarta Indonesia 701 p

118 Kamal N Q 1981 Suppression of whitebacked planto ner Sogatella fircaifra (Horvath) aLId rice lealfolder (nhal -cis licdb lis (Guenee) populations by natural cnclmie hlI) tiesis Grcgorio Araneta University Foundation Victoncta Park Metto Manila Philippines 179 Ip

119 Karim A N M Reaul 1975 A species ofliealy bug Trioni i70s ceres new oii rice in the Philippines Rice Entoniol Newsl 247

120 Katanyukul Wand C Chandaratat 1981 Insect pestsof uplaid rice and their control Report firstFAOUNlFi Tliailand training course on improved cultural practices for tplad rice Chiang Mai Thailand 5-23 Oct 1981 Food and Agriculture Organiation Rome Italy 22 p

121 Kern F 1955 lhletwioa(I)uninemts) bhhit Biturm(Coleoptera) a new pest of sugarcane inVenezuela [inSpanish with Etnglish sunmary] Agron Trop (Maracay) 527-29 (Rev Appl Eitoniol A4572)

122 Khan K M and S Ghai 1974 White grubs ald their control in India Pesticides (1Bombay) 819-25

123 Kiritani K 1975 Integrated pest management of rice pests Pages 66-70 itt Proceedings platning workshop ott cooperative hIeld research in pest managementP SMotooka ed East-West Center Honololu Hawaii USA

124 Kisimoto R 1967 Ecology of insect vectors forecasting and chemical control Pages 243-255 in The virus diseases of tle rice plant Proceedings of a symposium atthe International Rice Research Institute April 1967 The Johns Hopkins Press altimore Maryshyland USA

125 Kisimoto R and V A Dyck 1976 Climate and rice insects Pages 367-391 in Climate and rice International Rice Research Institute PO Box 933 Manila Philippines

126 Kureha S S Kobayashi SAsari and T Ishihara 1977 Life history and control of the mole cricket Grrllotalpa africana Palisot de Beauvois [in Japanese English translation 7520 1RRI library] Proc Kanto-Tosan Plant Prot Soc 21192-197

127 Kushwaha K S 1981 The black beetle as a pest of rice in Haryana India Int Rice Res Newsl 6(4)14

128 Lawani S M 1982 A review of tile effects of various agronomic practices on cereal stem borers Trop Pest Manage 28266-276

129 Leuck D B 1966 Biology of the lesser cornstalk borer in South Georgia J Econ Entomol 59 797-801

130 lever R J A W 1955 Insect pests of the rice plant in Malaya Int Rice Comm Newsl 1612-21

131 Li Ching-Sing and Shin-Foon Chiu 1951 The study ofthe rice gall midge Iachydiplosis orvza Wood Mason [in Chinese with English summary] 1Taiwan Agric Res 21-13

132 Iitsinger J A 1984 Assesament of the need-based application for rice Proceedings TechnologyTransfer Workshop 15-16 March 1984 International Rice Research Institute 1PO Box 933 Manila Philippines 6 p

133 Litsinger J A 1984 Environmental diversity and insect pest abundance with reference to the Pacific Pages 285-307 in Proceedings sub-regional training course on methods of controlling diseases insects and other pests of plants in the South Pacific 4-20 Oct 1982 Vaini Kingdom of Tonga Nukualofa

134 Litsinger J A RF Apostol and M BObusan 1983 White grub Leucopholis irrorata (Coleoptera Scarabaeidae) pest status popula-tion dynamics and chemical conllol in arice-maize cropping pattern in the Philippines J Econ Entoinol 761133-1138

135 Litsinger J A C de la Cruz F Raymundo A T Barrion M D Lumaban J P Bandong M S Venugopal F Paragna and E Balete 1982 Insect pests and insecticide reponse of dry-seeded bunded rice Pages 179-193 in Report of workshop on cropping systems research in Asia International Rice Research Institute Los Bafos Philippine

136 Litsinger J A M 1) Luinaban J P Bandong A T Barrion W Katanyukul and N Panda 1978 Insect pest management in rainfcd lowland rice agroecosystms of tropical Asia International Rice Research Conference 17-21 April 1978 International Rice Research Institute Los Bafios Philippines

137 Litsinger J A and K Moody 1976 Integrated pest management in multiple cropping systems Pages 293-317 in R I Papendick P A Sanchez and G B Triplett eds Multiple cropping American Society of Agronomy Madison Wisconsin USA

138 Loevinsohn M E 1984 The ecology and control of rice pests in relation to the intensity and synchrony of cultivation Ph D thesis University of London 36) p

139 Loevinsohn M E J A Litsinger N Panda M A Austria R F Apostol C Andrion and J Bandong 1982 A basis for predicting rice insect pest abundance in cropping systems Paper presented at the 1982 International rice research conference on rice and cropping systems 19-23 April 1982 International Rice Research Institute Los Bafios Iaguna Philippines 25 p

140 Lopez A W 1930 Repot of the Entomology Department Pages 145-172 in Annual report of the Philippine Sugar Association 1929-30 Manila

141 Lopez AW 1931 Entomology Department Pages 227-266 in Annual report of the director or research 1930-31 Philippine Sugar Association Manila

142 Lopez A W 1932 Entomology Deparment Pages 252-279 in Annual report of the Philippine Sugar Association 1931-32 Manila

143 Lungibill P and G G Ainslee 1917 The lessercornstalk borer US Dep Agric Agric Res Serv Tech Bull 539 27 p

IRPS No 123 January 1987 39

144 McGuire J U Jr and BS Crandall 1967 Survey of insect pests and plant diseases ofselected food crops of Mexico Central America and Panama U SDepartment of Agriculture International Agriculshytural Development Service Agricultuial Researrh Service October 1967 11 p

145 McNeill Sand T R ESouthwood 1978 The role of nitrogen in the development of insectplant relations Pages 77-9 in lliochemical aspects of plant and animal coevolution J B Ilarbornu ed Academic Press London UK

146 Maebara H 1955 Spatial distribution of lastinstiarlanaeof purplish stem borer (Sesamia iqferens Walker) hibernating in an upland rice field Bull Kagoshima Univ Fac Agric 438-41

147 Mallick E H and N G Hajra 1977 Response of rice plant to defoliation treatments at different developmental stages Indian Agric 2151-56

148 Mansingh A 1971 Physiological classification of dormancies in insects Canl Entomol 103983-10)9

149 Martell C G 1974 Catalogue of plant feeding insects of Mexico [in Spanish] Fitofilo 27(69)1-176

150 Martinez 0 J 1982 Principal rice pests in Ilanos orientalis [in Spanish] P~ages 80-94 in Curso deact nalization de conocimientos en arroz Villaniencis Instituto (olonmbiano Agropecuario

151 Martins J F da Silva F Ferrcira A S Prabhu F 1 P Zimmermann 1980 Use of pesticides for the control of the main subterranean upland rice pests esqui Agropecu Bras 1553-u

152 Martins I F da Silva F Fe eira rod BdamSilveira Pinheiro I Q2 Damage simulation caused by defoliators of upland rice [in Portuguese with English summary] Pesqui Agropecu Bras 171113-1119

153 Martins 1F da Silva N(iyuen van Tan and 1Ida Silveira Pinheiro 1981 Stem borer resistance in upland rice associated with plant morphological characters [in Portuuce Iesqui Agropecu Bras 16187-192

154 Masaki S 1980 Sumnmer diapause Ann Rev Entomiol 251-25 155 Masuzawa T 11 Suwa and F Nakasoji 1983 l)ifkleences of

oviposition preferencc and survival rate of two skipper butterflies Parnara guttata and PIchlpidai iathiais (Iepidoptera HIesperiidac) on rice plant and cogon grass New Entomol 3247-56

156 Maurya D M and C P Vaish 1984 Upland rice in India Pages 69-92 in An overview oftpland rice research Proceedings ofthe 1982 Bouakamp Ivory Coast upland rice workshop International Rice Research Institute IO Box 933 Mnila Philippines

157 Menon M G R R N Katiyai S Kuar and S S Misra 1970 Termites attacking the paddy crop Sci Cult 36294-295

158 Miyahara Y and 1 Nishikubo 1967 Behavior of young nymphs of the red rice root aphid Iholahsipiun ri liabhdontinalisSasaki in relation to movement from the leaf to the roots of upland rice [in Japanese] Pioc Assoc Plant Prot Kyushu 1363-66

159 Mo ljoa Tjien 1952 White grub of upland rice (Holotrichiahelleri Brsk) [in Indonesian with English summary] Contrib Agric Res Stn Bogor No 13134 p

160 Mochida 0 and T Suryana 1979 The occurrence of the brown planthopper and ragged stunt on lowland and upland rice ill Central Java and logyakarta in tile late of January 1979 Entomnolgy Survey Report No 15 Central Research Institute of Agriculture (CRIA) Sukamandi Indonesia I February 1979 9 p

161Monet C 1979 Varietal resistance to upland rice pests [inI French] Pages 35-39 in Annual report 1977 Institut des Savanes Department des Cultures Vivrieres Bouak6 Ivory Coast

162 Monte 0 1942 A caterpillar ofrice-fields[in Portuguese] Biol6gico (Sao Paulo) 8161-163 (Rev Appl Entomol A31414)

163 Munroe I) D 1979 Pest management strategy for hill rice in four resettlement areas in Sarawak Malaysia Pages 89-102 i1 Guidelines for integrated control of rice insect pests Food and Agriculture Organization Plant Production and Protection Paper 14 Rome Italy 115 p

40 IRPS No 123 January 1987

164 Nakasuji F M Ishii and Jen-Zon Ito 1984 Rice skippers in Taiwan and their life histories ZZrC (Taipei) Tech Bull 82 10 p

165 Navas I) 1976 Fall armyworum in rice Pages 99-106 il Ecological animal control by habitat management Proc Tall Timbers Con-ference 28 Feb-I Mar 1974 Fall Timbers Res Stn Misc Publ 6 Tallahassee Horida USA

166 Nickel 1 1 1961 Biological control of rice stem borer A feasibility study Tcch Bull 2 International Rice Rcarch Institute PO Box 933 Manila Philippines IIl p

167 Nigan P M and R A Vernia 1985 Insect pests of uplaild rice in Uttar Pradesh Int Rice Res Newsl 10(6)22

168 Nilakhe S S 1985 -cological observations on spittle bugs with emphasis on their occirence in rice Pcsqtui Agropecu Bras 20407-414

169 Nixon G EJ 195 1 lie association of ats with aphids and coccids Commonwealth Iusti lcte of Entonology Iondon UK 36 p

170 Nurdin F an Ishak Manti 1985 Study oit seedling fly of upland rice Interlational Upland Rice Moitoring Tour Report 26 lebruary-2 MarLI Sukaranti West Sumatra

171 Oka 1 N 1979 Ciltural conttol of the brown planthopper Pages 357-369 in Brown planthopper threat to rice production in Asia International Rice Research Institutc 1p0 Box 933 Manila Philippines

172 Ohni M and A Vilnaii 1975 Rice insect pests of Italy biology of Sipha grcria (Kaltenbach) and its control [in Italian] Riso (Milan) 2459-71

173 Otanes F Q atnd 1 I Sison 194 1 Pests of rice Philipp J Agric 12211-259

174 Painter R I 1951 Insect icsistance in crop plants University Press of Kansits Lawrence Kansas USA 520 p

175 Pantua P C and 1A litsinger 1981 Arthropod abundance in continuous vs biannual rice cropping systeis Int Rice Res Ncwsl 6(2)20-21

176 Pathak M ) and V A l)yck 1975 Sttudics ott insect pests of upland rice Pages 186-191 in Major research in upland rice International Rice Research Institute PO Box 933 Ilanila Philippines

177 Patil B R B M IHasabe and 1) S Ajri 198 I Chemical control of white gritb lIditriihinhorraia Fabr intfesting groundntut paddy atnd sorghum Indian 1Entomol 43232-236

178 Pawar A 1) 1974 (ollection and identification of insect pests of rice rlerminal repirt lInternational Rice Research Institute PO Box 933 Manila Philippines 72 p (tnimco)

179 IerrajU A aid A S Reedy 1965 Observations ott diapausc itt tie steti borer ohrice Rice News Teller (lai-Mar)9-13

180 Pont A C 1972 A review of the Oriental species of theri ona Rondani (I)iptera Muscidac) of econtomic importance Pages 27-102 in Control of sorglium shoot fly M G lomtani and W R Young eds Oxford atd IH1 Ptthl Co New I)clhi India 324 p

181 Prasad S C 1PChand aid M 1PSingh 1983 Incidence of rice gall midge in the upland plateau region of Hihar Oryza 20235-236

182 Rai P S 198 0 Nisaga simpex damage to rice in the hill tracts of South India Int Rice kles News[ 5(I)12

183 Rathorc Y S 1969 Gonocplhahli dcprssunt Fabr as itpredator of (tilo zonwIlhis (Swinhoc) and other moths in field cage at Patnagar Uttar Pradesh Indian 1Entotnol 3127

184 Reed W 1983 Crop losses causcd by insect pists in the developing world Proceedings 10th Internatial Congress of Plant Protection lrighton UK 174-80

185 Reedy M V 1981 Ateactogasterinhens(Walker)a new root-grub pest of tipland paddy and inaize Curr Sci 501076

186 Reveno M A 1968 Catalogue of the insect pests of economic crops in Colombia [in Spaiish] Asociacion Latino Americana de Entomo-logia (AIAE) 129-33

187 Reyes G M 1957 Rice dwarf disease in the Philippines Food and Agriculture Organiiation Plant Prot Bull 617-19

188 Rice S E A A Grigarick and M 0 Way 1982 Effect of leaf and panicle feeding byarmyworm (Lepidoptera Noctuidae) larvae on rice grain yield Environ Entomol 7593-595

189 Risch S J 1) Andow and M A Altieri 1983 Agioccosystem diversity arnd pest control data tentative conclusions and nizw research dirctions Environ Entomol 12625-629

190 Rivera C T 1956 Three organic insecticides in thecontrol of insects affecting upland rice Philipp Agric 39465-472

191 Roffrey 1 1972 Locusta outbreaks in the Philippines Acrida 1177-188

192 Rojas R 1) 1941 The effect of pruning at varying ages upon the yield of Ramairio Phipp Agric 29851-860

193 Roonwal M i 1979 Ternite life and termite control in tropical South Asia Scientilic Pnbl odhpr India 139 p

194 Rosetto C1 F da Silva Martins N C Schmidt and L EAzzini 1978 Damage caused by pasturespittle bugs 4Dvoi)lavopictaandDJ schach) on rice lBragantia Nota 6 (Matcli)25-27

195 Rosztto C1- S S Neto I- Amantc I) Iink 1Garcia-Vieira D M de SoLza N V Bansa t and A M de Olivcira 1971 Rice pests of Brazil [in Portuguese] Pages 149-238 inrFA()-International Rice Commission 2nd meeting of the Rice Committee for the Americas 6-11 l)ecember 1971 Pelotas Rio Grande do Sul Brazil

196 Rothschild G H L 1969 Obseratiois on the armyworm Spodoptera matritia acronrctoiiles Gn (Lep Noctuidae) Bull Entomol Res 59143-160

197 Rothschild G 1- L 1974 Parasites of the armworm Spolopiera mauritiaacroncitoihes in Malaysia Fntornopliaga 19293-299

198 Sachan i N S K Gangwar and 1N Katujar 1980 Field crickets damaging cages in North Eastern IHill Region Indian J Entomol 42526-527

199 Sands 1) P A 1977 The biology and ecology of Lepocorisa (Hemiptera Alydidac) in Papua New Guinea Res Bull 18 Departshyment of Primary Industry Port Moresby Papua New Guinea 104 p

200 Sarawak 1968 titomology division Pages 97-106 in Report of the research branch of the Department of Altriculture 1966 Kuching (Rev AIppl Eintomol A56603)

201 Satpathy JM M S Das and K Naik 1977 Effect of multiple and mixed cropping on the incidence of scine important pestsI Entomol Res 178-85

202 Sauer IHF G 1939 Notes Ott Elasmpnlpualtsligniosellths Zeller (Lep Pyr) a serious pest ofcereals in Sao Paolo [in Portuguese] Arq Inst Biol 10199-206 (Rev Appl Entotnol A28378)

203 Satinders 1L A 1 SKing and C L Vargas S 1983 Crop pests of Central Anerica with t bibliography [in Spanish] Centro Agroshynotnico Tropical de Investigacion y Ensenanza Turrialba Costa Rica (CATIE) Tec Bol 9 90 p

204 Sauphanor B 1985 Some factors of upland rice tolerttce to stem borers in West Africa Inseu Sci Appl 6429434

205 Sen P atnd S Chakravorty 1976 Stentflics in Haritighata Nadia West Bengal India [it Rice Res Newsl 1(12) 15

206 Senapai B and 1M Satpathy 1983 The seedling fly Atherigona orrzae Malloch is a pest of rice in Orissa (India) J Entomol Res 777-79

207 Shhjahan M S Alatn M Ah nad and BN Islam 1983 A new scarabaeid beetle pest of dryland rice in Bangladesh Int Rice Res Ncwsl 8(I)16-17

208 Sharma G L S C Modgol and R C Gautam 1979 Effect of depth of submergence ott incidence of bacterial blight and white grub infestation in transplanted rice Int Rice Res Newsl 4(6)12-13

209 Sharma S 1985 Scarabaeid beetle outbreak itt upland rice Int Rice Res Newsl 1(1(l)24

210 Shiraki T 1917 The paddy borer Schoenobhits incertulas Walker [achaku Agric Exp Stn Formosa 256 p

211 Silva A de Brito and 11P Magalhaes 1981 Insect pests of rice [in Portuguese] Circular Tecnica No 22 EMBRAPA Centro de Pesquisa t ecuaria de Tropico Unido (CPAJU) Belem Brazil 14 p

212 Singh S A 1979 Emergence of rice leaf butterfly as a rice pest in Manipur India int Rice Res Newsl 4(4)16-17

213 SinghS A 1983 Changing trends of insect pestsof rice in Manipur Int Rice Res Newsl 8(f)12

214 Siwi S S and P H van Doesburg 1984 Lenovorisa Latreille in Indonesia (Heteroptera Coreidae Alydinae) Zool Meded (Leiden) 58117-129

215 Soejitno J 1977 Reaction of some upland rice varieties toseedling fly eltherikonaexigua Stein [in Indonesian] Siri Hana I Penyakit 630-35 Lembga I osat Fenclilian Pertanian Bogor Indonesia

216 Socnardi 1967 Insect pests of rice in Indonesia Pages675-683 in The major inse pests of thu rice plant Proceedings of a symposium at the Internatital Rice Rec Wi itt S pternber 1964 The Johns Hopkins Press Baltimore Maryland USA

217 Soto P E A T Petcz and I W Buddenhagen 1976 Survey of insect pests and diseases of rice in different ecological zones in Nigeria Rice Entomol Ncwsl 435-46

218 Soto 11E and Z Ziddiqi 1976 Screening for resistance to African rice insect pests Varietal improsetnent sCmi n r Bouak Ivory Coast 13-17 Sept 1976 West Africa Rice l)evelopment Association Monrovia Iiberia

219 Squire F A 1972 Entomological problems in Bolivia PANS (PestArtic News Sumr) 18249-268

220 Stapely 1 II 1978 The rice leaf roller in the Solomon Islands Int Rice Res Newsl 3(l)13

221 Suenaga H 1937 On aphids attacking the roots of rice Oyo-I)oubts Zasshi 9150-152 [in Japanese] (Rev Appl Entomol A25745)

222 Suenaga H 1952 On the seasonal abundance of three subterranean aphids on upland rice Bull Kyushu Agric Exp Sti 1249-262

223 Sutynso Yas and Dandi Soekaria 1964 Beberapa hana yang ttenperang tana man padi tanah kering [in Indonesian] Rapat kerja Padi Tanah Kering 6 sd 8 Juily 1964 Kaliurang 9 p

224 Tallamy I) W and R F Denno 1983 Alternative life history patterns in risky environments in example from lace bugs Pages 129-147 in Insect life history patterns R F l)cnio and If Dingle eds Spring-Verlag 225 p

225 Tamura 1 1939 Observations on some insect pests of field crops [in Japanese] 1 Plant Prot 26414-420 (Rev Appl EntomolA27-556)

226 Tamura and Suzuki 1964 Influences of the early-transplanting practice in rice culture upon development of the rice stem borer Chilo siql)ressalisWalker Bull Ilokuriku Natil Agric Exp Stn 761-94

227 Tan J P 1924 The rice root aphid Drowhi ihirsoia Philipp Agric 13277-288

228 Tanaka 1 1957 Taxonomy and distribution of some subterranean aphids injurious to upland rice in Japan with description of a new species [in Japanese English summary] Botyu-Kagaku 22168-176

229 Tanaka T 1961 The rice root aphids their ecology and control Spec [full College of Agriculture Utsunomiya University Japan F3 p

230 Tauber M IC A Tauber R Nechols and 1Obrycki 1983 Seasonal activity of parasitoids control by external internal and genetic factors Pages 87-108 itt I)iapause and life cycle strategies in insects V K Brown and I Ilodek eds )r W Junk The Hague Netherlands

231 Taylor W F 1972 Effects of artificial defoliation (simulating pest damage) on varieties of upland rice Exp Agric 879-83

232 feotia T P S and S Natud 1966 Biotoomics of the rice skipper Parnaraniathiis Fabricius (Lepidoptera Hesperiidac) Indian J Entomol 28181-186

233 Teran F 0 1971 Rice pests of Santa Cruz [in Spanish Ministerio de Agricultura Sita Cruz Bolivia Bull 45 8 p

234 Than Itun 1976 Population dynamics of the yellow rice borer Tryporza itcertulas (Walker) and its damage to the rice plant MS thesis University of the Philippines at Ios Bafios Laguna Philippines 92 p

235 Uichanco L B 1911 Pests Pages 55-74 it Rice Philipp Agric 11-200

IRPS No 123 January 1987 41

236 van der Laan P A 195 1 Life history of the rice seedling fly Contrib Gen Agric Res Stn Boor Indonesia 118 15 p

237 van der Laan 11A 1959 Correlation between rainfall in the dry season and the occurrence of the white rice borer (Scirpolhaga innotala WIk) in lava Entomol Fxp Appl 212-20

238 Van Driesche R G 1983 Meaning of percent parasitismin studies of insect parasitism Fnviron Entomol 121 I1- 122

239 Venugopal M S and A litsingcr 1984 Fffect ofcarbofuran on rice growth Prot Ecol 7313-317

240 Vietnam Directorate of Rural Affairs 1962 Annual work progress report on crop improvement program of rice sugarcane vegetable and field crops (lor the period from July 1961 to June 1962) (11) 299 p (Rev Appl Entomol A51526-527)

241 Wan M T K 1972 Observations on rice leaf- and planthoppers in Sarawak (Malaysian Borneo) Malay Agric 148308-335

242 West Africa Rice I)esclopnient Association 1978 Annual research report Vol 2 Statistics Monrovia Liberia

243 West Africa Rice Development Association 1979 Annual research report p47 Monrovia Liberia

244 Williams ) F and CS lofgren 1981 Fli I illy l-468 anew bait toxicant for control of the red imported lire at Fla Entomol 64472477

245 Williams I) 1 1970 The mealybugs OFlomoptera Coccoidea Iscudococcidac) of sugar-cane rice and sorghum Bull Entomol Res 0109-188

246 Williams D) 1 1982 Pulinaria icervi (Signoret) and its allies ott sugar cane and other grasses Btll Entoinol Res 72111-117

247 Wirjosuiardjo S 1976 Study ofchenmical cont rot of insect pests ott upland rice Imiu Pertanian (Jogjakarta) 1323-329

248 Wirjosuhardjo S and N Wirarta 1976 Chemical control of the white grub tIolorithiatlltriin upland rice in M uneng Irobolinggo Indonesia) [in Indonesian] Imut Pertanian (Jogjakarta) 1330-332

249 Yano K T Miyake and V F Eastop 1983 The biology and economic importance of rice aphids (Henuiptera Aphid idae) a review Bull Entomol Res 73539-566

250 Yelp D and P Rhodes 19(4 lhe success of drilled dryland rice iii the Fiji Islands International Rice Commission Working Party on Rice Production and Protection Manila Philippines 3-10 March1964 Food and Agriculture Organization Rome Italy 13 p

251 Young W R 1981 Fifty-five years of research on the sorghum shootfly Insect Sci Appi 23-9

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No 106 Rice grassy stunt virus 2 a new strain of rice grassy stunt in tire literature

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Page 4: UPLANDRICE. INSECTl--1i M T -E!R.EOLOU t RTANCE ,AND …delphacid.s3.amazonaws.com › 6958.pdf · The significance of abiotic factors in upland rice insect ecology will be apparent

4 IRPS No 123 January 198

Table 1Insect pests of upland rice worldwide

Order Family Species

Isoptera Rhinotermitidae Coptotermesformosanus Shiraki

Termitidae Heterotermesindicola (Wasmann) Anacanthotermesviarum (Koenig) Anacanthotermesrugifrons Mathul et Sen-Sarma Captiterpnesnitobei (Shiraki)Corniterrnesstriatus (lHagen) Procapriterines mnushae Oshima Pericapritermes nigerianus Silvestri (=socialis)Macroternes nutalensis (Haviland)Macrotermes beiiicosus Smcat hman Macroterines gilius (Hagen) Macrotermnes spp Odontoterniesformosanus (Shiraki)Procornitermesarauloi Emerson Procornitermws triacifer (Siivestri) Trinerviterniesgenzinatus Wasmann

[= ebenerianus Sjostedt]MicroceroterynessppMicroterinesspp Synternes nolestits (Burmeister)

Dermaptera Forficulidae Termites unspecifiedDiaperasticuserythrocephalusOlivier

Orthoptera Gryllotalpidae Don lineare (Eschscholtz)GryllotalpaafricanaPalisot de Baeuvois Gryllotalpaorientalis (=africanu) Burmeistera

Gryllidae

Neocurtilla (= Gryllotalpa)hexadacryla (Perty) Scapteriscusdidact)hts(Latreille) Grjllusassinilis(= bimaculatus)(Fabricius)Gryllus (=Liogryllus) bimaculatusde Geer Brachytrupes portentosus (Lichtenstein) Brachytnmpes mnembranaceus DruryPlebeiogrylhsplebejus (Saussure) Teleogryllus testaceus (Walker)Teleogryllus occipitalis (Serville) Loxoblenimus haani Saussure

Pyrgomorphidae

Velarifictorusaspersus(Walker)Velarifictonis sp Euscyrtus concinnus (de Haan)Atractomorpha burri I Bolivar

Tettigoniidae A tractomorphapsittacinapsittacina(de Haan)Conocephahslongipennis(de Haan) Conocephalusrnaculatus(Lethierry) Conoceptahissaltator Saussure Euconocephah s varius (Walker)

Tetrigidae Acrididae

Caulopsiscuspidatus (Scudder) Caulopsis oberthuriScudder Phaneroptera furcifera (Stal) Amphinofus spp Aiolopus thalassinustamulus (Fabricius) Acrida ivilleinsei Dirsch Gesomida mundata zonocera Navas Gonistabicolor (de Haan)Oxyajaponicajaponica (Thunberg)Oxya hya intricata (Stal) Oxyafitscovittata (Marscliall)Oxya velox (Fabricius)Gas frimargus niarmoratusgrandis (Saussure) Xenocatantopshunilishumilis ServilleHierogiyphus banian (Fabricius) ltieroglyphusdaganensis Krauss Hicroglyphus nigrorepletus(1 Bolivar) Hieroglyphus oilziiorosCarl Heteropternisrespondens (Walker)Patangasuccincta (Linnaeus) Stenocatantops splendens (Thunberg) Chondracrisrosea brunneri Uvarov Trigonidium cicindeloides Rambur Locusta migratoriamanilensis (Meyen)Locusta migratoriacapito Saussure

Distribution

Asia Asia Asia Asia Asia Latin America Asia Africa Africa Africa Asia Africa Asia Latin America Latin America Africa

Asia Asia Africa Latin America Asia Africa Latin America Africa Asia Latin America Latin America Asia Asia Asia Africa Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Latin America Asia Latin America Latin America Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Africa Asia Asia Asia Asia Asia Asia Asia Asia Africa

Reference

85 82 190 190 82 85 151 82 85 7 85 85 178 12 82 85 85 195 195 85

193 11 12 157 60 85 195 72 36 186 12 32 59 72 76 84 81 150 195 203 211 81 76 178 b 48 b 76 178 b b b b b 55 178 178 178 65 178 68 81 68 178 178 178 178 178 55 72 178 178 78 178 178 55 55 72 6 82 82 55 54 55 59 130 55 178 55 78 55 82

IRPS No 123 January 1987 5

Table I continued

Order Family Species Distribution Reference

Locusta migratoria migratorioidesReische Africa 33 et Fairmaire

Schistocercagregaria (Forskal) Africa 33 Schistocerca americana (Drury) Latin America 82 195 Valanga nigricornis (Burmeister) Asia 82 Orphulella inticata Scudder Latin America 68

lomoptera Coccidae Pulvinaria sorghicola De Lotto Africa 246 Pulhinariaiceryi (Guerin) Latin America 186 246

Pseudococcidae taennococcus spp Asia 120 Lhorizococcus iu Williams Asia 245 Dysmicoccus boninsis (Kuwana)C Asia Africa 245

Latin AmericaDysmicoccus brevipes (Cockcrell)c Asia Africa 117 203245

Latin AmericaDysmicoccus oryzae (Wirjati)c Asia 245 Geococcus (=Ripersia) oryzae (Kuwana)C Asia 245Brevennia (= tererococcus)rehi (Lindinger) Asia 72 245Nipaccoccus graminis (Maskell) Africa 245 Planococcoides lingnani (Ferris) Asia 245 Pseudococcus saccharicola Takahashi Asia 245 Pseudococcus spp Latin America 60 Saccharicoccus sacchari (Cockerell) Africa Asia 245

Latin America Trionymus ceres Williamsc Asia 119245j rionymus sp Asia b Mealybug unspecified Asia 83 84

Aphididae Capitophorus[= Rhopalosiphumn] prunifoliae Asia 221 (Shinji)

Rhopalosiphum maidis (Fitch) Asia 249 Rhopalosiphum padi (Linnacus) Asia Africa 12 97 249 Rhopalosiphum rufiabdominalis (Sasaki) Asia Africa 66 76 120 195203

Latin America 221229240249 Tetraneura akinire Sasaki Asia 221249Tetraneurabasui ltillcRisLambers Asia 249Terraneura nigriabdominalis (Sasaki) Asia Africa 712 30 76 227229 (=Dryopeia hirsuta)

Tetraneuraradiciola Strand Asia 249 Paracleus ciniciformnis von Heyden Asia 249

[= Forda harukawai Tanaka]Pineus hanikawai (lnouye) Asia 228 Geoica hcifitga (Zehntner) Latin America 76 195 229 249

AsiaGeoica setulosa (Passerini) Asia 249 Anoecia fuliabdominalis (Sasaki) Asia 249 Anoecia comi (Fabricius) Asia 221229249Hysteroneura sctariae (Thomas) Africa Asia 7

Latin America Sipha glyceriae (Kaltenbach) Asia 249 Root aphid unspecified Asia 72 120

Cercopidae Deois schach (Fabricius) Latin America 61 68 194 Sepullia (=Denoplux) nigropunctata Stal Africa 29 Tomaspis ( Deois) fluxtuosa (Walker) Latin America 61 68 194 Tomaspis ( Dcois) flaiopicta (Stal) Latin America 61 61 194 To naspis (Dcois) complcta (Schmidt) Latin Aincrica 68 Tomaspis ( Zulia) entreriana (Berg) Latin America 68 Tomaspis ( Mahanarva) fimbriolata(Stal) Latin America 68 194 Tomaspis ( Aeneohna) spectabilis (Distant) Latin America 219 233 Aeneolamia varia Fciinai Latin America 81 111 Aeneolamia postica (Walker) Latin America 203 Abidama producta Walker Asia 54Delphacidae Sogatella furcifera (Horvath) Asia 59241 Niapanatalugens (Stal) Asia 30 59 72 84 241 Sogatodes oryzicola (Muir) Latin America 6167150Sogatodes cubanus (Crawford) Latin America 150 Sogatodes pusanus (Distant) Asia bPeregrinus maidis (Ashmead) Latin America 186 Laodelphax striatellus (Fallen) Asia 18

Cicadellidae Recilia dorsalis (Motschulsky) Asia 59 241 Nephotettiv nigropictus (Stal) Asia 59241

6 IRPS No 123 January 1987

Table I continued

Order Family

Hemiptera Lygaeidae

Cydnidae

Alydldae

Coreidae Corimelaenidae

Pentatomidae

Thysanoptera Thripidae

Species

Nephotettix virescens (Distant) Nephotettix modulatus Melichar Cofana spectra(Distant)Graphocephala spp Hortensiasimnilis (Walker) Exitianusobscurinervis (Stal)Balclutha spp Draeculacephalaclypeata Osborn Cicadulinabipunctella (Matsumura)Cymoninus turaensis(Paiva) Ninus insigniStal Pachybrachiusnenosus Horvath Caenoblissuspilosus Barber Blissus leucoptens (Say) Blissus spp Dlmnorphoptentscornutus novaeguineae GhauriDimorphopterus similis Slater Paroniuspiratoides (Costa)Pangaeus spp Aethus indicus (Westwood)Geotomnus pygmaeus (Dallas) Stibaropus inolginus Schiodte Cyrornenusbergi FroeschnerCyrtomnenus ciliatus (Palisot de Beauvois) Cyrtonenuscrassus Walker Tominetus spp

Riptortuslinearis (Fabricius) Riptorts spp Leptocorisaacuta (Thunberg)Leptocorisabiguttata (Walker)LeptocorisapalawanensisAhmad Leptocorisaoratorius(Fabricius)Leptocorisachinensis Dallas LeprocorisasolomonensisAhmad Leptocorisaspp Stenocorissouthiwoodi Ahmrad Stenocoris claviformis Ahmad MirperustorridusWestwood aetus trigonus (Thunberg)Alkindus atratusDistant ScaptocorisdivergensFroescher Scaptocoris castaneusPertyScotinopharatarsalis (Vollenhoven) Scotinopharascotti Horvath Eysarcoris (=Stollia) ventralis (Westwood) Dolycorisindicus Stal Macrinajuvenca Burmeister Tanda antiguensis (Westwood) Tantiageifi Schout Tantia perditor (Fabricius) Pygotnenidavaripennis (Westwood) Menida sppNezara viridula (Linnaeus)

Oebaluspoecilus (Dallas) Oebalusypsilon-gnseus (de Geer)Oebalusgrisescens (Sailer) Tibracea limbativentris (Stal)

Aspavia armigera(Fabricius)Diploysfissa Erichson Acrosternum marginatum (Palisot de Beauvois) Acrosternum sppFrankliniellarodeos Pergande Breginatothrips enustos Watson Stenchaetothrips(=Baliothrips)biformis

(Bagnall) (=(Jlloethripsoryzae (Williams)HaplothripspriesnerianusBagnall

Distribution

Asia Africa Asia Latin America Latin America Latin America Latin America Latin America Asia Asia Asia Asia Asia Latin America Asia Asia Africa Asia Latin America Asia Asia Asia Latin America Latin America Latin America Latin America

Asia Africa Asia Asia Asia Asia Asia Asia Asia Africa Africa Africa Asia Latin America Latin America Latin America Asia Asia Asia Asia Africa Latin America Africa Latin America Asia Asia Asia Africa

Latin America Latin America Latin America Latin America

Africa Africa Latin America Latin America Latin America Latin America Asia

Asia

Reference

59 241 217 59 241 6166 61 66 81 150 6166 66 81150 b 178 178 30 83 42 81150 203 30 80 62 178 195 178 178 178 203 203 203 203

55 178 12 72 78 120 214 214 b 59 120178114 178 54 636 82 12 6 12 82 178 81 150 60 is 195 59 120 178 78 36 186 36 186 b 55 12 30 54 55 78 82 120 61 66 81195 61 65 66 195 211 150 195 61 66 81 150 219 233 6 12 82 6 82 186 186 61 66 68 213235

178

7 IRPS No 123 January 1987

Table I continued

Order Family Species Distribution Reference

toleoptera Scarabacidae leteroligus (=Aphonoprocts) 15elcs Billberg Africa 36 Dynastinac (=Ileteroligusmees robustui Prell)

Alissonotum pauper Burnieiser Asia 82 Alissonotuon simile Arrow Asia 209 Dscinetusdubibs (Olivie) Latin America 28 195 233 Devscinetusgagates Burmncster Latin America 20 195 Dyscinctt spp Latin America 60 Eutheola Inimilis Bturmcistr Latin America 20 28 8 1 195 203

233Eutheola identarr(Biurnciter) Latin America 121 203 Heteron)ciwv aw rsonJack Africa 82 Hcreron)chusbtuterculatus Kolbe Africa 82 tleteronichtslicas (Klug) Africa 82 fhteron)chus mosamhicus Peringuey Africa 4 6 48 82 (=Veteronychus orzae Britton)

leteronychts lioderes Redtnbacher Asia 77 82 207 ltcteronchusarator (Fabricius) Africa 82 tleterontchusplebepus (Klug) A frica 57 82Ileteron chus pseuocongonnsis Ferreire Africa 32 Ileteronvchns rogiJrons Fairmaire Africa 82 Heteron chus nrsticus niger (K lug) Africa 82 Heteronvchus spp Africa 17 Lachnosterna longipennis (Blanchard) Asia 82 Lachnosternaspp Asia 120 Ligvros fossator Burmeister Latin America 82 Ligyrus nasutus Burmeister Latin America 203 Ligyrus (=Scarabaeus)ebenus (de Geer) Latin America 20 195

Dynastinac Ligtvros huonilis Burmeister Latin America 60 82 Aaladera castanea (Arrow) Asia 82 Maladera orientalis Motschulsky) Asia 82 Malaaerajaponica Motschulsky Asia 82 Phvlognathusdicynsius (Fabricius) Asia 82

Melolonthinae Exopholis h)ypoletca (Wiedemann) Asia 117 Lerlinosterna[= lolotrichia] consanguinea Asia 122

(Blanchard) Leucopholis rorida (Fabricius) Asia 82 117 Lepidiota blanchardi Dalla Torre Asia 76 82 Leucopholis irrorata (Chevrolat) Asia 76 134 Scrica internpta Walker Asia 76 Holotrichialonipennis (Blanchard) Asia 122 Holotrichia serrata Fabricius Asia 122 Holotrichia leucoph thabna (Wiedemann) Asia 82 Holotrichia seticollis Moser Asia 77 78 Holotrichia pnindanaoana Brenske Asia b Stenoeres spp Latin America 71 195 Atoenius spp Latin America Asia 203 Ph luplg aequata (Bates) Latin America 203 Piyvilophagacaraga Saylor Latin America 203 Phyllophagachiriquina (Bates) Latin America 203 Phyllophagadas)poda (Bates) Latin America 203 Phyllophagaclegans Saylor Latin Ameica 203 Phyllophagahondura Saylor Latin America 203 Phyllophagalatipes (Bates) Latin America 203 Phyllophagamontriesi (Blanchard) Latin America 203 Phyllophaga pamisetis (Bates) Latin America 203 Ph vllophaga sanfosicola Saylor Latin America 203 Phllophagasetifera (Burmeister) Latin America 203 Phyllophaga (=tlolotrichia)helleri Breuske Asia 82 117 159 Phvllophagaspp Latin America 81 150 White grub unspecified Asia 84

Rutclinae Lagochile trigona (Ilost) Latin America 82 Anomala dinidiata var barhata Burmeister Asia 77 78 82 Anomala huteralis(Burmeister) Asia 76 Anoimala lurida (Blanchard) Asia 76 Anomala sulcatula (Burmeister) Asia 76 Anontala varians (Olivier) Asia 82 Anomala antigua (Gyllenhal) Asia 82 Anoinala palida (Fabricius) Asia 82

8 IRPS No 123 January 1987

Table 1 continued

Order Family

Tenebrionidae

Elateridae

Languriidae Chrysomelidae

Chrysomelidae

Coccinellidae Curculionidae

Lepidoptera Lymantriidae

Amatidae

Arctiidae

Limacodidae Eupterotidac Noctuidac

Species

Anomaa volita (Blanchard) Adoretus caliginosus Burmeister Adoretus comprcssus (Weber) Papuana huhneri (Fairmaire) Papuana incrmnis Prell Popillia capricollis Hlope Gonocephalu depressur (Fabricius)Gonocephahm acuiangulum (Fairmaire)Gonocephalum sinphx (Fabricius) Epitragus sallei Champion Anaedus ptnctatissimnts ChampionAeolus spp4griotes spp Agriotes mancus (Say) Conoderts spp Aeoloderma brachmana (Candeze)Wireworni unspecified Anadastus filijf rmis (Fabricius)Dicladispa viridicanca(Kraat z)Trichispasericea (Guerin)Oediopalpastervalis (Weise) Oediopalpa guerini (Italy) Chaetocnema cvlindrica (Baly)Chactocnema denticulata S tephens Chaetocnemabasalis (Baly)Aulacophora sp nr similis Olivier Monolepta caripemne Ialy Monolepta bilasciata (Ilornstedt) Monolepta signata Olivier Cerotoma airojcsciata J acoby Diabrotica adelpha Ilarold Diabrotica balteata Leconte Diabrotica graminLa ltaly Diabrotica speciosa (Germar) Diabrotica limitafa quindecinmunctata(Germar)Diabrotica melanocephala (Fabricius)Diabrotica spp Altica (= Fondia)madagascariensis(Allard)Altica spp Epitrix cuctmeris (Harris) Epitrix spp Oulenjo or pae(Kuwayama) Disonchq (= Donacia) spp Flea beetle unspecified Chnootriba (= htIilachna)sinilis assimilis Mulsant Atactogaster iduriens Walker Hypomeces squtamosus (Fabricius) Tan vinecus discoidalis GyllenhalPhaulosoinusmusculinus (= mnus) Csiki Neobaridia anplitarsis CaseyPsalis pennatula (Fabricius) Laeliasufihsa (Walker) Euproctis irgtncula (Walker)Euproctis minor (Snellen)Euproctis xanthorrhoea (Kollar)Amata sp

Diacrisia obliqua (Walker) Creatonotusgangis Linnacus Latoia (= Parasa) bicolor (Walker)Nisaga simplex Walker Sesamia botanephaga Tams amp Bowden Sesamnia nonagroides botanephagaTfums et Bowden Sesamia calamistis llampson Sesamia inferens (Walker) Spodoptera eridania (Cramer)Spodoptera mauritia acron)ctoides Guenee Spodoptera frugiperda ( ESmith)

Distribution Reference

Asia 82 Asia 82 Asia 82 Asia 82 Asia 82 Asia 78 Asia 82 117 178 Asia 82 117 Africa 82 Latin America 203 Latin America 203 Latin America 195 Latin America 195 Latin America 203 Latin America 195 Asia h Asia Latin America 60 73 117 Asia 200 Africa 36 Africa II Latin America 6166 68 Latin America 6166 68 Asia 225 Latin America 61 66 68 81

Asia 178 Asia 55 Asia 55 Asia h Asia 55 Latin America 203 Latin Amefia 203 Latin America 203 Latin America 203 Latin America 6166 68 Latin Amcrie 195 Latin Amw 195 Latin Aierwca 61 81 Africa 57 Asia 178 Latin Americ 203 Latin America 81 Asia 82 Latin America 195 Asia 120 Africa 6 11 Asia 185 Asia 120 Africa 82 Africa 82 Latin America 66 68 195 233 Asia 117 Asia 117 Asia 117 Asia 117 Asia 117 Asia A S Pradhan

unpubl dataAsia 82 Asia 82 Asia 82 Asia 182 Africa 12 Africa 6 36 Africa 6 8 11 12 36 Asia 59 82 84240 Latin America 150 Asia -0 59 196 Latin America 28 61 66 68 81 150

152195203219233

9 IRPS No 123 January 1987

Table I continued

Order Family Species Distribution Reference

Spodoptera exigua (Hubner) Spodoptera exem pta (Walker) Spodoptera ornithogalli (Guenee)

Africa Africa Latin America

32 217 32 33 36217 150

Mythimna loreyi (Duponchel) Asia 30 Mythimna separata (Walker) Mythimna (- Pseudaletia)latifascia [= adultera

Asia Latin America

178 195

(Schaus)] (Walker)Mythimna (=Pseudaletia) sequax (Fabriclus) Latin America 68 Mythimna rosellinea (Walker) Asia b Mythimna yu (Guenee) Asia b Mocisfrugalis (Fabricius) Asia 30 117 Mocis latipes (Guenee) Latin America 66 68 149 150 203

Platysenta (=Spodoptera) compta (Walker) Asia 219233A Barrion unpubl data

Agrotis ipsilon (Hufnagel) Latin America Africa

33 68 81 150 1)5 203

Agrotis spp Latin America 60 Achaeajanata (Linnaeus) EldanasaccharinaWalker

Asia Africa

117 36

Armyworm unspecified Asia 72 84 Pyralidae Cnaphalocrocis medinalis (Guenee)

Marasmia bilinealisHampson Asia Africa

30 4559 72 84233 36

Marasmia trapezalis(Guenee) Africa 32 217 Marasmia (=Sumia) exigua (Butler) Asia 220 Marasnianmralis (Walker) Asia b Marasiniapatnalis Bradley Marasiniaspp

Asia Asia

b b

Leaf roller unspecified M2liarphaseparatellaRagont

Asia Africa

72 12 57

Maliarphasp Asia b Chilo auricilius Dudgeon Asia b Chilo partellus (Swinhoe) Chilo polychrysus (Meyrick)

Africa Asia

12 92 59 72

Qhilo diffusilineus J de Joannis Africa 6 1229 Chilosuppressalis(Walker)Cibozacconius Blezzinski

Asia Africa

59 72 6 11 12

Cravubus spp Latin America 203 Acigona loftini (Dyar) Latin America 203 Acigona chrysographella(Kollar)Scirpophagaincertulas(Walker)

Asia Asia

82 59 72

Scirpophagainnotata (Walker) Asia 59 200 Scirpophaganiuella (Fabricius) Asia b Elasinopalpuslignosellus (Zeller) Latin America 28 60 61 66 81 150

Diatraeasaccharalis(Fabricius) Latin America 162195203219233 28 66 81 111 150 195

Rupela albincllaCramer Latin America 111

Gelechlidae

Stem borer unspecified Undetermined stem borer nr Maliarpha BrachniaarctraeaMayr

Asia Asia Asia

72 b 55

Brachmi spp Africa 32 Hesperiidae Parnaraguttata Bremer et Grey Asia b

Parnaranaso Fabricius Asia 55 Pelopidasagnaagna (Moore) Pelopidasmathias (Fabricius)

Asia Asia Africa

164 36 155217

Pelopidasconjuncta conjuncta (Herrich-Schaffer) Asia b Borbo fanta Evans Africa 6

Satyridac Melanitisleda ismene Cramer Asia 84213 Mycalesis asophisHew Asia 30

Diptera Cecldomyiidae Mycalesis spp Orseoliaoryzae (Wood-Masn)

Asia Asia

b 72 88 131 181

Orseoliaoryzivora Harris et Gagne Africa 61132218 Chloropidae Steleocerellus (=Mepachymens) ensifer (Thomson)

Oscinella spp Asia Asia

205 205

Gaurax sppChloropsoryzae Matsuntura (= kuwonae Aldrich)

Asia Asia

205 82

Mucldae Atherigonaoryzae Malloch Asia 30 82 117130 Atherigona exigua Stein Asia 82 117 178

10 IRPS No 123 January 1987

Table I continued Order Family

Diopsidae

Agronyzidae

lHymenoptera Formicidae

Species

Atherigona oritentalisSchiner Atherigona indica MallochDiopsis longicornis(= thoracica)Macqurt Diopsis spp Pseudonapnyza asiatica Spencer Pseudonapomyza spicata (Malloch) Agronoza orvzae MunakataAcromyren landoltibalzani(Emery) Acromyrmex landoltifracticornis(Forel)Acromyrnex heyeri Forel)Atta bisphaericaFore Atta capiguaraGoncalves Atta laeLigata (F Smi h1) Atta opacicepsBorgmeierSolenopsisgeminata (abricius)Pheidole sp Pheidolegeton diversus Jerdon Monomorium pharaonis (Linnaeus) Ants unspecified

Distribution Reference

Asia 82 Asia Africa 82205Africa 11 1248 Africa 217 Asia 213 Asia h Asia bLatin America 6168 195 Latin America 61 195 Latin America 61 68 195 Latin America 61 68 195 Latin America 14 61 68 195Latin America 61 68 195 Latin America 61 195 Asia b Asia b Asia 1) Asia b Asia 120

aG orlentalls formerly recorded in Asia as G africana thPhilippines reported in this study CSubterranean root feeders

subterranean nests Foraging ants will feed on germinatedseeds only when the supply of ungerminated seed is low However these ant specialists feed on a wide varietyplants and thus help control grassy weeds (39) This dual

of

pest benefit role extends from tending ants which protecthoneydew-excreting pest species including root aphids and mealybugs (workers repel potential predators and even selectively kill parasitized aphids) to predators of leaf- and stem-feeding rice pests such as leaffolders armywormswhite grubs and stem borers (1 17)

The upland rice environment is particularly well suited to ants because of the lack of flooding Frequent tillage perpetuates grasses and granivorous species that are highlyadapted to constructing new nests in recently tilled fieldsAnt nests however are relatively shallow and tilling with a moldboard plow can destroy them Also upland ricefields are uually small and isolated and a given area in a smallfield has more field borders than it has in a larger fieldallowing high rates of infiltration from more sedentaryspecies such as Plieidole that do not colonize tilled fields but nest in soil under shrubs and trees along field bordersPheidole workers forage into fields from borders and store seed in their nests In slash-and-burn or other no-tillage rice cultures nests would be preserved in the field and possiblylead to high rates of seed predation However granivorous species do not prefer forest habitats

Damage is characterized by reduced usually patchy plantstand (120) Seed-thieving ants greatly prefer rice but they can be selective regarding plants they collect and store Theyharvest more seed than they can consume so patches of germip ing surplus seed (such as Rottboellia) will sproutfrom their nests during the rainy season

In Latin America leaf-cutting ants (Aita spp and Acromyrmex spp) occur only in upland fields and defoliate

young rice plants (47 68) They take leaves to fungal gardensin underground nests where fungi predigest the plant material

Termites Subterranean termites of the family Termitidae which lack symbiotic protozoa to help digest plants are the most frequently mentioned termite pests of upland rice (1185 120) This family cultures fungi in special undergroundcells --- fungal combs made of half-digested plant material (117) The fungi inoculated onto the combs break down plant material into food that termits can digest The termites then consume the fungal combs These grasslandtermites build nests below the plow layer in uplandricefields The colonies attack living plants only when dead plant material is not available They will attack a droughtshystressed crop but prefer older plants having greater cellulose content They will damage newly planted crops where clean culture has removed vegetation

Infestations are worse in deep light-textured soils with low moisture content The first sign of damage is yellowingof older leaves (195) Termites feed on roots (the plantyellows then wilts and finally collapses) and germinatingseed (loss of stand) or move above ground at night to cutseedlings at ground level which they cover with soil for later consumption Soft-bodied termites are highly sensitive to desiccation They live within a self-contained system of airtight chambers and build surface tunnels linei with nudand body secretions (carton) to maintain more than 90 relative humidity Termites can be located by their tunnels

Termites apparently can withstand limited submergence (157) and can be a pest of dry lowland seedbeds (117)perhaps by having nests below perched water tables

Termites appear to do more damage in Latin America where large ricefields are plowed with tractors deprivingtermites of preferred vegetation and forcing them to live on

rice plants In Asia upland ricefields are typically small with Iprge field border areas (ecotones) where termites can forage for preferred food other than growing rice plants All grassland termites do not feed on rice In an upland area in Batangas Province Philippines ricefields are commonly infested by Hosiititernesluzonensis (Oshima) however this termite does not damage rice In the same province though Mrroternesgivus(Hagen)feeds on rice seedlings (178)

In Africa termites may damage rice even though fields are small and surrounded by perennial vegetation These termite species perhaps prefer rice over other hosts

Mole crickets Nymphs and adults are nocturnal and burrow through soil feeding on roots of a wide variety of plants or forage above ground as predators of insects Adults live 3-5 mo and are even cannibalistic Mole crickets prefer low-lyin moist upland soils with high organic matter (II 81) In the lowlands they inhabit rice bunds or nonflooded fields Adults are highly mobile and can leave a flooded field to locate a more suitable habitat Losses show as wilted plants (from root feeding) or reduced stands (120) Damage is more common near field borders where mole crickets relocate after tillage (126)

Fiehl crickets Nymphs and adults of field crickets have similar nocturnal habits and damage rice as much as mole crickets do Piles of weeds removed from fields attract them They make subterranean nests and tunnel through the soil to feed on roots Some species prefer seed to roots Others feed at the base of stems causing deadhearts

Tunnel entrances surrounded by excavated soil are easily seen Young plants are cut at ground level and stored in underground cell Field crickets tre highly polyphagous as are all soil pests (117) Sachan et al (198) reported that field crickets prefer maize to upland rice Like mole crickets field crickets cannot survive in standing water and are therefore more prevalent in upland fields

Whitegrubsandhackbeetles Scarab beetles that feed on living roots as larvae but not as adults are called white grubs or chafers (Melolonthinae Rutellinae) and those that feed as adults but not as larvae on plant crowns are called black beetles (Dynastinac) In the tropics chafers or white grubs have a I-yr life cycle starting with adults emerging from the soil about I mo after the first downpour of the rainy season (134 159) They lay eggs at the same time farmers sow upland rice The rice passes its most susceptible stage and damage is mostly avoided when the white grubs are small After several months the long-lived larvae are large enough for two or three to denude the root system of mature rice (223) This intensity of damage is rare but wilting occurs when root loss is combined with drought stress

The larvae need damp (not saturated or dry) soil to survive and pass the unfavorable dry season 05-2 m underground in aestivation in the moist soil The first heavy rains of the season (20-30 amd) stimulate the grubs to resume activity and after several weeks they develop into

IRPS No 123 January 1987 11

pupae and adults and eventually dig their way to the soil surface and fly to nearby trees to seek food and mates The adults of most white grub species feed only on trees but in some species notably Eutheolahumilis the adults defoliate and cut off stems at ground level (28) Putheola hidentata damages sugarcane fields in Venezuela (121) Floods kill white grubs (208) Grasslands can support large populations and therefore white grubs can be more abundant in newly planted upland ricefields that were previously fallow (81)

Black beetles live up to a year adults can attack a crop at any stage by burrowing into the soil (127) and feeding at the base of stems to cause whiteheads (207) Adults emerge with the early rains and are normally more abundant and do more damage on a young rice crop with small root systems (38 81)

The larvae feed only on dead organic matter in dryland fields and do not attack rice Adults arc highly mobile and although sensitive to flooding can invade a field soon after it drains (115)

Root aplhids Several genera of migratory aphids feed on the roots of upland rice (249) Populations build up more in light-textured soils with high percolation rates Damage has also been reported from dry seed heds of lowland rice (52) Tending ants -- Pheidhle Cren aster Tetramoriuw lsius Tapinoma -- are necessary for root aphids to multiply Aiits harbor aphids in their nests over winter or during unfavorable periods and relocate them on rice plants digging tunnels along the root systems to allow these softshybodied insects to penetrate the soil Most intensive studies on aphid life histories have been done in temperate environments (158 222 229) Root aphids fly to rice plants at the beginning of the rice season and pass through several generations continually developing winged forms to relocate on alternate grassy hosts until winter when they move to perennial hosts usually trees Yield loss occurs mainly through reduced tillering but infested plants become yellow and stunted and in extreme cases wilt (227) Aphid populations build up gradually so damage usually begins in the late vegetative and reproductive growth stages

Root-feeding ineal tugs More than half of the mealybug species or rice feed on roots (245) Being soft bodied mealybugs are not adapted to living underground but they survive because tending ants (the same species that tend root aphids) dig burrows for them and move them from plant to plant Well-drained soils ihelp mcalybugs survive Although six species are recorded as subterranean root pests of rice we found no report citing damage Root-feeding mealybugs are probably more prevalent on perennial grasses than on short-lived annual rice

The most common rice mealyb ig Brevennia rehi is a foliar feeder favored by lack ol rain apparently it is responsible for high yield loss in Northeast India and Bangladesh (N Panda pers comm as stated by IRRI [1061) This report needs confirmation Mealybug damage was only briefly mentioned in another report (156)

12 IRPS No 123 January 1987

Root weevils A few species of upland weevils (IHporneces Donacia Atactogaster) attack rice roots as larvae and feed on stems and leaves as adults Damage ispatchy The larvae are not adapted to submergence and are true upland soil weevils isopposed to the so-called water weevils They rarely become sufficiently abundant to cause damage Adults on rice foliage or the soil surface show that the roots are being attacked by the grubs Root weevils are highly polyphagous (185)

Root heeths Soil-inhabiting false wireworms Gono-cephalumt spp are reported in upland ricefields The adults and larvae normally feed on decaying organic matter but land preparation for upland rice removes these sources The larvae feed on seedling roots and nocturnal adults roam on the soil surface to cut off seedlings at ground level (82) This group of ground beetles is highly omnivorous adults even prey on stem borer larvae (183) True wireworms (Elate-ridae) feed on living roots but rice as a host has been reported Infrequently (117) In Latin America the highlypclyphagous larvae of Diabrotica rootworms are also recorded on rice

Root htus Upland rice seedlings can be attacked by the adults anI nlViphs of chinch bugs Blissusand Caenoblissus and bro n bug Scaptocoris which feed on the roots with theirsucking nouthparts (80 83) Three genera of burrower bugs A tms (eotonhus and Stiharopus -are common iil grasslands in the Philippines and may attack the first crop (rice or maize) planted after the dry season Chinch bugs are most abundant after a dry spell These insects are large and can kill seedlings or stunt and reduce tillering on older plants I hey prefer other hosts and rarely damage rice Adults can also feed oii developing rice grains (81)

(utworms Soil-dwelling noctuid larvae hide under-ground during the day to avoid predation by birds and bccone active at night above grour cutting off you ing plants at ground level (hence tile naiie cvtworn) arvaedrag severed plants into hurrows llie larvae have wide host ranges arid pupatc in the soillgrotis ipsilon occurs worldwide and appears to be the most common cutworm pest of upland environments but is more prevalent inlowland rice particularly in dry seedbeds

IFoliar pests Other groups of insects are adapted to upland environments because they pass one growth stage underground either as egg (in heinrietabolous species) or pupa (in holomnetabolous groups)

Rice seedlibig maggots AitJeriota flies occur in non-fhoodcd environments in Asia and Africa (180) Feeding of larvae one per iller causes deadhearts Flies as opposed to moths only secondarily evolved as pliytophagous pestsfroni a saprophagous Origin Athrnonalarvae secondarihv feed onidecaying tillers whereas larvae of stem borer moths actively Iced on living tillers to cause deadlicarts Seedling maggots will die if they cannot sever a tiller on which to feed

as it decays Plants die under heavy attack whereas lighter infestations cause stunting delayed maturity and ragged leaves (236)

Eggs are laid only on actively growing plants (49) aid damage occurs wiihin the first week after crop emergence(170) Larvae pupate more in stems than in soil Seedling maggot attack is highly seasonal (206) normally peaking 2-3 mo after the onset of the rainy season These pests possiblyaestivatc over the dry season Rice isonly one of their many graminaceous crop and weed hosts

The rice stem maggot (hlorops orzae is adapted to temperate upland and lowland rice growing regions of Asia In Japan and China upland rice grows in mountainous regions a habitat of this chloropid fly The larvae tunnel into tilestems The stem maggot is more damaging during the seedling stage larvac hibernate in grasses

File several dipterans reported as stem maggots in dryshyseeded lowland seedlings inIndia (205) could presumably attack upland rio

Leaf urinrs Icaf miners are common on wheat and maize but are rarely reported on upland rice (213) Three leaf miners are more prevalent on upland than lowland rice inthe Philippines Pvudonalomy a asicanh Vficataand lIAgromrza orzae attack at the seedling stage The larvae develop inside the tunnels they construct in the parenchyma tissue P Vpicata and Aorvzac prefer maizeEehtsine and other grasses to rice spicatacan become abundant on wheat

Leaf beetles Chrysomelid and coccinellid eaf bcetles scrape or otherwise remove leaf tissue as larvae or adults or both This damage accelerates desiccation of plants Tile larvae of sonic species are root feeders Flea beetles Epitrix and (lraetocnetna and rootworms l)iahroticaspp feed on rice foliage only as adults Their root-feeding larvae have hosts other than rice The adults and larvae of the hispa-like Oediolmlpa spp beetles of Brazil and (hmootriha (=EPfishyachra) beetle of Africa feed on upland rice leaves Oealioshypapa spp larvae are leaf miners (mootriha is also pshyvalent in lowland environments (11) Leaf beetles prefer vegetative rice (81)

The rice hispa Dicladipaarnigera is not reported on upland rice and prcers more aquatic habitats In Central Africa however I)cladispm viridicyaiea attacks uplandrice in the vegetative stage as well as lowland seedbeds 7richipasericea is reported on upland rice in West Africa and Madagascar (57)

The rice leaf beetle 1d1ea 01izae riiuech like Chioropsor|rzae is adapted to temperate regions of Ashi and is usually a pest on lowland rice but seldom oi inpland rice in the mountains (82) The adult hibernates on grasses

Armmworms Species of grassland-adapted SpoptloeraPlatysemta and A ithimna larvae create widespread epishydemics They can defoliate ricefields generally in patchesfrom early vegetative growth to harvest (41) Armywornms pupate inthe soil and highly favor upland rice (34) Adults

can disperse long distances to colonize even remote upland areas They become more abundant than their natural enemies particularly after favorable rains following a prolonged drought (196) The drought kills the armyworm and natural enemies (mainly parasites) but the armyworm can recolonize rapidly

The natural increase insoil fertility from mineralization of soil N over the drought period promotes luxuriant plant growth to foster a rapid buildup of these highly fecund species Weeding rice increases tilelikelihood of damage The parasites normally return to control the armyworms but not until after serious damage

Outbreaks on upland rice have been reported in Ghana (3) inzibar (38) Central Africa (36) lPanama (165) Brazil (28) Malaysia (196) and India (167) Defoliation may be severe often leaving only the base of stems Armyworm larvae hide during the day under litter or in soil cracks At night they ascend plants to feed Larvae pupate in the soil

Thrips Thrips frequent upland rice but damage is less than in the lowlands Thrips larvae and adults feed on leaf blades by rasping causing leaf rolling and stippling Thrips prefer vegetative stage rice and their survival increases during drought (heavy rain washes them from foliage) although drought stress usually outweighs thrips damage Drought and thrips cause leaf rolling in upland rice (235) but thrips have been overlooked perhaps becaLse of their small size Thrips are most numerous indry season irrigated rice The combination of favorable plant growth and lack of rainfall is ideal for thrips In contrast the lack of rainfall in upland rice hurts the crop and eventually the thrips whose numbers must decline

In [atin America Frankinilla rodeos attacks panicles before they emerge from the boot calusing sterile grains

Stem hug~s Black bugs are the most common group of rice stem-feeding -lemiptera in Asia The most frequently mentioned species ----Scotinophara coarctaia and Shrida - prefer aquatic habitats and therefore could not occur in upland rice as sometimes reported Upland species in the same gerurs - notably S tarsalis and S scotti -- are morphologically similar and remove sap from stems of grasses and upland rice The upland species are notably more abundant in upland rice planted near forests

In Latin America tile Tibraca ima-large pentatomid tiventris causes deadhearts and whitehcads on upland rice Both adults and nymphs remove sap from internodes of plants more than 3 wk old Tillers are killed by mechanical damage and by the entry of secondary microbial infection This damage is often confused with that of lepidopterous stem borers (233) Eggs are laid on the leaves and stems Nymphs are gregarious This species can also be found on irrigated rice Drought reduces population buildup

Stiem borers Lepidopterous and dipterous stem borers are widely recorded on upland rice but cultural practices and alternate hosts determine abtndance Except for Elasmolmll)uS Iinosellts numbers usually build up toward

IRPS No 123 January 1987 13

the end of the crop growth cycle Therefore late-maturing varieties staggered planting and lack offloodingfavorstem borer buildup Many stem borer species can lie dormant in the stUbbleafter harvest and theirsurvival is encouraged by panicle harvesting (leaving tall stems) and lack of tillage after harvest

Stem borers disperse and lay many eggs Many species are polyphagous however the monophagous MaliarpiasepashyratellaDiopsis Ivmgicornis Scirpophaga incertlas and S innotata can also be abundant on upland rice (hilo Suplressalisdevelops fasterand becomes la rger when reared on upland than on lowland rice (226)

The lesser corn stalk borer lasmopal tt Ittoselhs is semisubterranean and is perhaps the stern borer most adapted to upland rice(129) Highly polyphagous it infests maize peanut and cowpea It attacks seedling rice (one larva can kill tip to four plants before maximum tillering) (195) (hiv and Maliarphastem borers prefer older plants But Scirpophagaspp will attack young plants even in the nursery lasn opalls larvae tunnel into the stems at or below the soil level mausing dcadhcarts Larvae are not found in their turnnels inside sterns beca use vhei disturbed they retreat into cases made from soil particles bound by silk The cases areattached to the tunnel entrance (143) This behavior probably evolved as protection from natural enemies Larvae pupate in the soil The lesser corn stalk borer is more abundant during drought damage is often confused with that of drought (233) This species prefers sandy soils arid its distribution within a field may relate to soil texture (143)

Diatraeasaccharalis aside from the lesser corn stalk borer is frequently a stem borer pest in Latin America According to Teran (233) rice and maize are more preferred hosts than sugarcaine its namesake

Atcitonacrvsograpla is a less known upland rice stem borer in Asia with habits similar to Easinopall s (82) because it is not found in lowlands Acutra has grassy weed alternate hosts As a moth it is often confused with the dark-headed stern borer(hi polur(standstriped stern borer (hi sup)lpsrcais

Other common upland rice stem borers are also reported in the lowlands The dominant species in upland rice vary greatly with location In Uttar lPradesh India (78) and in Japan (123) the pink stem borer Sesamia htfirens is more prevalent hIKenya the principal rice stem borer is Chi partelus

(92) The lowland African sten borers Aialiarpha sepashyratellaDiopsis vngicornis and Sesamia calatnisnis are less abundant Oimupland than on lowland rice(217) but perhaps cause greater yield loss because plant injury is greater if combined with drought css (92) The stalk-eyed stern borer D)ivpsis is more prevalent in wet habitats (6 1I217)

In Central and West Africa Maliarpha and Sesamia calanistis are the dominant stern borers of upland rice (8 9 10 II) (lilo dihlfilineui and Chio partellus inhabit the

14 IRPS No 123 January 1987

upland savannas whereas Chilo zacconio prefers the lowlands (II) Aaliarpha is most abundant in the mangrove swamp habitats (91) coastal regions (16) or rain forest zones (II) of Africa and appears to be the homologue ofScirlpophaga incertulas The yellow stem borr can also be prevalent in upland rice (120 167) The sugarcane borer Eldanasaccharinais so far only a potential pest of uplandrice (91) despite Grist and evers (82) warning

Grasshotl)ers andI locitas Short-horned grasshoppers and locusts -- Locvta Patanrga Schistocerca - layeggs in the soil inhabit grasslands and develop into swarms to seriously damage upland rice from time to time (173) Largenumbers can destroy a field leaving only the stubble In low cmuoroer s giassnoppeirs an cut panicles (173) Farmers may not plant if locust swarms are imminent Prolonged dryweather followed by favorable rain favors development of large swarms (173) Usually breeding in di) grasslands theydisperse toattack crops including upland rice(191) Eggsaresusceptible to desiccation d uring drough t and toinundation during seasonal rains With favorable soil moisture over consecutive seasons natural enemies ca nnot prevent a largepopulation increase leading to migratory swarms (64)

Most grasshoppers on rice are adapted to uplands (107)but lowland rice grasshoppers iervgl(rlhusspp and Ora spp occur occasionally on upland rice (2 82) They lay eggson rice foliage Nymphs are scmiaquatic

The gryllid Lsctrtus coucinnus is a pest of lowland and deep water rice in Bangladesh and Thailand In the Philippines it damages upland rice grown in isolated pockets near lakes The gryllid nymphs and ad ults feed on the central portions of folrage leasing only the midrib and leaf nargins

Leal ler Pvralid moths whose larvae fold leaves to make a feeding shelter occur worldwide (217) but arc appareritly most common in Asia Leaffolde rsattacking rice in Asia are a complex of species (26) (Ctalhahcrcisinedinalis the best-known rice Ia flolde r in Asia may prefer grassy weeds over rice (86) It is a late colonizer of uplandrice U1arasmja eviua a lowland species (220) nreflers rice to grass specres (86) and colonizes upland rice during il early growth stages llarasnria patnalis is the most corn-rnonly encountered species in upland rice in the Philippines Perennial grasses are their alternate hosts and they can survive year-round in upland rice environments The leaf-folders 11arasmia ruralisand Alarasmia spp are least aburndant Dormancy is unknown in leaffolders Cnaplha-ocrocis medhnalis however is known as a migrant (90)Leaffolders remove photosynthetic tissue attack during the flag leaf stage is particularly injurious Popurlations are normally held in check by natural enemies but leaffolders become parlicularly abundant in conditions of high plantfertility and shade

Butterflies Skippers PelopidasInathiasand Borhofanta are more prevalent in African lowland than in uplandhabitats (6) but extensive studies in Japan and China

showed that Pelopidas mathias prefers an upland habitat (155) Parnaragutata seasonally migrates between lowland and upland areas and is equally abundant in both environshyments (164)

The green horned caterpillar Melanitis leda ismene is more abundant in lowlands (212) but our experience in the Philippines suggests that Pelopidas niathiasand 11elanifis leda ismene are often more abundant in uplands than in lowlands Bamboo iaunbuiva spp annually sustains P mathiaspopulations in the Philippine uplands Adults feed on nectar from flowers and migrate to upland rice during the late vegetative stage

iPi erlled h i-1 1d dill L tIC i 7-0 l(become numerous probably because of high egg predationEgg and larval parasitization are often low Butterflies can readily disperse long distances to seek renlote uplandricefields However they have low biotic potential and even it predation rates are low the rarely become abundant Upland environments with their diverse microhabitats can provide more favorable sites than lowland rice plains

tutterfly pests of rice have broad host ranges and become dormant Juring urnfavorable times of the year

IolhagrotslaidopwraThe larvae of many polyshyphagous moths are reported to defoliate upland rice tussock moths lsaispelnnatula Iawlia sUifita luproctisirgtwltaE mnorand P vathorrhoeaslug caterpillar

litotiahicolorhairy caterpillars Nisaga sinp vandA mata spp and wooly bears l)iacrisiaohliqtia and Crvatonotts gangis Many of these polyphgous larvac migrate and are transients in rice including HelhWhis arm iera and 1t zea highly pol phagous species (82)

Gall nidges The Asian rice gall midge Orseoliaorrzae (88 131) and its African counterpart 0 ortzivora(6 11 12 32217) prelerlowland to upland rice Gall midges in rainfed areas are prevalent only during tle rainy season and dispersepoorly Their alternate hosts (wild rices LeersiaIschaetnuan and Pasahmt)are mostlyaquatic grasses and are less likelyto be near upland rice areas In an area of mixed uplandfloating and rainfed lowland rice cultures in northern Thailand upland rice which is ilanted a month earlier was attacked firstbut subsequcent populations were much lower than in rainfed lowla id or floating rice Low numbers of gallmidges passed the dry season orr perennial weeds such as Paspahim (88)

Smot (hid(s Reports of aphids attacking upland rice plants are few (249) Without maize the maize leaf aphidRho alosihhunm maidis attacks upland rice in the Philipshypines The plum peach aphid IIrsteroneuraselariae was reported in Sierra Leerne toinfest leavesand tnriperied grain(7) Siphaglrctriaecaused transverse linrea r necrotic striae iir rice leaves in Italy (172) I-lea v rains held populations in check Rice is not a preferred host for shoot aphids

Leafhoper andI )anthol)pers Cicadellids and delshyphacidsdispersecasilyand readilycolonie upland ricefrom nearby lowland areas A study in Sarawak (241) showed

IRPS No 123 January 1987 15

similar species in upland and lowland rice Nephoiettix The principal vectors of rice virus diseases generally have species dominated The most prevalent species accounting narrow host ranges and alternate hosts in the grass family for 90-95 of the sweep net samples wre N virescens N are mostly annuals dying in the dr3 season Some virus umropictus Nilaparvia hgens Soatellafiircifera Cfana vectors such as BPH can disperse lontg distances but few do

sectraand Recilia dorsalisLowlands had higher popula- Th restriction of upland rice and weed hosts to the rainy tions Katanyukul and Chandaratat (120) monitored hopper season the small area planted to upland rice and the hopper numbers in upland and lowland fields of northern Thaiiand vectors having narrow host ranges all work against virus and found similar results A comparative study of plant- diseases in upland rice hoppers in Fiji showed more brown planthoppe BPH) and In Latin America Sogctdhs o(rzicola and S cuhanus whitebacked planthopper (WBPH) in wetland rice than in transmit hoja blanca virus disease but reports on upland lowland rice and hopperburn was prevalent in lowland rice are rare except in the favored upland areas of Colombia areas with standing water (89) Peru and Venezuela (35 42) S rricolaocurs in Brazil

Hopperburn in upland ricefields in Asia is rare (84) In on upland rice but hojaI blanca has not been reported (67) Batangas Philippines isolated fields of upland rice become S orzicola has a wide host ramgc (10) similar to WBPI- in hopperburned infrequently from WBPH A report from Fiji Asia WBPH however is not known to transmit any virus cites hopperburn from BPH and WBPH (250) In these two disease situations upland rice was grown on highly fertic soil and In Africa pale yellow mosaic virus is endemic to swampy N fertilizer was applied Hoppers respond to better nutrition lowland areas where rice ratoons year-round (19) and is Normally upland rice is grown on poor soil and receives no mechanically vectored mainly by hispid beetles A minor fertilizer vector a flea beetle Chaeiocnema spp is an upland rice

In Asia upland rice ras are normally not far from pest and upland ricetields near endemic areas occasionally lowland ricefields-- a source of hoppers Otherwise hopper contract the virus species with greater polyphagy than BPH or Nelhotettiv Spittle bugs Cercopid nymphs producc a protective virescens would be dominant frothy covering that looks like saliva In Latin America

In Africa leathoppers and planthoppers do not become spittle bugs damage upland rice In parts of Brazil where abundant on upland rice (6 II) probably because lowland upland ricefields are surrounded by pastures adults rice is not widely rlanted nearby The only reported migrating from grasses can damage rice severely and cause hopperburn as from Nilapanrata maeander in breeder complete hopperburn in a young rice crop within a week of plots receiving high N rates in Nigeria (98) Hoppers colonization (194 195) Dispersal to rice occurs even if however are a greter problem in Latin America where pastures are verdant and spittle bugs prefer pasture grasses large grassland areas breed polyphagous species (Gra- to rice (168) Nymphs are rarely abundant enough to phocephala spp Hortensia spp lEvitianus ohscurunervis damage rice Regular rainfall favors their development Balclutha spp and Draeculacephala spp) vhich then (233) disperse to cause hopperburn on seedling rice (68) Rosetto et al (195) reported Graphocep7hala spp and Hortensia spp Seed pests to be equally abundant on upland and lowland rice Alydid and pentatomid bugs are seed pests of upland rice

The leafliopper and planthopper epidemics in Asia that worldwide In Asia Leptocorisa sp prevail Their habits caused severe losses from hopperburn and vijus dicases make them particularly suitable for upland rice 1)thev feed over the past several decades occurred principally on on many grasses (and therefore can survive the earl y wet lowland rice Except in a few isolated cases (160) upland season before rice sets its grain) 2)adults are long-ied and rice has been spared from tungro grassy stunt and ragged mobile allowing them to find isolated plantings 0 rice stunt virus diseases vectorcd by green lcafhopper (GLH) (besides seed bugs only birds and rats appear to have shyand BPI] (123) Virus diseases have occurred in upland rice capability) and 3) adults aestivate in forested areas or where lowland rice was nearby Rice dwarf disease however sugarcane fields during the dry sca son when neither rice nor was first recorded on upland rice in the Philippines (187) grassy weeds are present (I199) Rice seed bugs concentrate Virus diseases can be perpetuated only on livir plants or on small-scale upland riceficlds because they can actively insect vectors and no living rice - plans tooned or search them out The prevalence of groves of trees characshyvolunteer - grows year-round in upland arcas excert in a teristic of upland environments ensures nearby aostivation few with 12 mo of rainfall sites keeping the seed bugs close by

The smaller brown planthopper Laodcelphax striatellu Leptocorisae are uncommon in lowland rice plains occurs in upland rice in Japan (18) It spreads black- because 1) their populations are diluted in a sea of rice streaked dwarf and stripe viruses between winter wheat and planted more or less at the same time (they ca n only feed on barley to lowland rice Early planting of lowland rice rice during the milk to hard dough stages) 2) they lack nurseries spreads these viruses but upland ice has remained aestivation sites and 3) weedy fallow areas are limited uninfected The lowands of Asia are dominated by L oratorius

16 IRPS No 123 January 1987

which lays its eggs hign on tile foliage (199) L acuta and L solonlonlesis oviposit at ground level the former on litter and the latter loose on the ground These egg laying habis explain theirtenvironmental prefcrencecs(214) In Japan the postdiapause adults of L chinensis migrate from moun-tainous areas to upland ricefields during floweiing (1 14)

Pentatornid bugs prevail in Latin America (8182 195) and in Africa pentatomid and coreid bugs arc equally cited (6 82) l)amage is often characterstic of the species (5)rcoris ventralis and Mnida spp are the common

pentatomid seed bugs of Philippine upland rice but they prefer lEchinochva spp grass seed to rice Vearaviritda occurs worldwide and hoften more abundant in the uplands(54) We saw few reports of seed bugs on upland rice but see(d bugs apparently opelrate cqually between by land and upland rice Good dispersal is necessary ior species that call feed for only 2-3 xon a crop and must first find those crops at ripening stages

Other pct species feed n flowers In Mato Grosso Brazil (194) and Santa Cruz Bolivia the panicle weevil Aeo-hariuia amllitarsis attacks rice during flowering (68 233)The larva e rice sten horers adutlts feed at the bases of spikelets caIusing empty grains Also inLatin America thecydnid -lkiituttatralusisa seed bug as anad tilt and nymph (194)

A number of beetles ecd on poilen - coccinellids lialrotia Aulac lhora and Monohpl-a (55) - but becatise riCe is self-pollinating and fcrtiliation occurs before pikelets open they pose no threat to yield

IST ECOLOGY

We now look at characteiistics of the life cycles of upland rice insect pests to lea rn hew each plays an adaptive role in upland rice ecology

Life history strategies Orzasativadoes not tolerate drodgh well so upland rice is highly seasonal normally grown in the wettest months of the year Usually rice is present for less than half of the year The rice-fiee fallow poses serious problems of survival to upland rice insects which have evolved att least four mechanisms to ovrcolne tle cvcl ic-l lack of a hostI) polyphagy 2) longevity 3) dormancy and 4) vagility (dispersal)

Poldt)agy lolyphagy is defined as having hosts of at least two botanical families oligophagy of more than one gents and ionophagy of only one ventus (40) - inour case Orvza Most wild rices however are illi tiC and theieiirc are normally far removed from upland rice hahitat othewise more would be alternate hosts for uplanJ iice insects Thte highly mornophagous rice pests shySciriol)haita hiceritlas Nilal)arvaiahiens Iand Nealo--tettiyvircsctws in Asia and Aalirpha secparatella and 1ioj)sis lngicornis inAfrica - occur on upland rice but

are more abundant in lowland culture They can specialize in rice because they are highly vagile and can attack all growth stages By specializing in rice monophagous rice pests can outcompete related but polyphagous speciesIntensive andI extensive lowland rice culture has favored monophagous species Riqwla alhinella (Cramer) the onlymonophagous species in Latin America is rarely reported on upland ice (I1)

Because of its limited temporal and spatial existence upland rice has not favored the evolution of specialized species Clhang(44)shows evidence that upland rice has only recently been cultured by man from the lowlands thus pests have had little time to fully adapt to the crop All but the lowland-adapted insect pest species have alternate planthosts to rice Root pests with limited mobility are highly polyphagous Pests that specialize in one growth stage eg secdling maggots seed bugs are oligophagous or polyphagous

Most alternat hosts of rice pests are annual grasseswhich also are seasonal Upla_rice is also a host but not the preferred one of the most highly polypliagous spcciessoil pests grass hoppers armvwarrs and Vearaviirielda

The d ry season poses igreat obstacle to ani upland riceinsects survival and only sonic perennial plant hosts are suitable food during the off-seasoin Plolyphagy allows upland rice insects to suLrx-ixe during thC rainy season when plants- both annual and perennial -areactivelygrowing and therefore more nutritious

Loiugevitr The ability of aiinsect to live 2-3 mo without undergoing dormancy helps species sich as root-feedingwhite grbs (as larvae) and secd-feeding bugs (as adtilt) to survive unfavorable periods White grubs arc not highly fecund and their life strategy is to improve survivorship of limited progeny Their subterranean habitat hides them from manv natural enemies lBtut root tissue is not highly nutritious so they ltilst eat great quantities and need a largeinsect biomass to digest it Other soil-inhabiting rootshyfeeding insects have short life cycles but are more fecund

Seed pests need greater longcvi vas adults to locate ahost at early grain dcclopment arnd give sedentary offspring time to develop Few groups of upland rice insects live longin active development Polyphagy and longevity only increase their ability to survive during the rainy seasonOther mechanisms are needed to survive the dry season

orantc In the tropics the nonrice season is the dry season while intemperate areas tile off-+season is winter Many upland rice insects undergo dormancy inlsummer (aestivation) or winter (hibernation) Dormancy is simplyinactivity during winters with prolonged temperatures below 10 C insects find shelter arnd cease movement Activity is resu med with warmer t empcera tu res A deep state of irnactivity termed diapauise occurs in sole species that prepare physiologically (havxing low metabolism and convcrting gonad s to fatty fuel reserves) for the unfavorable season (148) They enter diapaUse inanticipation of unshy

IRPS No 123 January 1987 17

favorable weather and resume activity after favorable Armyworms skippers and locusts actively travel hundreds weather begins usually in response to photoperiod or of kilometers in air fronts such as intertropical convergencethrough a delayed response to a stimulus such as heavy zones to descend onto upland rice fields far from their rainfall (237) Not all individuals of a population have the breeding grounds (31) Several generations can build ip on same threshold for environmental responses for entering or alternate hosts in tile grass family before rice is plantedterminating diapause -- this is seen as a mechanism to These large insects are active fliers Smaller insects such as prevent total mortality if a false cue appears such as a thrips aphids and planthoppers passively migrate long sudden warm spell in winter or rain in the middle of the dry distances by the wind season (154) Otherspecies ean readily travel tens of kilometers - stem

Probably more upland rice insects undL go dormancy borers leaffolders leafhoppes cutworms spittle bugs leaf than we realize Candidates are Atherigona spp root beetles seed bugs ants and termites aphids root weevils crickets leaf beetles thrips and seed The least dispersive species are white grabs mealybugSbugs Insects known to entervarious states of aestivation are seedling maggots crickets weevils and gall midgesleptocorisaspp adults (199) Maliarphascparatellalarvae Each species has evolved a unique set of attributes to (9) Scirpophaga innotata larvae (117) Chilo partelhtsv enable it to survive and adapt to the changing uplandlarvae (21) ScirpopThagainccetulas (179) white grub larvae environments Several evolutionary avenues lead to fitness (134) Patangasuccincia adults (6 82) and Schistocerca (Table 2) but there is no reason to believe one set of gregariaeggs (82) attributes is better than another (224)

Insects known to enter hibernation are LodeIphaDstriatellus nymphs (124) Chtio Tvlpressalis larivae (125) DroughtSesaia inferens larvae (146) but not S calavisis(2) Well-drained rainfed upland soils are subject to drought ifIDiaraea saccharalis larvae (75) Scirpophaga- tcerlttas rains do not fall within 2-3 wk Prolonged drought followedlarae(5 lacchrli s lae (75)saSlOhagasshoperseggs (82)and felo pidas athias pOpae (232) by favorable rains stimulates armyworm and locust outshybreaks but mealybtigs root aphids and thrips become

Lcpmotorisa oratorisand L acuta adults aggregate on numerous onlyafter dry spells for two reasons First heavy trees sugarcane or other shady moist sites L paluwanmsis rain normally kills soft-bodied foliage-feeding thrips and confined to Sulu and Palawan Philippines lives on the mealybugs and soil-inhabiting root aphids and root mealyshygrass Brachiariamutica (Forsk) Stapf in the absence of bugs Drought eliminates this source of mortality Second rice The adults can live for 5 mo When aestivating their the rice plant responds to drought by breaking down rate of metabolism drops and they do not feed They will proteins into soluble N compounds which enter the phloem however puncture plant tissues of their aestivation host in and are taken up by these sap-feeding insects (245) Greater search of water This behavior has led to errors of the host nutrition therefore leads to greater survivorship and range of rice bugs (I) Aestivation quickly terminates with fecundity rain probably meaning aestivation is not deep Small rice area

Some stem borers aestivate as mature larvae in rice Except in the highly mechanized large landholdings of stubble or in straw stacked as livestock feed However Latin America upland ricefieldstypically are patchy within moriality increases proportionally with the duration of the a highly diverse flora The small fields result in higher ratios dry season even if the stubble remains undisturbed If the of perimeter to area than with the typically larger lowland stubble is plowed for a crop following rice few stem borers ricefields Upland ricefields themselves may be intercroppedsurvive Rainfall terminates dormancy but it normally with cereals legumes or root crops Mixed intercroppingtakes several weeks for moths to develop and emerge with a wide variety of species also occ in tribal slash-andshy

Last-instar white grub larvae tunnel 1-2 m deep in the soil burn agriculture (50 74) to construct pupal chambers in which they aestivate The Upland ricefields tend to be small because of the labor larvae are very sensitive to dryness so pupal chambers are needed to clear land in slash-and-burn areas and to till land sealed to coierve body moisture Aestivation terminates to minimize weeds Rice requires more tillage than does with the first heavy rains but as in stem borer 3-4 wk pass maize because rice competes less well with weeds Uplandbefore adults develop rice is more a subsistence than a cash crop and small areas

Insects may respond differently in each region to cold (I ha) can feed a family temperature or drought Biotypes or loal populations may Small fields favor some insect pests such as some ant evolve (56) Local populations of Scirpophagaincertulasor species that can forage from more permanent field borders Patangasuccincta may have become adapted to prevailing Trees provide food and mating sites for white grubs (134) as conditions well as aestivation sites for LetTocorisa spp (199) Field

iagiliti The ability to disperse combined with high border grasses plus tIhose within a ricefield provide alternate fecundity orshort life cycles enable some upland rice pests to food for all but the highly monophagous upland rice better exploit the temporarily favorable upland habitat colonizers and a habitat for natural enemies

0 z

Table 2 Life history patterns of upland rice insects or insect groups

Host range Longevity Dormancy Vagility FertilityMonophagy Oligophagy Polyphagy Low Medium High Yes No Low Dispersive Migratory Low Medium High(Oryza spp) (grasses sedges) (angiosperms) (lt3 wk) (gt2 mo) (lt1 kin) (lt50 eggs) (gt500 eggs)

Ants xTermites x x x

Mole crickets x x x xx x xField crickets x xx

White grubs x x x xx xRoot aphids x x xxMealybugs x

x x x xxSeedling maggots x

x x x x xLeaf beetles

x xx xArmyworms x x xx x

Thrips x x xx xChilo spp x x xx xMaliarpha x x x x

xLocusts x x Xx x xGall midges x x xxiVephotettL iirescens x x xx x x xLeprocorisaspp x

x xLeaftolders x x x

Sogatellafurcifera x

xx x x x x x x

00

IRPS No 123 January 1Q87 19

All this appeals to favor insect pest buildup but the notwithstanding the release of stored nutrients in response contrary is true Loevinsohn (138) showed that insect pest to drought that temporarily benefits philoem feeders populations respond exponentially with the proportion of land devoted to rice up to about 75 of the area then the M POR ANCE I I P1A 1) RWI P1St S

rate of response declines In most upland rico areas tie rice crop occupies less than 50i of the area- therefore the Visits to upland ricefields normally reveal 1kw insect pests potential for population buildup is low This relationship Loevinsohn (138) and Locvinsohn ct al (139) compared may not hold for seed pests that can locate small ricefields insect abundance using annual light trap atchcs in upland

Another outcome of the highly diverse flora and phical rainfed lowland and irrigated lowland sites in the Philipshyenvironment of upland rice isthateach upland ricearca has pines Although only one upland site was studied data a unique composition of insect pests showed equal or fewer insects at the upland site than at the

For example in Tanauan Batangas Philippines the rainfcd lowland sites and fewer than at the irrigated sites main upland rice pests are rice leaffolder(RLF) armvworm (Table 3) M Lltiple regression analysis showed that the key Aythntnaseparata WBPH ants Soli psis gentitataand factors explaining insect abundance among a set of cropping white grub Leucopholis irrorata Sixty kilometers away in intensity variables were the number of rice crops grown per Real Quezon the main pests are seedling maggot Athe- year followed by area devoted to rice Table 3 shows that rigona orae flea beetle (Thacitcmnra hasalis thrips the upland rice site had only I rice crop perym rand that rice Stenchaetothrips hiurnmis and rice bug Leptocorisa was planted in only 20Vi of the area Fertiliter and oratorius In a third Philippine upland rice site in Claveria insecticide use had little bea ring on the trend The armters at Misamis Oriental Mindanao the main pests are ants the upland site applied 60 kg N ha higher than at most Solenopsis gemtinata seedling maggot Alieriona orvzae irrigated sites white grubs Holotrichia in)iUaiuoan t and Lucopholiv Insect occurrence at the upland site was omparable to irrorata stem borers Sesamia ifiArelr (hio auriciliusC that at the Cagayan rainfed lowland site parlicularly in 1982stpessallis Scirpophaga incertulas S innoiata and when drought prevented many farmers fron planting(only

4c ona chrrsurapella root aphids Tetraneura niriah- 40(of the rice area eventually was planted that year) M ore dunrinalisand Rhupaosiphunm riahbdminalis lcaffolders favorable weather occurred in 1980 and 1981 which Marasniapataltis and Chaphalocrocis Medinalis army- supported greater rice areas and consequently more insects worm MIthinna separata and rice bug Leltucorisa The most intensive site for insect pests was South Cotabato oratoriusThe great diversity of insect problems is typical Mindanao where farmers plant 23-25 rice crops per year fer upland rice and makes control efforts more difficult with irrigation The Zaragoa Nutva Ecija site also had particularly regarding breeding for insect resistance high pest abundance and at this site the fields were highly

asynchronous Asynchrony was the third most important Low yield potential Iactor in explaining rice insect abundance Most upland rice is grown on low-nutrient and mineral- Rice cropping intensity in upland areas is bound to toxic soils which when combined with eiratic rainfall remain low Only one rainid upland crop is possible per make upland rice a highly riskycrop for the farmerto invest year (Farmers in Claveria Misamis Oriental Mindanao in costly soil amendments and land management Often Philippines tried asecond upland ricecrop i 1984a ycarof upland rice isgrown far from markets making inputs even favorable rains but no harvest was possible b-cause of more expensive Farmers growing a subsistence crop hosi- drought blast and birds) Rice area isaso bound to remain tate to invest in costly inputs to raise yields low because of the high labor and power rcquirelmenlts to

Even having varieties that could double existing yield prepare the land (because of weeds)and because upland rice potential under present management levels would probably wvill not compete with lowland rice as a cash crop Some not prompt many farmers to use purchased insecticide upland rice sites with prolonged rainy seasons howeverare unless tile risk of crop failure were reduced such as it was planted asynchronously such as the Clacra site with irrigation systems in lowland rice The seasonal abundance of Upland rice insect pests

The Tanauan Batangas site is atypical for upland rice attracted toa light trap wasgrapied for crop year 1980-81 in The soils and rainfall pattern are favorable and because of Tanauan FBatangas (Fig I 2) W13P11 was he most its nearness to Manila farmers have cash resources ftm common rice hopper and its population peaked at the sales of vegetables to purchase tertilier They applied an midgrowth stage as it does in lowland rice D-Vac suction average of 60 kg Nha after particle initiation and obtained samples showed an earlier peak than that from a light trap 25-35 tha yields The high fertility isone reason why RLF indicating emigration W13PI usually emigrate after the armyworms and WBPH developed into large numbers vegetative stage Populations in 1980 reached levels that Farmers however used no insecticide on upland rice caused patches of slight yellowing in some fields WBPH although they did on vegetables probably cause hopperburn in Batangas They immigrate

The poor growing conditions fOr rice generally mean year-round as shown by low levels in the light trap BPH poor growth and low fecundity for insect pests fIeding on it however were scarce in the light trap and not collected with

Table 3 Abundance of rice insect pests and factors in rice cropping intensity of Philippine upland rainfed lowland and irrigated lowland locations where light traps were operated daily for atleast I yr -I

0Rainfed Rainfed lowland Irrigated lowlandupland

Pangasinan Cagayan lloilo Santa Maria Nueva Ecija South Cotabato Tanauan Caaringayan Caaringayan Bangag Bangag Bangag Buray

LagunaCale

Batangas Manaoag 1980-81 Santa Motiica 1982-83 Cabanatuan Zaragoza Avancena NaniamanSolana 1981-82 1982-83 Oton

1980-81 1979-80 Oton 1981-82 1981-82 Koronadal Koronadal1980-81 1979-80 1979-80 1983 1983

Insect (kerosene light trapNephotcttixcatch nolyr)virescens 650 1240 500 1800 330 210 3400 1600 160N nigropictus 210 - 360 18400 5500 2900- 140 90 20 - shy 1700 -Reciliadorsalis 710 - 3200 1700180 890 630 140 170 2400 800Nilaparratalugens 130 690 - - 4200 4600300 520 750 180 60 3300 1600 740 310Sogatellafiircifera 1700 31000 13000 8200310 210 3300 610 270 980 1600 6300 -Cyrrorhinus - 12000 6700600 540 3200 290 160 330 120 410Scirpophaga spp 50 410 620 - - 21000 12000150 180 220 270 30 1000Cnaphalocrocismedinalis 70 70 540 610 270 11000 6100200 140 100 6 60 250Chilo suppressalis 0 60 60 1900 300 200- - 30 20 0 6 3 160 -C auricilius - 0 00 - - 20 3 60 20 3 0Sesania inferens 0 - 80 60 3 80 10 51 -Cropping intensity _ -

Rice crops (noyr) 10 10 12 10 10 10 13Rice area (7c) 17 19 20 20 25 2320 85 85 60 70 40 85 85 90Fertilizer (kg Nha) 60 20 85 80 70 7020 0 0 0 30 30Insecticide (no of 40 60 40 300 04 04 0 03 300 13 13 30 50 40 33 33applicationscrop)

__________

IRPS No 123 January 1987 21

Light trapped Insects Drytnd __ 2(notrap-nght) Suctioned Insects (nom )

50 -)o 9 jO40- SOW ffurciWO Light trp

D-Vc suction

20 to 4

20

Egg porasitizahion (M) 75 60

300 SOOampReOftwciffr

45 30shy

15

-insects (notrop-night) 4 Nileporvata Iuen5

Egg parastization()

Light trapped insects (no trop-nght) Suchoned insects (no m 2 )

6 8 Cytorh~nuls hvidipennii 0

4 05 2

50 0 Insects (noltrop-night)40-ui Ipanh pe g

Moohase 1 Seasonal abundance measured by light trap

Rainfall(etod) and D-Vac suction sampler of whitebacked

to ~ hopper Nilaparvata lugens and lea ffolder60)-t Cnaphalocrocis mnedinalis andplthpe g20 L I II I I I parasites and mtrid predator Ortorhjntus livi-Apr May Jun Jul Aug Sep Oct Nov Dec JaIn Feb Mar dipennis set against cropping pattern daily1 9 80o 19 8 1rainfall and moon phase Tanauan Batangas

1980-81

the D- Vac Cyrtorhinus build up on the crop coterminous are collected in light traps during full moon (109)with WBPH buildup Cyrtorhinus disperses during the Because upland rice shares most of the lowland rice pestsrainy season RLF not readily collected in light traps was and those pest species are generally less impotant to uplandprevalent only during the rice crop rice than lowland rice there is a general belief that insect

Nephotettix virescens was more abundant than Nnigro- pests are less important onpicLus in the light trap collections and on the crop GLH upland than lowland rice Arcview of limited data on upland rice shows an averag of population peaked at the reproductive stage as did zigzag 10-21 yield loss from insect pests on a par with that ofleafhopper (ZLH) Crtorhinus tracked the populations of lowland uee determined by the same methodology (132) planthoppers and leafhoppers Both GLH species were (Table 4) McGuire and Crandall (144) made a similarcollected year-round in the light trap indicating immigra- estimate for Central America Cramer (51) howevertion from nearby irrigated rice areas Possibly because of the estimated 35 yieldlow populations loss in South America based on ainsect number did not correlate with similar pest complex - stem borers and seed bugs shymoon phase although reports indicate that more hoppers attacking irrigated rice in North America This estimate did

22 IRPS No 123 January 1987

Light trapped insects D~~nd (no trap-night )

St plht Suctined insects (nom 2

)

D-Voc suction 1520

1o

0 Lih trap 5 ~105 o5

00 Eggparositizotion()5040 NephOtgtha virneens

30 20

I010 A-Insects (notrop -night)

4 Nephatelh nigropccent i

2 o I vV^J - -- ^ VA____ f _

Light trapped insects (no trap-night) Suctioned insects (no rnE

10 -1208 ReCdO doRsol

6 15

4 10 052 iwxALAiamp 0

Light trapped insects (notrap-night) 2Suc oned nsects (nom )

8 CyrtorhmnusI6o nn]10 i0075

4 05

2 0 0

025

2 Seasonal abundance measured by light trap Roinfall (mmd) Moon phase and D-Vac suction sampler of green leafhoppersNephotettix virescens and N ngropicrus and I zigzag leafhopper Recilia dorsalis their egg 60

parasites and micid predator Cyrtorhinus livi-dipennis set against cropping pattern dailyrainfall and moon phase Tanauan Batangas1980-81

40

5 Lf Apr May Jun

1980 Jul Aug Sp Oct Nov I

-aMar

1981

not account for soil pests and was not based on field data in the absence of pests Carbofuran granules are phytotonicAkinsola (I1) cited mainly data from irrigated rice areas in to rice (239) The prevailing hypothesis is that carbofuranAfrica but estimated yield losses of upland rice between 15 prevents soil bacteria from consuming fertilizer The phytoshyand 30 from insects However losses in Thailand over a tonic effect is through greater availability of fertilizer to the4-yr period ranged from I to 13 averaging 5(106) plant Denitrifying bacteria are known to be more active inYield loss data however should be interpreted with aerobic than anaerobic soils therefore it may not be merecaution (184) The Philippine data were all derived from coincidence that the highest recorded yield losses (22-69)trials on farmers fields under farmers normal agronomic occurred in those trials with 12-30 kg basal Nha On themanagement practices except for Bukidnon and Capiz other hand contro with carbofuran granules may not be aswhere researchers used fertilizer Batangas farmers are good as a seed treatment against seed pests and perhapsatypical because ofhigh levels of fertilizer used The trials in yields would have been higher if a systemic seed treatmentBrazil were carried out at an experiment station but insecticide had been used conditions were similar to those of local farmers The 1979 and 1980 trials in Batangas had two full-

Carbofuran granules were used in every trial as a broad- protection treatments one using carbofuran granules andspectrum insecticide applied basally with the seed An the other using bendiocarb WP as a seed treatment Plotsobjection to using insecticides to measure yield losses isthat protected with bendiocarb yielded higher than those proshyinsecticidescandirectlyorindirectlystimulateplantgrowth tected with carbofuran granules The other trials in the

Table 4 Yield losses to insect pests and yield responses to insecticide in upland rice in the Philippines Brazil and Upper Volta

Yield (tha) Yield loss (5) Location Year Cultivar Protected

with Protected

without Check Total Vegetative Reproductive Ripening

Basal N (kgha) Reference

carbofuran carbofuran Protected No with carbofuran

carbofuran

Philippines Pili Camarines Sur Tanauan Batangas

Pangantucan Bukidnon

Dumarao t-apiz

BrazilGoias Minas Ge~ais

Upper Volta6

1975 1974 1974 1975 1975 1976 1977 1978

1979

1980

1979 1980 1979

1977 1978 1979

Bursiging puti Kinanda Dagge Kinanda Dagge Dagge Dagge Dagge UPL Ri5 Dagge UPL Ri5 Dagge UPL RiS UPL Ri5 UPL Ri5 UPL RiS

IAC47 IAC47 IAC47

14 33 11 36 35 31 30 27 37 26 40 30 40 42 46 35

19 21 09

--

-34 24 24 27 --

29 43 31 42 -

47 25

---

10 28 05 27 27 30 26 29 33 28 33 29 43 23 36 11

15 15 06

29 15 55 25 23

3 13 -7 11

-7 18

3 -7 45 22 69

24 28 35

--

21 -11 -20

4 -

-3

23 7

-2 -23 56

-

-

-__0

6 --

-

-23 14

5 0 8 2

22

-

--

0

-

--18 11

3 -10

4 10

7

--7 -

14 -

-1 6

10 -5

8 9 7

-

--

0 0 0 0 0 0 0 0 0 0 0

30 30 30

12 12 12

99 99

100 100 101 101 102 102 102 102 103 103 104 104 104

66 66 66

Farako-Ba

Mean

1977 1978

IRAT10 IRATIO

38 66

--

40 43

-5 35

21

-

10

--

--

--

- 242 243

Z

tJ

-w

24 IRPS No 123 January 1987

Philippines which had two treatments offering protection In the 9 Philippine trials where yield losses were measuredwith and without carbofuran granules relied on further by growth stage 10 of the yield loss (21) occurred at thecontrol with foliar spray and not on seed dressings The vegetative stage and 517 occurred at each of the reproshycarbofuran-protected plots had an 18yield los compared ductive and ripening stages Ibis result is similar to thewith only 10 with foliar sprayS This yield difference is pattern of yield loss in lowland rice measured by successiveprobably due to differences in protection against sown-seed treatments where the crop is unprotected at each growthpests (foliar spray do not control sown-seed pests) rather stage but proticted during the other growth stagesthan phytotonic effects became basal fertilizer wa used in However the insect complexes r sponsible for vegetativeonly 2 of the 10 comparisons stage yield losses in upland rice are entirely diftfrent than in

The recorded yield losses were not related to yield lowland rice The key vegetative stage pests in lowland ricepotential (Fig 3) One might expect higheryielding crops to arc aquatic whereas those in upland rice are sown--seedhave higher percentage of yield losses but this ws not tile (sown in soil) root and seedling pests This dift rence maycase One reason is that insect damage exacerbates plant explain why agricul torists have overlooked tile ilmportanccinjury from drought stress low yielding fields from d ought of upland rice insect pests Many reports of upland rice pestsstress would show disproportionate differences between in Asia ha Ve focused on major lowland species (46 108)insect-protected and -unprotected treatments Shoot and It isalso surprising that tei vegctative stage of upland rice root pests would be more responsible for losses from would record the highest yield losses becatuseseeral studiesdrought stress than sown-scd pests A second reason have shown that upland rice can readily recover from foliageconcerns seed pests which would have a greater impact on reinoval during that stage (117 152 165 188) Upland ricelow tillering low yielding varieties such as )aggc Low poscsses a hitgh obilitV to compensate for ca rly loss oftillering varieties nnot fill in the space created by removal foliage and will even be stinulated by foliage removal toof seeds oweer 1) R i 5 is high yielding because it tillersP1 produce a higher viel than plants ithout any foliageactively which can close the canopy to provide greater weed removed (63 192 231) [ie reason for this apparentcontrol discrepancy hovce is that yield loss at the vegetative

stage does not octUr frol leat arca loss but fron pestsYield loss () removing sown seed and feeding on devcloping tillers and roots -- damage that cannot be readily conpLtnsatCd for

70 - Measuring yield losses in upland rice cnvironrnents is further complicated by a report from Brazil that greater yield losses occurred in response to better insect control60 - =-09 as Insecticide seed treatments resuled in denser plant stands

(n=21) which in turn created a selectively more favorable environshyment for blast (69) This resuIt was parlticurla rlxexacerbated50 when cirbofuran granules were used in seed furrows Aside

ffrom a greater plant stand ca rbofuran iay have auigmented the N lecl and stimulated blast If so ca rbofuran should be40 - replaced with another insecticide or the seeding or fertilizerI rates decreased

Other methods to determine the impact of upland rice30 shy pests wvcrc to interview farners or to develop single speciesbull bull correlations of population levels and yield loss Feneira (66)2 NIM)st (75(1) Golas farmers reporteddid both in Brazil

20- insect losses in ufpland rice and 251 said those lossesS avcragcd 37(1 [his result is consistent with tie yield losses 0 0 measured by field trials in the same statc(Table4) For every

I0 4 of seedlings removC d by leaf-cutting ants a I 7yield loss is predicted Similarly Ferreira (66) found that ai

0 - average of 53 thrips per panicle before panicle initiation meant 3 times more unfillh d grains

-IO0 I 1 1 I I-101 0 2 3 4 5 6 7

Yield (tho) C)N [ROt IM FI IODS 3 Correlation between upland rice yield and yield lols caused by Upland rice insect pests may be controlled by culturalinsects measured in 21 trials in the Philippines Ivory Coast andBrazil genetic biological and chemical methods

IRPS No 123 January 1987 25

Cultural control dry season plowing will have little effect on them Plowing Crop husbandry to reduce upland insect pest populations when the soil is wet from the first monsoon rains will expose can be classified into practices effective in a single field and soil insects that tunnel close to the soil surface to predatory those effective only at the community level Cultural birds chickens dogs and even man (field crickets nole controls should come first in a pest management program crickets and white grubs a re delicacies in man tra ditional They have broad stable effects as pests have little pos- diets) sibility of overcoming them through selection of biotyps The greatest degree i1 inIscct control fron tillage comes (138) On the other hand some cultu ral controls work only from plowing soon after rice ha rvest when most soil insects with high labor or power inputs and connunity-wilt lie close to the soil surface anrd stern borers are in the rice methods require coordination of iany farniters stuble Whitegrubsare mature and tillage exposesthem to

Pantin time The luxury of being able to shift the time of predators Tirnelv control will prtec the crop following pla ntring upland rice is open onl hr la rniers in areas with a rice and lowr the populationIfor succeedinig years Plowingprolonged rainifall pa ttern lI copping intensity or underthe ice stulble helps it decompose killing sten borer rnechanied land preparatiot Gcnerally planlting as soon larvae inside as the rFiinv season begins will lower populatiois of 1110st [requent tillage destroys ait nests bit not termites insect pests Insect populationsarelo aftcra dlvseoll or Tillage and clean culture howes er rcliove food for winter fallow delayed plaiting lets them build tip on termites which then mtay attack a young rice crop Scarab alIternate hosts that gi vigorotsly with the first seasonial beetles also prefer to miposit in eceitly tilled fields Noshyratiris This first flush iof weds cros lItxuttriartlv with iceCult is dibbling faors the survival oftillace Clre Such rineralied N released during the dr season aid made stils i aels tnd insects thtt pttpate or lay eggs- in the soil

available to the plants with the rains It iliso igrtmitcally Zero tilla hio eerl conserves predators ofthe GIH (37) advanttgeous for the rice ciop to tap this trlLItlaI fertility nt it mcttoi lo lessen the impact o0 soil insects and grow vigorotsly to compete with wrus and to tolerate brotdcasting seed or planting in ftltrrows is prefc ired over pest infestation dihlirig in hills Crickets are large enotlgh to eat seedlings in

Larly plantngs terd to escape seedling maggolts Upland a hill atrd tillering normnally ca iriot fill in tlhe missing space rice in Bitangas is planted in Ma and ine itli the Iirst Oftcn dibbll rd is not well covered and iseasily found by rains Fairners are highly mtotiated to plant early tocscape foraging insets ioIdents and birds drlitght and typlotns ricar harct Arn Atrgtst planting Plan uit sii Incrcatsini sced density protects tile cropwould be evcely attacked by It riim orivzo At tti againrst seed and seelling pests especially ants particularly other Philippiie tipland sites Rea l Qteot antd (laelit if the crop will germinate and ermerge quickly M isainis Oriental seedlig mnaggots arnrige ric boe t se hr trftinll itI India rice intcrclopped witi cotton or farnes have a longer growing season and plnit 2 rin or pigeon pci had lower 6IIt and WBIlII poptrlations than mote after the ortset of the rainy season liaaiangas ftrmners rice alone (211) NfailC arid Urplald ir_ 2re coilltori in the plant ca rly becia use ofa shorter raiinfall period arnd the desi re Philippines he chatngC in pest status folii iniercropping is to lliarvet before the typhoor sea son peaks ltighly locatioll-specific (133) cmianarid the net elfc be either

Early plantings cornbind with early-iatiuring varictics nil bcnelficial ortdetriential (189) For example BIatangias provide higher crp tolerartce for white grubs Aestiatior farmers intercrip Itro in their ipland ricefields A liornshyterminates with the first rains but 5-6 moare ineeded for the nrii iotd-Ahis i (I iiiacius) which norlially feeds resting larvae to develop into beetles emerge and lay eggs on taro ill feed or miarbv rice plants Intercropping and for larvae to reach the danriigii g ird-instar stage probably would not afft0 seed pests A crluanion crop (134 In lataigas this normually iseiough tiite for harvest planted ith rice and haing gleater tolerarnce for root

It Brazil howecver delayed planting is a suggested dariage or being toxic nayact isi trapcrop to root-feeditig cultural control for the lesser cornstalk borer and insect pests Acial-fei i ig pests voultId be most affected by pests favored l drought (66 195) Cropping internsity in intrer ropping Shoot aphids leaf beetles sten borers and Brail is low and delayed plraing places the rice crop in the hopplers would be likely ca ntidates to note effcts On the niost stable rainfall period reducing likelihood old rought othlt halnd the corlipalior crop stch~ sitniaie Iiiv be more

In rilniy arca s rinmirsare roi atl to plant other crops benehited than rice from initecropping ( 137) before rice and althNgh it woultl be desirabh to avoid 11echi Weeuling luring the first month after clop infestations by planting rice cuatl lariners lack the labor cstablishent will force pcts such is anyvworris which and time to do so ol a low priority crp prefer grassy cetds ontor tle vorling rice Soil iampsecs will

77age Uplild rice Stils ae light textured and Call be mvnye to the rice crop Cleani citrltirc also lay lore territes plowed duiring the d rv seasor notl only to grin time for to attack I u cetincti inrice ceds i ay he le optinal planting with lie fitst Inilsoon riiains but also to desiccate solition 13) soil insects arid weeds such is ntsedge that foim rhizomes Irtlizr Itneatsing N Iciiliier oin hlwland rice has Htowever becta use most soil insects lie below die plow layer fAtwvored Irigher populains of pliithoppeiv Imlhoppers

26 IRPS No 123 January 1987

and leaffolde rs However because of the need to temper fertilizer use on upland rice o prevent blast fIrtilizer rates will be lower than is comonia r for lowlad rice causing relatixely less hopper and IaIfTfoide r population buildup However inupland rice areas without a history of blast such Is Batangas farmers apply high rates of N and conscqtrently the crop has high infcstation levels of WBPI]and RIF The fhrmrcrs leec insect buildup by splitting the fertili11r in0to two to three applicatiois bit do not applV fertilizer at the vegetati estage because this prodrces a tall crop which illeasily lodge during typhoons Greater crop fertility leads to greater insect survival (better nutrition) larger insects (to cat no re foliage) arid greater fec titd t (to lav nmre eggs) (128 145 171) Not all insect species respondequally hovr Increasing rates (if N l anrid K increased Dialraa s-acchralis and (iaeiurema flea beetle but reduced thrips (66) Zn rcdLuced deadheart density from the lesser cornIstalk borer

Fnldingi Although most Upland rice areas do nIot hawxcaccess to wa ter floodiig the fields is rRrrnriierdcd in areas of Brazil or root bug and white gib control ( 195)

Crop rotatioti Tillage before planting a crop after rc harvest will control stern boreis and other pests renain n the stubble arid will unearth soil pests Planting a non-gratriiaceous crop after rice is recommncrded for termite control (66) Crop rotation is less nttdcd in upland rice culture than in lowlands because atcrop-free period is assured even after harvest of crops following rice

t1ulchilt iryl plastic irli irig designed for weedcoitrol ii upland rice inJapan resulted in ntr highanri liy infestation of (ihilo sulytresvais (93 9-1 95) NIulched Upland rice grew more luxuriantlv and attracted more (Avipositing mothis than u lched rice Also the rice plant and stein borers imatoured 10rC quickly and a partialgeneratiori af vouing ista iNoverwintcred in the stubbleLotili--malitrt varieti Quickly maturing crops reduce the tmbCr of pCst generations that Calt build up Rapid crop establishment olan ca rl v-natu ring vaitvty if)pa 11icLa r will lessen white gib and stem borer damiage White grub larvae intle last iistar are highly destIruCtive so aii early-iiatri ng variety will escape serious root loss Stem borers build 1ip slowlyx in rice to ecormic abundant in a late-Maturing crop anrd caiuse whi tlieads Numibers of somespecies increase exponeitially with each succeeding genera-ti so a xarietyexhat atures I trowh earier thian another will hac less insect damage Early-naturing vaiicties however will not lessen seed pest daiage either ii the soil or oina standing crop

Strwhoi hAIJttam Inrscts tcd irig on thlie aerial portions of rice plats disperse from carly to late plantedfields AS iiiibCirs increase exponentially with eachinsect

gtscnrattm e eiglbrig f rmc~s1(uil upp wh rc generation pests build tIP where neighboring fihrTC e9r1sstagger their plantings II Batangas farmers plant uplandrice within I iro use 120-d s-arieties and plant at the start of the rainy season Ini Misarnis Oriental laricrs stagger their

plantings up to 3 mo apart and use a set of atieties that mature in 4-7 mo Insect pest danage particularly from stem borers is greater in Misarmis Oriental Also seedling maggots damage rice there because rainfall favors frequent planting of maic In Batangas rice escapes seedling maggots bCca usC of distinct drs season and synchron0us early planting with the onset of ains

Therefore combinations of cultural practices ea rlyplanting synchronous planing crop rotation and earlyshymaturing vaneties protect the rice crop against most insect pests

Ilmant resistance Beca tse upland rice has few specialized insect pests and many insect types attack the crop worldwide regional breeding and strong internatiornal cooperation are required

Sown-sced pests arc economically controlled by insectishycide seed treatment and should the relore receive lowxpriori ty inbreeding objectives

Root pests are normally cost ly to control with insecticides but finding resista t sonres mayvbe di fficult Tanaka (229) for exa -ple fi led to find varieties resistant to root aphids Selecting for large root bioniass perhaps should be the strategy for pests sLuch as Whlite grubs that renliove roots Fainter(174) howcvcr cites varietiesof crops tolerant ofor resistant t(i root pests Some sorghun varieties are high in cyanide so rice sorghun might incorporate broadshyspcctruri chemical resistance

Seedling vigor and drought tolerance are high prioritybreeding objectives so tupland rices should have large r root systernsand be able to tolerate ligherlevels of root loss Sonie root loss inolder plants is beneficial if it stimulates new root developnrut and therefore crihances intake of soil nutrients (53) The rice seedling inaggot rtiLotia spp may be

controlled thirough resistant aittics A hi rgc breeding effort on sorghniu has had fair success inideveloping arreties esrstait to Asoccata Rodani (251) A field trial comparing 10 upland ricc varieties showed differences aniong varieties va ring from II to 4(- damaged tilleis (215) Hovver resistarce to 1Iill-0r gtia species may rot cross ox-rti otier

Shiraki (210) reported that upland i-ice stens wcrc harder than lows-land rice stemsperhaps becaIusc (f a higher silicoricontent and thus were more resistant to yellow stern borer hc tortia Iow tillering (f uplaid iice xarneties prodtres a

higher percentage of infested tillers than would occor in higher tillering lowland iice under a siiilar egg density The corollary therefore is to select for high tilleriig upland rices

Stemii bor amcstheh main targes dobrecders inAfrica (571 204 )but nticl oifth e sc reen irig is dotie unde r lo wland co d t osrm s wih A h r~ o l pe i s n acorditions (11 96 218) As with lthrinrnwspecies mayhave to be dealt with independently however initial results show cross resistance between African and Asian species(92)

In Brazil screening Ior insect resistance focttses oil lou I pest problems Varieties tolerant of lasmolwalpus ligi-sellus and resistant to )iatraea saccharalis have been identified (70 153) Current work also emphasies 7iraca limativetntris and Thmialspis (=Iosis) flavouicta (66) These insect pests Ilt onlV citise IIigh yield losses tihey also ire difficult to control chemically ofr by other ilealls

ManV upland rice vaieties aic early maturing and are resistant to pests such as gall midge (I Xl ) Early mIturity is often a highly desirable trait to esuilpe pest buildup but it is not true resista nce bca usc ilthose va reties are plinted late they will be itsdamaged as ain early planted susc ible variety

P~erhaps tile cosiiiopolitall of all upland rice pestsimost are the sced pests (ripenling stagc) which pose a particular problem beta use scuds re ised IsIO for hitnit4ans Ayltoxic substatice introd tcld geticticllyilmay also be toxic to nal iniless it caln he placed old in thfe IdhuffSelectmi fo hairiness or lotng wns kill mIlct sthilrohleiiis dutilig threshing pa rtictfarl maitial threshinfl [hireshers ill complain of itcfimin atid itteascd (fust levels

Attempts itIRRI to fitid tcistatncc to the rice bug plorisa have ilotbeCnl prmitisilg lIIetIattonlids iorc

through the lenilna or palca hillhairs sedstVIor seeds wilh awns that deter bugs promote complaints from threshers Seed bugs liptocorisa and Stcnocoris enter the seed through the opening between tile lemna and palC but varieties with narrow openings are diifficult to mill

Breeding priorities should concentrate ott inotCI gruLtLIps that are most difficult to c nttrol by other meains Steil) horers appear to be the most widespread group that sigiiificantly dalag s rice aind is hard to manage by other Ilnca us

Biological control Very fe stutdic of mitatt~l eneties of rice pests in upland envirollnletits wee in tile wefouiid literature Therefore have not compiled a list of species Ve present a list of natural enemies iOtid in upland ricefields tuic latialln Batangas for some conitno iOldiar pest species ([itble 5)

Muich ofl thc lollowinle itfor ittiol cotlliCs rol Stutdies in the lowlatnds or roi crops otithcr thlin tice

Qu(ailtr o 11ll t11t( Nat trrtf lii hcoeits ight Ile used agaitist ile social insects tilt and termites Sollie predator specice live ill theirf nests n ilnic theirnllt ip-pearanIlCe aMRidbehavior IJtrouMdCing a mote aggressise ant species to displace pest ant codonies is a possibility (169) Patihogens as potential natural encnies have reccived little atteition (I13) Natural ncnuics asa group however hac not been toIl iporta it in Iegulating aintshown be and ternite numbers

The nocturnal behavior aid subteitmien habitat of lole crickets and field crickets protect them from many ntllral enemies There are only limited records of natural cnelics for these groups of pests some pathogens (87) but mainly

IRPS No 123 JattuIny I 271987

parasites - scelionids on eggsand sphecids on ilymphs and adults Sphecid wasps Larra carutmaria (SImitI hI L huzwuensis Rohwer and lsaltiuhta Willi ams speciali in mole crickets pa ralyzing them nd dragging the ninto thcir nests as food lot their voting ()tier sphCcid species Lins aurulenoa (Iabricius) lot maiac (Ashmead ) 1 lhshylssellaus (Smith) and Ml lal)oriosu (Snithf specialie in field clickets

White grUbsO the other hand hae Icti scriuns enuigh to have been extensively studied for cotttroi 1W nati al enemies( 112 140 141) Scoliid parasites C(ajsoMcispp have beetn intrtduced itto tilePhilippites for control of Ielxl ]udisiro la The rcsearchcts w Intttrdced themle during ill era claimed that the white gttionthreak as controlled (142) howevcr it isnormal for pest epidemics to subside and there was no diect evidence to indictte that tfe parasite was responsible A tmore promisitg titctllod might he to identify iso ltcttass prt~f tcartd apply hiactelaln fingal pathogens to tile soil Ihe lalattese beetle in North America has hcci coitrolled hv tite conmnleciall aailable hactcrial prcpa ration itacihlpu pav ofr milksdiscas (71) Spote prtyefott the soil itloted ilrtier reitiain actic fr Veas

Root aphid sardtnctly bugs readilslic tilib to predatots and parlsites once their tetndine ants are controlled The larvac and adtilts of a coccinellid beetle 1Scynms sp prey on llriadlouais in slash and burn upland rice Baits treated with insecticide may be used to control tending ants by killing their ourig (244)

Root-tkcditig blugs false wireoris wireorms and root wccils aic r ofsuch little ifilpoltatice that thenomttll role of liatural Cticniics hias tiot beei asscssed A possibility hoevrl tiglt he tit ItaIIitic niaCIMtodcs lintt1g11en Itese soil-itlhlfbitine mCtitodcs W5uhtfhave to be imiass Produiced

large miltibcr of niatltrll cnemics ircknown ft CltIoriS IC IpblCeii is thtt clttvortnlls ofonlie the crop soott alter land ptel)tritiOti a1tithe bcgit iniig of the raittv

Citsoln hfien tiatllral leniiiy populations aric lo Colltloil WOtuld iii ole mass prodtticing andirclcaslng key spccies

le egg stage is the fpirt (olthe Cedlitgt Ialot life cycle tI hich to f)ctis ftor tittlril elctit cottntrol hplraites o1 pleditors ttsit elleggs highly expoIsYCl I Sgg pailsites or predators stolld probahly e ittass andtbe produced released Littlh i toruialioll is a ailable ott what species to trV o ho to Iniss podLice thiCIi A etiloplhid wasp Icmihlaol sp effctiicly regushy

lates the leatf lcloalapolm2a spioanoiiitr heat andc rice antid should be collsctvcd

leaf beetles whCllose e am] eggs ate laid in the grutindhirO ould be motC difficutlt to control by niatulral enelV Mnll ipuhltiOn But species sUici IS c_gsthispid beetles ith

and larVle oilleascs ate vtilcratble to iparaites and pledators

Natural enemisc of tlirips otl rice are little known

28 IRPS No 123 January 1987

Table S Parasites recorded from upland rice insect pests in Tanauan Batangas Philippines 1977-84

Order Family Species Host stage

I lymttenoptera Braconidae Leaffolders Cnaphalocrocis Marasinia spy

Cardiochilesphilipplinensis Ashmead a Larva

Cotesia angustillasis(Gahan)Cotesia or cypris (Nixon)

Larva Larva

Cotesia or taeniaticornis(Nixon) Larva Cilonusinotakatae Munakata Chelonits SppAlacrocetrts philippinensis (Ashtmead)

Egg to larva Egg to larva Larva

Iclineumonidac Orgilts spp Isclutojppa hireator(Fabricius)

Larva Larva

Itoplectis naraigav(Ashmcad) Larva Tenelhtcha philippiensis (Ashmead) Larva Tenelchha stangli (Ashmead) a

irichomma cnaphalocrosis Uchida Larva Larva

Bethylidac Elasmidac

Goniozms or trianguli r Kieffer lasmtusalhopictus Crawford

Larva Larva

Diptera

Encyrtidac Chalcididac

Tachinidac

Elasmtus spp Copidosomopsisnacohiae (Fidy) Brach nteria evcarinata Gahan Biachyieria lasus (Walker) Ztgohothria ciliata (Wulp)

Larva Egg to larva Pupa Pupa Larva

Argyrophylx nigrotihialis(Blaranov) Larva

l-ymenoptera Braconidac Yellow stern borer Scirpopiaga incertulas

Braco chinensisSzeplegeti Larva CIthelotis spp Chelonus niunakatae Munakata Cotesi flaripes Cameron Stenoracon nicerilli (Bingham) a

Larva Egg to larva Larva Larva

Ichneumonidac Tropobracon schoenobii (Viereck) Amatroinorpiaaccepta metathoracica(Ashmiead)Frihorus sinicus (I lolmgren)

Larva Larva Larva

Ischnojoppaluteator (Fabricius) Larva Isotima nr dammeratiti Rohwer Larva Tenelucha philippinensis (Ashmead) a

Larva Tenehicha stangli (Ashmcad)Trichomma enaphalocrosis Uchida

Larva Lara

Chalcididac Eulophidac Trichogranmmatidae

Xanthopimnpla stemmator (Thunberg) Charops hrachYpterumn Gupta and Malteswary Brachtm eria sppTetrasticttus schoenohii Ferriere Trichorigranm chihunis Ishi

Larva to pupa Larva Pupa Egg Egg

Diptera

Scelionidac Pteromalidae Tachinidae

Tt ichograintnta aponiicuitt (Ashniead) Telenomits rowani (Gahan) a

Trichoinalopsis apattelctena (Crawford) Jeirhaea spp

Egg Fgg Egg pupa Larva

Z4gobothria ciliata (Wulp) Larva

Ilymenoptera lecneumonirlac Gol fringed borer (hilo auricilius

Trichomm enoptalocrosisUchida Larva

Ieioenoptera Braconidae

Aantholimlh steiniator (Thtinberg) a

Pink stet Ioret Sesamia inferens StoInacoil nicrilhi (Bingh am)

Loarva to pupa

larva

Diptera Ilyinenoptera

Armvtnyorms Mythimna separata Spodoptera mauritia acronyetoidesTachinidae Zvgohothriaatropiora( Robinea i -Desvoid y)Eulophidae It-phctrtiscitapadoe (Ashimead)

Larva Larva

(halcididae Braconidac

Brachpin ria spip (oesia spp

Larva larva

IHyntenoptera I)iptera

Rice skipper Ielop ida (halcididae Tachinidae

math ias aidgreen iortedci ciliarNIelan itis eda Brat lhincria sp iii matuqitatt Cameron Irgyrolhirlaxi trotlhiais(marumtv)a1

ismtne Larva Larva

Diptera Ilymenoptera

Tachnidae lelineutionidae

fuown se tilo per Mois frugalisIrgtrophrlax 1igrothiahiv 3 a rat ov) anthopimpa pmnctata (Iabricius)

Larva Larva to pupa

aMost dominant

IRPS No 123 January 1987 29

Normally predatory thrips prey on herbivorous thrips strepsiptera on both Spiders dominate tilenymphaladultNatural enemies of stem bugs --Scolinoplara and Tibraa predators Spittle bugs move to rice from pasture grasses - are in order of importance egg par~isites and predators and biocontrol would be successful only ifcarried out before fungal pathogens and nymphal adult parasites Egg tiley immigrate into ricefields parasites could either be introduced fron other areas or Biological control tactics for hemipterous seed bugs mass produced and released ould follow those outlined for stein bugs

Armyworms are normally lwld in hcck by the activities Coml)arison of environnewtns Natural enemies are of egg and larval parasites When these natural enemies fail perhaps faced with even greaterconstraints than insect pestsusually because of drought armyworns become epidemic to survive ind reproduce in upland environments TheyIt may be worthwhile therefore to release parasites during have adapted ani survived by having wide host ranges the the rainy months following a drought Virus diseases of ability to aestivate or hibernate over unfavorable seasonslarvae would beanotheravnue to exploreforarnyworms and the ability to disperse or by other mechanisms

Parasites have ben traditionallyconsidered for biolegical Comparing the most dominant and therefore the most control attempts against stein borers usually by rekasing adapted species of natural enemies collected in various exotic species( 166) This approach has not met with success environments in the Philippines we can understand how and new avenues should be explored The role of predators different rice environments are arid pathogens is little inderstood and deserves greater Anagqrms olptahiisa mymarid planthopper egg parasiteattention Egg parasitism rates are normally reasonably is cquallyadapted to all Philippine rice environments (Tablehigh but should be supplemented with effective egg 6) It is selective for planthoppers and even parasitizes eggspredators Orthopterans are potential egg predators of the n-iize planthopper Peregrinitsmaidis itt upland areas Metioche vittaticollis (Sta ) Anavipha nifiwnnis (Saus- Alavohis also parasitizes maize planthopper eggs but is sure) and Conocephahs hongipnni (de i-laan) feed on most prevalent on BPH in the irrigated wetlands (onashystem borer eggs etioche and Anaxil)ha specialize in eggs tocerus spp like A optalhilis on BPH and WBPI Iattacks with no hair such as Chilo spp The omnivorous (ono- the eggs of GLH in ell rie environments It is also found cephahsspecializes in eggs cov red with hairs it is a proven parasitizing the eggs ef the white rice licahopper (oimna egg predator of Scirlouphaga spp but has the discouraging spetra Oirosita naias a trichogrammatid egg parasitehabit of eating rice guin (65) attacks BIPH and WBPH in the wetlands and has no other

Grasshoppers have egg (scelionid) and nymphalad ult host A related species 0 aesopi specializes in G1-H in (nemestrinid tachinid and sarcophagid) v irasites Among wetland environments Dryinid strepsipt-an and pipunshypathogens protozoans have been recorded most frequently culid nymphal and adult parasites of planthoppers and Control of locust species by natural enemies could con- leafhoppers also specialize either in hopper species or centrate on their habitual breeding areas environments Among the dryinids Pseuhdgonatoipsflavi-

Leaffolders have rich complexes of natural enemies some feimur prefers the lowlands and BPH while P nuch has no adapted to upland rice (22 27) ranging from egg predators environmental prefirences but is adapted only to WBPH (gryllids coccinellids) to larval parasites (braconids ichncu- The main d ryinid species on GLH aptogonatop)usspp is monids) to larval prcdators (ants and carabids) to larval most dominant in rainfed lowlands and also attacks other pathogeiis (vira and fungal diseases) Efforts to augment leafhoppers within Amnrasca Cicadulinaand Bachaltuathese natural enemies should focus on the egg predators Strepsiptera are more dominant on plarithoppers than (rearing releasing)and viral or fungal pathogens (culturing lealhoppers The Elenchus species in the Philippines attack spraying) all species of planthoppers within the genera Sogawilla

Parasitization of the large conspicuous larvae of Sogatodels NilaparsataHarualia and Opiconsiva Elenshyskippers green horned caterpillars and other polyphagous cusrvasuntaisuiis adapted to the major rice environments Lepidoptera by tachinids and chalcids is normally low ELg but prefers BPH in rainfed arid WBPIH in wetland environshyparasites and predators are perhaps the key to their control ments Ilalictoph(utsmtnroeiand HVpectrus attack G LH

Gall midges are parasitized by pterormalids and platy- and COfimravcctra respectivelygasterids that aestivate within their larval host between Pipunculids are mainly parasitic oii lealhoppers Each seasons Shoot aphids are preyed upon by coccinellids species has a unique environmental preference Tomosshysyrphids and chrysopids varireIa oraetoraprefers Ne7 otetltix nigropictrs and

Planthoppers arid leafhoppers have egg parasites and Pilunwulhtsjavanensisattacks Dehoceplhahls spp which predators as well as nymphialad nlt parasitesand predators are more abundant in upland than irrigated lowland their rich complex of natural enemies includes fungal environments T suhvirescensattacks Nephotettix virescens pathogens The principal egg parasites are myrnarids and and prefers the lowlands Pilpuiulus nttillattls is an trichogramniatids Egg predators are norinally mirid plant upland rice parasite arid attacks only Nephotetix speciesbugs that suck out the yolk Nymphal ad nIt parasites are More than 17 recorded species of leaffolder parasites in dryinids on planthoppers pipunculids on lealhoppers and the Philippines attack species of C(naplwocrocis and

30 IRPS No 123 January 1987

Table 6 Dominant natural enemies of rice pests in upland rainfed lowland and irrigated lowland rice environments of the Philippines

Natural enemy

Host stage Species

Brown planthop-Parasite Egg

MymaridaeAnagrus optabilis(Perkins) Anagrusflaveolus Waterhouse

TrichogrammatidacOligosita naiasGirault

NymphAdult Dryinidac

Pseudogotatopuslaiifemttr Esaki et Hashimoto Elenchidae

Elenchus yasurnatsui Kifune and Hirashima

Environment

Upand Rainfed lowland Irrigated lowland

Ailaparata lugens

x x X x

x

x x

x X X Whitebacked planthoppers Sogatella furcifera and Sogatodes pusanusEgg

MymaridaeAnagrus optabilis(Perkins)

TrichogrammatidacOl~gositanaias Girault

NymphAdult Dryinidae

Pseudogonatopusnudus Perkins Elenchidae

Elenchus yasutatsui Kifune et Hirashima

Parasite Egg

MymaridaeGonatocerus spp

TrichogrammatidaeOligositaaesopi Girault

NymphAdult Pipunculidac

Tomosvaryella subviresccns Loew Tomosvaryella oryzaetora (Koizumi) Pipunclusmutillatus (de Meijere)

DryinidaeIlaplogonatopusspp

Egg Copidosomopsisnacoleiae (Eady)

Larva BraLonidae

Cardiochiles philipphnensisAshnead lchneumonidac

Trichiomnia cnaphalocrosisUchida Temelucha stangli (Ashmead)

Pupa Chalcididae

BrachymeriaexcarinataGahan Brachymerialasus (Walker) Brachymeria nr tachardiaegroup

Parasite Egg

Scelionidae Telenomus rowani (Gahan)

EulophidacTettasticusschoenobhi Ferriere

Larva Braconidae

Stenobraconnicevillei (Bingham) Iclhncurnonidac

Temelucha phiiippinensis(Ashinead)Temelucha stangli (Ashmead)

x x x

x x

X x X

x x Green leafhopper Nephotettix virescens

X x x

x x

X X X X

X

X Rice leaffolders CnaphalocrocisMarasmia

x X x

X

X X

x x x x x X x x x

Yellow stem borer Scirpophaga incertulas

X X x

x

x

x X XAinauromorphaaccepta metathoracica(Ashmead) x

continued on next page

IRPS No 123 January 1987 31

Table 6 cntinued

Natural enemy Environment

Host stage Species Upland Rainfed lowland Irrigated lowland

Larva-pupa Gold fringed borer Chilo attriciiuslchneumonidae

Xanthopimpla stemmator (Thunberg) x aPupa x

Pteromalidae Trichoinalopsis apanteloctena (Crawford) ax

Pink stem borerb Sesamnia inferensParasite

Larva Braconidae

Stenobracon nicevillei (Bingham) x b b Armyworms Mvthinna SpodopteraLarva

Tachinidac Zygobothriaarroptvora (Robineau-Desvoidy) x x

Braconidac Cotesia spp

X Skipper Pelopidas mathias and green horned caterpillar Melanitis leda ismeneF-gg

TrichogramrnatidacTrichogrannaspp x

Scelionidae Undetermined Scelionid

x Larva

Tachinidae Argyroph lax nitrotibialis (Baranov) X x

Pteromalidae Trichomnalopsis apanteloctena (Crawford)

X Predator

CoenagrionidaeAgriocnenis femina femina Brauer x xIschnura senegalensis tambur

Miridae CrtorhinsliidipennisReuter x x

Nabidac Stcnonabis tagalicus (Stal) x

VeliidaeAlicrovelia douglasi atrolineata (Bergroth) x

Coccinellidae Micraspis crocea (Mulsant) X x x

Vespidae Eumenes campaniformis (Fabricius) x

Formicidac Diaecamna spp xOdontoponera transversa (F Smith) x

Carabidae Chaenius spp OphioneaW- aJasciataSchmidt-Goebel

x

Ophionea ishii ishii Habu x x X

Gryllidav Anaxipta longipennis (Saussur-) x xMetioche ittaticollis (Sial) x x x

Tettigoniidae Conocephalus longipennis (de Haan) x x x

aNo rearing made on rainfed lowland bNo rearing made on rainfed lowland or Irrigated lowland

MrasmiaAmong the Laval parasites the braconid Cardio- lowlands and parasitizes oriental maize borer OsIriniachilesphilijpphensis is most adapted to an upland environ- furnacalis (Guenee) larvae The leaffolder pupal parasitesment It also parasitizes Hydelepta indicata a common not only occur equally in all environments but also haveleaffolder of legumes The ichneumonids Trichomma cna- wvide host ranges- parasitizing species of OsIrinia Marucaphalocrosis and Temehcha stangli occur more in the Hydelepta and Honiona - in maize and legumeslowlands T stangli is most adapted to irrigated lowlands The principal egg parasite of yeltow stem borer (YSB)and also parasitizes Chilo and Scirpophaga stem borer Telenomus rowani is widely adapted to all environmentslarvae T cnaphalocrosis is most prevalent in rainfed and has an unusual alternate host -Tabanus eggs (25)

32 IRPS No 123 January 1987

Tetrastictu schoenobii specializes only in YSB and occurs mainly in irrigated lowlands -- further proof that YSB is most adapted to the wetter environments

Larval parasites of YSB are highly environment-specific The braconid Stenohracon nicevillei occurs only in upland rice but also attacks the pink stem borer (PSB) Tieinchwa phiiiplPwnsis an ichneumonid occurs in upland rice and also parasitizes the larvae of (7hiu stqpresaisinatro-morpha accepta melaihoracica like 7 stangi is mnost adapted to the irrigated lowlands

The dark-headed stem borer(l)HSB) (ih pochit3sus was not collected in all rice environments and the gold fringed borer Chilo auricilius nd PSB were collected only in upland rice Xaantiholiplastenrnaloran ichneumonid larval parasite appears to be widely adapted whereas TrihomaslopisapanitI)cteiiaa pteromalid pupal parasite occurs mostly in wetlands and also parasitizes larvae of Chilo suppressais and Pelhuidas mathias

Armyworms Myit hinin and Spodujipera have two main larval parasites - a tachinid Zrgohothriaatropivora in rainfed environments and a braconid Cotesia spp in irrigated lowlands

The butterfly pests Pelopidas and 4elanitishave tricho-grammatid egg parasites found only in irrigated lowland environments

Two larval parasites show environmental preferences --Argrroph av nigrotihialis a tachinid in rainfed habitats and Trichonialopsisapantelkcnela in irrigated ricefields A limgrotibialisalsoattacks the sweet potato hornwormAgrius convolvtdi and T apanteloctena parasitizes DH S1B

Predators as a group show more distinct environmental preferences The aquatic damselflies Coenagrionidae are most abundant in the lowlands Cvriorhinus is also most adapted to the lowlands The nabid Stenonabis tagalicus occurs only in the uplands and isalso prevalent on legumes The coccinellid lady beetles wasps and ants are more prevalent in upland areas AlicrasIpis cit-ea larvae prey ona variety of aphids attacking legumes and maize Etunienes campanonia vespid mud wasp makes nests in trees and is therefore most adapted to the more botanically diverse uplands Soil-dwelling ants cannot tolerate flooding

Upland environments are habitats to arboreal carabid beetles Three species of Chlaeniusbeetles prey on leaffolder larvae and are perhaps more important than parasites (27)

The gryllids and tettigoniids are widely adapted egg predators not only on rice but also on maize and legumes They feed on eggs of most insects that are laid on leaves

The spidercommunity of upland rice environments isrich in species At one site (Batangas Philippines) 31 species have been recorded (23) Of the 176 spider species recorded in Philippine ricefields about one-half (82) occur in upland rice The spiderspecies of upland rice environments overlap more with those of rainfed lowland than with those of irrigated lowland rice (24) The same study showed that of the three environments irrigated rice has (ie greatest spider species diversity foiiowed by rainfed lowland and upland

However of the 10 most prevalent ricefield spiders 9 were abundant in upland rice showing wide adaptation All three spider guilds -- orb-web space-web and huting spiders shywere prevalent in upland ricefields There were however differences in environmental prelerences for some spider species

Of the orb-wcavers Tetragniathamanlihulatawas partishycila rlv abundant 7 japonica was low and Ltucatge dehcorata was absent in upland rice

As in other environments Aiypeiwnaifirtosana was the most dominant space-web spider however its relative numbers were lowerthan for lowland sites Among hunting spiders Ox|opes javanus and Lcosa leatcost fgla were more abundant and Lvcosa lpseudoanmnlata was less abundant in upland compared with lowland rice envishyroniments

Extensive rearing of rice insect pests in upland rainfed lowland and irrigated sites over an 8-yr period in the Philippines provided insight into the effectiveness of parasites as natural enemies (Table 7 8) Most of toe data are from large sample sizes taken over at least one crop which overcomes the pitfalls expressed by Van Driesche (238)

VSB egg parasitization was surprisingly similar across all environments Larval parasitization was highly dynamic among sites and even years within the same site but showed somewhat lowerlevels in uplandsthan in irrigated wetlands This would have been predicted for egg parasites as well since YSB is adapted to the lowlands and so must be its parasites This isevidence that Telenomus rowani is as good at dispersing as its main hosts Trowani clings onto the anal tufts of female YSB and parasitizes eggs as they are laid This phoresy may explain its wide adaptation The more specialized lowland YSB larval parasites therefore appear more effective than their upland counterparts with more alternate hosts

Low collection levels of PSB larvae especially in the wetlands make comparisons between environments difficult Low levels of larval parasifization were recorded in both rainfed environments Except in the Cagayan rainfed wetland site in 1980 low levels ef larval parasitization were evident at irrigated and rainfed sitesalike No differences by environment were evident Higher levels of parasitization occurred in Sarawak during a 1967 outbreak also from tachinids Rothschild (197) indicated that higher parasishytization occurred in upland rice areas

Skipper and green horned caterpillar larval parasitization rates were generally low and showed no environmental effect htighest activity occurred in Cagayan in 1980 (48 parasitization) Although larval parasitization levels of RLF were variable year to year activity appeared greater in the irrigated lowlands than in the uplands Again the highest levels were recorded in Cagayan in 1980 Cagayan has extensive grasslands and fallow around the rice areas and a relatively long rainfall period Rainfall in 1930 was good for crop growth and apparently for increase of larval parasites

IRPS No 123 January 1987 33

Table 7 Parasitization rates of rice planthoppers and green leafhopper in upland rainfed lowland and irrigated lowland environments Philipshypines 1976-840

Environment and site Year Egg parasitization () Nymphaladult parasitization ()

BPH W13P GLH BPII WBPH GLIH

UplandBatangas

Rainfed lowland

1976 1979 1980 1982

15 17 12

2(47)8(39)

4(51)

9(1800)14 (212)

12(304)

8 (101)6 (127)

9(116)

Iloilo

Cagayan

1976 1977 1978 1981 1980 1981 1982

0 (69)

11

44 48 40

-

-

46 55 44

0 (471)

11

52 56 49

0 (200)9 (214)

10 (1259)8(235)6 (900) 3(800)

-7 (462)

11 (270)3 (313)5 (900) 1 (800)

4 (600)10 (725)10 (362)12(870)5 (900) 3 (800)

Pangasinan

Irrigatedlowland

1984 1976 1978 1979 1982

13 (436) 13(379) 14 (457)

6(14)7 (197)

17(198) 11 (486) 18 (275)

3 (81)10 (274)3 (116)

16(871) 19(88)

28(211)9 (764)5 (647)7 (1204) 8(305)

Laguna 1977 11 7 21 1978 31 1979 14 4 4 28 1980 7 8 20 1981 1983

56 7(278)

26 16 (1650)

16 27(1921)

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R I Apostol unpublished 1lo1lo and Pangasinan 136 Cagayan BCanapi unpublished Liuna 175 I C Pantua unpublished 118 234 43 Carino and Shepard unpublished BPH = brown planthopper Nila parvara htgens W131Pi =whitebacked planthopper Sogatelia furcifera GLH = green leafhopper Nephotettrix virescens Figiires in parentheses are sample sizes

Tauher et al (230) reported that internal external and genetic factors influence the seasonal activity of parasitoids

Hopper egg parasites but not nymphaladult parasites were also parlicularly abundant at the Cagayan site possibly the egg parasites have more alternative hopper prey in the weedy areas than nymphaladult paisites

The seasonal dynamics of hopper egg parasites can be seen for upland rice in Batangas(Fig I 2) and for rainfed lowland lice in Pangasinan( 135)and Cagayan( 105) Levels of BPHWBPI-L and GLH egg parasitization were corn-parable in upland and irrigated lowland environments Except for some high rates of paraMsitization early in the season the parasitization rate tended to be steady in all envilonments However hopper nymphaladult parasi-tizution levels were more similar in the rainfed environments and lower than in irrigated rice

Overall parasites do not appear highly effective by themselves against upland or even lowland pests Among natural enemies parasites have attracted the most attention because they can be more readily assesscd

Predators and pathogens lend themselves better to management practices Predators the most important group of natural enemies in riccfields are difficult to quantify On the other hand pathogens appear less im-portant but can be readily cultured and disseminated

The natural enemy commulnity of upland rice is rich in species and differs significantly from that of lowland rice

These beneficial organisms must be conserved by applying insecticides judiciously particularly sprays Seed or soil placement of chemicals minimizes exposure to natural enemies The strategy to derive the greatest benefit from natural enemies isto allow the greatest number of beneficial species to thrive Then perhaps at least one species will be effective against each pest at any tine overcoming the variable abundance of each species within and between years

Programs to introduce exotic species or mass produce indigenous ones are ambitious and expensive because they require trained people on a sustained not ad hoc basis

Chemical control Insecticides offer rapid and efficient control of upland insect pests (190) but-should be used only after other control measures have been considered Insecticides are rarely used on upland rice because of cost (32) Upland rice yields are normally too low to justify the expenditure Also most upland rice is grown as a cash crop Spraying upland rice is more difficult than lowland rice because water is less accessible Government subsidy programs to provide pestishycides to tribal peoples for upland rice production have created problems of toxicity to humans because isolated people have had no experience to be able to handle pesticides safely

However ins-eticide use has beenjustified when a crop is

0Z

Table 8 Parasitization rates of lepidopterous pests in upland rainfed lowland and irrigated lowland environments Philippines 1976-84a

Parasitization (5)

Environment and site Year YSB GFB PSB Earcutting Swarming Skipper Greenborned RLF Brown larva larva caterpillar caterpillar caterpillar caterpillar larva semilooperEgg Larva larva larva larva larva larva

UplandBatangas 1977 - 0 (57) - 0 (63) - 0 (71) - 0 (106) 11(142) 2 (101)1978 43 (4697) 8(451) 5 (38) 3 (40) 6 (61) 2(58) 15 (57) 13 (129) 13 (1286) 4 (30)1979 59(4135) 12(320) 13 (11) 5(28) 9(45) 6(46) 7(23) 8(160) 9(1152) 8(26)

Rainfed lowland Iloilo 1976 - 0 (10) 0(10) 0 (7) 0 (19) 2 13) 15 (180) 5 (76)1978 0 (54) 0 (3) 0 (17) 9 (64) 4 (78) 3 (73) 0 (121) 0 (191)1979 64 (2350) 16 (84) 6 (17) 0(6) 18 (68) 25 (104)

1981 56 (1322) 18(304) 0(5) 12(51) 18(138)Pangasinan 1976 60 (7 7 )b 62(82) 0(111) 5 (39)

1978 58 (53)b 5(51) Cagayan 1979 59(713) 8 (101) 6 (34) 0(7)

1980 0(11) 4 (28) 15 (286)

62 ( 48(177) 61(140)1981 0(5) 12 (2) 13(45) 14 (92)

Irrigatedlowland Laguna 1974 62 8

1974-75 50 41 1975 43 72 1978 96 (623) 21(24) 33 (60)1979 83(2017) 18(110) 8(62) 11 (89) 22(400)1980 78 (1625) 20(75) 0 (16) 4 (72)1983 60

9 (80) 7 (270) 30

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R F Apostol unpublished Iloilo and Pangasnan 136 Cagayan B Canapi unpublished Laguna 175 P CPantua unpublished 118 234 143 Carino and Shepard unpublished YSB = yellow stem borer ScirpophagaincertulasGFB = gold fringed borer Chilo auricilius PSB = pink stem borerSesamia inferens earcutting caterpillar Alythimna separata swarming caterpillar Spcdoptera mauritia acronyctoides skipper Pelopidas mathias reen horned caterpillar Melanitls ledaismene RLF = rice leaffoldersCnaphalocrocismedinalis and Marasmiaspp brown semilooper Mocisfrugalis Figures in parentheses are sample sizes Egg masses

IRPS No 123 January 1987 35

Table 9 Effect of seeding rate and insecticide on plant density and Yield losses from insect pests in upland rice often areyield of Dagge upland rice Batangas Philippines 1976a comparable to those of lowland rainfed or irrigated rice

Seeding rate Insecticide Dosage Plant density Yield (0-Y30o) but usually from different pests The principal (kg aiha) seed treatment (kg aiha) (noIra-row) (tha) groups are soil pests feeding on sown seed and rootsat 14 DE followed by seedling pests These groups have largely gone

50 None - 35 a 20 a unnoticed leading researchers to conclude that stem borers50 Carbofuran F 10 38 ab 23 a and seed bugs are the most important pests (17) Yield loss 10 beCabofuranF100 None - 4551 c 23 a trials that pinpoint major pests need to includc seed

100 Dieldrin WP 10 52 cd 22 a treatments Granules applied in the furrow may not give150 None - 59 d 22 a adequate protection from many seed pests

aAv of 8 fields DE = days after crop emergence In a column means The floristic and pedological diversity of upland rice followed by a common letter are not significantly different at the environments is matched by the diversity of arthropods5level by D)MRT F = flowable Wi = wettable powder both pests and their natural enemies Irrigation has homoshy

genized lowland rice environments Flooded soils tend tothreatened by epidemics of locusts or armyworms New have the same properties and to eliminate soil pests The technology stressing efficiency has minimized insecticide spatial and temporal dominance of cultivated rice has cost benefited specialist pests

Seed treatment is normally inexpensive and can be Upland rice has no specialist arthropod species The economicallyjustified in -nanycases(66) Treating seed with monophagous pests on upland rice prefer the lowlands systemic insecticide (216) but not chlorinated hydrocarbons where rice evolved Rice in the uplands exists only because (247) is most effective in controlling seedling maggots of man and even though lowland rice predates upland riceFoliar sprays have to be repeated to obtain control and specialist species evolved Wheat maize and sorghum do often are washed off by frequent early season rains not have monophagous pests leading us to conclude that Granular insecticides are effective but required dosages are there is no evolutionary advantage for inssects to have too expensive to justify except under heavy infestation narrow host ranges in upland graminaceae(247) When directed against first-instar larvae banding Pests have adapted to the uplands not only by having granules in seed furrows (177) can be effective against white wider plant host ranges but also by having long life cyclesgrubs at low dosages (025 kg aii ha) (134) Seed treatments undergoing dormancy or being dispersive Each pest has a are not promising for white grub control (248) unique life history strategy suited to highly unpredictable

Seed treatment is also effective ainst ants and may be environments In agivenyear someinsect specics an better cheaper than increasing the seeding rate In Batangas exploit upland rice as a food source but no pest is a senious treating 50 kg seed ha was as effective as increasing the one every year on upland rice seeding rate to 100 kgha and using untreated seeds (Table The low yielding potential of upland rice means less 9) In Brazil seed treatments control termites and other soil quantity and lower quality of food for pests to exploit but insects including the lessercornstalk borer(66) Domiciano also less incentive for farmers to undertake control (58) however found seed treatments inconsistent in their The most effective control would be early and synchroshyeffect against high populations of lesser corn stalk borer nous planting of early-maturing upland rice varieties to

Baiting can be an inexpensive way of controlling ants prevent insect pest buildup Breeding for insect resistance seedling maggot flies mole crickets and field crickets Baits should initially focus on stem borers because they are verymade of locally available material and impregnated with difficult to control by other means Biocontrol efforts insecticide can be sparingly distributed taking adantage of should first stress conservation of natural enemies preshythe pests ability to disperse and encounter bait sites dators and pathogens rather than parasites Efficient

The new low-volume hand-held controlled-droplet chemical control includes seed treatment or baiting for seed sprayers may offer an advantage over high-volume knap- seedling and some root pests and using controlled dropletsack sprayes for controlling foliar pests Upland rice sprayers for spot treatments against foliar pests and seed canopies are more open than those in lowland rice and bugs when economic thresholds are reached better droplet penetration should result Controlled droplet applicators would be ideal for seedbugs and defoliators

CONCLUSION

Upland rice is attacked by a wider array of insect pests than is lowland rice mainly beca use of the addition of soil pests Generally population levels of lowland rice pests are lower in upland than lowland rice leading many people to conclude that insect pests are not important on upland rice

36 IRPS No 123 January 1987

REFERENCES CITED

1Abraham C C and K S R Mony 1978 Occurrence ofLeptocorisa acuta Fabr (Coreidae Hemiptera) as a pest of nutmeg trees J Bombay Nat ttistSoc 74553

2 Agyen-Sampong M 1975 HlierogrlphusdiaganensisKrauss (Orthop-tera Acrididae) a new pest of rice in Northern Ghana Ghana J Agric Sci 8249-253

3 Agyen-Samnpong M 1976 Varietal resistance and other inle-ratcd pest control components as factors of rice varietal improvement in West Africa Prcsented at the Second varietal improvement serinar 14-18 Sep 1976 Bouak Ivory Coast West Africa Rice Development Association Monrovia Liberia 18 p

4 Agyen-Sampong M 1977 Outbreak of Ihccrorrdnisorae in rice seedlings near Rokupr Sierra Leone lit Rice Res Newsl 2(6)16-17

5 Agyen-Sampong M 1980 Dirty panicles and rice yield reduction caused by hugs IntRice Res Newsl 5(l)11-12

6 Agyen-Sampong M 1982 The major pest problems of irrigated upland and mangrove swanip rice ecosystems ir the humid tropical Guinea savanna and Salel climate zones Pages 318-328 in Pro-ceedings training course cn concepts techniques and application of irtegrated pest management in rice inWest Africa 10-20 Jan 1982 Fendall Liberia West Africa Rice l)evelopioent Association Monrovia Liberia 505 p

7 Akibo-Betts V I and S A Raymundo 1978 Aphids as rice pests in Sierra Leone lit Rice Res Newsl 3(6)15-16

8 Akinsola F A 1975 Present status of different rice stemborers in parts of Nigeria iicL Entomo Newsl 328

9 Akinsola FA 1979 Ihe biology and ecology of rice stemborers in Nigeria Ph 1) thesis University ol lbadan Nigeria 217 p

10 Akinsola E A 1979 Guidelines of integrated management of insect pests of upland rice in West Africa Presented at the Seminar on integrated management of rice diseases and insect pests at Bobo-Dioulasso Upper Volta 17-22 Sep 1979 West Africa Rice Develop-ment Association Monrovia Liberia

II Akinsola IA 1984 Insect pests of upland rice in Africa Pages 301-305 in An overview of upland rice research Proceedings of the 1982 lIouak Iwsrv Coast upland rice workshop International Rice Research Institute 13O Box 933 Manila Philippines

12 Alam M S V TJohn and Kaung Zan 1985 Insect pests and diseases of rice in Africa Pages 67-82 in Rice improvement in Eastern Central and Southern Africa proceedings of the international rice workshop at Iusaka Zambia 9-19 April 1984

13 Alticri M A A van Schoonhovcn and J D Doll 1977 The ecological role of weeds in insect pest management systems a review illustiated with bean (Phaseohsvulgaris L)cropping systems PANS (Pest Artie News Summ) 23195-205

14 Anmante F 1967 The leaf-cutting ant Atia capiguaraa pest of pastures [in Portuguese] Biol6gico (Sao Paulo) 33113-120 (Rev Appl Entomol A56207)

15Andrade A C and D Puzzi 1951 Preliminary results of experi-merits on the control of Sculptcoris iastantus on sugarcane [in Portuguese] Bioulgico (Sao Paulo) 1744-49 (Rev Appl Entomol A41421422)

16 Appert J 1970 Maliarplia separaelha (rice stem borer) new observations of entomological problems of rice in Madagascar [in French] Agron Trop (Paris) 25329-334

17 Ariadeau Mand Z Harahap 1986 Relevant upland rice breeding objectives Pages 189-197 in Progress in upland rice research Proceedings of the 1985 Jakarta conference International Rice Research Institute 1O Box 933 Manila Philippines

18Asar S and Kisimoto 1975 Seasonal occurrence of the smaller brown planthopper in upland rice [in Japanese] Proc Kanto-Tosan Plant Prot Soc 2228

19 Bakker W 1974 Characterization and ecological aspects of rice yellow mottle virus in Kenya Meded Landbouwhogesh Wage-ningen 152 p

20 Balut F F 1970 White grubs attacking rice Oryza ativa L [in Portuguese] Biol6gico (So Paulo) 36321-322

21 Banerjee S N and L M Pramanik 1967 The lepidopterous stalk borers of rice and their life cycles in the tropics pages 103-124 in The major insect pests ofthe rice plant Proceedings ofa symposium at the International Rice Research Institute September 1964 The Johns Hopkins Press Baltimore Maryland USA

22 Barrion A T and J A Litsinger 1980 Ants a natural enemy of cliaiarac in dryland rice IntRice Res Newsl

5(4)22-23 23 Barrion A T and J A litsinger 1981 The spider fauna of

Philippine rice agroecosystem 1Dryland Philipp Entomnol 5139-166

24 Barrion A T and 1 A Iitsinger 1984 The spider fauna of Philippine rice agroccosystcms II Welland Philipp Eiitomol 611-37

25 Barrion A T1 and J A Litsinger 1984 lihanus (Diptera Tabanidae) eggs an alternative host of rice stei borer (SI3) egg parasite Telenoonis digns (Iliymenoptcra Scelionidae) lit Rice Res Newvsl 9(6)19

26 Barrion A T and J A Litsinger 1985 Identilication of rice leaffolders by wing markings Int Rice Res Newsl 10( 1)24

27 Barrion A T and J A Litsinger 1985 Chctiosspp (Coleoptera Carabidae) a leaffolder ILF)predator lit Rice Res Newsl 10(1)21

28 Bertels A 1970 Insect pests of rice and theircontrol [in Portuguese] Pelotas Brazil Instituto de Expcrimentacao Agropccuararias do Sul Circular 43 24 p

29 Bonzi S M 1982 Ctilo (flisilineus J de Joannis (Lepidoptera Pyralidae) stem borer of irrigated cereals in Upper Volta LioFrench] Agron Trop Riu Rizic Cult Vivrieres Trop 37207-209

30 Bourke T V T L Fenner J N LStihick G Baker EHassan D F OSullivan and C S li 1977 Insect pest survey for the year ending 30th June 1969 Papta New Guinea )epartment of Agrishyculture Stock and Fisheries Port Moresby 57 p

31 Bowden 1 1973 Migration of pests in the tropics Meded Rijksfac Landbouwwet Gent 38785-796

32 Breniere J 1973 The main rice parasites in West Africa and their control Pages 27-42 in Proceedings seminar on plant protection for the rice crop West Africa Rice Development Association Monrovia Liberia

33 Breniere J 1976 The principal insect pests of rice in West Africa and their control West Africa Rice Development Association Monrovia Liberia 52 p

34 Brown ES 1962 The African arinyworn Spoiolptera eXI)Ina (Walker) (Lepidoptera Noctuidae) a review of the literature Cornshymonwealth Institute of Entomology London UK57 p

35 Brown F B 1968 Upland rice in Latin America FAO-International Rice Commission 12th Session of the Working Party on Rice Production and Protection 2-14 Sept Peradeniya Sri lanka Food and Agriculture Organization Rome Italy 6 p

36 Buyckx F J 1 1962 Disease and insect pests of crops inCongo Rwanda and Burundi [iil French] Publications dc LInstilut National pour lestude Agrononiquedu Congo M Weissenbruck S A Brussels Belgium 708 p

37 Cariiio F 0 1981 Role of natural enentics inpopulation suppression and pest management of green rice lealhoppers Ph D thesis University ofthe Philippines at Los Bafitos PhilippinesLaguna 136 -

38 Carpente A J 1983 The development of rice cultivation and extension of vegetable productin in Zanzibar Final Report Vol II1 October 1983 Food and Agricultme Organization Ronte Italy

39 Carroll C P and S1 Risch 1983 Tropical annual cropping systems ant ecology Environ Manage 751-57

4(0 Cates R G 198 1 Host plant predictability and the feeding patterns of monophagous oligophagous and polyphagous insect herbivores Oecologia (Berl) 48319-326

IRPS No 123 January 1987 37

41 Cendcafia S M and F B Caora 1967 Insect pests of rice in the 62 Ene 1 C 1981 I)iiorldhol)ierusihilis Slater (licteroplera Philippines Pages 591-616 in The najor insect pests ofthe rice plant lygacidue Illissinac) a new rice pest front Nigeria lit Rice Res Proceedings of a symposium at the International Rice Research Newsl N3)15 Institute September 1964 The Johns Hopkins Press Baltimore 63 Fugenio T S 1965 rhe effect oi e-vCgetation on rice production Maryland USA 729 p and the possibility of overcoming it by leaf pruning MS thesis

42 Centro Ihternacional de Agricklittira Tropical 1984 Upland rice in University ol the Philippines at Los Baijos ILaguna Ihilippins 69 p Iatin Anerica Pages 93-119 hi An os ervies of uplaind rice research 04 Farrow R A 1975 The Alrican nmigratory locust ii is main Proceedings of the 1982 Boinakamp Ivory Coast upland rice workshop outbreak area of the middle Niger qlinlitatise studies of solitary International Rice Research Institute P0 Blox 933 Manila populations in relation to environmental factors Locusla No II Philippines 198 p

43 Chandra G 1979 Taxonomy ad hn riononlics ofthe imect parasites of 65 Fennal R G 1943 Food crop investigation Windward and rice Icalloppers and planthoppers in the Pihilippines and their Ieeward Islands First report Report onl work dunring the periodiiiportairce iii natural biol)gical cointrol Philipp Fntoltol 4119-139 April 1942-April 1943 Imperial College of Tropical Agriculture

44 Chang T T 1976 The originii colut ion enlIivation dissemination Trinidad 16 p (Rev of Appl Fintornol A3222-25) and diersificition of Asian and African rices -uphytica 25425-44 1 66 Ferreira F 1983 Integrated pest ritaageiret of upland rice [ill

45 Chatierjee P B 1979 Rice leaffolder attacks in India lit Rice Res Portuguese English summary] Pages 323-341 in Upland riceculttre Newsl 4(3)21 factors affecting production E Ferreira T Yamada and F-

46 Cheng Cbien-pan 19(4 Upland rice in Taiwan International Rice Malmolta eds Institutoda Potassae Foslato Institnto Internacional Cnmmission Wiorking Party oit Rice Production ard Protection I()th da Potassa Piracicaha Sao Paulo lraid Meeting 3-10 March 1964 Manila Philippines Food and Agricul- 67 Ferreira E arid F dl Silva Martins 1979 Occurrence of ture Organiaion Rome Italy 5 p Sogatohs orvziola (Muir) inupland rice in Goias Ilrail lit Rice

47 Chcrrett 1 M and 1) 1Peregrine 1976 A review of the status of Rcs Ncwssl 4(4)18 leaf-cutting ants and their control Ann Appl Biol 84124-128 68 Ferreira E and 1F da Silva Martiris 1984 Insect pestof rice ill

48 Chipeta F M 1974 Pests of rice in Malawi Pages 90-95 in Brazil and their control Jill Portugucsej FMIlRAtPA-CNPAF Proceedings Sith eastern Africa cereals research conference 10-15 Goiania Brmiil 67 p March 1974 DRB Mialda ed Ministry of Agriculture lilongwe 69 Ferreira FI F ca Silha Miartins S S Nero and 1 It P de Malawi Carvallio 1982 M tltiplc regression analysis todeterimine the effect of

49 Chu Yau-i 1979 The d atiaging period of the rice seedling fly insect populatiois ott upland rice yield [inIPortuguese Pesqui (Atlterigotiavxigu Stcin) (NItiscide Diptera) oil cori) at Fast Java Agropecu Bras 17671-675 Indonesia Plant Prot Bill 21403413 70 Ferreira F 1F da Silva Martins and F 1PZimmermann 1979

50 Conklin H C 1957 IHaninoo agriculture a report on aln integral Rice varietal resistance to tle lesser corn stalk borer [in Port uguese] systent of shi fti rig cult ivationi in the Pltilippincs Food and Agriculture Pesqui Agropccu Biras 14317-321 Organization Rome Italy 209 p 71 Fleming W F 1968 Biological control of lie hapanese beetle U S

51 Cramer H II 1967 Plant proection and world crop production Dep Agric RLs Sers [ech Bull 1383 78 p Pilaniensclhutz-Nachr 201-524 72 Food and Agriculture Organiation of the United Nations 1982

52 l)ani R C and Majunidar 1978 Prelininary observations on tie Country report surmimary Pages 27-37 in Report on the first incidence of root aphids on differeit rice cultnrcs Sci Cult 4488-89 FAOIUN)IPhail and regional training coiuirse in iniproved ctltural

53 Davidson R I 1979 lffects of root feeding on foliage yield Pages practices for upland rice 5-23 Oct 1981 Northern Region Agrculturd117-120 in T K Ci osb and R B Pottinger eds Proceedings of the Developririt Center Chiang Mai Food and Agricuilture Organiiashy2nd Australian conlerence on grassland invertebrate ecology Govt tion Rome Italy 37 n Printer Wellington New Zealand 73 Franco do Ariaral S 195 1 Residual action of insecticides in turfy

54 l)ean G 1W 1976 Rice insect pests inlaos Int Rice Res Newsl soil--the results of a field experiment [in Portuguese] Biol6gico (Sao 1(2)15 Paulo) 17209-214 (Rev Appl Entomol A41423)

55 I)ean G J W 1978 Insect pests of rice in Laos PANS 24(3)280-289 74 Freeman ) 1970 Report onithe Iban london School of -coroiics 56 )ingle H 1984 Behavior genes and lifc histories conplex adapta- Monographs on Social Anthropology No 41 Athlane Press

tions to uncertain environments Pages 169-194 in A new ecology London UK 317 p novel approaches to interactive systems P W Price C N Slobod- 75 Fuchs T W1 A Harding and 1W Smithlr 1979 Induction and chikoff and W S Gaud cdsl ohn Wiley New York USA termination oldiapause in the sugarcane borer Ann Flitonlol Soc

57 Dobelmann-JP and M Falais 1968 Upland rice cultivation ill tie Am 72271-274 North-western part of Madagascar in 1966-1967 [in French English 76 (hbriel B1) 1975 A review tf the major insect pests of some tpland summary] Trop Agron 2313-21 crops in tlc Philippines Pages 86-111 in Ad Htoc panel of experts on

58 Doniiciano N L 1979 Chemical control of tie lesser corn stalk pest disease arnd weed problems in some rainfed crops Bangkok (lolsonno~paloIgnowslhts) (Zeller 1848) (I-epid(ptera Ihycitid ae) Thailand Food aid Agriculture Organization Roire Italy

on rice in Ponta Gressa Parara 1977-1978 [in Portuguese] Pages 77 Garg D K and N K Shal 1983 White grub outbreak on rainfed 291-303 in Anais 1 Reuniao de tecnicos cmiirizicultura do estado de dryland rice in Uttar Pradeslh Int Rice Res Ncwsl 81(4)16-17 Sao Paulo-Campinas Cati Brazil 78 Garg I) K and IP Tandon 1982 Major insect pests of rice on hilly

59 Dunsmore 1 R 1970 Investigations on tlie varieties pests and tracts of Uttar Pradesh India lilt Rice Res Newsl 7(11)11-12diseases of upland rice (hill padi) in Sarawak Malaysia Int Rice 79 Garrity 1) P 1983 Upland rice research an international bibliograpy Comm Newsl 1929-35 1975-1982 Iternational Rice Research hnstitnte PO Box 933

60 Elias 1R1965 The new decline of rice seedlings is not caused by Manila Philippines 121 p parasites [in Portuguese] Biol6gico (Sao Paulo) 313-6 80 Ghauri M S K 1982 A new subspecies of Iinorh)rone bull

61 Empresa Brasiliera de Pesquisa Agropecuaria 1984 Upland rice itt cornlus Slater (Hemiptera Iygaeidae) front Papua New Guinea on Brazil Pages 121-134 it Av overview of upland rice research rice and carpet grass Ihll lEntoniol Res 72133-137 Proceedings ofthe 1982 Blouak Ivory Coast Upland rice workshop 81 Gonzales F 0 PArregoces R I ihernandef and OT Parada International Rice Research Institute PO Box 933 Manila 1983 Insect and imite pests of rice arid their control in Latin America Philippines [in Spanish] la Federacion Nacional de Arrocerts (Fedearroz)

Bogota Colombia 60 n

38 IRPS No 123 January 1987

82 Grist D H and R J A W Lever 1969 Pests of rice Longmans London UK520 p

83 Hale P R 1979 Two subterranean pests of upland rice in Papua New Guirnca IntRice Res Newsl 4(2)16-17

84 Hale P R and J T Hale 1975 Insect pests of rice in the New Guinea Islands Rice Entomol Newsl 34

85 Harris W V 1969 Termites as pests of crops and trees Common-wealth Institute of Entomology London UK 41 p

86 Hattori 1 1968 A method to distinguish between grass leaf roller (Ctaplha rocis medinalisGuenee) and the rice leaf roller (Susurtia exigua Butler) [in Japanese] Plant Prot (Japan) 23167-170

87 Henry I E and 1A Onsager 1984 Experimental control of the mormon cricket Iahrosshnph by NJsena locustae(Aficrospora Microsporidae) a protozoan parasite of grasshoppers (Ort Acri-didae) Entomophaga 27197-201

88 Hidaka 1 P Vungsilabulr and SKadkao 1974 Studies on ecology and control of the rice gall midge in Thailand Tech Bull 6 TropicalAgricultural Research Center Tokyo Japan 113 p

89 Hincklcy A 1) 1963 Ecology and control of rice planthoppers in Fiji Bull Entomrol Res 53467 481

90 Hirao J 1982 Ecology and chtInical control of the rice leaffolder Jpn Pestic Inf 4114-17

91 Ho DT 1984 Sten borer incifience in rice ecosystems in KenyaEast Africa Pages 307-320 in A overview of upland rice research Proceedings of the 1982 Bouac Ivory Coast upland rice workshop International Rice Research Institute PO Box 933 Manila Philippines

92 Ho )T J 1Njokah and J GKibuka 1983 Studies on rice sten borers in Kenya with emphasis on ifaliarpiaipiraellRag Insect Sci Appl 465-73

93 Ichihara I T Ito A NIatsuo and M Sawada 1971 Chemical control of the rice striped stem borer in upland rice grown in plastic mulch [iii Japanese] Proc Kanto-Tosan Plant Prot Soc 1882

94 IclhiharaIand A Matsuo 1969 Damage caused by rice striped stett borer to upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1686-87

95 Ichhara I atid A MIatsun11970 Conside ration oft lie extraordinary prevalence of rice striped stem borer in upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1778-79

96 Institut de Recherches Agrolnliqes Iropicale et des Cultures Vivrieres 1978 Insect resistance Pages 12-13 in Upland rice 1978 atnual report Bouak Ivory Coast 27 p

97 International Institute for Tropical Agriculture 1983 Aphids itt rice roots I ITA Res Briefs 4(1-3)4-5

98 International Institite fi Tropical Agriculture 1983 Hopperburn in rice caused by brown plathopper IITA Res Briefs 4(l-3)6-7

99 International Rice Research Institute 1975 Annual report for 1974 PO Box 933 Manila Philippines p 216

100 International Rice Restarch Institute 1976 Annual repiort for 1975 PO Box 933 Manila Philippines p 228

101 International Rice Research Institute 1979 Cropping systems program Component technology development and evaluation Page 404 i Annual report for 1978 11O Box 933 Manila Philippines

102 International Rice Research Institute Department of Entomology 1981) Insecticid evaluation 1979 Tables 73 74 PO Box 933Manila Philipes

103 International Rice Research Institute Department of Entonmology 1981 Insecticide evalomtion 1980 Table 81 POBox 933 Manila Philippines

104 International Rice Research Institute 1981 Report of a meeting of Asian rice-based cropping systems entomologists Eleventh cropping systems working group meeting 18-22 May 1981 PO Box 933 Manila Philippines p 132 133 144

105 International Rice Research Institute 1984 Cropping systems program Pages 368-373 in Annual report for 1983 PO Box 933 Manila Philippines

106 International Rice Research Institutc 1984 Upland rice in Asia Pages 45-68 in An overview of upland rice research Proceedings of the 1982 Bouakamp Ivory Coast upland rice workiiop loternational Rice Research Institute 11O Box 933 Manila Philippines

107 Irshad M 1977 Grasshoppers in rice in Pakistan lit Rice Res Newsl 2(3)8-9

108 Ito R and TOno 1964 Upland lice its culture and varieties in Japan lIternational Rice Cormmission loth Meeting of the Working Party on Rice Production and Protection 3-10 March 1964 Manila Philippines Food and Agriculture Organization Rome Italy 12 p

109 Jeffrey M I and V A Dyck 1983 Effect of moonlight on lightshytrap catches of brown planthopper lit Rice Res Newsl 8(6) 13

110 Jennings P R and T Alicia Pineda 1983 Management ofthe rice plnthoppers Sogatodsorivi(coIl in Latin America Presented at the meeting on stable plant resistance FAO and lIndocsian I)inectorate of Food Crops Protection Bali lndonesia 29 Novcniber-2 l)ccemshyber 9 p

IllJimenez Gomez J A 1979 Rice insect pests i lla ios ()riCntales lil Spanish] Pages 99-103 in Rice ciltivation Instituto (olotibiatto Agropecuario Compndio No 29

112 Jongeleen F JJ Soesanto and A Mockidjo 1979 Studies oti pathogens of two species of Scarabacidac froinIndonesia Z Angew Enlomnol 87412-417

113 Jouvenaz 1) PC S lofgren and W A Baiks 1981 Biological control of imported fire antsa review of cirrent knowledge Bull Entomol Soc Am 27203-208

114 Kainoh Y K Shiniiu S MarUani Y Taiaki 1980 lost finding behavior of tie rice bu lplotori+JchinvittjI)allas (liciniptera Corcidac) with special reference to diet pattern ol aggregation and feeding on the rice plant Appl Entomol Zool 15225-233

115 Kakkar K 0 P Sharma and P I Sharma 1977 Note on Ihteronyrchus liodresRedt (Coleopicra I)ynastidae) as asporadic pest of paddy and maie inHimachal Pradesh Corr Agric 156-57

116 Kalode M131 79 Major insect pest problems and their control in upland rice (Unpublishcd mineogriph CWI oJNaaiat lni Chandaratat (198 1)original not seen)

117 Kalshoven L i 98 I The pests of crops in I ldoniesia (translated by P A vatn der Laan) 1 T Ichtiar B3aru-vati Iloeve lakarta Indonesia 701 p

118 Kamal N Q 1981 Suppression of whitebacked planto ner Sogatella fircaifra (Horvath) aLId rice lealfolder (nhal -cis licdb lis (Guenee) populations by natural cnclmie hlI) tiesis Grcgorio Araneta University Foundation Victoncta Park Metto Manila Philippines 179 Ip

119 Karim A N M Reaul 1975 A species ofliealy bug Trioni i70s ceres new oii rice in the Philippines Rice Entoniol Newsl 247

120 Katanyukul Wand C Chandaratat 1981 Insect pestsof uplaid rice and their control Report firstFAOUNlFi Tliailand training course on improved cultural practices for tplad rice Chiang Mai Thailand 5-23 Oct 1981 Food and Agriculture Organiation Rome Italy 22 p

121 Kern F 1955 lhletwioa(I)uninemts) bhhit Biturm(Coleoptera) a new pest of sugarcane inVenezuela [inSpanish with Etnglish sunmary] Agron Trop (Maracay) 527-29 (Rev Appl Eitoniol A4572)

122 Khan K M and S Ghai 1974 White grubs ald their control in India Pesticides (1Bombay) 819-25

123 Kiritani K 1975 Integrated pest management of rice pests Pages 66-70 itt Proceedings platning workshop ott cooperative hIeld research in pest managementP SMotooka ed East-West Center Honololu Hawaii USA

124 Kisimoto R 1967 Ecology of insect vectors forecasting and chemical control Pages 243-255 in The virus diseases of tle rice plant Proceedings of a symposium atthe International Rice Research Institute April 1967 The Johns Hopkins Press altimore Maryshyland USA

125 Kisimoto R and V A Dyck 1976 Climate and rice insects Pages 367-391 in Climate and rice International Rice Research Institute PO Box 933 Manila Philippines

126 Kureha S S Kobayashi SAsari and T Ishihara 1977 Life history and control of the mole cricket Grrllotalpa africana Palisot de Beauvois [in Japanese English translation 7520 1RRI library] Proc Kanto-Tosan Plant Prot Soc 21192-197

127 Kushwaha K S 1981 The black beetle as a pest of rice in Haryana India Int Rice Res Newsl 6(4)14

128 Lawani S M 1982 A review of tile effects of various agronomic practices on cereal stem borers Trop Pest Manage 28266-276

129 Leuck D B 1966 Biology of the lesser cornstalk borer in South Georgia J Econ Entomol 59 797-801

130 lever R J A W 1955 Insect pests of the rice plant in Malaya Int Rice Comm Newsl 1612-21

131 Li Ching-Sing and Shin-Foon Chiu 1951 The study ofthe rice gall midge Iachydiplosis orvza Wood Mason [in Chinese with English summary] 1Taiwan Agric Res 21-13

132 Iitsinger J A 1984 Assesament of the need-based application for rice Proceedings TechnologyTransfer Workshop 15-16 March 1984 International Rice Research Institute 1PO Box 933 Manila Philippines 6 p

133 Litsinger J A 1984 Environmental diversity and insect pest abundance with reference to the Pacific Pages 285-307 in Proceedings sub-regional training course on methods of controlling diseases insects and other pests of plants in the South Pacific 4-20 Oct 1982 Vaini Kingdom of Tonga Nukualofa

134 Litsinger J A RF Apostol and M BObusan 1983 White grub Leucopholis irrorata (Coleoptera Scarabaeidae) pest status popula-tion dynamics and chemical conllol in arice-maize cropping pattern in the Philippines J Econ Entoinol 761133-1138

135 Litsinger J A C de la Cruz F Raymundo A T Barrion M D Lumaban J P Bandong M S Venugopal F Paragna and E Balete 1982 Insect pests and insecticide reponse of dry-seeded bunded rice Pages 179-193 in Report of workshop on cropping systems research in Asia International Rice Research Institute Los Bafos Philippine

136 Litsinger J A M 1) Luinaban J P Bandong A T Barrion W Katanyukul and N Panda 1978 Insect pest management in rainfcd lowland rice agroecosystms of tropical Asia International Rice Research Conference 17-21 April 1978 International Rice Research Institute Los Bafios Philippines

137 Litsinger J A and K Moody 1976 Integrated pest management in multiple cropping systems Pages 293-317 in R I Papendick P A Sanchez and G B Triplett eds Multiple cropping American Society of Agronomy Madison Wisconsin USA

138 Loevinsohn M E 1984 The ecology and control of rice pests in relation to the intensity and synchrony of cultivation Ph D thesis University of London 36) p

139 Loevinsohn M E J A Litsinger N Panda M A Austria R F Apostol C Andrion and J Bandong 1982 A basis for predicting rice insect pest abundance in cropping systems Paper presented at the 1982 International rice research conference on rice and cropping systems 19-23 April 1982 International Rice Research Institute Los Bafios Iaguna Philippines 25 p

140 Lopez A W 1930 Repot of the Entomology Department Pages 145-172 in Annual report of the Philippine Sugar Association 1929-30 Manila

141 Lopez AW 1931 Entomology Department Pages 227-266 in Annual report of the director or research 1930-31 Philippine Sugar Association Manila

142 Lopez A W 1932 Entomology Deparment Pages 252-279 in Annual report of the Philippine Sugar Association 1931-32 Manila

143 Lungibill P and G G Ainslee 1917 The lessercornstalk borer US Dep Agric Agric Res Serv Tech Bull 539 27 p

IRPS No 123 January 1987 39

144 McGuire J U Jr and BS Crandall 1967 Survey of insect pests and plant diseases ofselected food crops of Mexico Central America and Panama U SDepartment of Agriculture International Agriculshytural Development Service Agricultuial Researrh Service October 1967 11 p

145 McNeill Sand T R ESouthwood 1978 The role of nitrogen in the development of insectplant relations Pages 77-9 in lliochemical aspects of plant and animal coevolution J B Ilarbornu ed Academic Press London UK

146 Maebara H 1955 Spatial distribution of lastinstiarlanaeof purplish stem borer (Sesamia iqferens Walker) hibernating in an upland rice field Bull Kagoshima Univ Fac Agric 438-41

147 Mallick E H and N G Hajra 1977 Response of rice plant to defoliation treatments at different developmental stages Indian Agric 2151-56

148 Mansingh A 1971 Physiological classification of dormancies in insects Canl Entomol 103983-10)9

149 Martell C G 1974 Catalogue of plant feeding insects of Mexico [in Spanish] Fitofilo 27(69)1-176

150 Martinez 0 J 1982 Principal rice pests in Ilanos orientalis [in Spanish] P~ages 80-94 in Curso deact nalization de conocimientos en arroz Villaniencis Instituto (olonmbiano Agropecuario

151 Martins J F da Silva F Ferrcira A S Prabhu F 1 P Zimmermann 1980 Use of pesticides for the control of the main subterranean upland rice pests esqui Agropecu Bras 1553-u

152 Martins I F da Silva F Fe eira rod BdamSilveira Pinheiro I Q2 Damage simulation caused by defoliators of upland rice [in Portuguese with English summary] Pesqui Agropecu Bras 171113-1119

153 Martins 1F da Silva N(iyuen van Tan and 1Ida Silveira Pinheiro 1981 Stem borer resistance in upland rice associated with plant morphological characters [in Portuuce Iesqui Agropecu Bras 16187-192

154 Masaki S 1980 Sumnmer diapause Ann Rev Entomiol 251-25 155 Masuzawa T 11 Suwa and F Nakasoji 1983 l)ifkleences of

oviposition preferencc and survival rate of two skipper butterflies Parnara guttata and PIchlpidai iathiais (Iepidoptera HIesperiidac) on rice plant and cogon grass New Entomol 3247-56

156 Maurya D M and C P Vaish 1984 Upland rice in India Pages 69-92 in An overview oftpland rice research Proceedings ofthe 1982 Bouakamp Ivory Coast upland rice workshop International Rice Research Institute IO Box 933 Mnila Philippines

157 Menon M G R R N Katiyai S Kuar and S S Misra 1970 Termites attacking the paddy crop Sci Cult 36294-295

158 Miyahara Y and 1 Nishikubo 1967 Behavior of young nymphs of the red rice root aphid Iholahsipiun ri liabhdontinalisSasaki in relation to movement from the leaf to the roots of upland rice [in Japanese] Pioc Assoc Plant Prot Kyushu 1363-66

159 Mo ljoa Tjien 1952 White grub of upland rice (Holotrichiahelleri Brsk) [in Indonesian with English summary] Contrib Agric Res Stn Bogor No 13134 p

160 Mochida 0 and T Suryana 1979 The occurrence of the brown planthopper and ragged stunt on lowland and upland rice ill Central Java and logyakarta in tile late of January 1979 Entomnolgy Survey Report No 15 Central Research Institute of Agriculture (CRIA) Sukamandi Indonesia I February 1979 9 p

161Monet C 1979 Varietal resistance to upland rice pests [inI French] Pages 35-39 in Annual report 1977 Institut des Savanes Department des Cultures Vivrieres Bouak6 Ivory Coast

162 Monte 0 1942 A caterpillar ofrice-fields[in Portuguese] Biol6gico (Sao Paulo) 8161-163 (Rev Appl Entomol A31414)

163 Munroe I) D 1979 Pest management strategy for hill rice in four resettlement areas in Sarawak Malaysia Pages 89-102 i1 Guidelines for integrated control of rice insect pests Food and Agriculture Organization Plant Production and Protection Paper 14 Rome Italy 115 p

40 IRPS No 123 January 1987

164 Nakasuji F M Ishii and Jen-Zon Ito 1984 Rice skippers in Taiwan and their life histories ZZrC (Taipei) Tech Bull 82 10 p

165 Navas I) 1976 Fall armyworum in rice Pages 99-106 il Ecological animal control by habitat management Proc Tall Timbers Con-ference 28 Feb-I Mar 1974 Fall Timbers Res Stn Misc Publ 6 Tallahassee Horida USA

166 Nickel 1 1 1961 Biological control of rice stem borer A feasibility study Tcch Bull 2 International Rice Rcarch Institute PO Box 933 Manila Philippines IIl p

167 Nigan P M and R A Vernia 1985 Insect pests of uplaild rice in Uttar Pradesh Int Rice Res Newsl 10(6)22

168 Nilakhe S S 1985 -cological observations on spittle bugs with emphasis on their occirence in rice Pcsqtui Agropecu Bras 20407-414

169 Nixon G EJ 195 1 lie association of ats with aphids and coccids Commonwealth Iusti lcte of Entonology Iondon UK 36 p

170 Nurdin F an Ishak Manti 1985 Study oit seedling fly of upland rice Interlational Upland Rice Moitoring Tour Report 26 lebruary-2 MarLI Sukaranti West Sumatra

171 Oka 1 N 1979 Ciltural conttol of the brown planthopper Pages 357-369 in Brown planthopper threat to rice production in Asia International Rice Research Institutc 1p0 Box 933 Manila Philippines

172 Ohni M and A Vilnaii 1975 Rice insect pests of Italy biology of Sipha grcria (Kaltenbach) and its control [in Italian] Riso (Milan) 2459-71

173 Otanes F Q atnd 1 I Sison 194 1 Pests of rice Philipp J Agric 12211-259

174 Painter R I 1951 Insect icsistance in crop plants University Press of Kansits Lawrence Kansas USA 520 p

175 Pantua P C and 1A litsinger 1981 Arthropod abundance in continuous vs biannual rice cropping systeis Int Rice Res Ncwsl 6(2)20-21

176 Pathak M ) and V A l)yck 1975 Sttudics ott insect pests of upland rice Pages 186-191 in Major research in upland rice International Rice Research Institute PO Box 933 Ilanila Philippines

177 Patil B R B M IHasabe and 1) S Ajri 198 I Chemical control of white gritb lIditriihinhorraia Fabr intfesting groundntut paddy atnd sorghum Indian 1Entomol 43232-236

178 Pawar A 1) 1974 (ollection and identification of insect pests of rice rlerminal repirt lInternational Rice Research Institute PO Box 933 Manila Philippines 72 p (tnimco)

179 IerrajU A aid A S Reedy 1965 Observations ott diapausc itt tie steti borer ohrice Rice News Teller (lai-Mar)9-13

180 Pont A C 1972 A review of the Oriental species of theri ona Rondani (I)iptera Muscidac) of econtomic importance Pages 27-102 in Control of sorglium shoot fly M G lomtani and W R Young eds Oxford atd IH1 Ptthl Co New I)clhi India 324 p

181 Prasad S C 1PChand aid M 1PSingh 1983 Incidence of rice gall midge in the upland plateau region of Hihar Oryza 20235-236

182 Rai P S 198 0 Nisaga simpex damage to rice in the hill tracts of South India Int Rice kles News[ 5(I)12

183 Rathorc Y S 1969 Gonocplhahli dcprssunt Fabr as itpredator of (tilo zonwIlhis (Swinhoc) and other moths in field cage at Patnagar Uttar Pradesh Indian 1Entotnol 3127

184 Reed W 1983 Crop losses causcd by insect pists in the developing world Proceedings 10th Internatial Congress of Plant Protection lrighton UK 174-80

185 Reedy M V 1981 Ateactogasterinhens(Walker)a new root-grub pest of tipland paddy and inaize Curr Sci 501076

186 Reveno M A 1968 Catalogue of the insect pests of economic crops in Colombia [in Spaiish] Asociacion Latino Americana de Entomo-logia (AIAE) 129-33

187 Reyes G M 1957 Rice dwarf disease in the Philippines Food and Agriculture Organiiation Plant Prot Bull 617-19

188 Rice S E A A Grigarick and M 0 Way 1982 Effect of leaf and panicle feeding byarmyworm (Lepidoptera Noctuidae) larvae on rice grain yield Environ Entomol 7593-595

189 Risch S J 1) Andow and M A Altieri 1983 Agioccosystem diversity arnd pest control data tentative conclusions and nizw research dirctions Environ Entomol 12625-629

190 Rivera C T 1956 Three organic insecticides in thecontrol of insects affecting upland rice Philipp Agric 39465-472

191 Roffrey 1 1972 Locusta outbreaks in the Philippines Acrida 1177-188

192 Rojas R 1) 1941 The effect of pruning at varying ages upon the yield of Ramairio Phipp Agric 29851-860

193 Roonwal M i 1979 Ternite life and termite control in tropical South Asia Scientilic Pnbl odhpr India 139 p

194 Rosetto C1 F da Silva Martins N C Schmidt and L EAzzini 1978 Damage caused by pasturespittle bugs 4Dvoi)lavopictaandDJ schach) on rice lBragantia Nota 6 (Matcli)25-27

195 Rosztto C1- S S Neto I- Amantc I) Iink 1Garcia-Vieira D M de SoLza N V Bansa t and A M de Olivcira 1971 Rice pests of Brazil [in Portuguese] Pages 149-238 inrFA()-International Rice Commission 2nd meeting of the Rice Committee for the Americas 6-11 l)ecember 1971 Pelotas Rio Grande do Sul Brazil

196 Rothschild G H L 1969 Obseratiois on the armyworm Spodoptera matritia acronrctoiiles Gn (Lep Noctuidae) Bull Entomol Res 59143-160

197 Rothschild G 1- L 1974 Parasites of the armworm Spolopiera mauritiaacroncitoihes in Malaysia Fntornopliaga 19293-299

198 Sachan i N S K Gangwar and 1N Katujar 1980 Field crickets damaging cages in North Eastern IHill Region Indian J Entomol 42526-527

199 Sands 1) P A 1977 The biology and ecology of Lepocorisa (Hemiptera Alydidac) in Papua New Guinea Res Bull 18 Departshyment of Primary Industry Port Moresby Papua New Guinea 104 p

200 Sarawak 1968 titomology division Pages 97-106 in Report of the research branch of the Department of Altriculture 1966 Kuching (Rev AIppl Eintomol A56603)

201 Satpathy JM M S Das and K Naik 1977 Effect of multiple and mixed cropping on the incidence of scine important pestsI Entomol Res 178-85

202 Sauer IHF G 1939 Notes Ott Elasmpnlpualtsligniosellths Zeller (Lep Pyr) a serious pest ofcereals in Sao Paolo [in Portuguese] Arq Inst Biol 10199-206 (Rev Appl Entotnol A28378)

203 Satinders 1L A 1 SKing and C L Vargas S 1983 Crop pests of Central Anerica with t bibliography [in Spanish] Centro Agroshynotnico Tropical de Investigacion y Ensenanza Turrialba Costa Rica (CATIE) Tec Bol 9 90 p

204 Sauphanor B 1985 Some factors of upland rice tolerttce to stem borers in West Africa Inseu Sci Appl 6429434

205 Sen P atnd S Chakravorty 1976 Stentflics in Haritighata Nadia West Bengal India [it Rice Res Newsl 1(12) 15

206 Senapai B and 1M Satpathy 1983 The seedling fly Atherigona orrzae Malloch is a pest of rice in Orissa (India) J Entomol Res 777-79

207 Shhjahan M S Alatn M Ah nad and BN Islam 1983 A new scarabaeid beetle pest of dryland rice in Bangladesh Int Rice Res Ncwsl 8(I)16-17

208 Sharma G L S C Modgol and R C Gautam 1979 Effect of depth of submergence ott incidence of bacterial blight and white grub infestation in transplanted rice Int Rice Res Newsl 4(6)12-13

209 Sharma S 1985 Scarabaeid beetle outbreak itt upland rice Int Rice Res Newsl 1(1(l)24

210 Shiraki T 1917 The paddy borer Schoenobhits incertulas Walker [achaku Agric Exp Stn Formosa 256 p

211 Silva A de Brito and 11P Magalhaes 1981 Insect pests of rice [in Portuguese] Circular Tecnica No 22 EMBRAPA Centro de Pesquisa t ecuaria de Tropico Unido (CPAJU) Belem Brazil 14 p

212 Singh S A 1979 Emergence of rice leaf butterfly as a rice pest in Manipur India int Rice Res Newsl 4(4)16-17

213 SinghS A 1983 Changing trends of insect pestsof rice in Manipur Int Rice Res Newsl 8(f)12

214 Siwi S S and P H van Doesburg 1984 Lenovorisa Latreille in Indonesia (Heteroptera Coreidae Alydinae) Zool Meded (Leiden) 58117-129

215 Soejitno J 1977 Reaction of some upland rice varieties toseedling fly eltherikonaexigua Stein [in Indonesian] Siri Hana I Penyakit 630-35 Lembga I osat Fenclilian Pertanian Bogor Indonesia

216 Socnardi 1967 Insect pests of rice in Indonesia Pages675-683 in The major inse pests of thu rice plant Proceedings of a symposium at the Internatital Rice Rec Wi itt S pternber 1964 The Johns Hopkins Press Baltimore Maryland USA

217 Soto P E A T Petcz and I W Buddenhagen 1976 Survey of insect pests and diseases of rice in different ecological zones in Nigeria Rice Entomol Ncwsl 435-46

218 Soto 11E and Z Ziddiqi 1976 Screening for resistance to African rice insect pests Varietal improsetnent sCmi n r Bouak Ivory Coast 13-17 Sept 1976 West Africa Rice l)evelopment Association Monrovia Iiberia

219 Squire F A 1972 Entomological problems in Bolivia PANS (PestArtic News Sumr) 18249-268

220 Stapely 1 II 1978 The rice leaf roller in the Solomon Islands Int Rice Res Newsl 3(l)13

221 Suenaga H 1937 On aphids attacking the roots of rice Oyo-I)oubts Zasshi 9150-152 [in Japanese] (Rev Appl Entomol A25745)

222 Suenaga H 1952 On the seasonal abundance of three subterranean aphids on upland rice Bull Kyushu Agric Exp Sti 1249-262

223 Sutynso Yas and Dandi Soekaria 1964 Beberapa hana yang ttenperang tana man padi tanah kering [in Indonesian] Rapat kerja Padi Tanah Kering 6 sd 8 Juily 1964 Kaliurang 9 p

224 Tallamy I) W and R F Denno 1983 Alternative life history patterns in risky environments in example from lace bugs Pages 129-147 in Insect life history patterns R F l)cnio and If Dingle eds Spring-Verlag 225 p

225 Tamura 1 1939 Observations on some insect pests of field crops [in Japanese] 1 Plant Prot 26414-420 (Rev Appl EntomolA27-556)

226 Tamura and Suzuki 1964 Influences of the early-transplanting practice in rice culture upon development of the rice stem borer Chilo siql)ressalisWalker Bull Ilokuriku Natil Agric Exp Stn 761-94

227 Tan J P 1924 The rice root aphid Drowhi ihirsoia Philipp Agric 13277-288

228 Tanaka 1 1957 Taxonomy and distribution of some subterranean aphids injurious to upland rice in Japan with description of a new species [in Japanese English summary] Botyu-Kagaku 22168-176

229 Tanaka T 1961 The rice root aphids their ecology and control Spec [full College of Agriculture Utsunomiya University Japan F3 p

230 Tauber M IC A Tauber R Nechols and 1Obrycki 1983 Seasonal activity of parasitoids control by external internal and genetic factors Pages 87-108 itt I)iapause and life cycle strategies in insects V K Brown and I Ilodek eds )r W Junk The Hague Netherlands

231 Taylor W F 1972 Effects of artificial defoliation (simulating pest damage) on varieties of upland rice Exp Agric 879-83

232 feotia T P S and S Natud 1966 Biotoomics of the rice skipper Parnaraniathiis Fabricius (Lepidoptera Hesperiidac) Indian J Entomol 28181-186

233 Teran F 0 1971 Rice pests of Santa Cruz [in Spanish Ministerio de Agricultura Sita Cruz Bolivia Bull 45 8 p

234 Than Itun 1976 Population dynamics of the yellow rice borer Tryporza itcertulas (Walker) and its damage to the rice plant MS thesis University of the Philippines at Ios Bafios Laguna Philippines 92 p

235 Uichanco L B 1911 Pests Pages 55-74 it Rice Philipp Agric 11-200

IRPS No 123 January 1987 41

236 van der Laan P A 195 1 Life history of the rice seedling fly Contrib Gen Agric Res Stn Boor Indonesia 118 15 p

237 van der Laan 11A 1959 Correlation between rainfall in the dry season and the occurrence of the white rice borer (Scirpolhaga innotala WIk) in lava Entomol Fxp Appl 212-20

238 Van Driesche R G 1983 Meaning of percent parasitismin studies of insect parasitism Fnviron Entomol 121 I1- 122

239 Venugopal M S and A litsingcr 1984 Fffect ofcarbofuran on rice growth Prot Ecol 7313-317

240 Vietnam Directorate of Rural Affairs 1962 Annual work progress report on crop improvement program of rice sugarcane vegetable and field crops (lor the period from July 1961 to June 1962) (11) 299 p (Rev Appl Entomol A51526-527)

241 Wan M T K 1972 Observations on rice leaf- and planthoppers in Sarawak (Malaysian Borneo) Malay Agric 148308-335

242 West Africa Rice I)esclopnient Association 1978 Annual research report Vol 2 Statistics Monrovia Liberia

243 West Africa Rice Development Association 1979 Annual research report p47 Monrovia Liberia

244 Williams ) F and CS lofgren 1981 Fli I illy l-468 anew bait toxicant for control of the red imported lire at Fla Entomol 64472477

245 Williams I) 1 1970 The mealybugs OFlomoptera Coccoidea Iscudococcidac) of sugar-cane rice and sorghum Bull Entomol Res 0109-188

246 Williams D) 1 1982 Pulinaria icervi (Signoret) and its allies ott sugar cane and other grasses Btll Entoinol Res 72111-117

247 Wirjosuiardjo S 1976 Study ofchenmical cont rot of insect pests ott upland rice Imiu Pertanian (Jogjakarta) 1323-329

248 Wirjosuhardjo S and N Wirarta 1976 Chemical control of the white grub tIolorithiatlltriin upland rice in M uneng Irobolinggo Indonesia) [in Indonesian] Imut Pertanian (Jogjakarta) 1330-332

249 Yano K T Miyake and V F Eastop 1983 The biology and economic importance of rice aphids (Henuiptera Aphid idae) a review Bull Entomol Res 73539-566

250 Yelp D and P Rhodes 19(4 lhe success of drilled dryland rice iii the Fiji Islands International Rice Commission Working Party on Rice Production and Protection Manila Philippines 3-10 March1964 Food and Agriculture Organization Rome Italy 13 p

251 Young W R 1981 Fifty-five years of research on the sorghum shootfly Insect Sci Appi 23-9

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IRPS No 123 January 1987 5

Table I continued

Order Family Species Distribution Reference

Locusta migratoria migratorioidesReische Africa 33 et Fairmaire

Schistocercagregaria (Forskal) Africa 33 Schistocerca americana (Drury) Latin America 82 195 Valanga nigricornis (Burmeister) Asia 82 Orphulella inticata Scudder Latin America 68

lomoptera Coccidae Pulvinaria sorghicola De Lotto Africa 246 Pulhinariaiceryi (Guerin) Latin America 186 246

Pseudococcidae taennococcus spp Asia 120 Lhorizococcus iu Williams Asia 245 Dysmicoccus boninsis (Kuwana)C Asia Africa 245

Latin AmericaDysmicoccus brevipes (Cockcrell)c Asia Africa 117 203245

Latin AmericaDysmicoccus oryzae (Wirjati)c Asia 245 Geococcus (=Ripersia) oryzae (Kuwana)C Asia 245Brevennia (= tererococcus)rehi (Lindinger) Asia 72 245Nipaccoccus graminis (Maskell) Africa 245 Planococcoides lingnani (Ferris) Asia 245 Pseudococcus saccharicola Takahashi Asia 245 Pseudococcus spp Latin America 60 Saccharicoccus sacchari (Cockerell) Africa Asia 245

Latin America Trionymus ceres Williamsc Asia 119245j rionymus sp Asia b Mealybug unspecified Asia 83 84

Aphididae Capitophorus[= Rhopalosiphumn] prunifoliae Asia 221 (Shinji)

Rhopalosiphum maidis (Fitch) Asia 249 Rhopalosiphum padi (Linnacus) Asia Africa 12 97 249 Rhopalosiphum rufiabdominalis (Sasaki) Asia Africa 66 76 120 195203

Latin America 221229240249 Tetraneura akinire Sasaki Asia 221249Tetraneurabasui ltillcRisLambers Asia 249Terraneura nigriabdominalis (Sasaki) Asia Africa 712 30 76 227229 (=Dryopeia hirsuta)

Tetraneuraradiciola Strand Asia 249 Paracleus ciniciformnis von Heyden Asia 249

[= Forda harukawai Tanaka]Pineus hanikawai (lnouye) Asia 228 Geoica hcifitga (Zehntner) Latin America 76 195 229 249

AsiaGeoica setulosa (Passerini) Asia 249 Anoecia fuliabdominalis (Sasaki) Asia 249 Anoecia comi (Fabricius) Asia 221229249Hysteroneura sctariae (Thomas) Africa Asia 7

Latin America Sipha glyceriae (Kaltenbach) Asia 249 Root aphid unspecified Asia 72 120

Cercopidae Deois schach (Fabricius) Latin America 61 68 194 Sepullia (=Denoplux) nigropunctata Stal Africa 29 Tomaspis ( Deois) fluxtuosa (Walker) Latin America 61 68 194 Tomaspis ( Dcois) flaiopicta (Stal) Latin America 61 61 194 To naspis (Dcois) complcta (Schmidt) Latin Aincrica 68 Tomaspis ( Zulia) entreriana (Berg) Latin America 68 Tomaspis ( Mahanarva) fimbriolata(Stal) Latin America 68 194 Tomaspis ( Aeneohna) spectabilis (Distant) Latin America 219 233 Aeneolamia varia Fciinai Latin America 81 111 Aeneolamia postica (Walker) Latin America 203 Abidama producta Walker Asia 54Delphacidae Sogatella furcifera (Horvath) Asia 59241 Niapanatalugens (Stal) Asia 30 59 72 84 241 Sogatodes oryzicola (Muir) Latin America 6167150Sogatodes cubanus (Crawford) Latin America 150 Sogatodes pusanus (Distant) Asia bPeregrinus maidis (Ashmead) Latin America 186 Laodelphax striatellus (Fallen) Asia 18

Cicadellidae Recilia dorsalis (Motschulsky) Asia 59 241 Nephotettiv nigropictus (Stal) Asia 59241

6 IRPS No 123 January 1987

Table I continued

Order Family

Hemiptera Lygaeidae

Cydnidae

Alydldae

Coreidae Corimelaenidae

Pentatomidae

Thysanoptera Thripidae

Species

Nephotettix virescens (Distant) Nephotettix modulatus Melichar Cofana spectra(Distant)Graphocephala spp Hortensiasimnilis (Walker) Exitianusobscurinervis (Stal)Balclutha spp Draeculacephalaclypeata Osborn Cicadulinabipunctella (Matsumura)Cymoninus turaensis(Paiva) Ninus insigniStal Pachybrachiusnenosus Horvath Caenoblissuspilosus Barber Blissus leucoptens (Say) Blissus spp Dlmnorphoptentscornutus novaeguineae GhauriDimorphopterus similis Slater Paroniuspiratoides (Costa)Pangaeus spp Aethus indicus (Westwood)Geotomnus pygmaeus (Dallas) Stibaropus inolginus Schiodte Cyrornenusbergi FroeschnerCyrtomnenus ciliatus (Palisot de Beauvois) Cyrtonenuscrassus Walker Tominetus spp

Riptortuslinearis (Fabricius) Riptorts spp Leptocorisaacuta (Thunberg)Leptocorisabiguttata (Walker)LeptocorisapalawanensisAhmad Leptocorisaoratorius(Fabricius)Leptocorisachinensis Dallas LeprocorisasolomonensisAhmad Leptocorisaspp Stenocorissouthiwoodi Ahmrad Stenocoris claviformis Ahmad MirperustorridusWestwood aetus trigonus (Thunberg)Alkindus atratusDistant ScaptocorisdivergensFroescher Scaptocoris castaneusPertyScotinopharatarsalis (Vollenhoven) Scotinopharascotti Horvath Eysarcoris (=Stollia) ventralis (Westwood) Dolycorisindicus Stal Macrinajuvenca Burmeister Tanda antiguensis (Westwood) Tantiageifi Schout Tantia perditor (Fabricius) Pygotnenidavaripennis (Westwood) Menida sppNezara viridula (Linnaeus)

Oebaluspoecilus (Dallas) Oebalusypsilon-gnseus (de Geer)Oebalusgrisescens (Sailer) Tibracea limbativentris (Stal)

Aspavia armigera(Fabricius)Diploysfissa Erichson Acrosternum marginatum (Palisot de Beauvois) Acrosternum sppFrankliniellarodeos Pergande Breginatothrips enustos Watson Stenchaetothrips(=Baliothrips)biformis

(Bagnall) (=(Jlloethripsoryzae (Williams)HaplothripspriesnerianusBagnall

Distribution

Asia Africa Asia Latin America Latin America Latin America Latin America Latin America Asia Asia Asia Asia Asia Latin America Asia Asia Africa Asia Latin America Asia Asia Asia Latin America Latin America Latin America Latin America

Asia Africa Asia Asia Asia Asia Asia Asia Asia Africa Africa Africa Asia Latin America Latin America Latin America Asia Asia Asia Asia Africa Latin America Africa Latin America Asia Asia Asia Africa

Latin America Latin America Latin America Latin America

Africa Africa Latin America Latin America Latin America Latin America Asia

Asia

Reference

59 241 217 59 241 6166 61 66 81 150 6166 66 81150 b 178 178 30 83 42 81150 203 30 80 62 178 195 178 178 178 203 203 203 203

55 178 12 72 78 120 214 214 b 59 120178114 178 54 636 82 12 6 12 82 178 81 150 60 is 195 59 120 178 78 36 186 36 186 b 55 12 30 54 55 78 82 120 61 66 81195 61 65 66 195 211 150 195 61 66 81 150 219 233 6 12 82 6 82 186 186 61 66 68 213235

178

7 IRPS No 123 January 1987

Table I continued

Order Family Species Distribution Reference

toleoptera Scarabacidae leteroligus (=Aphonoprocts) 15elcs Billberg Africa 36 Dynastinac (=Ileteroligusmees robustui Prell)

Alissonotum pauper Burnieiser Asia 82 Alissonotuon simile Arrow Asia 209 Dscinetusdubibs (Olivie) Latin America 28 195 233 Devscinetusgagates Burmncster Latin America 20 195 Dyscinctt spp Latin America 60 Eutheola Inimilis Bturmcistr Latin America 20 28 8 1 195 203

233Eutheola identarr(Biurnciter) Latin America 121 203 Heteron)ciwv aw rsonJack Africa 82 Hcreron)chusbtuterculatus Kolbe Africa 82 tleteronichtslicas (Klug) Africa 82 fhteron)chus mosamhicus Peringuey Africa 4 6 48 82 (=Veteronychus orzae Britton)

leteronychts lioderes Redtnbacher Asia 77 82 207 ltcteronchusarator (Fabricius) Africa 82 tleterontchusplebepus (Klug) A frica 57 82Ileteron chus pseuocongonnsis Ferreire Africa 32 Ileteronvchns rogiJrons Fairmaire Africa 82 Heteron chus nrsticus niger (K lug) Africa 82 Heteronvchus spp Africa 17 Lachnosterna longipennis (Blanchard) Asia 82 Lachnosternaspp Asia 120 Ligvros fossator Burmeister Latin America 82 Ligyrus nasutus Burmeister Latin America 203 Ligyrus (=Scarabaeus)ebenus (de Geer) Latin America 20 195

Dynastinac Ligtvros huonilis Burmeister Latin America 60 82 Aaladera castanea (Arrow) Asia 82 Maladera orientalis Motschulsky) Asia 82 Malaaerajaponica Motschulsky Asia 82 Phvlognathusdicynsius (Fabricius) Asia 82

Melolonthinae Exopholis h)ypoletca (Wiedemann) Asia 117 Lerlinosterna[= lolotrichia] consanguinea Asia 122

(Blanchard) Leucopholis rorida (Fabricius) Asia 82 117 Lepidiota blanchardi Dalla Torre Asia 76 82 Leucopholis irrorata (Chevrolat) Asia 76 134 Scrica internpta Walker Asia 76 Holotrichialonipennis (Blanchard) Asia 122 Holotrichia serrata Fabricius Asia 122 Holotrichia leucoph thabna (Wiedemann) Asia 82 Holotrichia seticollis Moser Asia 77 78 Holotrichia pnindanaoana Brenske Asia b Stenoeres spp Latin America 71 195 Atoenius spp Latin America Asia 203 Ph luplg aequata (Bates) Latin America 203 Piyvilophagacaraga Saylor Latin America 203 Phyllophagachiriquina (Bates) Latin America 203 Phyllophagadas)poda (Bates) Latin America 203 Phyllophagaclegans Saylor Latin Ameica 203 Phyllophagahondura Saylor Latin America 203 Phyllophagalatipes (Bates) Latin America 203 Phyllophagamontriesi (Blanchard) Latin America 203 Phyllophaga pamisetis (Bates) Latin America 203 Ph vllophaga sanfosicola Saylor Latin America 203 Phllophagasetifera (Burmeister) Latin America 203 Phyllophaga (=tlolotrichia)helleri Breuske Asia 82 117 159 Phvllophagaspp Latin America 81 150 White grub unspecified Asia 84

Rutclinae Lagochile trigona (Ilost) Latin America 82 Anomala dinidiata var barhata Burmeister Asia 77 78 82 Anomala huteralis(Burmeister) Asia 76 Anoimala lurida (Blanchard) Asia 76 Anomala sulcatula (Burmeister) Asia 76 Anontala varians (Olivier) Asia 82 Anomala antigua (Gyllenhal) Asia 82 Anoinala palida (Fabricius) Asia 82

8 IRPS No 123 January 1987

Table 1 continued

Order Family

Tenebrionidae

Elateridae

Languriidae Chrysomelidae

Chrysomelidae

Coccinellidae Curculionidae

Lepidoptera Lymantriidae

Amatidae

Arctiidae

Limacodidae Eupterotidac Noctuidac

Species

Anomaa volita (Blanchard) Adoretus caliginosus Burmeister Adoretus comprcssus (Weber) Papuana huhneri (Fairmaire) Papuana incrmnis Prell Popillia capricollis Hlope Gonocephalu depressur (Fabricius)Gonocephahm acuiangulum (Fairmaire)Gonocephalum sinphx (Fabricius) Epitragus sallei Champion Anaedus ptnctatissimnts ChampionAeolus spp4griotes spp Agriotes mancus (Say) Conoderts spp Aeoloderma brachmana (Candeze)Wireworni unspecified Anadastus filijf rmis (Fabricius)Dicladispa viridicanca(Kraat z)Trichispasericea (Guerin)Oediopalpastervalis (Weise) Oediopalpa guerini (Italy) Chaetocnema cvlindrica (Baly)Chactocnema denticulata S tephens Chaetocnemabasalis (Baly)Aulacophora sp nr similis Olivier Monolepta caripemne Ialy Monolepta bilasciata (Ilornstedt) Monolepta signata Olivier Cerotoma airojcsciata J acoby Diabrotica adelpha Ilarold Diabrotica balteata Leconte Diabrotica graminLa ltaly Diabrotica speciosa (Germar) Diabrotica limitafa quindecinmunctata(Germar)Diabrotica melanocephala (Fabricius)Diabrotica spp Altica (= Fondia)madagascariensis(Allard)Altica spp Epitrix cuctmeris (Harris) Epitrix spp Oulenjo or pae(Kuwayama) Disonchq (= Donacia) spp Flea beetle unspecified Chnootriba (= htIilachna)sinilis assimilis Mulsant Atactogaster iduriens Walker Hypomeces squtamosus (Fabricius) Tan vinecus discoidalis GyllenhalPhaulosoinusmusculinus (= mnus) Csiki Neobaridia anplitarsis CaseyPsalis pennatula (Fabricius) Laeliasufihsa (Walker) Euproctis irgtncula (Walker)Euproctis minor (Snellen)Euproctis xanthorrhoea (Kollar)Amata sp

Diacrisia obliqua (Walker) Creatonotusgangis Linnacus Latoia (= Parasa) bicolor (Walker)Nisaga simplex Walker Sesamia botanephaga Tams amp Bowden Sesamnia nonagroides botanephagaTfums et Bowden Sesamia calamistis llampson Sesamia inferens (Walker) Spodoptera eridania (Cramer)Spodoptera mauritia acron)ctoides Guenee Spodoptera frugiperda ( ESmith)

Distribution Reference

Asia 82 Asia 82 Asia 82 Asia 82 Asia 82 Asia 78 Asia 82 117 178 Asia 82 117 Africa 82 Latin America 203 Latin America 203 Latin America 195 Latin America 195 Latin America 203 Latin America 195 Asia h Asia Latin America 60 73 117 Asia 200 Africa 36 Africa II Latin America 6166 68 Latin America 6166 68 Asia 225 Latin America 61 66 68 81

Asia 178 Asia 55 Asia 55 Asia h Asia 55 Latin America 203 Latin Amefia 203 Latin America 203 Latin America 203 Latin America 6166 68 Latin Amcrie 195 Latin Amw 195 Latin Aierwca 61 81 Africa 57 Asia 178 Latin Americ 203 Latin America 81 Asia 82 Latin America 195 Asia 120 Africa 6 11 Asia 185 Asia 120 Africa 82 Africa 82 Latin America 66 68 195 233 Asia 117 Asia 117 Asia 117 Asia 117 Asia 117 Asia A S Pradhan

unpubl dataAsia 82 Asia 82 Asia 82 Asia 182 Africa 12 Africa 6 36 Africa 6 8 11 12 36 Asia 59 82 84240 Latin America 150 Asia -0 59 196 Latin America 28 61 66 68 81 150

152195203219233

9 IRPS No 123 January 1987

Table I continued

Order Family Species Distribution Reference

Spodoptera exigua (Hubner) Spodoptera exem pta (Walker) Spodoptera ornithogalli (Guenee)

Africa Africa Latin America

32 217 32 33 36217 150

Mythimna loreyi (Duponchel) Asia 30 Mythimna separata (Walker) Mythimna (- Pseudaletia)latifascia [= adultera

Asia Latin America

178 195

(Schaus)] (Walker)Mythimna (=Pseudaletia) sequax (Fabriclus) Latin America 68 Mythimna rosellinea (Walker) Asia b Mythimna yu (Guenee) Asia b Mocisfrugalis (Fabricius) Asia 30 117 Mocis latipes (Guenee) Latin America 66 68 149 150 203

Platysenta (=Spodoptera) compta (Walker) Asia 219233A Barrion unpubl data

Agrotis ipsilon (Hufnagel) Latin America Africa

33 68 81 150 1)5 203

Agrotis spp Latin America 60 Achaeajanata (Linnaeus) EldanasaccharinaWalker

Asia Africa

117 36

Armyworm unspecified Asia 72 84 Pyralidae Cnaphalocrocis medinalis (Guenee)

Marasmia bilinealisHampson Asia Africa

30 4559 72 84233 36

Marasmia trapezalis(Guenee) Africa 32 217 Marasmia (=Sumia) exigua (Butler) Asia 220 Marasnianmralis (Walker) Asia b Marasiniapatnalis Bradley Marasiniaspp

Asia Asia

b b

Leaf roller unspecified M2liarphaseparatellaRagont

Asia Africa

72 12 57

Maliarphasp Asia b Chilo auricilius Dudgeon Asia b Chilo partellus (Swinhoe) Chilo polychrysus (Meyrick)

Africa Asia

12 92 59 72

Qhilo diffusilineus J de Joannis Africa 6 1229 Chilosuppressalis(Walker)Cibozacconius Blezzinski

Asia Africa

59 72 6 11 12

Cravubus spp Latin America 203 Acigona loftini (Dyar) Latin America 203 Acigona chrysographella(Kollar)Scirpophagaincertulas(Walker)

Asia Asia

82 59 72

Scirpophagainnotata (Walker) Asia 59 200 Scirpophaganiuella (Fabricius) Asia b Elasinopalpuslignosellus (Zeller) Latin America 28 60 61 66 81 150

Diatraeasaccharalis(Fabricius) Latin America 162195203219233 28 66 81 111 150 195

Rupela albincllaCramer Latin America 111

Gelechlidae

Stem borer unspecified Undetermined stem borer nr Maliarpha BrachniaarctraeaMayr

Asia Asia Asia

72 b 55

Brachmi spp Africa 32 Hesperiidae Parnaraguttata Bremer et Grey Asia b

Parnaranaso Fabricius Asia 55 Pelopidasagnaagna (Moore) Pelopidasmathias (Fabricius)

Asia Asia Africa

164 36 155217

Pelopidasconjuncta conjuncta (Herrich-Schaffer) Asia b Borbo fanta Evans Africa 6

Satyridac Melanitisleda ismene Cramer Asia 84213 Mycalesis asophisHew Asia 30

Diptera Cecldomyiidae Mycalesis spp Orseoliaoryzae (Wood-Masn)

Asia Asia

b 72 88 131 181

Orseoliaoryzivora Harris et Gagne Africa 61132218 Chloropidae Steleocerellus (=Mepachymens) ensifer (Thomson)

Oscinella spp Asia Asia

205 205

Gaurax sppChloropsoryzae Matsuntura (= kuwonae Aldrich)

Asia Asia

205 82

Mucldae Atherigonaoryzae Malloch Asia 30 82 117130 Atherigona exigua Stein Asia 82 117 178

10 IRPS No 123 January 1987

Table I continued Order Family

Diopsidae

Agronyzidae

lHymenoptera Formicidae

Species

Atherigona oritentalisSchiner Atherigona indica MallochDiopsis longicornis(= thoracica)Macqurt Diopsis spp Pseudonapnyza asiatica Spencer Pseudonapomyza spicata (Malloch) Agronoza orvzae MunakataAcromyren landoltibalzani(Emery) Acromyrmex landoltifracticornis(Forel)Acromyrnex heyeri Forel)Atta bisphaericaFore Atta capiguaraGoncalves Atta laeLigata (F Smi h1) Atta opacicepsBorgmeierSolenopsisgeminata (abricius)Pheidole sp Pheidolegeton diversus Jerdon Monomorium pharaonis (Linnaeus) Ants unspecified

Distribution Reference

Asia 82 Asia Africa 82205Africa 11 1248 Africa 217 Asia 213 Asia h Asia bLatin America 6168 195 Latin America 61 195 Latin America 61 68 195 Latin America 61 68 195 Latin America 14 61 68 195Latin America 61 68 195 Latin America 61 195 Asia b Asia b Asia 1) Asia b Asia 120

aG orlentalls formerly recorded in Asia as G africana thPhilippines reported in this study CSubterranean root feeders

subterranean nests Foraging ants will feed on germinatedseeds only when the supply of ungerminated seed is low However these ant specialists feed on a wide varietyplants and thus help control grassy weeds (39) This dual

of

pest benefit role extends from tending ants which protecthoneydew-excreting pest species including root aphids and mealybugs (workers repel potential predators and even selectively kill parasitized aphids) to predators of leaf- and stem-feeding rice pests such as leaffolders armywormswhite grubs and stem borers (1 17)

The upland rice environment is particularly well suited to ants because of the lack of flooding Frequent tillage perpetuates grasses and granivorous species that are highlyadapted to constructing new nests in recently tilled fieldsAnt nests however are relatively shallow and tilling with a moldboard plow can destroy them Also upland ricefields are uually small and isolated and a given area in a smallfield has more field borders than it has in a larger fieldallowing high rates of infiltration from more sedentaryspecies such as Plieidole that do not colonize tilled fields but nest in soil under shrubs and trees along field bordersPheidole workers forage into fields from borders and store seed in their nests In slash-and-burn or other no-tillage rice cultures nests would be preserved in the field and possiblylead to high rates of seed predation However granivorous species do not prefer forest habitats

Damage is characterized by reduced usually patchy plantstand (120) Seed-thieving ants greatly prefer rice but they can be selective regarding plants they collect and store Theyharvest more seed than they can consume so patches of germip ing surplus seed (such as Rottboellia) will sproutfrom their nests during the rainy season

In Latin America leaf-cutting ants (Aita spp and Acromyrmex spp) occur only in upland fields and defoliate

young rice plants (47 68) They take leaves to fungal gardensin underground nests where fungi predigest the plant material

Termites Subterranean termites of the family Termitidae which lack symbiotic protozoa to help digest plants are the most frequently mentioned termite pests of upland rice (1185 120) This family cultures fungi in special undergroundcells --- fungal combs made of half-digested plant material (117) The fungi inoculated onto the combs break down plant material into food that termits can digest The termites then consume the fungal combs These grasslandtermites build nests below the plow layer in uplandricefields The colonies attack living plants only when dead plant material is not available They will attack a droughtshystressed crop but prefer older plants having greater cellulose content They will damage newly planted crops where clean culture has removed vegetation

Infestations are worse in deep light-textured soils with low moisture content The first sign of damage is yellowingof older leaves (195) Termites feed on roots (the plantyellows then wilts and finally collapses) and germinatingseed (loss of stand) or move above ground at night to cutseedlings at ground level which they cover with soil for later consumption Soft-bodied termites are highly sensitive to desiccation They live within a self-contained system of airtight chambers and build surface tunnels linei with nudand body secretions (carton) to maintain more than 90 relative humidity Termites can be located by their tunnels

Termites apparently can withstand limited submergence (157) and can be a pest of dry lowland seedbeds (117)perhaps by having nests below perched water tables

Termites appear to do more damage in Latin America where large ricefields are plowed with tractors deprivingtermites of preferred vegetation and forcing them to live on

rice plants In Asia upland ricefields are typically small with Iprge field border areas (ecotones) where termites can forage for preferred food other than growing rice plants All grassland termites do not feed on rice In an upland area in Batangas Province Philippines ricefields are commonly infested by Hosiititernesluzonensis (Oshima) however this termite does not damage rice In the same province though Mrroternesgivus(Hagen)feeds on rice seedlings (178)

In Africa termites may damage rice even though fields are small and surrounded by perennial vegetation These termite species perhaps prefer rice over other hosts

Mole crickets Nymphs and adults are nocturnal and burrow through soil feeding on roots of a wide variety of plants or forage above ground as predators of insects Adults live 3-5 mo and are even cannibalistic Mole crickets prefer low-lyin moist upland soils with high organic matter (II 81) In the lowlands they inhabit rice bunds or nonflooded fields Adults are highly mobile and can leave a flooded field to locate a more suitable habitat Losses show as wilted plants (from root feeding) or reduced stands (120) Damage is more common near field borders where mole crickets relocate after tillage (126)

Fiehl crickets Nymphs and adults of field crickets have similar nocturnal habits and damage rice as much as mole crickets do Piles of weeds removed from fields attract them They make subterranean nests and tunnel through the soil to feed on roots Some species prefer seed to roots Others feed at the base of stems causing deadhearts

Tunnel entrances surrounded by excavated soil are easily seen Young plants are cut at ground level and stored in underground cell Field crickets tre highly polyphagous as are all soil pests (117) Sachan et al (198) reported that field crickets prefer maize to upland rice Like mole crickets field crickets cannot survive in standing water and are therefore more prevalent in upland fields

Whitegrubsandhackbeetles Scarab beetles that feed on living roots as larvae but not as adults are called white grubs or chafers (Melolonthinae Rutellinae) and those that feed as adults but not as larvae on plant crowns are called black beetles (Dynastinac) In the tropics chafers or white grubs have a I-yr life cycle starting with adults emerging from the soil about I mo after the first downpour of the rainy season (134 159) They lay eggs at the same time farmers sow upland rice The rice passes its most susceptible stage and damage is mostly avoided when the white grubs are small After several months the long-lived larvae are large enough for two or three to denude the root system of mature rice (223) This intensity of damage is rare but wilting occurs when root loss is combined with drought stress

The larvae need damp (not saturated or dry) soil to survive and pass the unfavorable dry season 05-2 m underground in aestivation in the moist soil The first heavy rains of the season (20-30 amd) stimulate the grubs to resume activity and after several weeks they develop into

IRPS No 123 January 1987 11

pupae and adults and eventually dig their way to the soil surface and fly to nearby trees to seek food and mates The adults of most white grub species feed only on trees but in some species notably Eutheolahumilis the adults defoliate and cut off stems at ground level (28) Putheola hidentata damages sugarcane fields in Venezuela (121) Floods kill white grubs (208) Grasslands can support large populations and therefore white grubs can be more abundant in newly planted upland ricefields that were previously fallow (81)

Black beetles live up to a year adults can attack a crop at any stage by burrowing into the soil (127) and feeding at the base of stems to cause whiteheads (207) Adults emerge with the early rains and are normally more abundant and do more damage on a young rice crop with small root systems (38 81)

The larvae feed only on dead organic matter in dryland fields and do not attack rice Adults arc highly mobile and although sensitive to flooding can invade a field soon after it drains (115)

Root aplhids Several genera of migratory aphids feed on the roots of upland rice (249) Populations build up more in light-textured soils with high percolation rates Damage has also been reported from dry seed heds of lowland rice (52) Tending ants -- Pheidhle Cren aster Tetramoriuw lsius Tapinoma -- are necessary for root aphids to multiply Aiits harbor aphids in their nests over winter or during unfavorable periods and relocate them on rice plants digging tunnels along the root systems to allow these softshybodied insects to penetrate the soil Most intensive studies on aphid life histories have been done in temperate environments (158 222 229) Root aphids fly to rice plants at the beginning of the rice season and pass through several generations continually developing winged forms to relocate on alternate grassy hosts until winter when they move to perennial hosts usually trees Yield loss occurs mainly through reduced tillering but infested plants become yellow and stunted and in extreme cases wilt (227) Aphid populations build up gradually so damage usually begins in the late vegetative and reproductive growth stages

Root-feeding ineal tugs More than half of the mealybug species or rice feed on roots (245) Being soft bodied mealybugs are not adapted to living underground but they survive because tending ants (the same species that tend root aphids) dig burrows for them and move them from plant to plant Well-drained soils ihelp mcalybugs survive Although six species are recorded as subterranean root pests of rice we found no report citing damage Root-feeding mealybugs are probably more prevalent on perennial grasses than on short-lived annual rice

The most common rice mealyb ig Brevennia rehi is a foliar feeder favored by lack ol rain apparently it is responsible for high yield loss in Northeast India and Bangladesh (N Panda pers comm as stated by IRRI [1061) This report needs confirmation Mealybug damage was only briefly mentioned in another report (156)

12 IRPS No 123 January 1987

Root weevils A few species of upland weevils (IHporneces Donacia Atactogaster) attack rice roots as larvae and feed on stems and leaves as adults Damage ispatchy The larvae are not adapted to submergence and are true upland soil weevils isopposed to the so-called water weevils They rarely become sufficiently abundant to cause damage Adults on rice foliage or the soil surface show that the roots are being attacked by the grubs Root weevils are highly polyphagous (185)

Root heeths Soil-inhabiting false wireworms Gono-cephalumt spp are reported in upland ricefields The adults and larvae normally feed on decaying organic matter but land preparation for upland rice removes these sources The larvae feed on seedling roots and nocturnal adults roam on the soil surface to cut off seedlings at ground level (82) This group of ground beetles is highly omnivorous adults even prey on stem borer larvae (183) True wireworms (Elate-ridae) feed on living roots but rice as a host has been reported Infrequently (117) In Latin America the highlypclyphagous larvae of Diabrotica rootworms are also recorded on rice

Root htus Upland rice seedlings can be attacked by the adults anI nlViphs of chinch bugs Blissusand Caenoblissus and bro n bug Scaptocoris which feed on the roots with theirsucking nouthparts (80 83) Three genera of burrower bugs A tms (eotonhus and Stiharopus -are common iil grasslands in the Philippines and may attack the first crop (rice or maize) planted after the dry season Chinch bugs are most abundant after a dry spell These insects are large and can kill seedlings or stunt and reduce tillering on older plants I hey prefer other hosts and rarely damage rice Adults can also feed oii developing rice grains (81)

(utworms Soil-dwelling noctuid larvae hide under-ground during the day to avoid predation by birds and bccone active at night above grour cutting off you ing plants at ground level (hence tile naiie cvtworn) arvaedrag severed plants into hurrows llie larvae have wide host ranges arid pupatc in the soillgrotis ipsilon occurs worldwide and appears to be the most common cutworm pest of upland environments but is more prevalent inlowland rice particularly in dry seedbeds

IFoliar pests Other groups of insects are adapted to upland environments because they pass one growth stage underground either as egg (in heinrietabolous species) or pupa (in holomnetabolous groups)

Rice seedlibig maggots AitJeriota flies occur in non-fhoodcd environments in Asia and Africa (180) Feeding of larvae one per iller causes deadhearts Flies as opposed to moths only secondarily evolved as pliytophagous pestsfroni a saprophagous Origin Athrnonalarvae secondarihv feed onidecaying tillers whereas larvae of stem borer moths actively Iced on living tillers to cause deadlicarts Seedling maggots will die if they cannot sever a tiller on which to feed

as it decays Plants die under heavy attack whereas lighter infestations cause stunting delayed maturity and ragged leaves (236)

Eggs are laid only on actively growing plants (49) aid damage occurs wiihin the first week after crop emergence(170) Larvae pupate more in stems than in soil Seedling maggot attack is highly seasonal (206) normally peaking 2-3 mo after the onset of the rainy season These pests possiblyaestivatc over the dry season Rice isonly one of their many graminaceous crop and weed hosts

The rice stem maggot (hlorops orzae is adapted to temperate upland and lowland rice growing regions of Asia In Japan and China upland rice grows in mountainous regions a habitat of this chloropid fly The larvae tunnel into tilestems The stem maggot is more damaging during the seedling stage larvac hibernate in grasses

File several dipterans reported as stem maggots in dryshyseeded lowland seedlings inIndia (205) could presumably attack upland rio

Leaf urinrs Icaf miners are common on wheat and maize but are rarely reported on upland rice (213) Three leaf miners are more prevalent on upland than lowland rice inthe Philippines Pvudonalomy a asicanh Vficataand lIAgromrza orzae attack at the seedling stage The larvae develop inside the tunnels they construct in the parenchyma tissue P Vpicata and Aorvzac prefer maizeEehtsine and other grasses to rice spicatacan become abundant on wheat

Leaf beetles Chrysomelid and coccinellid eaf bcetles scrape or otherwise remove leaf tissue as larvae or adults or both This damage accelerates desiccation of plants Tile larvae of sonic species are root feeders Flea beetles Epitrix and (lraetocnetna and rootworms l)iahroticaspp feed on rice foliage only as adults Their root-feeding larvae have hosts other than rice The adults and larvae of the hispa-like Oediolmlpa spp beetles of Brazil and (hmootriha (=EPfishyachra) beetle of Africa feed on upland rice leaves Oealioshypapa spp larvae are leaf miners (mootriha is also pshyvalent in lowland environments (11) Leaf beetles prefer vegetative rice (81)

The rice hispa Dicladipaarnigera is not reported on upland rice and prcers more aquatic habitats In Central Africa however I)cladispm viridicyaiea attacks uplandrice in the vegetative stage as well as lowland seedbeds 7richipasericea is reported on upland rice in West Africa and Madagascar (57)

The rice leaf beetle 1d1ea 01izae riiuech like Chioropsor|rzae is adapted to temperate regions of Ashi and is usually a pest on lowland rice but seldom oi inpland rice in the mountains (82) The adult hibernates on grasses

Armmworms Species of grassland-adapted SpoptloeraPlatysemta and A ithimna larvae create widespread epishydemics They can defoliate ricefields generally in patchesfrom early vegetative growth to harvest (41) Armywornms pupate inthe soil and highly favor upland rice (34) Adults

can disperse long distances to colonize even remote upland areas They become more abundant than their natural enemies particularly after favorable rains following a prolonged drought (196) The drought kills the armyworm and natural enemies (mainly parasites) but the armyworm can recolonize rapidly

The natural increase insoil fertility from mineralization of soil N over the drought period promotes luxuriant plant growth to foster a rapid buildup of these highly fecund species Weeding rice increases tilelikelihood of damage The parasites normally return to control the armyworms but not until after serious damage

Outbreaks on upland rice have been reported in Ghana (3) inzibar (38) Central Africa (36) lPanama (165) Brazil (28) Malaysia (196) and India (167) Defoliation may be severe often leaving only the base of stems Armyworm larvae hide during the day under litter or in soil cracks At night they ascend plants to feed Larvae pupate in the soil

Thrips Thrips frequent upland rice but damage is less than in the lowlands Thrips larvae and adults feed on leaf blades by rasping causing leaf rolling and stippling Thrips prefer vegetative stage rice and their survival increases during drought (heavy rain washes them from foliage) although drought stress usually outweighs thrips damage Drought and thrips cause leaf rolling in upland rice (235) but thrips have been overlooked perhaps becaLse of their small size Thrips are most numerous indry season irrigated rice The combination of favorable plant growth and lack of rainfall is ideal for thrips In contrast the lack of rainfall in upland rice hurts the crop and eventually the thrips whose numbers must decline

In [atin America Frankinilla rodeos attacks panicles before they emerge from the boot calusing sterile grains

Stem hug~s Black bugs are the most common group of rice stem-feeding -lemiptera in Asia The most frequently mentioned species ----Scotinophara coarctaia and Shrida - prefer aquatic habitats and therefore could not occur in upland rice as sometimes reported Upland species in the same gerurs - notably S tarsalis and S scotti -- are morphologically similar and remove sap from stems of grasses and upland rice The upland species are notably more abundant in upland rice planted near forests

In Latin America tile Tibraca ima-large pentatomid tiventris causes deadhearts and whitehcads on upland rice Both adults and nymphs remove sap from internodes of plants more than 3 wk old Tillers are killed by mechanical damage and by the entry of secondary microbial infection This damage is often confused with that of lepidopterous stem borers (233) Eggs are laid on the leaves and stems Nymphs are gregarious This species can also be found on irrigated rice Drought reduces population buildup

Stiem borers Lepidopterous and dipterous stem borers are widely recorded on upland rice but cultural practices and alternate hosts determine abtndance Except for Elasmolmll)uS Iinosellts numbers usually build up toward

IRPS No 123 January 1987 13

the end of the crop growth cycle Therefore late-maturing varieties staggered planting and lack offloodingfavorstem borer buildup Many stem borer species can lie dormant in the stUbbleafter harvest and theirsurvival is encouraged by panicle harvesting (leaving tall stems) and lack of tillage after harvest

Stem borers disperse and lay many eggs Many species are polyphagous however the monophagous MaliarpiasepashyratellaDiopsis Ivmgicornis Scirpophaga incertlas and S innotata can also be abundant on upland rice (hilo Suplressalisdevelops fasterand becomes la rger when reared on upland than on lowland rice (226)

The lesser corn stalk borer lasmopal tt Ittoselhs is semisubterranean and is perhaps the stern borer most adapted to upland rice(129) Highly polyphagous it infests maize peanut and cowpea It attacks seedling rice (one larva can kill tip to four plants before maximum tillering) (195) (hiv and Maliarphastem borers prefer older plants But Scirpophagaspp will attack young plants even in the nursery lasn opalls larvae tunnel into the stems at or below the soil level mausing dcadhcarts Larvae are not found in their turnnels inside sterns beca use vhei disturbed they retreat into cases made from soil particles bound by silk The cases areattached to the tunnel entrance (143) This behavior probably evolved as protection from natural enemies Larvae pupate in the soil The lesser corn stalk borer is more abundant during drought damage is often confused with that of drought (233) This species prefers sandy soils arid its distribution within a field may relate to soil texture (143)

Diatraeasaccharalis aside from the lesser corn stalk borer is frequently a stem borer pest in Latin America According to Teran (233) rice and maize are more preferred hosts than sugarcaine its namesake

Atcitonacrvsograpla is a less known upland rice stem borer in Asia with habits similar to Easinopall s (82) because it is not found in lowlands Acutra has grassy weed alternate hosts As a moth it is often confused with the dark-headed stern borer(hi polur(standstriped stern borer (hi sup)lpsrcais

Other common upland rice stem borers are also reported in the lowlands The dominant species in upland rice vary greatly with location In Uttar lPradesh India (78) and in Japan (123) the pink stem borer Sesamia htfirens is more prevalent hIKenya the principal rice stem borer is Chi partelus

(92) The lowland African sten borers Aialiarpha sepashyratellaDiopsis vngicornis and Sesamia calatnisnis are less abundant Oimupland than on lowland rice(217) but perhaps cause greater yield loss because plant injury is greater if combined with drought css (92) The stalk-eyed stern borer D)ivpsis is more prevalent in wet habitats (6 1I217)

In Central and West Africa Maliarpha and Sesamia calanistis are the dominant stern borers of upland rice (8 9 10 II) (lilo dihlfilineui and Chio partellus inhabit the

14 IRPS No 123 January 1987

upland savannas whereas Chilo zacconio prefers the lowlands (II) Aaliarpha is most abundant in the mangrove swamp habitats (91) coastal regions (16) or rain forest zones (II) of Africa and appears to be the homologue ofScirlpophaga incertulas The yellow stem borr can also be prevalent in upland rice (120 167) The sugarcane borer Eldanasaccharinais so far only a potential pest of uplandrice (91) despite Grist and evers (82) warning

Grasshotl)ers andI locitas Short-horned grasshoppers and locusts -- Locvta Patanrga Schistocerca - layeggs in the soil inhabit grasslands and develop into swarms to seriously damage upland rice from time to time (173) Largenumbers can destroy a field leaving only the stubble In low cmuoroer s giassnoppeirs an cut panicles (173) Farmers may not plant if locust swarms are imminent Prolonged dryweather followed by favorable rain favors development of large swarms (173) Usually breeding in di) grasslands theydisperse toattack crops including upland rice(191) Eggsaresusceptible to desiccation d uring drough t and toinundation during seasonal rains With favorable soil moisture over consecutive seasons natural enemies ca nnot prevent a largepopulation increase leading to migratory swarms (64)

Most grasshoppers on rice are adapted to uplands (107)but lowland rice grasshoppers iervgl(rlhusspp and Ora spp occur occasionally on upland rice (2 82) They lay eggson rice foliage Nymphs are scmiaquatic

The gryllid Lsctrtus coucinnus is a pest of lowland and deep water rice in Bangladesh and Thailand In the Philippines it damages upland rice grown in isolated pockets near lakes The gryllid nymphs and ad ults feed on the central portions of folrage leasing only the midrib and leaf nargins

Leal ler Pvralid moths whose larvae fold leaves to make a feeding shelter occur worldwide (217) but arc appareritly most common in Asia Leaffolde rsattacking rice in Asia are a complex of species (26) (Ctalhahcrcisinedinalis the best-known rice Ia flolde r in Asia may prefer grassy weeds over rice (86) It is a late colonizer of uplandrice U1arasmja eviua a lowland species (220) nreflers rice to grass specres (86) and colonizes upland rice during il early growth stages llarasnria patnalis is the most corn-rnonly encountered species in upland rice in the Philippines Perennial grasses are their alternate hosts and they can survive year-round in upland rice environments The leaf-folders 11arasmia ruralisand Alarasmia spp are least aburndant Dormancy is unknown in leaffolders Cnaplha-ocrocis medhnalis however is known as a migrant (90)Leaffolders remove photosynthetic tissue attack during the flag leaf stage is particularly injurious Popurlations are normally held in check by natural enemies but leaffolders become parlicularly abundant in conditions of high plantfertility and shade

Butterflies Skippers PelopidasInathiasand Borhofanta are more prevalent in African lowland than in uplandhabitats (6) but extensive studies in Japan and China

showed that Pelopidas mathias prefers an upland habitat (155) Parnaragutata seasonally migrates between lowland and upland areas and is equally abundant in both environshyments (164)

The green horned caterpillar Melanitis leda ismene is more abundant in lowlands (212) but our experience in the Philippines suggests that Pelopidas niathiasand 11elanifis leda ismene are often more abundant in uplands than in lowlands Bamboo iaunbuiva spp annually sustains P mathiaspopulations in the Philippine uplands Adults feed on nectar from flowers and migrate to upland rice during the late vegetative stage

iPi erlled h i-1 1d dill L tIC i 7-0 l(become numerous probably because of high egg predationEgg and larval parasitization are often low Butterflies can readily disperse long distances to seek renlote uplandricefields However they have low biotic potential and even it predation rates are low the rarely become abundant Upland environments with their diverse microhabitats can provide more favorable sites than lowland rice plains

tutterfly pests of rice have broad host ranges and become dormant Juring urnfavorable times of the year

IolhagrotslaidopwraThe larvae of many polyshyphagous moths are reported to defoliate upland rice tussock moths lsaispelnnatula Iawlia sUifita luproctisirgtwltaE mnorand P vathorrhoeaslug caterpillar

litotiahicolorhairy caterpillars Nisaga sinp vandA mata spp and wooly bears l)iacrisiaohliqtia and Crvatonotts gangis Many of these polyphgous larvac migrate and are transients in rice including HelhWhis arm iera and 1t zea highly pol phagous species (82)

Gall nidges The Asian rice gall midge Orseoliaorrzae (88 131) and its African counterpart 0 ortzivora(6 11 12 32217) prelerlowland to upland rice Gall midges in rainfed areas are prevalent only during tle rainy season and dispersepoorly Their alternate hosts (wild rices LeersiaIschaetnuan and Pasahmt)are mostlyaquatic grasses and are less likelyto be near upland rice areas In an area of mixed uplandfloating and rainfed lowland rice cultures in northern Thailand upland rice which is ilanted a month earlier was attacked firstbut subsequcent populations were much lower than in rainfed lowla id or floating rice Low numbers of gallmidges passed the dry season orr perennial weeds such as Paspahim (88)

Smot (hid(s Reports of aphids attacking upland rice plants are few (249) Without maize the maize leaf aphidRho alosihhunm maidis attacks upland rice in the Philipshypines The plum peach aphid IIrsteroneuraselariae was reported in Sierra Leerne toinfest leavesand tnriperied grain(7) Siphaglrctriaecaused transverse linrea r necrotic striae iir rice leaves in Italy (172) I-lea v rains held populations in check Rice is not a preferred host for shoot aphids

Leafhoper andI )anthol)pers Cicadellids and delshyphacidsdispersecasilyand readilycolonie upland ricefrom nearby lowland areas A study in Sarawak (241) showed

IRPS No 123 January 1987 15

similar species in upland and lowland rice Nephoiettix The principal vectors of rice virus diseases generally have species dominated The most prevalent species accounting narrow host ranges and alternate hosts in the grass family for 90-95 of the sweep net samples wre N virescens N are mostly annuals dying in the dr3 season Some virus umropictus Nilaparvia hgens Soatellafiircifera Cfana vectors such as BPH can disperse lontg distances but few do

sectraand Recilia dorsalisLowlands had higher popula- Th restriction of upland rice and weed hosts to the rainy tions Katanyukul and Chandaratat (120) monitored hopper season the small area planted to upland rice and the hopper numbers in upland and lowland fields of northern Thaiiand vectors having narrow host ranges all work against virus and found similar results A comparative study of plant- diseases in upland rice hoppers in Fiji showed more brown planthoppe BPH) and In Latin America Sogctdhs o(rzicola and S cuhanus whitebacked planthopper (WBPH) in wetland rice than in transmit hoja blanca virus disease but reports on upland lowland rice and hopperburn was prevalent in lowland rice are rare except in the favored upland areas of Colombia areas with standing water (89) Peru and Venezuela (35 42) S rricolaocurs in Brazil

Hopperburn in upland ricefields in Asia is rare (84) In on upland rice but hojaI blanca has not been reported (67) Batangas Philippines isolated fields of upland rice become S orzicola has a wide host ramgc (10) similar to WBPI- in hopperburned infrequently from WBPH A report from Fiji Asia WBPH however is not known to transmit any virus cites hopperburn from BPH and WBPH (250) In these two disease situations upland rice was grown on highly fertic soil and In Africa pale yellow mosaic virus is endemic to swampy N fertilizer was applied Hoppers respond to better nutrition lowland areas where rice ratoons year-round (19) and is Normally upland rice is grown on poor soil and receives no mechanically vectored mainly by hispid beetles A minor fertilizer vector a flea beetle Chaeiocnema spp is an upland rice

In Asia upland rice ras are normally not far from pest and upland ricetields near endemic areas occasionally lowland ricefields-- a source of hoppers Otherwise hopper contract the virus species with greater polyphagy than BPH or Nelhotettiv Spittle bugs Cercopid nymphs producc a protective virescens would be dominant frothy covering that looks like saliva In Latin America

In Africa leathoppers and planthoppers do not become spittle bugs damage upland rice In parts of Brazil where abundant on upland rice (6 II) probably because lowland upland ricefields are surrounded by pastures adults rice is not widely rlanted nearby The only reported migrating from grasses can damage rice severely and cause hopperburn as from Nilapanrata maeander in breeder complete hopperburn in a young rice crop within a week of plots receiving high N rates in Nigeria (98) Hoppers colonization (194 195) Dispersal to rice occurs even if however are a greter problem in Latin America where pastures are verdant and spittle bugs prefer pasture grasses large grassland areas breed polyphagous species (Gra- to rice (168) Nymphs are rarely abundant enough to phocephala spp Hortensia spp lEvitianus ohscurunervis damage rice Regular rainfall favors their development Balclutha spp and Draeculacephala spp) vhich then (233) disperse to cause hopperburn on seedling rice (68) Rosetto et al (195) reported Graphocep7hala spp and Hortensia spp Seed pests to be equally abundant on upland and lowland rice Alydid and pentatomid bugs are seed pests of upland rice

The leafliopper and planthopper epidemics in Asia that worldwide In Asia Leptocorisa sp prevail Their habits caused severe losses from hopperburn and vijus dicases make them particularly suitable for upland rice 1)thev feed over the past several decades occurred principally on on many grasses (and therefore can survive the earl y wet lowland rice Except in a few isolated cases (160) upland season before rice sets its grain) 2)adults are long-ied and rice has been spared from tungro grassy stunt and ragged mobile allowing them to find isolated plantings 0 rice stunt virus diseases vectorcd by green lcafhopper (GLH) (besides seed bugs only birds and rats appear to have shyand BPI] (123) Virus diseases have occurred in upland rice capability) and 3) adults aestivate in forested areas or where lowland rice was nearby Rice dwarf disease however sugarcane fields during the dry sca son when neither rice nor was first recorded on upland rice in the Philippines (187) grassy weeds are present (I199) Rice seed bugs concentrate Virus diseases can be perpetuated only on livir plants or on small-scale upland riceficlds because they can actively insect vectors and no living rice - plans tooned or search them out The prevalence of groves of trees characshyvolunteer - grows year-round in upland arcas excert in a teristic of upland environments ensures nearby aostivation few with 12 mo of rainfall sites keeping the seed bugs close by

The smaller brown planthopper Laodcelphax striatellu Leptocorisae are uncommon in lowland rice plains occurs in upland rice in Japan (18) It spreads black- because 1) their populations are diluted in a sea of rice streaked dwarf and stripe viruses between winter wheat and planted more or less at the same time (they ca n only feed on barley to lowland rice Early planting of lowland rice rice during the milk to hard dough stages) 2) they lack nurseries spreads these viruses but upland ice has remained aestivation sites and 3) weedy fallow areas are limited uninfected The lowands of Asia are dominated by L oratorius

16 IRPS No 123 January 1987

which lays its eggs hign on tile foliage (199) L acuta and L solonlonlesis oviposit at ground level the former on litter and the latter loose on the ground These egg laying habis explain theirtenvironmental prefcrencecs(214) In Japan the postdiapause adults of L chinensis migrate from moun-tainous areas to upland ricefields during floweiing (1 14)

Pentatornid bugs prevail in Latin America (8182 195) and in Africa pentatomid and coreid bugs arc equally cited (6 82) l)amage is often characterstic of the species (5)rcoris ventralis and Mnida spp are the common

pentatomid seed bugs of Philippine upland rice but they prefer lEchinochva spp grass seed to rice Vearaviritda occurs worldwide and hoften more abundant in the uplands(54) We saw few reports of seed bugs on upland rice but see(d bugs apparently opelrate cqually between by land and upland rice Good dispersal is necessary ior species that call feed for only 2-3 xon a crop and must first find those crops at ripening stages

Other pct species feed n flowers In Mato Grosso Brazil (194) and Santa Cruz Bolivia the panicle weevil Aeo-hariuia amllitarsis attacks rice during flowering (68 233)The larva e rice sten horers adutlts feed at the bases of spikelets caIusing empty grains Also inLatin America thecydnid -lkiituttatralusisa seed bug as anad tilt and nymph (194)

A number of beetles ecd on poilen - coccinellids lialrotia Aulac lhora and Monohpl-a (55) - but becatise riCe is self-pollinating and fcrtiliation occurs before pikelets open they pose no threat to yield

IST ECOLOGY

We now look at characteiistics of the life cycles of upland rice insect pests to lea rn hew each plays an adaptive role in upland rice ecology

Life history strategies Orzasativadoes not tolerate drodgh well so upland rice is highly seasonal normally grown in the wettest months of the year Usually rice is present for less than half of the year The rice-fiee fallow poses serious problems of survival to upland rice insects which have evolved att least four mechanisms to ovrcolne tle cvcl ic-l lack of a hostI) polyphagy 2) longevity 3) dormancy and 4) vagility (dispersal)

Poldt)agy lolyphagy is defined as having hosts of at least two botanical families oligophagy of more than one gents and ionophagy of only one ventus (40) - inour case Orvza Most wild rices however are illi tiC and theieiirc are normally far removed from upland rice hahitat othewise more would be alternate hosts for uplanJ iice insects Thte highly mornophagous rice pests shySciriol)haita hiceritlas Nilal)arvaiahiens Iand Nealo--tettiyvircsctws in Asia and Aalirpha secparatella and 1ioj)sis lngicornis inAfrica - occur on upland rice but

are more abundant in lowland culture They can specialize in rice because they are highly vagile and can attack all growth stages By specializing in rice monophagous rice pests can outcompete related but polyphagous speciesIntensive andI extensive lowland rice culture has favored monophagous species Riqwla alhinella (Cramer) the onlymonophagous species in Latin America is rarely reported on upland ice (I1)

Because of its limited temporal and spatial existence upland rice has not favored the evolution of specialized species Clhang(44)shows evidence that upland rice has only recently been cultured by man from the lowlands thus pests have had little time to fully adapt to the crop All but the lowland-adapted insect pest species have alternate planthosts to rice Root pests with limited mobility are highly polyphagous Pests that specialize in one growth stage eg secdling maggots seed bugs are oligophagous or polyphagous

Most alternat hosts of rice pests are annual grasseswhich also are seasonal Upla_rice is also a host but not the preferred one of the most highly polypliagous spcciessoil pests grass hoppers armvwarrs and Vearaviirielda

The d ry season poses igreat obstacle to ani upland riceinsects survival and only sonic perennial plant hosts are suitable food during the off-seasoin Plolyphagy allows upland rice insects to suLrx-ixe during thC rainy season when plants- both annual and perennial -areactivelygrowing and therefore more nutritious

Loiugevitr The ability of aiinsect to live 2-3 mo without undergoing dormancy helps species sich as root-feedingwhite grbs (as larvae) and secd-feeding bugs (as adtilt) to survive unfavorable periods White grubs arc not highly fecund and their life strategy is to improve survivorship of limited progeny Their subterranean habitat hides them from manv natural enemies lBtut root tissue is not highly nutritious so they ltilst eat great quantities and need a largeinsect biomass to digest it Other soil-inhabiting rootshyfeeding insects have short life cycles but are more fecund

Seed pests need greater longcvi vas adults to locate ahost at early grain dcclopment arnd give sedentary offspring time to develop Few groups of upland rice insects live longin active development Polyphagy and longevity only increase their ability to survive during the rainy seasonOther mechanisms are needed to survive the dry season

orantc In the tropics the nonrice season is the dry season while intemperate areas tile off-+season is winter Many upland rice insects undergo dormancy inlsummer (aestivation) or winter (hibernation) Dormancy is simplyinactivity during winters with prolonged temperatures below 10 C insects find shelter arnd cease movement Activity is resu med with warmer t empcera tu res A deep state of irnactivity termed diapauise occurs in sole species that prepare physiologically (havxing low metabolism and convcrting gonad s to fatty fuel reserves) for the unfavorable season (148) They enter diapaUse inanticipation of unshy

IRPS No 123 January 1987 17

favorable weather and resume activity after favorable Armyworms skippers and locusts actively travel hundreds weather begins usually in response to photoperiod or of kilometers in air fronts such as intertropical convergencethrough a delayed response to a stimulus such as heavy zones to descend onto upland rice fields far from their rainfall (237) Not all individuals of a population have the breeding grounds (31) Several generations can build ip on same threshold for environmental responses for entering or alternate hosts in tile grass family before rice is plantedterminating diapause -- this is seen as a mechanism to These large insects are active fliers Smaller insects such as prevent total mortality if a false cue appears such as a thrips aphids and planthoppers passively migrate long sudden warm spell in winter or rain in the middle of the dry distances by the wind season (154) Otherspecies ean readily travel tens of kilometers - stem

Probably more upland rice insects undL go dormancy borers leaffolders leafhoppes cutworms spittle bugs leaf than we realize Candidates are Atherigona spp root beetles seed bugs ants and termites aphids root weevils crickets leaf beetles thrips and seed The least dispersive species are white grabs mealybugSbugs Insects known to entervarious states of aestivation are seedling maggots crickets weevils and gall midgesleptocorisaspp adults (199) Maliarphascparatellalarvae Each species has evolved a unique set of attributes to (9) Scirpophaga innotata larvae (117) Chilo partelhtsv enable it to survive and adapt to the changing uplandlarvae (21) ScirpopThagainccetulas (179) white grub larvae environments Several evolutionary avenues lead to fitness (134) Patangasuccincia adults (6 82) and Schistocerca (Table 2) but there is no reason to believe one set of gregariaeggs (82) attributes is better than another (224)

Insects known to enter hibernation are LodeIphaDstriatellus nymphs (124) Chtio Tvlpressalis larivae (125) DroughtSesaia inferens larvae (146) but not S calavisis(2) Well-drained rainfed upland soils are subject to drought ifIDiaraea saccharalis larvae (75) Scirpophaga- tcerlttas rains do not fall within 2-3 wk Prolonged drought followedlarae(5 lacchrli s lae (75)saSlOhagasshoperseggs (82)and felo pidas athias pOpae (232) by favorable rains stimulates armyworm and locust outshybreaks but mealybtigs root aphids and thrips become

Lcpmotorisa oratorisand L acuta adults aggregate on numerous onlyafter dry spells for two reasons First heavy trees sugarcane or other shady moist sites L paluwanmsis rain normally kills soft-bodied foliage-feeding thrips and confined to Sulu and Palawan Philippines lives on the mealybugs and soil-inhabiting root aphids and root mealyshygrass Brachiariamutica (Forsk) Stapf in the absence of bugs Drought eliminates this source of mortality Second rice The adults can live for 5 mo When aestivating their the rice plant responds to drought by breaking down rate of metabolism drops and they do not feed They will proteins into soluble N compounds which enter the phloem however puncture plant tissues of their aestivation host in and are taken up by these sap-feeding insects (245) Greater search of water This behavior has led to errors of the host nutrition therefore leads to greater survivorship and range of rice bugs (I) Aestivation quickly terminates with fecundity rain probably meaning aestivation is not deep Small rice area

Some stem borers aestivate as mature larvae in rice Except in the highly mechanized large landholdings of stubble or in straw stacked as livestock feed However Latin America upland ricefieldstypically are patchy within moriality increases proportionally with the duration of the a highly diverse flora The small fields result in higher ratios dry season even if the stubble remains undisturbed If the of perimeter to area than with the typically larger lowland stubble is plowed for a crop following rice few stem borers ricefields Upland ricefields themselves may be intercroppedsurvive Rainfall terminates dormancy but it normally with cereals legumes or root crops Mixed intercroppingtakes several weeks for moths to develop and emerge with a wide variety of species also occ in tribal slash-andshy

Last-instar white grub larvae tunnel 1-2 m deep in the soil burn agriculture (50 74) to construct pupal chambers in which they aestivate The Upland ricefields tend to be small because of the labor larvae are very sensitive to dryness so pupal chambers are needed to clear land in slash-and-burn areas and to till land sealed to coierve body moisture Aestivation terminates to minimize weeds Rice requires more tillage than does with the first heavy rains but as in stem borer 3-4 wk pass maize because rice competes less well with weeds Uplandbefore adults develop rice is more a subsistence than a cash crop and small areas

Insects may respond differently in each region to cold (I ha) can feed a family temperature or drought Biotypes or loal populations may Small fields favor some insect pests such as some ant evolve (56) Local populations of Scirpophagaincertulasor species that can forage from more permanent field borders Patangasuccincta may have become adapted to prevailing Trees provide food and mating sites for white grubs (134) as conditions well as aestivation sites for LetTocorisa spp (199) Field

iagiliti The ability to disperse combined with high border grasses plus tIhose within a ricefield provide alternate fecundity orshort life cycles enable some upland rice pests to food for all but the highly monophagous upland rice better exploit the temporarily favorable upland habitat colonizers and a habitat for natural enemies

0 z

Table 2 Life history patterns of upland rice insects or insect groups

Host range Longevity Dormancy Vagility FertilityMonophagy Oligophagy Polyphagy Low Medium High Yes No Low Dispersive Migratory Low Medium High(Oryza spp) (grasses sedges) (angiosperms) (lt3 wk) (gt2 mo) (lt1 kin) (lt50 eggs) (gt500 eggs)

Ants xTermites x x x

Mole crickets x x x xx x xField crickets x xx

White grubs x x x xx xRoot aphids x x xxMealybugs x

x x x xxSeedling maggots x

x x x x xLeaf beetles

x xx xArmyworms x x xx x

Thrips x x xx xChilo spp x x xx xMaliarpha x x x x

xLocusts x x Xx x xGall midges x x xxiVephotettL iirescens x x xx x x xLeprocorisaspp x

x xLeaftolders x x x

Sogatellafurcifera x

xx x x x x x x

00

IRPS No 123 January 1Q87 19

All this appeals to favor insect pest buildup but the notwithstanding the release of stored nutrients in response contrary is true Loevinsohn (138) showed that insect pest to drought that temporarily benefits philoem feeders populations respond exponentially with the proportion of land devoted to rice up to about 75 of the area then the M POR ANCE I I P1A 1) RWI P1St S

rate of response declines In most upland rico areas tie rice crop occupies less than 50i of the area- therefore the Visits to upland ricefields normally reveal 1kw insect pests potential for population buildup is low This relationship Loevinsohn (138) and Locvinsohn ct al (139) compared may not hold for seed pests that can locate small ricefields insect abundance using annual light trap atchcs in upland

Another outcome of the highly diverse flora and phical rainfed lowland and irrigated lowland sites in the Philipshyenvironment of upland rice isthateach upland ricearca has pines Although only one upland site was studied data a unique composition of insect pests showed equal or fewer insects at the upland site than at the

For example in Tanauan Batangas Philippines the rainfcd lowland sites and fewer than at the irrigated sites main upland rice pests are rice leaffolder(RLF) armvworm (Table 3) M Lltiple regression analysis showed that the key Aythntnaseparata WBPH ants Soli psis gentitataand factors explaining insect abundance among a set of cropping white grub Leucopholis irrorata Sixty kilometers away in intensity variables were the number of rice crops grown per Real Quezon the main pests are seedling maggot Athe- year followed by area devoted to rice Table 3 shows that rigona orae flea beetle (Thacitcmnra hasalis thrips the upland rice site had only I rice crop perym rand that rice Stenchaetothrips hiurnmis and rice bug Leptocorisa was planted in only 20Vi of the area Fertiliter and oratorius In a third Philippine upland rice site in Claveria insecticide use had little bea ring on the trend The armters at Misamis Oriental Mindanao the main pests are ants the upland site applied 60 kg N ha higher than at most Solenopsis gemtinata seedling maggot Alieriona orvzae irrigated sites white grubs Holotrichia in)iUaiuoan t and Lucopholiv Insect occurrence at the upland site was omparable to irrorata stem borers Sesamia ifiArelr (hio auriciliusC that at the Cagayan rainfed lowland site parlicularly in 1982stpessallis Scirpophaga incertulas S innoiata and when drought prevented many farmers fron planting(only

4c ona chrrsurapella root aphids Tetraneura niriah- 40(of the rice area eventually was planted that year) M ore dunrinalisand Rhupaosiphunm riahbdminalis lcaffolders favorable weather occurred in 1980 and 1981 which Marasniapataltis and Chaphalocrocis Medinalis army- supported greater rice areas and consequently more insects worm MIthinna separata and rice bug Leltucorisa The most intensive site for insect pests was South Cotabato oratoriusThe great diversity of insect problems is typical Mindanao where farmers plant 23-25 rice crops per year fer upland rice and makes control efforts more difficult with irrigation The Zaragoa Nutva Ecija site also had particularly regarding breeding for insect resistance high pest abundance and at this site the fields were highly

asynchronous Asynchrony was the third most important Low yield potential Iactor in explaining rice insect abundance Most upland rice is grown on low-nutrient and mineral- Rice cropping intensity in upland areas is bound to toxic soils which when combined with eiratic rainfall remain low Only one rainid upland crop is possible per make upland rice a highly riskycrop for the farmerto invest year (Farmers in Claveria Misamis Oriental Mindanao in costly soil amendments and land management Often Philippines tried asecond upland ricecrop i 1984a ycarof upland rice isgrown far from markets making inputs even favorable rains but no harvest was possible b-cause of more expensive Farmers growing a subsistence crop hosi- drought blast and birds) Rice area isaso bound to remain tate to invest in costly inputs to raise yields low because of the high labor and power rcquirelmenlts to

Even having varieties that could double existing yield prepare the land (because of weeds)and because upland rice potential under present management levels would probably wvill not compete with lowland rice as a cash crop Some not prompt many farmers to use purchased insecticide upland rice sites with prolonged rainy seasons howeverare unless tile risk of crop failure were reduced such as it was planted asynchronously such as the Clacra site with irrigation systems in lowland rice The seasonal abundance of Upland rice insect pests

The Tanauan Batangas site is atypical for upland rice attracted toa light trap wasgrapied for crop year 1980-81 in The soils and rainfall pattern are favorable and because of Tanauan FBatangas (Fig I 2) W13P11 was he most its nearness to Manila farmers have cash resources ftm common rice hopper and its population peaked at the sales of vegetables to purchase tertilier They applied an midgrowth stage as it does in lowland rice D-Vac suction average of 60 kg Nha after particle initiation and obtained samples showed an earlier peak than that from a light trap 25-35 tha yields The high fertility isone reason why RLF indicating emigration W13PI usually emigrate after the armyworms and WBPH developed into large numbers vegetative stage Populations in 1980 reached levels that Farmers however used no insecticide on upland rice caused patches of slight yellowing in some fields WBPH although they did on vegetables probably cause hopperburn in Batangas They immigrate

The poor growing conditions fOr rice generally mean year-round as shown by low levels in the light trap BPH poor growth and low fecundity for insect pests fIeding on it however were scarce in the light trap and not collected with

Table 3 Abundance of rice insect pests and factors in rice cropping intensity of Philippine upland rainfed lowland and irrigated lowland locations where light traps were operated daily for atleast I yr -I

0Rainfed Rainfed lowland Irrigated lowlandupland

Pangasinan Cagayan lloilo Santa Maria Nueva Ecija South Cotabato Tanauan Caaringayan Caaringayan Bangag Bangag Bangag Buray

LagunaCale

Batangas Manaoag 1980-81 Santa Motiica 1982-83 Cabanatuan Zaragoza Avancena NaniamanSolana 1981-82 1982-83 Oton

1980-81 1979-80 Oton 1981-82 1981-82 Koronadal Koronadal1980-81 1979-80 1979-80 1983 1983

Insect (kerosene light trapNephotcttixcatch nolyr)virescens 650 1240 500 1800 330 210 3400 1600 160N nigropictus 210 - 360 18400 5500 2900- 140 90 20 - shy 1700 -Reciliadorsalis 710 - 3200 1700180 890 630 140 170 2400 800Nilaparratalugens 130 690 - - 4200 4600300 520 750 180 60 3300 1600 740 310Sogatellafiircifera 1700 31000 13000 8200310 210 3300 610 270 980 1600 6300 -Cyrrorhinus - 12000 6700600 540 3200 290 160 330 120 410Scirpophaga spp 50 410 620 - - 21000 12000150 180 220 270 30 1000Cnaphalocrocismedinalis 70 70 540 610 270 11000 6100200 140 100 6 60 250Chilo suppressalis 0 60 60 1900 300 200- - 30 20 0 6 3 160 -C auricilius - 0 00 - - 20 3 60 20 3 0Sesania inferens 0 - 80 60 3 80 10 51 -Cropping intensity _ -

Rice crops (noyr) 10 10 12 10 10 10 13Rice area (7c) 17 19 20 20 25 2320 85 85 60 70 40 85 85 90Fertilizer (kg Nha) 60 20 85 80 70 7020 0 0 0 30 30Insecticide (no of 40 60 40 300 04 04 0 03 300 13 13 30 50 40 33 33applicationscrop)

__________

IRPS No 123 January 1987 21

Light trapped Insects Drytnd __ 2(notrap-nght) Suctioned Insects (nom )

50 -)o 9 jO40- SOW ffurciWO Light trp

D-Vc suction

20 to 4

20

Egg porasitizahion (M) 75 60

300 SOOampReOftwciffr

45 30shy

15

-insects (notrop-night) 4 Nileporvata Iuen5

Egg parastization()

Light trapped insects (no trop-nght) Suchoned insects (no m 2 )

6 8 Cytorh~nuls hvidipennii 0

4 05 2

50 0 Insects (noltrop-night)40-ui Ipanh pe g

Moohase 1 Seasonal abundance measured by light trap

Rainfall(etod) and D-Vac suction sampler of whitebacked

to ~ hopper Nilaparvata lugens and lea ffolder60)-t Cnaphalocrocis mnedinalis andplthpe g20 L I II I I I parasites and mtrid predator Ortorhjntus livi-Apr May Jun Jul Aug Sep Oct Nov Dec JaIn Feb Mar dipennis set against cropping pattern daily1 9 80o 19 8 1rainfall and moon phase Tanauan Batangas

1980-81

the D- Vac Cyrtorhinus build up on the crop coterminous are collected in light traps during full moon (109)with WBPH buildup Cyrtorhinus disperses during the Because upland rice shares most of the lowland rice pestsrainy season RLF not readily collected in light traps was and those pest species are generally less impotant to uplandprevalent only during the rice crop rice than lowland rice there is a general belief that insect

Nephotettix virescens was more abundant than Nnigro- pests are less important onpicLus in the light trap collections and on the crop GLH upland than lowland rice Arcview of limited data on upland rice shows an averag of population peaked at the reproductive stage as did zigzag 10-21 yield loss from insect pests on a par with that ofleafhopper (ZLH) Crtorhinus tracked the populations of lowland uee determined by the same methodology (132) planthoppers and leafhoppers Both GLH species were (Table 4) McGuire and Crandall (144) made a similarcollected year-round in the light trap indicating immigra- estimate for Central America Cramer (51) howevertion from nearby irrigated rice areas Possibly because of the estimated 35 yieldlow populations loss in South America based on ainsect number did not correlate with similar pest complex - stem borers and seed bugs shymoon phase although reports indicate that more hoppers attacking irrigated rice in North America This estimate did

22 IRPS No 123 January 1987

Light trapped insects D~~nd (no trap-night )

St plht Suctined insects (nom 2

)

D-Voc suction 1520

1o

0 Lih trap 5 ~105 o5

00 Eggparositizotion()5040 NephOtgtha virneens

30 20

I010 A-Insects (notrop -night)

4 Nephatelh nigropccent i

2 o I vV^J - -- ^ VA____ f _

Light trapped insects (no trap-night) Suctioned insects (no rnE

10 -1208 ReCdO doRsol

6 15

4 10 052 iwxALAiamp 0

Light trapped insects (notrap-night) 2Suc oned nsects (nom )

8 CyrtorhmnusI6o nn]10 i0075

4 05

2 0 0

025

2 Seasonal abundance measured by light trap Roinfall (mmd) Moon phase and D-Vac suction sampler of green leafhoppersNephotettix virescens and N ngropicrus and I zigzag leafhopper Recilia dorsalis their egg 60

parasites and micid predator Cyrtorhinus livi-dipennis set against cropping pattern dailyrainfall and moon phase Tanauan Batangas1980-81

40

5 Lf Apr May Jun

1980 Jul Aug Sp Oct Nov I

-aMar

1981

not account for soil pests and was not based on field data in the absence of pests Carbofuran granules are phytotonicAkinsola (I1) cited mainly data from irrigated rice areas in to rice (239) The prevailing hypothesis is that carbofuranAfrica but estimated yield losses of upland rice between 15 prevents soil bacteria from consuming fertilizer The phytoshyand 30 from insects However losses in Thailand over a tonic effect is through greater availability of fertilizer to the4-yr period ranged from I to 13 averaging 5(106) plant Denitrifying bacteria are known to be more active inYield loss data however should be interpreted with aerobic than anaerobic soils therefore it may not be merecaution (184) The Philippine data were all derived from coincidence that the highest recorded yield losses (22-69)trials on farmers fields under farmers normal agronomic occurred in those trials with 12-30 kg basal Nha On themanagement practices except for Bukidnon and Capiz other hand contro with carbofuran granules may not be aswhere researchers used fertilizer Batangas farmers are good as a seed treatment against seed pests and perhapsatypical because ofhigh levels of fertilizer used The trials in yields would have been higher if a systemic seed treatmentBrazil were carried out at an experiment station but insecticide had been used conditions were similar to those of local farmers The 1979 and 1980 trials in Batangas had two full-

Carbofuran granules were used in every trial as a broad- protection treatments one using carbofuran granules andspectrum insecticide applied basally with the seed An the other using bendiocarb WP as a seed treatment Plotsobjection to using insecticides to measure yield losses isthat protected with bendiocarb yielded higher than those proshyinsecticidescandirectlyorindirectlystimulateplantgrowth tected with carbofuran granules The other trials in the

Table 4 Yield losses to insect pests and yield responses to insecticide in upland rice in the Philippines Brazil and Upper Volta

Yield (tha) Yield loss (5) Location Year Cultivar Protected

with Protected

without Check Total Vegetative Reproductive Ripening

Basal N (kgha) Reference

carbofuran carbofuran Protected No with carbofuran

carbofuran

Philippines Pili Camarines Sur Tanauan Batangas

Pangantucan Bukidnon

Dumarao t-apiz

BrazilGoias Minas Ge~ais

Upper Volta6

1975 1974 1974 1975 1975 1976 1977 1978

1979

1980

1979 1980 1979

1977 1978 1979

Bursiging puti Kinanda Dagge Kinanda Dagge Dagge Dagge Dagge UPL Ri5 Dagge UPL Ri5 Dagge UPL RiS UPL Ri5 UPL Ri5 UPL RiS

IAC47 IAC47 IAC47

14 33 11 36 35 31 30 27 37 26 40 30 40 42 46 35

19 21 09

--

-34 24 24 27 --

29 43 31 42 -

47 25

---

10 28 05 27 27 30 26 29 33 28 33 29 43 23 36 11

15 15 06

29 15 55 25 23

3 13 -7 11

-7 18

3 -7 45 22 69

24 28 35

--

21 -11 -20

4 -

-3

23 7

-2 -23 56

-

-

-__0

6 --

-

-23 14

5 0 8 2

22

-

--

0

-

--18 11

3 -10

4 10

7

--7 -

14 -

-1 6

10 -5

8 9 7

-

--

0 0 0 0 0 0 0 0 0 0 0

30 30 30

12 12 12

99 99

100 100 101 101 102 102 102 102 103 103 104 104 104

66 66 66

Farako-Ba

Mean

1977 1978

IRAT10 IRATIO

38 66

--

40 43

-5 35

21

-

10

--

--

--

- 242 243

Z

tJ

-w

24 IRPS No 123 January 1987

Philippines which had two treatments offering protection In the 9 Philippine trials where yield losses were measuredwith and without carbofuran granules relied on further by growth stage 10 of the yield loss (21) occurred at thecontrol with foliar spray and not on seed dressings The vegetative stage and 517 occurred at each of the reproshycarbofuran-protected plots had an 18yield los compared ductive and ripening stages Ibis result is similar to thewith only 10 with foliar sprayS This yield difference is pattern of yield loss in lowland rice measured by successiveprobably due to differences in protection against sown-seed treatments where the crop is unprotected at each growthpests (foliar spray do not control sown-seed pests) rather stage but proticted during the other growth stagesthan phytotonic effects became basal fertilizer wa used in However the insect complexes r sponsible for vegetativeonly 2 of the 10 comparisons stage yield losses in upland rice are entirely diftfrent than in

The recorded yield losses were not related to yield lowland rice The key vegetative stage pests in lowland ricepotential (Fig 3) One might expect higheryielding crops to arc aquatic whereas those in upland rice are sown--seedhave higher percentage of yield losses but this ws not tile (sown in soil) root and seedling pests This dift rence maycase One reason is that insect damage exacerbates plant explain why agricul torists have overlooked tile ilmportanccinjury from drought stress low yielding fields from d ought of upland rice insect pests Many reports of upland rice pestsstress would show disproportionate differences between in Asia ha Ve focused on major lowland species (46 108)insect-protected and -unprotected treatments Shoot and It isalso surprising that tei vegctative stage of upland rice root pests would be more responsible for losses from would record the highest yield losses becatuseseeral studiesdrought stress than sown-scd pests A second reason have shown that upland rice can readily recover from foliageconcerns seed pests which would have a greater impact on reinoval during that stage (117 152 165 188) Upland ricelow tillering low yielding varieties such as )aggc Low poscsses a hitgh obilitV to compensate for ca rly loss oftillering varieties nnot fill in the space created by removal foliage and will even be stinulated by foliage removal toof seeds oweer 1) R i 5 is high yielding because it tillersP1 produce a higher viel than plants ithout any foliageactively which can close the canopy to provide greater weed removed (63 192 231) [ie reason for this apparentcontrol discrepancy hovce is that yield loss at the vegetative

stage does not octUr frol leat arca loss but fron pestsYield loss () removing sown seed and feeding on devcloping tillers and roots -- damage that cannot be readily conpLtnsatCd for

70 - Measuring yield losses in upland rice cnvironrnents is further complicated by a report from Brazil that greater yield losses occurred in response to better insect control60 - =-09 as Insecticide seed treatments resuled in denser plant stands

(n=21) which in turn created a selectively more favorable environshyment for blast (69) This resuIt was parlticurla rlxexacerbated50 when cirbofuran granules were used in seed furrows Aside

ffrom a greater plant stand ca rbofuran iay have auigmented the N lecl and stimulated blast If so ca rbofuran should be40 - replaced with another insecticide or the seeding or fertilizerI rates decreased

Other methods to determine the impact of upland rice30 shy pests wvcrc to interview farners or to develop single speciesbull bull correlations of population levels and yield loss Feneira (66)2 NIM)st (75(1) Golas farmers reporteddid both in Brazil

20- insect losses in ufpland rice and 251 said those lossesS avcragcd 37(1 [his result is consistent with tie yield losses 0 0 measured by field trials in the same statc(Table4) For every

I0 4 of seedlings removC d by leaf-cutting ants a I 7yield loss is predicted Similarly Ferreira (66) found that ai

0 - average of 53 thrips per panicle before panicle initiation meant 3 times more unfillh d grains

-IO0 I 1 1 I I-101 0 2 3 4 5 6 7

Yield (tho) C)N [ROt IM FI IODS 3 Correlation between upland rice yield and yield lols caused by Upland rice insect pests may be controlled by culturalinsects measured in 21 trials in the Philippines Ivory Coast andBrazil genetic biological and chemical methods

IRPS No 123 January 1987 25

Cultural control dry season plowing will have little effect on them Plowing Crop husbandry to reduce upland insect pest populations when the soil is wet from the first monsoon rains will expose can be classified into practices effective in a single field and soil insects that tunnel close to the soil surface to predatory those effective only at the community level Cultural birds chickens dogs and even man (field crickets nole controls should come first in a pest management program crickets and white grubs a re delicacies in man tra ditional They have broad stable effects as pests have little pos- diets) sibility of overcoming them through selection of biotyps The greatest degree i1 inIscct control fron tillage comes (138) On the other hand some cultu ral controls work only from plowing soon after rice ha rvest when most soil insects with high labor or power inputs and connunity-wilt lie close to the soil surface anrd stern borers are in the rice methods require coordination of iany farniters stuble Whitegrubsare mature and tillage exposesthem to

Pantin time The luxury of being able to shift the time of predators Tirnelv control will prtec the crop following pla ntring upland rice is open onl hr la rniers in areas with a rice and lowr the populationIfor succeedinig years Plowingprolonged rainifall pa ttern lI copping intensity or underthe ice stulble helps it decompose killing sten borer rnechanied land preparatiot Gcnerally planlting as soon larvae inside as the rFiinv season begins will lower populatiois of 1110st [requent tillage destroys ait nests bit not termites insect pests Insect populationsarelo aftcra dlvseoll or Tillage and clean culture howes er rcliove food for winter fallow delayed plaiting lets them build tip on termites which then mtay attack a young rice crop Scarab alIternate hosts that gi vigorotsly with the first seasonial beetles also prefer to miposit in eceitly tilled fields Noshyratiris This first flush iof weds cros lItxuttriartlv with iceCult is dibbling faors the survival oftillace Clre Such rineralied N released during the dr season aid made stils i aels tnd insects thtt pttpate or lay eggs- in the soil

available to the plants with the rains It iliso igrtmitcally Zero tilla hio eerl conserves predators ofthe GIH (37) advanttgeous for the rice ciop to tap this trlLItlaI fertility nt it mcttoi lo lessen the impact o0 soil insects and grow vigorotsly to compete with wrus and to tolerate brotdcasting seed or planting in ftltrrows is prefc ired over pest infestation dihlirig in hills Crickets are large enotlgh to eat seedlings in

Larly plantngs terd to escape seedling maggolts Upland a hill atrd tillering normnally ca iriot fill in tlhe missing space rice in Bitangas is planted in Ma and ine itli the Iirst Oftcn dibbll rd is not well covered and iseasily found by rains Fairners are highly mtotiated to plant early tocscape foraging insets ioIdents and birds drlitght and typlotns ricar harct Arn Atrgtst planting Plan uit sii Incrcatsini sced density protects tile cropwould be evcely attacked by It riim orivzo At tti againrst seed and seelling pests especially ants particularly other Philippiie tipland sites Rea l Qteot antd (laelit if the crop will germinate and ermerge quickly M isainis Oriental seedlig mnaggots arnrige ric boe t se hr trftinll itI India rice intcrclopped witi cotton or farnes have a longer growing season and plnit 2 rin or pigeon pci had lower 6IIt and WBIlII poptrlations than mote after the ortset of the rainy season liaaiangas ftrmners rice alone (211) NfailC arid Urplald ir_ 2re coilltori in the plant ca rly becia use ofa shorter raiinfall period arnd the desi re Philippines he chatngC in pest status folii iniercropping is to lliarvet before the typhoor sea son peaks ltighly locatioll-specific (133) cmianarid the net elfc be either

Early plantings cornbind with early-iatiuring varictics nil bcnelficial ortdetriential (189) For example BIatangias provide higher crp tolerartce for white grubs Aestiatior farmers intercrip Itro in their ipland ricefields A liornshyterminates with the first rains but 5-6 moare ineeded for the nrii iotd-Ahis i (I iiiacius) which norlially feeds resting larvae to develop into beetles emerge and lay eggs on taro ill feed or miarbv rice plants Intercropping and for larvae to reach the danriigii g ird-instar stage probably would not afft0 seed pests A crluanion crop (134 In lataigas this normually iseiough tiite for harvest planted ith rice and haing gleater tolerarnce for root

It Brazil howecver delayed planting is a suggested dariage or being toxic nayact isi trapcrop to root-feeditig cultural control for the lesser cornstalk borer and insect pests Acial-fei i ig pests voultId be most affected by pests favored l drought (66 195) Cropping internsity in intrer ropping Shoot aphids leaf beetles sten borers and Brail is low and delayed plraing places the rice crop in the hopplers would be likely ca ntidates to note effcts On the niost stable rainfall period reducing likelihood old rought othlt halnd the corlipalior crop stch~ sitniaie Iiiv be more

In rilniy arca s rinmirsare roi atl to plant other crops benehited than rice from initecropping ( 137) before rice and althNgh it woultl be desirabh to avoid 11echi Weeuling luring the first month after clop infestations by planting rice cuatl lariners lack the labor cstablishent will force pcts such is anyvworris which and time to do so ol a low priority crp prefer grassy cetds ontor tle vorling rice Soil iampsecs will

77age Uplild rice Stils ae light textured and Call be mvnye to the rice crop Cleani citrltirc also lay lore territes plowed duiring the d rv seasor notl only to grin time for to attack I u cetincti inrice ceds i ay he le optinal planting with lie fitst Inilsoon riiains but also to desiccate solition 13) soil insects arid weeds such is ntsedge that foim rhizomes Irtlizr Itneatsing N Iciiliier oin hlwland rice has Htowever becta use most soil insects lie below die plow layer fAtwvored Irigher populains of pliithoppeiv Imlhoppers

26 IRPS No 123 January 1987

and leaffolde rs However because of the need to temper fertilizer use on upland rice o prevent blast fIrtilizer rates will be lower than is comonia r for lowlad rice causing relatixely less hopper and IaIfTfoide r population buildup However inupland rice areas without a history of blast such Is Batangas farmers apply high rates of N and conscqtrently the crop has high infcstation levels of WBPI]and RIF The fhrmrcrs leec insect buildup by splitting the fertili11r in0to two to three applicatiois bit do not applV fertilizer at the vegetati estage because this prodrces a tall crop which illeasily lodge during typhoons Greater crop fertility leads to greater insect survival (better nutrition) larger insects (to cat no re foliage) arid greater fec titd t (to lav nmre eggs) (128 145 171) Not all insect species respondequally hovr Increasing rates (if N l anrid K increased Dialraa s-acchralis and (iaeiurema flea beetle but reduced thrips (66) Zn rcdLuced deadheart density from the lesser cornIstalk borer

Fnldingi Although most Upland rice areas do nIot hawxcaccess to wa ter floodiig the fields is rRrrnriierdcd in areas of Brazil or root bug and white gib control ( 195)

Crop rotatioti Tillage before planting a crop after rc harvest will control stern boreis and other pests renain n the stubble arid will unearth soil pests Planting a non-gratriiaceous crop after rice is recommncrded for termite control (66) Crop rotation is less nttdcd in upland rice culture than in lowlands because atcrop-free period is assured even after harvest of crops following rice

t1ulchilt iryl plastic irli irig designed for weedcoitrol ii upland rice inJapan resulted in ntr highanri liy infestation of (ihilo sulytresvais (93 9-1 95) NIulched Upland rice grew more luxuriantlv and attracted more (Avipositing mothis than u lched rice Also the rice plant and stein borers imatoured 10rC quickly and a partialgeneratiori af vouing ista iNoverwintcred in the stubbleLotili--malitrt varieti Quickly maturing crops reduce the tmbCr of pCst generations that Calt build up Rapid crop establishment olan ca rl v-natu ring vaitvty if)pa 11icLa r will lessen white gib and stem borer damiage White grub larvae intle last iistar are highly destIruCtive so aii early-iiatri ng variety will escape serious root loss Stem borers build 1ip slowlyx in rice to ecormic abundant in a late-Maturing crop anrd caiuse whi tlieads Numibers of somespecies increase exponeitially with each succeeding genera-ti so a xarietyexhat atures I trowh earier thian another will hac less insect damage Early-naturing vaiicties however will not lessen seed pest daiage either ii the soil or oina standing crop

Strwhoi hAIJttam Inrscts tcd irig on thlie aerial portions of rice plats disperse from carly to late plantedfields AS iiiibCirs increase exponentially with eachinsect

gtscnrattm e eiglbrig f rmc~s1(uil upp wh rc generation pests build tIP where neighboring fihrTC e9r1sstagger their plantings II Batangas farmers plant uplandrice within I iro use 120-d s-arieties and plant at the start of the rainy season Ini Misarnis Oriental laricrs stagger their

plantings up to 3 mo apart and use a set of atieties that mature in 4-7 mo Insect pest danage particularly from stem borers is greater in Misarmis Oriental Also seedling maggots damage rice there because rainfall favors frequent planting of maic In Batangas rice escapes seedling maggots bCca usC of distinct drs season and synchron0us early planting with the onset of ains

Therefore combinations of cultural practices ea rlyplanting synchronous planing crop rotation and earlyshymaturing vaneties protect the rice crop against most insect pests

Ilmant resistance Beca tse upland rice has few specialized insect pests and many insect types attack the crop worldwide regional breeding and strong internatiornal cooperation are required

Sown-sced pests arc economically controlled by insectishycide seed treatment and should the relore receive lowxpriori ty inbreeding objectives

Root pests are normally cost ly to control with insecticides but finding resista t sonres mayvbe di fficult Tanaka (229) for exa -ple fi led to find varieties resistant to root aphids Selecting for large root bioniass perhaps should be the strategy for pests sLuch as Whlite grubs that renliove roots Fainter(174) howcvcr cites varietiesof crops tolerant ofor resistant t(i root pests Some sorghun varieties are high in cyanide so rice sorghun might incorporate broadshyspcctruri chemical resistance

Seedling vigor and drought tolerance are high prioritybreeding objectives so tupland rices should have large r root systernsand be able to tolerate ligherlevels of root loss Sonie root loss inolder plants is beneficial if it stimulates new root developnrut and therefore crihances intake of soil nutrients (53) The rice seedling inaggot rtiLotia spp may be

controlled thirough resistant aittics A hi rgc breeding effort on sorghniu has had fair success inideveloping arreties esrstait to Asoccata Rodani (251) A field trial comparing 10 upland ricc varieties showed differences aniong varieties va ring from II to 4(- damaged tilleis (215) Hovver resistarce to 1Iill-0r gtia species may rot cross ox-rti otier

Shiraki (210) reported that upland i-ice stens wcrc harder than lows-land rice stemsperhaps becaIusc (f a higher silicoricontent and thus were more resistant to yellow stern borer hc tortia Iow tillering (f uplaid iice xarneties prodtres a

higher percentage of infested tillers than would occor in higher tillering lowland iice under a siiilar egg density The corollary therefore is to select for high tilleriig upland rices

Stemii bor amcstheh main targes dobrecders inAfrica (571 204 )but nticl oifth e sc reen irig is dotie unde r lo wland co d t osrm s wih A h r~ o l pe i s n acorditions (11 96 218) As with lthrinrnwspecies mayhave to be dealt with independently however initial results show cross resistance between African and Asian species(92)

In Brazil screening Ior insect resistance focttses oil lou I pest problems Varieties tolerant of lasmolwalpus ligi-sellus and resistant to )iatraea saccharalis have been identified (70 153) Current work also emphasies 7iraca limativetntris and Thmialspis (=Iosis) flavouicta (66) These insect pests Ilt onlV citise IIigh yield losses tihey also ire difficult to control chemically ofr by other ilealls

ManV upland rice vaieties aic early maturing and are resistant to pests such as gall midge (I Xl ) Early mIturity is often a highly desirable trait to esuilpe pest buildup but it is not true resista nce bca usc ilthose va reties are plinted late they will be itsdamaged as ain early planted susc ible variety

P~erhaps tile cosiiiopolitall of all upland rice pestsimost are the sced pests (ripenling stagc) which pose a particular problem beta use scuds re ised IsIO for hitnit4ans Ayltoxic substatice introd tcld geticticllyilmay also be toxic to nal iniless it caln he placed old in thfe IdhuffSelectmi fo hairiness or lotng wns kill mIlct sthilrohleiiis dutilig threshing pa rtictfarl maitial threshinfl [hireshers ill complain of itcfimin atid itteascd (fust levels

Attempts itIRRI to fitid tcistatncc to the rice bug plorisa have ilotbeCnl prmitisilg lIIetIattonlids iorc

through the lenilna or palca hillhairs sedstVIor seeds wilh awns that deter bugs promote complaints from threshers Seed bugs liptocorisa and Stcnocoris enter the seed through the opening between tile lemna and palC but varieties with narrow openings are diifficult to mill

Breeding priorities should concentrate ott inotCI gruLtLIps that are most difficult to c nttrol by other meains Steil) horers appear to be the most widespread group that sigiiificantly dalag s rice aind is hard to manage by other Ilnca us

Biological control Very fe stutdic of mitatt~l eneties of rice pests in upland envirollnletits wee in tile wefouiid literature Therefore have not compiled a list of species Ve present a list of natural enemies iOtid in upland ricefields tuic latialln Batangas for some conitno iOldiar pest species ([itble 5)

Muich ofl thc lollowinle itfor ittiol cotlliCs rol Stutdies in the lowlatnds or roi crops otithcr thlin tice

Qu(ailtr o 11ll t11t( Nat trrtf lii hcoeits ight Ile used agaitist ile social insects tilt and termites Sollie predator specice live ill theirf nests n ilnic theirnllt ip-pearanIlCe aMRidbehavior IJtrouMdCing a mote aggressise ant species to displace pest ant codonies is a possibility (169) Patihogens as potential natural encnies have reccived little atteition (I13) Natural ncnuics asa group however hac not been toIl iporta it in Iegulating aintshown be and ternite numbers

The nocturnal behavior aid subteitmien habitat of lole crickets and field crickets protect them from many ntllral enemies There are only limited records of natural cnelics for these groups of pests some pathogens (87) but mainly

IRPS No 123 JattuIny I 271987

parasites - scelionids on eggsand sphecids on ilymphs and adults Sphecid wasps Larra carutmaria (SImitI hI L huzwuensis Rohwer and lsaltiuhta Willi ams speciali in mole crickets pa ralyzing them nd dragging the ninto thcir nests as food lot their voting ()tier sphCcid species Lins aurulenoa (Iabricius) lot maiac (Ashmead ) 1 lhshylssellaus (Smith) and Ml lal)oriosu (Snithf specialie in field clickets

White grUbsO the other hand hae Icti scriuns enuigh to have been extensively studied for cotttroi 1W nati al enemies( 112 140 141) Scoliid parasites C(ajsoMcispp have beetn intrtduced itto tilePhilippites for control of Ielxl ]udisiro la The rcsearchcts w Intttrdced themle during ill era claimed that the white gttionthreak as controlled (142) howevcr it isnormal for pest epidemics to subside and there was no diect evidence to indictte that tfe parasite was responsible A tmore promisitg titctllod might he to identify iso ltcttass prt~f tcartd apply hiactelaln fingal pathogens to tile soil Ihe lalattese beetle in North America has hcci coitrolled hv tite conmnleciall aailable hactcrial prcpa ration itacihlpu pav ofr milksdiscas (71) Spote prtyefott the soil itloted ilrtier reitiain actic fr Veas

Root aphid sardtnctly bugs readilslic tilib to predatots and parlsites once their tetndine ants are controlled The larvac and adtilts of a coccinellid beetle 1Scynms sp prey on llriadlouais in slash and burn upland rice Baits treated with insecticide may be used to control tending ants by killing their ourig (244)

Root-tkcditig blugs false wireoris wireorms and root wccils aic r ofsuch little ifilpoltatice that thenomttll role of liatural Cticniics hias tiot beei asscssed A possibility hoevrl tiglt he tit ItaIIitic niaCIMtodcs lintt1g11en Itese soil-itlhlfbitine mCtitodcs W5uhtfhave to be imiass Produiced

large miltibcr of niatltrll cnemics ircknown ft CltIoriS IC IpblCeii is thtt clttvortnlls ofonlie the crop soott alter land ptel)tritiOti a1tithe bcgit iniig of the raittv

Citsoln hfien tiatllral leniiiy populations aric lo Colltloil WOtuld iii ole mass prodtticing andirclcaslng key spccies

le egg stage is the fpirt (olthe Cedlitgt Ialot life cycle tI hich to f)ctis ftor tittlril elctit cottntrol hplraites o1 pleditors ttsit elleggs highly expoIsYCl I Sgg pailsites or predators stolld probahly e ittass andtbe produced released Littlh i toruialioll is a ailable ott what species to trV o ho to Iniss podLice thiCIi A etiloplhid wasp Icmihlaol sp effctiicly regushy

lates the leatf lcloalapolm2a spioanoiiitr heat andc rice antid should be collsctvcd

leaf beetles whCllose e am] eggs ate laid in the grutindhirO ould be motC difficutlt to control by niatulral enelV Mnll ipuhltiOn But species sUici IS c_gsthispid beetles ith

and larVle oilleascs ate vtilcratble to iparaites and pledators

Natural enemisc of tlirips otl rice are little known

28 IRPS No 123 January 1987

Table S Parasites recorded from upland rice insect pests in Tanauan Batangas Philippines 1977-84

Order Family Species Host stage

I lymttenoptera Braconidae Leaffolders Cnaphalocrocis Marasinia spy

Cardiochilesphilipplinensis Ashmead a Larva

Cotesia angustillasis(Gahan)Cotesia or cypris (Nixon)

Larva Larva

Cotesia or taeniaticornis(Nixon) Larva Cilonusinotakatae Munakata Chelonits SppAlacrocetrts philippinensis (Ashtmead)

Egg to larva Egg to larva Larva

Iclineumonidac Orgilts spp Isclutojppa hireator(Fabricius)

Larva Larva

Itoplectis naraigav(Ashmcad) Larva Tenelhtcha philippiensis (Ashmead) Larva Tenelchha stangli (Ashmead) a

irichomma cnaphalocrosis Uchida Larva Larva

Bethylidac Elasmidac

Goniozms or trianguli r Kieffer lasmtusalhopictus Crawford

Larva Larva

Diptera

Encyrtidac Chalcididac

Tachinidac

Elasmtus spp Copidosomopsisnacohiae (Fidy) Brach nteria evcarinata Gahan Biachyieria lasus (Walker) Ztgohothria ciliata (Wulp)

Larva Egg to larva Pupa Pupa Larva

Argyrophylx nigrotihialis(Blaranov) Larva

l-ymenoptera Braconidac Yellow stern borer Scirpopiaga incertulas

Braco chinensisSzeplegeti Larva CIthelotis spp Chelonus niunakatae Munakata Cotesi flaripes Cameron Stenoracon nicerilli (Bingham) a

Larva Egg to larva Larva Larva

Ichneumonidac Tropobracon schoenobii (Viereck) Amatroinorpiaaccepta metathoracica(Ashmiead)Frihorus sinicus (I lolmgren)

Larva Larva Larva

Ischnojoppaluteator (Fabricius) Larva Isotima nr dammeratiti Rohwer Larva Tenelucha philippinensis (Ashmead) a

Larva Tenehicha stangli (Ashmcad)Trichomma enaphalocrosis Uchida

Larva Lara

Chalcididac Eulophidac Trichogranmmatidae

Xanthopimnpla stemmator (Thunberg) Charops hrachYpterumn Gupta and Malteswary Brachtm eria sppTetrasticttus schoenohii Ferriere Trichorigranm chihunis Ishi

Larva to pupa Larva Pupa Egg Egg

Diptera

Scelionidac Pteromalidae Tachinidae

Tt ichograintnta aponiicuitt (Ashniead) Telenomits rowani (Gahan) a

Trichoinalopsis apattelctena (Crawford) Jeirhaea spp

Egg Fgg Egg pupa Larva

Z4gobothria ciliata (Wulp) Larva

Ilymenoptera lecneumonirlac Gol fringed borer (hilo auricilius

Trichomm enoptalocrosisUchida Larva

Ieioenoptera Braconidae

Aantholimlh steiniator (Thtinberg) a

Pink stet Ioret Sesamia inferens StoInacoil nicrilhi (Bingh am)

Loarva to pupa

larva

Diptera Ilyinenoptera

Armvtnyorms Mythimna separata Spodoptera mauritia acronyetoidesTachinidae Zvgohothriaatropiora( Robinea i -Desvoid y)Eulophidae It-phctrtiscitapadoe (Ashimead)

Larva Larva

(halcididae Braconidac

Brachpin ria spip (oesia spp

Larva larva

IHyntenoptera I)iptera

Rice skipper Ielop ida (halcididae Tachinidae

math ias aidgreen iortedci ciliarNIelan itis eda Brat lhincria sp iii matuqitatt Cameron Irgyrolhirlaxi trotlhiais(marumtv)a1

ismtne Larva Larva

Diptera Ilymenoptera

Tachnidae lelineutionidae

fuown se tilo per Mois frugalisIrgtrophrlax 1igrothiahiv 3 a rat ov) anthopimpa pmnctata (Iabricius)

Larva Larva to pupa

aMost dominant

IRPS No 123 January 1987 29

Normally predatory thrips prey on herbivorous thrips strepsiptera on both Spiders dominate tilenymphaladultNatural enemies of stem bugs --Scolinoplara and Tibraa predators Spittle bugs move to rice from pasture grasses - are in order of importance egg par~isites and predators and biocontrol would be successful only ifcarried out before fungal pathogens and nymphal adult parasites Egg tiley immigrate into ricefields parasites could either be introduced fron other areas or Biological control tactics for hemipterous seed bugs mass produced and released ould follow those outlined for stein bugs

Armyworms are normally lwld in hcck by the activities Coml)arison of environnewtns Natural enemies are of egg and larval parasites When these natural enemies fail perhaps faced with even greaterconstraints than insect pestsusually because of drought armyworns become epidemic to survive ind reproduce in upland environments TheyIt may be worthwhile therefore to release parasites during have adapted ani survived by having wide host ranges the the rainy months following a drought Virus diseases of ability to aestivate or hibernate over unfavorable seasonslarvae would beanotheravnue to exploreforarnyworms and the ability to disperse or by other mechanisms

Parasites have ben traditionallyconsidered for biolegical Comparing the most dominant and therefore the most control attempts against stein borers usually by rekasing adapted species of natural enemies collected in various exotic species( 166) This approach has not met with success environments in the Philippines we can understand how and new avenues should be explored The role of predators different rice environments are arid pathogens is little inderstood and deserves greater Anagqrms olptahiisa mymarid planthopper egg parasiteattention Egg parasitism rates are normally reasonably is cquallyadapted to all Philippine rice environments (Tablehigh but should be supplemented with effective egg 6) It is selective for planthoppers and even parasitizes eggspredators Orthopterans are potential egg predators of the n-iize planthopper Peregrinitsmaidis itt upland areas Metioche vittaticollis (Sta ) Anavipha nifiwnnis (Saus- Alavohis also parasitizes maize planthopper eggs but is sure) and Conocephahs hongipnni (de i-laan) feed on most prevalent on BPH in the irrigated wetlands (onashystem borer eggs etioche and Anaxil)ha specialize in eggs tocerus spp like A optalhilis on BPH and WBPI Iattacks with no hair such as Chilo spp The omnivorous (ono- the eggs of GLH in ell rie environments It is also found cephahsspecializes in eggs cov red with hairs it is a proven parasitizing the eggs ef the white rice licahopper (oimna egg predator of Scirlouphaga spp but has the discouraging spetra Oirosita naias a trichogrammatid egg parasitehabit of eating rice guin (65) attacks BIPH and WBPH in the wetlands and has no other

Grasshoppers have egg (scelionid) and nymphalad ult host A related species 0 aesopi specializes in G1-H in (nemestrinid tachinid and sarcophagid) v irasites Among wetland environments Dryinid strepsipt-an and pipunshypathogens protozoans have been recorded most frequently culid nymphal and adult parasites of planthoppers and Control of locust species by natural enemies could con- leafhoppers also specialize either in hopper species or centrate on their habitual breeding areas environments Among the dryinids Pseuhdgonatoipsflavi-

Leaffolders have rich complexes of natural enemies some feimur prefers the lowlands and BPH while P nuch has no adapted to upland rice (22 27) ranging from egg predators environmental prefirences but is adapted only to WBPH (gryllids coccinellids) to larval parasites (braconids ichncu- The main d ryinid species on GLH aptogonatop)usspp is monids) to larval prcdators (ants and carabids) to larval most dominant in rainfed lowlands and also attacks other pathogeiis (vira and fungal diseases) Efforts to augment leafhoppers within Amnrasca Cicadulinaand Bachaltuathese natural enemies should focus on the egg predators Strepsiptera are more dominant on plarithoppers than (rearing releasing)and viral or fungal pathogens (culturing lealhoppers The Elenchus species in the Philippines attack spraying) all species of planthoppers within the genera Sogawilla

Parasitization of the large conspicuous larvae of Sogatodels NilaparsataHarualia and Opiconsiva Elenshyskippers green horned caterpillars and other polyphagous cusrvasuntaisuiis adapted to the major rice environments Lepidoptera by tachinids and chalcids is normally low ELg but prefers BPH in rainfed arid WBPIH in wetland environshyparasites and predators are perhaps the key to their control ments Ilalictoph(utsmtnroeiand HVpectrus attack G LH

Gall midges are parasitized by pterormalids and platy- and COfimravcctra respectivelygasterids that aestivate within their larval host between Pipunculids are mainly parasitic oii lealhoppers Each seasons Shoot aphids are preyed upon by coccinellids species has a unique environmental preference Tomosshysyrphids and chrysopids varireIa oraetoraprefers Ne7 otetltix nigropictrs and

Planthoppers arid leafhoppers have egg parasites and Pilunwulhtsjavanensisattacks Dehoceplhahls spp which predators as well as nymphialad nlt parasitesand predators are more abundant in upland than irrigated lowland their rich complex of natural enemies includes fungal environments T suhvirescensattacks Nephotettix virescens pathogens The principal egg parasites are myrnarids and and prefers the lowlands Pilpuiulus nttillattls is an trichogramniatids Egg predators are norinally mirid plant upland rice parasite arid attacks only Nephotetix speciesbugs that suck out the yolk Nymphal ad nIt parasites are More than 17 recorded species of leaffolder parasites in dryinids on planthoppers pipunculids on lealhoppers and the Philippines attack species of C(naplwocrocis and

30 IRPS No 123 January 1987

Table 6 Dominant natural enemies of rice pests in upland rainfed lowland and irrigated lowland rice environments of the Philippines

Natural enemy

Host stage Species

Brown planthop-Parasite Egg

MymaridaeAnagrus optabilis(Perkins) Anagrusflaveolus Waterhouse

TrichogrammatidacOligosita naiasGirault

NymphAdult Dryinidac

Pseudogotatopuslaiifemttr Esaki et Hashimoto Elenchidae

Elenchus yasurnatsui Kifune and Hirashima

Environment

Upand Rainfed lowland Irrigated lowland

Ailaparata lugens

x x X x

x

x x

x X X Whitebacked planthoppers Sogatella furcifera and Sogatodes pusanusEgg

MymaridaeAnagrus optabilis(Perkins)

TrichogrammatidacOl~gositanaias Girault

NymphAdult Dryinidae

Pseudogonatopusnudus Perkins Elenchidae

Elenchus yasutatsui Kifune et Hirashima

Parasite Egg

MymaridaeGonatocerus spp

TrichogrammatidaeOligositaaesopi Girault

NymphAdult Pipunculidac

Tomosvaryella subviresccns Loew Tomosvaryella oryzaetora (Koizumi) Pipunclusmutillatus (de Meijere)

DryinidaeIlaplogonatopusspp

Egg Copidosomopsisnacoleiae (Eady)

Larva BraLonidae

Cardiochiles philipphnensisAshnead lchneumonidac

Trichiomnia cnaphalocrosisUchida Temelucha stangli (Ashmead)

Pupa Chalcididae

BrachymeriaexcarinataGahan Brachymerialasus (Walker) Brachymeria nr tachardiaegroup

Parasite Egg

Scelionidae Telenomus rowani (Gahan)

EulophidacTettasticusschoenobhi Ferriere

Larva Braconidae

Stenobraconnicevillei (Bingham) Iclhncurnonidac

Temelucha phiiippinensis(Ashinead)Temelucha stangli (Ashmead)

x x x

x x

X x X

x x Green leafhopper Nephotettix virescens

X x x

x x

X X X X

X

X Rice leaffolders CnaphalocrocisMarasmia

x X x

X

X X

x x x x x X x x x

Yellow stem borer Scirpophaga incertulas

X X x

x

x

x X XAinauromorphaaccepta metathoracica(Ashmead) x

continued on next page

IRPS No 123 January 1987 31

Table 6 cntinued

Natural enemy Environment

Host stage Species Upland Rainfed lowland Irrigated lowland

Larva-pupa Gold fringed borer Chilo attriciiuslchneumonidae

Xanthopimpla stemmator (Thunberg) x aPupa x

Pteromalidae Trichoinalopsis apanteloctena (Crawford) ax

Pink stem borerb Sesamnia inferensParasite

Larva Braconidae

Stenobracon nicevillei (Bingham) x b b Armyworms Mvthinna SpodopteraLarva

Tachinidac Zygobothriaarroptvora (Robineau-Desvoidy) x x

Braconidac Cotesia spp

X Skipper Pelopidas mathias and green horned caterpillar Melanitis leda ismeneF-gg

TrichogramrnatidacTrichogrannaspp x

Scelionidae Undetermined Scelionid

x Larva

Tachinidae Argyroph lax nitrotibialis (Baranov) X x

Pteromalidae Trichomnalopsis apanteloctena (Crawford)

X Predator

CoenagrionidaeAgriocnenis femina femina Brauer x xIschnura senegalensis tambur

Miridae CrtorhinsliidipennisReuter x x

Nabidac Stcnonabis tagalicus (Stal) x

VeliidaeAlicrovelia douglasi atrolineata (Bergroth) x

Coccinellidae Micraspis crocea (Mulsant) X x x

Vespidae Eumenes campaniformis (Fabricius) x

Formicidac Diaecamna spp xOdontoponera transversa (F Smith) x

Carabidae Chaenius spp OphioneaW- aJasciataSchmidt-Goebel

x

Ophionea ishii ishii Habu x x X

Gryllidav Anaxipta longipennis (Saussur-) x xMetioche ittaticollis (Sial) x x x

Tettigoniidae Conocephalus longipennis (de Haan) x x x

aNo rearing made on rainfed lowland bNo rearing made on rainfed lowland or Irrigated lowland

MrasmiaAmong the Laval parasites the braconid Cardio- lowlands and parasitizes oriental maize borer OsIriniachilesphilijpphensis is most adapted to an upland environ- furnacalis (Guenee) larvae The leaffolder pupal parasitesment It also parasitizes Hydelepta indicata a common not only occur equally in all environments but also haveleaffolder of legumes The ichneumonids Trichomma cna- wvide host ranges- parasitizing species of OsIrinia Marucaphalocrosis and Temehcha stangli occur more in the Hydelepta and Honiona - in maize and legumeslowlands T stangli is most adapted to irrigated lowlands The principal egg parasite of yeltow stem borer (YSB)and also parasitizes Chilo and Scirpophaga stem borer Telenomus rowani is widely adapted to all environmentslarvae T cnaphalocrosis is most prevalent in rainfed and has an unusual alternate host -Tabanus eggs (25)

32 IRPS No 123 January 1987

Tetrastictu schoenobii specializes only in YSB and occurs mainly in irrigated lowlands -- further proof that YSB is most adapted to the wetter environments

Larval parasites of YSB are highly environment-specific The braconid Stenohracon nicevillei occurs only in upland rice but also attacks the pink stem borer (PSB) Tieinchwa phiiiplPwnsis an ichneumonid occurs in upland rice and also parasitizes the larvae of (7hiu stqpresaisinatro-morpha accepta melaihoracica like 7 stangi is mnost adapted to the irrigated lowlands

The dark-headed stem borer(l)HSB) (ih pochit3sus was not collected in all rice environments and the gold fringed borer Chilo auricilius nd PSB were collected only in upland rice Xaantiholiplastenrnaloran ichneumonid larval parasite appears to be widely adapted whereas TrihomaslopisapanitI)cteiiaa pteromalid pupal parasite occurs mostly in wetlands and also parasitizes larvae of Chilo suppressais and Pelhuidas mathias

Armyworms Myit hinin and Spodujipera have two main larval parasites - a tachinid Zrgohothriaatropivora in rainfed environments and a braconid Cotesia spp in irrigated lowlands

The butterfly pests Pelopidas and 4elanitishave tricho-grammatid egg parasites found only in irrigated lowland environments

Two larval parasites show environmental preferences --Argrroph av nigrotihialis a tachinid in rainfed habitats and Trichonialopsisapantelkcnela in irrigated ricefields A limgrotibialisalsoattacks the sweet potato hornwormAgrius convolvtdi and T apanteloctena parasitizes DH S1B

Predators as a group show more distinct environmental preferences The aquatic damselflies Coenagrionidae are most abundant in the lowlands Cvriorhinus is also most adapted to the lowlands The nabid Stenonabis tagalicus occurs only in the uplands and isalso prevalent on legumes The coccinellid lady beetles wasps and ants are more prevalent in upland areas AlicrasIpis cit-ea larvae prey ona variety of aphids attacking legumes and maize Etunienes campanonia vespid mud wasp makes nests in trees and is therefore most adapted to the more botanically diverse uplands Soil-dwelling ants cannot tolerate flooding

Upland environments are habitats to arboreal carabid beetles Three species of Chlaeniusbeetles prey on leaffolder larvae and are perhaps more important than parasites (27)

The gryllids and tettigoniids are widely adapted egg predators not only on rice but also on maize and legumes They feed on eggs of most insects that are laid on leaves

The spidercommunity of upland rice environments isrich in species At one site (Batangas Philippines) 31 species have been recorded (23) Of the 176 spider species recorded in Philippine ricefields about one-half (82) occur in upland rice The spiderspecies of upland rice environments overlap more with those of rainfed lowland than with those of irrigated lowland rice (24) The same study showed that of the three environments irrigated rice has (ie greatest spider species diversity foiiowed by rainfed lowland and upland

However of the 10 most prevalent ricefield spiders 9 were abundant in upland rice showing wide adaptation All three spider guilds -- orb-web space-web and huting spiders shywere prevalent in upland ricefields There were however differences in environmental prelerences for some spider species

Of the orb-wcavers Tetragniathamanlihulatawas partishycila rlv abundant 7 japonica was low and Ltucatge dehcorata was absent in upland rice

As in other environments Aiypeiwnaifirtosana was the most dominant space-web spider however its relative numbers were lowerthan for lowland sites Among hunting spiders Ox|opes javanus and Lcosa leatcost fgla were more abundant and Lvcosa lpseudoanmnlata was less abundant in upland compared with lowland rice envishyroniments

Extensive rearing of rice insect pests in upland rainfed lowland and irrigated sites over an 8-yr period in the Philippines provided insight into the effectiveness of parasites as natural enemies (Table 7 8) Most of toe data are from large sample sizes taken over at least one crop which overcomes the pitfalls expressed by Van Driesche (238)

VSB egg parasitization was surprisingly similar across all environments Larval parasitization was highly dynamic among sites and even years within the same site but showed somewhat lowerlevels in uplandsthan in irrigated wetlands This would have been predicted for egg parasites as well since YSB is adapted to the lowlands and so must be its parasites This isevidence that Telenomus rowani is as good at dispersing as its main hosts Trowani clings onto the anal tufts of female YSB and parasitizes eggs as they are laid This phoresy may explain its wide adaptation The more specialized lowland YSB larval parasites therefore appear more effective than their upland counterparts with more alternate hosts

Low collection levels of PSB larvae especially in the wetlands make comparisons between environments difficult Low levels of larval parasifization were recorded in both rainfed environments Except in the Cagayan rainfed wetland site in 1980 low levels ef larval parasitization were evident at irrigated and rainfed sitesalike No differences by environment were evident Higher levels of parasitization occurred in Sarawak during a 1967 outbreak also from tachinids Rothschild (197) indicated that higher parasishytization occurred in upland rice areas

Skipper and green horned caterpillar larval parasitization rates were generally low and showed no environmental effect htighest activity occurred in Cagayan in 1980 (48 parasitization) Although larval parasitization levels of RLF were variable year to year activity appeared greater in the irrigated lowlands than in the uplands Again the highest levels were recorded in Cagayan in 1980 Cagayan has extensive grasslands and fallow around the rice areas and a relatively long rainfall period Rainfall in 1930 was good for crop growth and apparently for increase of larval parasites

IRPS No 123 January 1987 33

Table 7 Parasitization rates of rice planthoppers and green leafhopper in upland rainfed lowland and irrigated lowland environments Philipshypines 1976-840

Environment and site Year Egg parasitization () Nymphaladult parasitization ()

BPH W13P GLH BPII WBPH GLIH

UplandBatangas

Rainfed lowland

1976 1979 1980 1982

15 17 12

2(47)8(39)

4(51)

9(1800)14 (212)

12(304)

8 (101)6 (127)

9(116)

Iloilo

Cagayan

1976 1977 1978 1981 1980 1981 1982

0 (69)

11

44 48 40

-

-

46 55 44

0 (471)

11

52 56 49

0 (200)9 (214)

10 (1259)8(235)6 (900) 3(800)

-7 (462)

11 (270)3 (313)5 (900) 1 (800)

4 (600)10 (725)10 (362)12(870)5 (900) 3 (800)

Pangasinan

Irrigatedlowland

1984 1976 1978 1979 1982

13 (436) 13(379) 14 (457)

6(14)7 (197)

17(198) 11 (486) 18 (275)

3 (81)10 (274)3 (116)

16(871) 19(88)

28(211)9 (764)5 (647)7 (1204) 8(305)

Laguna 1977 11 7 21 1978 31 1979 14 4 4 28 1980 7 8 20 1981 1983

56 7(278)

26 16 (1650)

16 27(1921)

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R I Apostol unpublished 1lo1lo and Pangasinan 136 Cagayan BCanapi unpublished Liuna 175 I C Pantua unpublished 118 234 43 Carino and Shepard unpublished BPH = brown planthopper Nila parvara htgens W131Pi =whitebacked planthopper Sogatelia furcifera GLH = green leafhopper Nephotettrix virescens Figiires in parentheses are sample sizes

Tauher et al (230) reported that internal external and genetic factors influence the seasonal activity of parasitoids

Hopper egg parasites but not nymphaladult parasites were also parlicularly abundant at the Cagayan site possibly the egg parasites have more alternative hopper prey in the weedy areas than nymphaladult paisites

The seasonal dynamics of hopper egg parasites can be seen for upland rice in Batangas(Fig I 2) and for rainfed lowland lice in Pangasinan( 135)and Cagayan( 105) Levels of BPHWBPI-L and GLH egg parasitization were corn-parable in upland and irrigated lowland environments Except for some high rates of paraMsitization early in the season the parasitization rate tended to be steady in all envilonments However hopper nymphaladult parasi-tizution levels were more similar in the rainfed environments and lower than in irrigated rice

Overall parasites do not appear highly effective by themselves against upland or even lowland pests Among natural enemies parasites have attracted the most attention because they can be more readily assesscd

Predators and pathogens lend themselves better to management practices Predators the most important group of natural enemies in riccfields are difficult to quantify On the other hand pathogens appear less im-portant but can be readily cultured and disseminated

The natural enemy commulnity of upland rice is rich in species and differs significantly from that of lowland rice

These beneficial organisms must be conserved by applying insecticides judiciously particularly sprays Seed or soil placement of chemicals minimizes exposure to natural enemies The strategy to derive the greatest benefit from natural enemies isto allow the greatest number of beneficial species to thrive Then perhaps at least one species will be effective against each pest at any tine overcoming the variable abundance of each species within and between years

Programs to introduce exotic species or mass produce indigenous ones are ambitious and expensive because they require trained people on a sustained not ad hoc basis

Chemical control Insecticides offer rapid and efficient control of upland insect pests (190) but-should be used only after other control measures have been considered Insecticides are rarely used on upland rice because of cost (32) Upland rice yields are normally too low to justify the expenditure Also most upland rice is grown as a cash crop Spraying upland rice is more difficult than lowland rice because water is less accessible Government subsidy programs to provide pestishycides to tribal peoples for upland rice production have created problems of toxicity to humans because isolated people have had no experience to be able to handle pesticides safely

However ins-eticide use has beenjustified when a crop is

0Z

Table 8 Parasitization rates of lepidopterous pests in upland rainfed lowland and irrigated lowland environments Philippines 1976-84a

Parasitization (5)

Environment and site Year YSB GFB PSB Earcutting Swarming Skipper Greenborned RLF Brown larva larva caterpillar caterpillar caterpillar caterpillar larva semilooperEgg Larva larva larva larva larva larva

UplandBatangas 1977 - 0 (57) - 0 (63) - 0 (71) - 0 (106) 11(142) 2 (101)1978 43 (4697) 8(451) 5 (38) 3 (40) 6 (61) 2(58) 15 (57) 13 (129) 13 (1286) 4 (30)1979 59(4135) 12(320) 13 (11) 5(28) 9(45) 6(46) 7(23) 8(160) 9(1152) 8(26)

Rainfed lowland Iloilo 1976 - 0 (10) 0(10) 0 (7) 0 (19) 2 13) 15 (180) 5 (76)1978 0 (54) 0 (3) 0 (17) 9 (64) 4 (78) 3 (73) 0 (121) 0 (191)1979 64 (2350) 16 (84) 6 (17) 0(6) 18 (68) 25 (104)

1981 56 (1322) 18(304) 0(5) 12(51) 18(138)Pangasinan 1976 60 (7 7 )b 62(82) 0(111) 5 (39)

1978 58 (53)b 5(51) Cagayan 1979 59(713) 8 (101) 6 (34) 0(7)

1980 0(11) 4 (28) 15 (286)

62 ( 48(177) 61(140)1981 0(5) 12 (2) 13(45) 14 (92)

Irrigatedlowland Laguna 1974 62 8

1974-75 50 41 1975 43 72 1978 96 (623) 21(24) 33 (60)1979 83(2017) 18(110) 8(62) 11 (89) 22(400)1980 78 (1625) 20(75) 0 (16) 4 (72)1983 60

9 (80) 7 (270) 30

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R F Apostol unpublished Iloilo and Pangasnan 136 Cagayan B Canapi unpublished Laguna 175 P CPantua unpublished 118 234 143 Carino and Shepard unpublished YSB = yellow stem borer ScirpophagaincertulasGFB = gold fringed borer Chilo auricilius PSB = pink stem borerSesamia inferens earcutting caterpillar Alythimna separata swarming caterpillar Spcdoptera mauritia acronyctoides skipper Pelopidas mathias reen horned caterpillar Melanitls ledaismene RLF = rice leaffoldersCnaphalocrocismedinalis and Marasmiaspp brown semilooper Mocisfrugalis Figures in parentheses are sample sizes Egg masses

IRPS No 123 January 1987 35

Table 9 Effect of seeding rate and insecticide on plant density and Yield losses from insect pests in upland rice often areyield of Dagge upland rice Batangas Philippines 1976a comparable to those of lowland rainfed or irrigated rice

Seeding rate Insecticide Dosage Plant density Yield (0-Y30o) but usually from different pests The principal (kg aiha) seed treatment (kg aiha) (noIra-row) (tha) groups are soil pests feeding on sown seed and rootsat 14 DE followed by seedling pests These groups have largely gone

50 None - 35 a 20 a unnoticed leading researchers to conclude that stem borers50 Carbofuran F 10 38 ab 23 a and seed bugs are the most important pests (17) Yield loss 10 beCabofuranF100 None - 4551 c 23 a trials that pinpoint major pests need to includc seed

100 Dieldrin WP 10 52 cd 22 a treatments Granules applied in the furrow may not give150 None - 59 d 22 a adequate protection from many seed pests

aAv of 8 fields DE = days after crop emergence In a column means The floristic and pedological diversity of upland rice followed by a common letter are not significantly different at the environments is matched by the diversity of arthropods5level by D)MRT F = flowable Wi = wettable powder both pests and their natural enemies Irrigation has homoshy

genized lowland rice environments Flooded soils tend tothreatened by epidemics of locusts or armyworms New have the same properties and to eliminate soil pests The technology stressing efficiency has minimized insecticide spatial and temporal dominance of cultivated rice has cost benefited specialist pests

Seed treatment is normally inexpensive and can be Upland rice has no specialist arthropod species The economicallyjustified in -nanycases(66) Treating seed with monophagous pests on upland rice prefer the lowlands systemic insecticide (216) but not chlorinated hydrocarbons where rice evolved Rice in the uplands exists only because (247) is most effective in controlling seedling maggots of man and even though lowland rice predates upland riceFoliar sprays have to be repeated to obtain control and specialist species evolved Wheat maize and sorghum do often are washed off by frequent early season rains not have monophagous pests leading us to conclude that Granular insecticides are effective but required dosages are there is no evolutionary advantage for inssects to have too expensive to justify except under heavy infestation narrow host ranges in upland graminaceae(247) When directed against first-instar larvae banding Pests have adapted to the uplands not only by having granules in seed furrows (177) can be effective against white wider plant host ranges but also by having long life cyclesgrubs at low dosages (025 kg aii ha) (134) Seed treatments undergoing dormancy or being dispersive Each pest has a are not promising for white grub control (248) unique life history strategy suited to highly unpredictable

Seed treatment is also effective ainst ants and may be environments In agivenyear someinsect specics an better cheaper than increasing the seeding rate In Batangas exploit upland rice as a food source but no pest is a senious treating 50 kg seed ha was as effective as increasing the one every year on upland rice seeding rate to 100 kgha and using untreated seeds (Table The low yielding potential of upland rice means less 9) In Brazil seed treatments control termites and other soil quantity and lower quality of food for pests to exploit but insects including the lessercornstalk borer(66) Domiciano also less incentive for farmers to undertake control (58) however found seed treatments inconsistent in their The most effective control would be early and synchroshyeffect against high populations of lesser corn stalk borer nous planting of early-maturing upland rice varieties to

Baiting can be an inexpensive way of controlling ants prevent insect pest buildup Breeding for insect resistance seedling maggot flies mole crickets and field crickets Baits should initially focus on stem borers because they are verymade of locally available material and impregnated with difficult to control by other means Biocontrol efforts insecticide can be sparingly distributed taking adantage of should first stress conservation of natural enemies preshythe pests ability to disperse and encounter bait sites dators and pathogens rather than parasites Efficient

The new low-volume hand-held controlled-droplet chemical control includes seed treatment or baiting for seed sprayers may offer an advantage over high-volume knap- seedling and some root pests and using controlled dropletsack sprayes for controlling foliar pests Upland rice sprayers for spot treatments against foliar pests and seed canopies are more open than those in lowland rice and bugs when economic thresholds are reached better droplet penetration should result Controlled droplet applicators would be ideal for seedbugs and defoliators

CONCLUSION

Upland rice is attacked by a wider array of insect pests than is lowland rice mainly beca use of the addition of soil pests Generally population levels of lowland rice pests are lower in upland than lowland rice leading many people to conclude that insect pests are not important on upland rice

36 IRPS No 123 January 1987

REFERENCES CITED

1Abraham C C and K S R Mony 1978 Occurrence ofLeptocorisa acuta Fabr (Coreidae Hemiptera) as a pest of nutmeg trees J Bombay Nat ttistSoc 74553

2 Agyen-Sampong M 1975 HlierogrlphusdiaganensisKrauss (Orthop-tera Acrididae) a new pest of rice in Northern Ghana Ghana J Agric Sci 8249-253

3 Agyen-Samnpong M 1976 Varietal resistance and other inle-ratcd pest control components as factors of rice varietal improvement in West Africa Prcsented at the Second varietal improvement serinar 14-18 Sep 1976 Bouak Ivory Coast West Africa Rice Development Association Monrovia Liberia 18 p

4 Agyen-Sampong M 1977 Outbreak of Ihccrorrdnisorae in rice seedlings near Rokupr Sierra Leone lit Rice Res Newsl 2(6)16-17

5 Agyen-Sampong M 1980 Dirty panicles and rice yield reduction caused by hugs IntRice Res Newsl 5(l)11-12

6 Agyen-Sampong M 1982 The major pest problems of irrigated upland and mangrove swanip rice ecosystems ir the humid tropical Guinea savanna and Salel climate zones Pages 318-328 in Pro-ceedings training course cn concepts techniques and application of irtegrated pest management in rice inWest Africa 10-20 Jan 1982 Fendall Liberia West Africa Rice l)evelopioent Association Monrovia Liberia 505 p

7 Akibo-Betts V I and S A Raymundo 1978 Aphids as rice pests in Sierra Leone lit Rice Res Newsl 3(6)15-16

8 Akinsola F A 1975 Present status of different rice stemborers in parts of Nigeria iicL Entomo Newsl 328

9 Akinsola FA 1979 Ihe biology and ecology of rice stemborers in Nigeria Ph 1) thesis University ol lbadan Nigeria 217 p

10 Akinsola E A 1979 Guidelines of integrated management of insect pests of upland rice in West Africa Presented at the Seminar on integrated management of rice diseases and insect pests at Bobo-Dioulasso Upper Volta 17-22 Sep 1979 West Africa Rice Develop-ment Association Monrovia Liberia

II Akinsola IA 1984 Insect pests of upland rice in Africa Pages 301-305 in An overview of upland rice research Proceedings of the 1982 lIouak Iwsrv Coast upland rice workshop International Rice Research Institute 13O Box 933 Manila Philippines

12 Alam M S V TJohn and Kaung Zan 1985 Insect pests and diseases of rice in Africa Pages 67-82 in Rice improvement in Eastern Central and Southern Africa proceedings of the international rice workshop at Iusaka Zambia 9-19 April 1984

13 Alticri M A A van Schoonhovcn and J D Doll 1977 The ecological role of weeds in insect pest management systems a review illustiated with bean (Phaseohsvulgaris L)cropping systems PANS (Pest Artie News Summ) 23195-205

14 Anmante F 1967 The leaf-cutting ant Atia capiguaraa pest of pastures [in Portuguese] Biol6gico (Sao Paulo) 33113-120 (Rev Appl Entomol A56207)

15Andrade A C and D Puzzi 1951 Preliminary results of experi-merits on the control of Sculptcoris iastantus on sugarcane [in Portuguese] Bioulgico (Sao Paulo) 1744-49 (Rev Appl Entomol A41421422)

16 Appert J 1970 Maliarplia separaelha (rice stem borer) new observations of entomological problems of rice in Madagascar [in French] Agron Trop (Paris) 25329-334

17 Ariadeau Mand Z Harahap 1986 Relevant upland rice breeding objectives Pages 189-197 in Progress in upland rice research Proceedings of the 1985 Jakarta conference International Rice Research Institute 1O Box 933 Manila Philippines

18Asar S and Kisimoto 1975 Seasonal occurrence of the smaller brown planthopper in upland rice [in Japanese] Proc Kanto-Tosan Plant Prot Soc 2228

19 Bakker W 1974 Characterization and ecological aspects of rice yellow mottle virus in Kenya Meded Landbouwhogesh Wage-ningen 152 p

20 Balut F F 1970 White grubs attacking rice Oryza ativa L [in Portuguese] Biol6gico (So Paulo) 36321-322

21 Banerjee S N and L M Pramanik 1967 The lepidopterous stalk borers of rice and their life cycles in the tropics pages 103-124 in The major insect pests ofthe rice plant Proceedings ofa symposium at the International Rice Research Institute September 1964 The Johns Hopkins Press Baltimore Maryland USA

22 Barrion A T and J A Litsinger 1980 Ants a natural enemy of cliaiarac in dryland rice IntRice Res Newsl

5(4)22-23 23 Barrion A T and J A litsinger 1981 The spider fauna of

Philippine rice agroecosystem 1Dryland Philipp Entomnol 5139-166

24 Barrion A T and 1 A Iitsinger 1984 The spider fauna of Philippine rice agroccosystcms II Welland Philipp Eiitomol 611-37

25 Barrion A T1 and J A Litsinger 1984 lihanus (Diptera Tabanidae) eggs an alternative host of rice stei borer (SI3) egg parasite Telenoonis digns (Iliymenoptcra Scelionidae) lit Rice Res Newvsl 9(6)19

26 Barrion A T and J A Litsinger 1985 Identilication of rice leaffolders by wing markings Int Rice Res Newsl 10( 1)24

27 Barrion A T and J A Litsinger 1985 Chctiosspp (Coleoptera Carabidae) a leaffolder ILF)predator lit Rice Res Newsl 10(1)21

28 Bertels A 1970 Insect pests of rice and theircontrol [in Portuguese] Pelotas Brazil Instituto de Expcrimentacao Agropccuararias do Sul Circular 43 24 p

29 Bonzi S M 1982 Ctilo (flisilineus J de Joannis (Lepidoptera Pyralidae) stem borer of irrigated cereals in Upper Volta LioFrench] Agron Trop Riu Rizic Cult Vivrieres Trop 37207-209

30 Bourke T V T L Fenner J N LStihick G Baker EHassan D F OSullivan and C S li 1977 Insect pest survey for the year ending 30th June 1969 Papta New Guinea )epartment of Agrishyculture Stock and Fisheries Port Moresby 57 p

31 Bowden 1 1973 Migration of pests in the tropics Meded Rijksfac Landbouwwet Gent 38785-796

32 Breniere J 1973 The main rice parasites in West Africa and their control Pages 27-42 in Proceedings seminar on plant protection for the rice crop West Africa Rice Development Association Monrovia Liberia

33 Breniere J 1976 The principal insect pests of rice in West Africa and their control West Africa Rice Development Association Monrovia Liberia 52 p

34 Brown ES 1962 The African arinyworn Spoiolptera eXI)Ina (Walker) (Lepidoptera Noctuidae) a review of the literature Cornshymonwealth Institute of Entomology London UK57 p

35 Brown F B 1968 Upland rice in Latin America FAO-International Rice Commission 12th Session of the Working Party on Rice Production and Protection 2-14 Sept Peradeniya Sri lanka Food and Agriculture Organization Rome Italy 6 p

36 Buyckx F J 1 1962 Disease and insect pests of crops inCongo Rwanda and Burundi [iil French] Publications dc LInstilut National pour lestude Agrononiquedu Congo M Weissenbruck S A Brussels Belgium 708 p

37 Cariiio F 0 1981 Role of natural enentics inpopulation suppression and pest management of green rice lealhoppers Ph D thesis University ofthe Philippines at Los Bafitos PhilippinesLaguna 136 -

38 Carpente A J 1983 The development of rice cultivation and extension of vegetable productin in Zanzibar Final Report Vol II1 October 1983 Food and Agricultme Organization Ronte Italy

39 Carroll C P and S1 Risch 1983 Tropical annual cropping systems ant ecology Environ Manage 751-57

4(0 Cates R G 198 1 Host plant predictability and the feeding patterns of monophagous oligophagous and polyphagous insect herbivores Oecologia (Berl) 48319-326

IRPS No 123 January 1987 37

41 Cendcafia S M and F B Caora 1967 Insect pests of rice in the 62 Ene 1 C 1981 I)iiorldhol)ierusihilis Slater (licteroplera Philippines Pages 591-616 in The najor insect pests ofthe rice plant lygacidue Illissinac) a new rice pest front Nigeria lit Rice Res Proceedings of a symposium at the International Rice Research Newsl N3)15 Institute September 1964 The Johns Hopkins Press Baltimore 63 Fugenio T S 1965 rhe effect oi e-vCgetation on rice production Maryland USA 729 p and the possibility of overcoming it by leaf pruning MS thesis

42 Centro Ihternacional de Agricklittira Tropical 1984 Upland rice in University ol the Philippines at Los Baijos ILaguna Ihilippins 69 p Iatin Anerica Pages 93-119 hi An os ervies of uplaind rice research 04 Farrow R A 1975 The Alrican nmigratory locust ii is main Proceedings of the 1982 Boinakamp Ivory Coast upland rice workshop outbreak area of the middle Niger qlinlitatise studies of solitary International Rice Research Institute P0 Blox 933 Manila populations in relation to environmental factors Locusla No II Philippines 198 p

43 Chandra G 1979 Taxonomy ad hn riononlics ofthe imect parasites of 65 Fennal R G 1943 Food crop investigation Windward and rice Icalloppers and planthoppers in the Pihilippines and their Ieeward Islands First report Report onl work dunring the periodiiiportairce iii natural biol)gical cointrol Philipp Fntoltol 4119-139 April 1942-April 1943 Imperial College of Tropical Agriculture

44 Chang T T 1976 The originii colut ion enlIivation dissemination Trinidad 16 p (Rev of Appl Fintornol A3222-25) and diersificition of Asian and African rices -uphytica 25425-44 1 66 Ferreira F 1983 Integrated pest ritaageiret of upland rice [ill

45 Chatierjee P B 1979 Rice leaffolder attacks in India lit Rice Res Portuguese English summary] Pages 323-341 in Upland riceculttre Newsl 4(3)21 factors affecting production E Ferreira T Yamada and F-

46 Cheng Cbien-pan 19(4 Upland rice in Taiwan International Rice Malmolta eds Institutoda Potassae Foslato Institnto Internacional Cnmmission Wiorking Party oit Rice Production ard Protection I()th da Potassa Piracicaha Sao Paulo lraid Meeting 3-10 March 1964 Manila Philippines Food and Agricul- 67 Ferreira E arid F dl Silva Martins 1979 Occurrence of ture Organiaion Rome Italy 5 p Sogatohs orvziola (Muir) inupland rice in Goias Ilrail lit Rice

47 Chcrrett 1 M and 1) 1Peregrine 1976 A review of the status of Rcs Ncwssl 4(4)18 leaf-cutting ants and their control Ann Appl Biol 84124-128 68 Ferreira E and 1F da Silva Martiris 1984 Insect pestof rice ill

48 Chipeta F M 1974 Pests of rice in Malawi Pages 90-95 in Brazil and their control Jill Portugucsej FMIlRAtPA-CNPAF Proceedings Sith eastern Africa cereals research conference 10-15 Goiania Brmiil 67 p March 1974 DRB Mialda ed Ministry of Agriculture lilongwe 69 Ferreira FI F ca Silha Miartins S S Nero and 1 It P de Malawi Carvallio 1982 M tltiplc regression analysis todeterimine the effect of

49 Chu Yau-i 1979 The d atiaging period of the rice seedling fly insect populatiois ott upland rice yield [inIPortuguese Pesqui (Atlterigotiavxigu Stcin) (NItiscide Diptera) oil cori) at Fast Java Agropecu Bras 17671-675 Indonesia Plant Prot Bill 21403413 70 Ferreira F 1F da Silva Martins and F 1PZimmermann 1979

50 Conklin H C 1957 IHaninoo agriculture a report on aln integral Rice varietal resistance to tle lesser corn stalk borer [in Port uguese] systent of shi fti rig cult ivationi in the Pltilippincs Food and Agriculture Pesqui Agropccu Biras 14317-321 Organization Rome Italy 209 p 71 Fleming W F 1968 Biological control of lie hapanese beetle U S

51 Cramer H II 1967 Plant proection and world crop production Dep Agric RLs Sers [ech Bull 1383 78 p Pilaniensclhutz-Nachr 201-524 72 Food and Agriculture Organiation of the United Nations 1982

52 l)ani R C and Majunidar 1978 Prelininary observations on tie Country report surmimary Pages 27-37 in Report on the first incidence of root aphids on differeit rice cultnrcs Sci Cult 4488-89 FAOIUN)IPhail and regional training coiuirse in iniproved ctltural

53 Davidson R I 1979 lffects of root feeding on foliage yield Pages practices for upland rice 5-23 Oct 1981 Northern Region Agrculturd117-120 in T K Ci osb and R B Pottinger eds Proceedings of the Developririt Center Chiang Mai Food and Agricuilture Organiiashy2nd Australian conlerence on grassland invertebrate ecology Govt tion Rome Italy 37 n Printer Wellington New Zealand 73 Franco do Ariaral S 195 1 Residual action of insecticides in turfy

54 l)ean G 1W 1976 Rice insect pests inlaos Int Rice Res Newsl soil--the results of a field experiment [in Portuguese] Biol6gico (Sao 1(2)15 Paulo) 17209-214 (Rev Appl Entomol A41423)

55 I)ean G J W 1978 Insect pests of rice in Laos PANS 24(3)280-289 74 Freeman ) 1970 Report onithe Iban london School of -coroiics 56 )ingle H 1984 Behavior genes and lifc histories conplex adapta- Monographs on Social Anthropology No 41 Athlane Press

tions to uncertain environments Pages 169-194 in A new ecology London UK 317 p novel approaches to interactive systems P W Price C N Slobod- 75 Fuchs T W1 A Harding and 1W Smithlr 1979 Induction and chikoff and W S Gaud cdsl ohn Wiley New York USA termination oldiapause in the sugarcane borer Ann Flitonlol Soc

57 Dobelmann-JP and M Falais 1968 Upland rice cultivation ill tie Am 72271-274 North-western part of Madagascar in 1966-1967 [in French English 76 (hbriel B1) 1975 A review tf the major insect pests of some tpland summary] Trop Agron 2313-21 crops in tlc Philippines Pages 86-111 in Ad Htoc panel of experts on

58 Doniiciano N L 1979 Chemical control of tie lesser corn stalk pest disease arnd weed problems in some rainfed crops Bangkok (lolsonno~paloIgnowslhts) (Zeller 1848) (I-epid(ptera Ihycitid ae) Thailand Food aid Agriculture Organization Roire Italy

on rice in Ponta Gressa Parara 1977-1978 [in Portuguese] Pages 77 Garg D K and N K Shal 1983 White grub outbreak on rainfed 291-303 in Anais 1 Reuniao de tecnicos cmiirizicultura do estado de dryland rice in Uttar Pradeslh Int Rice Res Ncwsl 81(4)16-17 Sao Paulo-Campinas Cati Brazil 78 Garg I) K and IP Tandon 1982 Major insect pests of rice on hilly

59 Dunsmore 1 R 1970 Investigations on tlie varieties pests and tracts of Uttar Pradesh India lilt Rice Res Newsl 7(11)11-12diseases of upland rice (hill padi) in Sarawak Malaysia Int Rice 79 Garrity 1) P 1983 Upland rice research an international bibliograpy Comm Newsl 1929-35 1975-1982 Iternational Rice Research hnstitnte PO Box 933

60 Elias 1R1965 The new decline of rice seedlings is not caused by Manila Philippines 121 p parasites [in Portuguese] Biol6gico (Sao Paulo) 313-6 80 Ghauri M S K 1982 A new subspecies of Iinorh)rone bull

61 Empresa Brasiliera de Pesquisa Agropecuaria 1984 Upland rice itt cornlus Slater (Hemiptera Iygaeidae) front Papua New Guinea on Brazil Pages 121-134 it Av overview of upland rice research rice and carpet grass Ihll lEntoniol Res 72133-137 Proceedings ofthe 1982 Blouak Ivory Coast Upland rice workshop 81 Gonzales F 0 PArregoces R I ihernandef and OT Parada International Rice Research Institute PO Box 933 Manila 1983 Insect and imite pests of rice arid their control in Latin America Philippines [in Spanish] la Federacion Nacional de Arrocerts (Fedearroz)

Bogota Colombia 60 n

38 IRPS No 123 January 1987

82 Grist D H and R J A W Lever 1969 Pests of rice Longmans London UK520 p

83 Hale P R 1979 Two subterranean pests of upland rice in Papua New Guirnca IntRice Res Newsl 4(2)16-17

84 Hale P R and J T Hale 1975 Insect pests of rice in the New Guinea Islands Rice Entomol Newsl 34

85 Harris W V 1969 Termites as pests of crops and trees Common-wealth Institute of Entomology London UK 41 p

86 Hattori 1 1968 A method to distinguish between grass leaf roller (Ctaplha rocis medinalisGuenee) and the rice leaf roller (Susurtia exigua Butler) [in Japanese] Plant Prot (Japan) 23167-170

87 Henry I E and 1A Onsager 1984 Experimental control of the mormon cricket Iahrosshnph by NJsena locustae(Aficrospora Microsporidae) a protozoan parasite of grasshoppers (Ort Acri-didae) Entomophaga 27197-201

88 Hidaka 1 P Vungsilabulr and SKadkao 1974 Studies on ecology and control of the rice gall midge in Thailand Tech Bull 6 TropicalAgricultural Research Center Tokyo Japan 113 p

89 Hincklcy A 1) 1963 Ecology and control of rice planthoppers in Fiji Bull Entomrol Res 53467 481

90 Hirao J 1982 Ecology and chtInical control of the rice leaffolder Jpn Pestic Inf 4114-17

91 Ho DT 1984 Sten borer incifience in rice ecosystems in KenyaEast Africa Pages 307-320 in A overview of upland rice research Proceedings of the 1982 Bouac Ivory Coast upland rice workshop International Rice Research Institute PO Box 933 Manila Philippines

92 Ho )T J 1Njokah and J GKibuka 1983 Studies on rice sten borers in Kenya with emphasis on ifaliarpiaipiraellRag Insect Sci Appl 465-73

93 Ichihara I T Ito A NIatsuo and M Sawada 1971 Chemical control of the rice striped stem borer in upland rice grown in plastic mulch [iii Japanese] Proc Kanto-Tosan Plant Prot Soc 1882

94 IclhiharaIand A Matsuo 1969 Damage caused by rice striped stett borer to upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1686-87

95 Ichhara I atid A MIatsun11970 Conside ration oft lie extraordinary prevalence of rice striped stem borer in upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1778-79

96 Institut de Recherches Agrolnliqes Iropicale et des Cultures Vivrieres 1978 Insect resistance Pages 12-13 in Upland rice 1978 atnual report Bouak Ivory Coast 27 p

97 International Institute for Tropical Agriculture 1983 Aphids itt rice roots I ITA Res Briefs 4(1-3)4-5

98 International Institite fi Tropical Agriculture 1983 Hopperburn in rice caused by brown plathopper IITA Res Briefs 4(l-3)6-7

99 International Rice Research Institute 1975 Annual report for 1974 PO Box 933 Manila Philippines p 216

100 International Rice Restarch Institute 1976 Annual repiort for 1975 PO Box 933 Manila Philippines p 228

101 International Rice Research Institute 1979 Cropping systems program Component technology development and evaluation Page 404 i Annual report for 1978 11O Box 933 Manila Philippines

102 International Rice Research Institute Department of Entomology 1981) Insecticid evaluation 1979 Tables 73 74 PO Box 933Manila Philipes

103 International Rice Research Institute Department of Entonmology 1981 Insecticide evalomtion 1980 Table 81 POBox 933 Manila Philippines

104 International Rice Research Institute 1981 Report of a meeting of Asian rice-based cropping systems entomologists Eleventh cropping systems working group meeting 18-22 May 1981 PO Box 933 Manila Philippines p 132 133 144

105 International Rice Research Institute 1984 Cropping systems program Pages 368-373 in Annual report for 1983 PO Box 933 Manila Philippines

106 International Rice Research Institutc 1984 Upland rice in Asia Pages 45-68 in An overview of upland rice research Proceedings of the 1982 Bouakamp Ivory Coast upland rice workiiop loternational Rice Research Institute 11O Box 933 Manila Philippines

107 Irshad M 1977 Grasshoppers in rice in Pakistan lit Rice Res Newsl 2(3)8-9

108 Ito R and TOno 1964 Upland lice its culture and varieties in Japan lIternational Rice Cormmission loth Meeting of the Working Party on Rice Production and Protection 3-10 March 1964 Manila Philippines Food and Agriculture Organization Rome Italy 12 p

109 Jeffrey M I and V A Dyck 1983 Effect of moonlight on lightshytrap catches of brown planthopper lit Rice Res Newsl 8(6) 13

110 Jennings P R and T Alicia Pineda 1983 Management ofthe rice plnthoppers Sogatodsorivi(coIl in Latin America Presented at the meeting on stable plant resistance FAO and lIndocsian I)inectorate of Food Crops Protection Bali lndonesia 29 Novcniber-2 l)ccemshyber 9 p

IllJimenez Gomez J A 1979 Rice insect pests i lla ios ()riCntales lil Spanish] Pages 99-103 in Rice ciltivation Instituto (olotibiatto Agropecuario Compndio No 29

112 Jongeleen F JJ Soesanto and A Mockidjo 1979 Studies oti pathogens of two species of Scarabacidac froinIndonesia Z Angew Enlomnol 87412-417

113 Jouvenaz 1) PC S lofgren and W A Baiks 1981 Biological control of imported fire antsa review of cirrent knowledge Bull Entomol Soc Am 27203-208

114 Kainoh Y K Shiniiu S MarUani Y Taiaki 1980 lost finding behavior of tie rice bu lplotori+JchinvittjI)allas (liciniptera Corcidac) with special reference to diet pattern ol aggregation and feeding on the rice plant Appl Entomol Zool 15225-233

115 Kakkar K 0 P Sharma and P I Sharma 1977 Note on Ihteronyrchus liodresRedt (Coleopicra I)ynastidae) as asporadic pest of paddy and maie inHimachal Pradesh Corr Agric 156-57

116 Kalode M131 79 Major insect pest problems and their control in upland rice (Unpublishcd mineogriph CWI oJNaaiat lni Chandaratat (198 1)original not seen)

117 Kalshoven L i 98 I The pests of crops in I ldoniesia (translated by P A vatn der Laan) 1 T Ichtiar B3aru-vati Iloeve lakarta Indonesia 701 p

118 Kamal N Q 1981 Suppression of whitebacked planto ner Sogatella fircaifra (Horvath) aLId rice lealfolder (nhal -cis licdb lis (Guenee) populations by natural cnclmie hlI) tiesis Grcgorio Araneta University Foundation Victoncta Park Metto Manila Philippines 179 Ip

119 Karim A N M Reaul 1975 A species ofliealy bug Trioni i70s ceres new oii rice in the Philippines Rice Entoniol Newsl 247

120 Katanyukul Wand C Chandaratat 1981 Insect pestsof uplaid rice and their control Report firstFAOUNlFi Tliailand training course on improved cultural practices for tplad rice Chiang Mai Thailand 5-23 Oct 1981 Food and Agriculture Organiation Rome Italy 22 p

121 Kern F 1955 lhletwioa(I)uninemts) bhhit Biturm(Coleoptera) a new pest of sugarcane inVenezuela [inSpanish with Etnglish sunmary] Agron Trop (Maracay) 527-29 (Rev Appl Eitoniol A4572)

122 Khan K M and S Ghai 1974 White grubs ald their control in India Pesticides (1Bombay) 819-25

123 Kiritani K 1975 Integrated pest management of rice pests Pages 66-70 itt Proceedings platning workshop ott cooperative hIeld research in pest managementP SMotooka ed East-West Center Honololu Hawaii USA

124 Kisimoto R 1967 Ecology of insect vectors forecasting and chemical control Pages 243-255 in The virus diseases of tle rice plant Proceedings of a symposium atthe International Rice Research Institute April 1967 The Johns Hopkins Press altimore Maryshyland USA

125 Kisimoto R and V A Dyck 1976 Climate and rice insects Pages 367-391 in Climate and rice International Rice Research Institute PO Box 933 Manila Philippines

126 Kureha S S Kobayashi SAsari and T Ishihara 1977 Life history and control of the mole cricket Grrllotalpa africana Palisot de Beauvois [in Japanese English translation 7520 1RRI library] Proc Kanto-Tosan Plant Prot Soc 21192-197

127 Kushwaha K S 1981 The black beetle as a pest of rice in Haryana India Int Rice Res Newsl 6(4)14

128 Lawani S M 1982 A review of tile effects of various agronomic practices on cereal stem borers Trop Pest Manage 28266-276

129 Leuck D B 1966 Biology of the lesser cornstalk borer in South Georgia J Econ Entomol 59 797-801

130 lever R J A W 1955 Insect pests of the rice plant in Malaya Int Rice Comm Newsl 1612-21

131 Li Ching-Sing and Shin-Foon Chiu 1951 The study ofthe rice gall midge Iachydiplosis orvza Wood Mason [in Chinese with English summary] 1Taiwan Agric Res 21-13

132 Iitsinger J A 1984 Assesament of the need-based application for rice Proceedings TechnologyTransfer Workshop 15-16 March 1984 International Rice Research Institute 1PO Box 933 Manila Philippines 6 p

133 Litsinger J A 1984 Environmental diversity and insect pest abundance with reference to the Pacific Pages 285-307 in Proceedings sub-regional training course on methods of controlling diseases insects and other pests of plants in the South Pacific 4-20 Oct 1982 Vaini Kingdom of Tonga Nukualofa

134 Litsinger J A RF Apostol and M BObusan 1983 White grub Leucopholis irrorata (Coleoptera Scarabaeidae) pest status popula-tion dynamics and chemical conllol in arice-maize cropping pattern in the Philippines J Econ Entoinol 761133-1138

135 Litsinger J A C de la Cruz F Raymundo A T Barrion M D Lumaban J P Bandong M S Venugopal F Paragna and E Balete 1982 Insect pests and insecticide reponse of dry-seeded bunded rice Pages 179-193 in Report of workshop on cropping systems research in Asia International Rice Research Institute Los Bafos Philippine

136 Litsinger J A M 1) Luinaban J P Bandong A T Barrion W Katanyukul and N Panda 1978 Insect pest management in rainfcd lowland rice agroecosystms of tropical Asia International Rice Research Conference 17-21 April 1978 International Rice Research Institute Los Bafios Philippines

137 Litsinger J A and K Moody 1976 Integrated pest management in multiple cropping systems Pages 293-317 in R I Papendick P A Sanchez and G B Triplett eds Multiple cropping American Society of Agronomy Madison Wisconsin USA

138 Loevinsohn M E 1984 The ecology and control of rice pests in relation to the intensity and synchrony of cultivation Ph D thesis University of London 36) p

139 Loevinsohn M E J A Litsinger N Panda M A Austria R F Apostol C Andrion and J Bandong 1982 A basis for predicting rice insect pest abundance in cropping systems Paper presented at the 1982 International rice research conference on rice and cropping systems 19-23 April 1982 International Rice Research Institute Los Bafios Iaguna Philippines 25 p

140 Lopez A W 1930 Repot of the Entomology Department Pages 145-172 in Annual report of the Philippine Sugar Association 1929-30 Manila

141 Lopez AW 1931 Entomology Department Pages 227-266 in Annual report of the director or research 1930-31 Philippine Sugar Association Manila

142 Lopez A W 1932 Entomology Deparment Pages 252-279 in Annual report of the Philippine Sugar Association 1931-32 Manila

143 Lungibill P and G G Ainslee 1917 The lessercornstalk borer US Dep Agric Agric Res Serv Tech Bull 539 27 p

IRPS No 123 January 1987 39

144 McGuire J U Jr and BS Crandall 1967 Survey of insect pests and plant diseases ofselected food crops of Mexico Central America and Panama U SDepartment of Agriculture International Agriculshytural Development Service Agricultuial Researrh Service October 1967 11 p

145 McNeill Sand T R ESouthwood 1978 The role of nitrogen in the development of insectplant relations Pages 77-9 in lliochemical aspects of plant and animal coevolution J B Ilarbornu ed Academic Press London UK

146 Maebara H 1955 Spatial distribution of lastinstiarlanaeof purplish stem borer (Sesamia iqferens Walker) hibernating in an upland rice field Bull Kagoshima Univ Fac Agric 438-41

147 Mallick E H and N G Hajra 1977 Response of rice plant to defoliation treatments at different developmental stages Indian Agric 2151-56

148 Mansingh A 1971 Physiological classification of dormancies in insects Canl Entomol 103983-10)9

149 Martell C G 1974 Catalogue of plant feeding insects of Mexico [in Spanish] Fitofilo 27(69)1-176

150 Martinez 0 J 1982 Principal rice pests in Ilanos orientalis [in Spanish] P~ages 80-94 in Curso deact nalization de conocimientos en arroz Villaniencis Instituto (olonmbiano Agropecuario

151 Martins J F da Silva F Ferrcira A S Prabhu F 1 P Zimmermann 1980 Use of pesticides for the control of the main subterranean upland rice pests esqui Agropecu Bras 1553-u

152 Martins I F da Silva F Fe eira rod BdamSilveira Pinheiro I Q2 Damage simulation caused by defoliators of upland rice [in Portuguese with English summary] Pesqui Agropecu Bras 171113-1119

153 Martins 1F da Silva N(iyuen van Tan and 1Ida Silveira Pinheiro 1981 Stem borer resistance in upland rice associated with plant morphological characters [in Portuuce Iesqui Agropecu Bras 16187-192

154 Masaki S 1980 Sumnmer diapause Ann Rev Entomiol 251-25 155 Masuzawa T 11 Suwa and F Nakasoji 1983 l)ifkleences of

oviposition preferencc and survival rate of two skipper butterflies Parnara guttata and PIchlpidai iathiais (Iepidoptera HIesperiidac) on rice plant and cogon grass New Entomol 3247-56

156 Maurya D M and C P Vaish 1984 Upland rice in India Pages 69-92 in An overview oftpland rice research Proceedings ofthe 1982 Bouakamp Ivory Coast upland rice workshop International Rice Research Institute IO Box 933 Mnila Philippines

157 Menon M G R R N Katiyai S Kuar and S S Misra 1970 Termites attacking the paddy crop Sci Cult 36294-295

158 Miyahara Y and 1 Nishikubo 1967 Behavior of young nymphs of the red rice root aphid Iholahsipiun ri liabhdontinalisSasaki in relation to movement from the leaf to the roots of upland rice [in Japanese] Pioc Assoc Plant Prot Kyushu 1363-66

159 Mo ljoa Tjien 1952 White grub of upland rice (Holotrichiahelleri Brsk) [in Indonesian with English summary] Contrib Agric Res Stn Bogor No 13134 p

160 Mochida 0 and T Suryana 1979 The occurrence of the brown planthopper and ragged stunt on lowland and upland rice ill Central Java and logyakarta in tile late of January 1979 Entomnolgy Survey Report No 15 Central Research Institute of Agriculture (CRIA) Sukamandi Indonesia I February 1979 9 p

161Monet C 1979 Varietal resistance to upland rice pests [inI French] Pages 35-39 in Annual report 1977 Institut des Savanes Department des Cultures Vivrieres Bouak6 Ivory Coast

162 Monte 0 1942 A caterpillar ofrice-fields[in Portuguese] Biol6gico (Sao Paulo) 8161-163 (Rev Appl Entomol A31414)

163 Munroe I) D 1979 Pest management strategy for hill rice in four resettlement areas in Sarawak Malaysia Pages 89-102 i1 Guidelines for integrated control of rice insect pests Food and Agriculture Organization Plant Production and Protection Paper 14 Rome Italy 115 p

40 IRPS No 123 January 1987

164 Nakasuji F M Ishii and Jen-Zon Ito 1984 Rice skippers in Taiwan and their life histories ZZrC (Taipei) Tech Bull 82 10 p

165 Navas I) 1976 Fall armyworum in rice Pages 99-106 il Ecological animal control by habitat management Proc Tall Timbers Con-ference 28 Feb-I Mar 1974 Fall Timbers Res Stn Misc Publ 6 Tallahassee Horida USA

166 Nickel 1 1 1961 Biological control of rice stem borer A feasibility study Tcch Bull 2 International Rice Rcarch Institute PO Box 933 Manila Philippines IIl p

167 Nigan P M and R A Vernia 1985 Insect pests of uplaild rice in Uttar Pradesh Int Rice Res Newsl 10(6)22

168 Nilakhe S S 1985 -cological observations on spittle bugs with emphasis on their occirence in rice Pcsqtui Agropecu Bras 20407-414

169 Nixon G EJ 195 1 lie association of ats with aphids and coccids Commonwealth Iusti lcte of Entonology Iondon UK 36 p

170 Nurdin F an Ishak Manti 1985 Study oit seedling fly of upland rice Interlational Upland Rice Moitoring Tour Report 26 lebruary-2 MarLI Sukaranti West Sumatra

171 Oka 1 N 1979 Ciltural conttol of the brown planthopper Pages 357-369 in Brown planthopper threat to rice production in Asia International Rice Research Institutc 1p0 Box 933 Manila Philippines

172 Ohni M and A Vilnaii 1975 Rice insect pests of Italy biology of Sipha grcria (Kaltenbach) and its control [in Italian] Riso (Milan) 2459-71

173 Otanes F Q atnd 1 I Sison 194 1 Pests of rice Philipp J Agric 12211-259

174 Painter R I 1951 Insect icsistance in crop plants University Press of Kansits Lawrence Kansas USA 520 p

175 Pantua P C and 1A litsinger 1981 Arthropod abundance in continuous vs biannual rice cropping systeis Int Rice Res Ncwsl 6(2)20-21

176 Pathak M ) and V A l)yck 1975 Sttudics ott insect pests of upland rice Pages 186-191 in Major research in upland rice International Rice Research Institute PO Box 933 Ilanila Philippines

177 Patil B R B M IHasabe and 1) S Ajri 198 I Chemical control of white gritb lIditriihinhorraia Fabr intfesting groundntut paddy atnd sorghum Indian 1Entomol 43232-236

178 Pawar A 1) 1974 (ollection and identification of insect pests of rice rlerminal repirt lInternational Rice Research Institute PO Box 933 Manila Philippines 72 p (tnimco)

179 IerrajU A aid A S Reedy 1965 Observations ott diapausc itt tie steti borer ohrice Rice News Teller (lai-Mar)9-13

180 Pont A C 1972 A review of the Oriental species of theri ona Rondani (I)iptera Muscidac) of econtomic importance Pages 27-102 in Control of sorglium shoot fly M G lomtani and W R Young eds Oxford atd IH1 Ptthl Co New I)clhi India 324 p

181 Prasad S C 1PChand aid M 1PSingh 1983 Incidence of rice gall midge in the upland plateau region of Hihar Oryza 20235-236

182 Rai P S 198 0 Nisaga simpex damage to rice in the hill tracts of South India Int Rice kles News[ 5(I)12

183 Rathorc Y S 1969 Gonocplhahli dcprssunt Fabr as itpredator of (tilo zonwIlhis (Swinhoc) and other moths in field cage at Patnagar Uttar Pradesh Indian 1Entotnol 3127

184 Reed W 1983 Crop losses causcd by insect pists in the developing world Proceedings 10th Internatial Congress of Plant Protection lrighton UK 174-80

185 Reedy M V 1981 Ateactogasterinhens(Walker)a new root-grub pest of tipland paddy and inaize Curr Sci 501076

186 Reveno M A 1968 Catalogue of the insect pests of economic crops in Colombia [in Spaiish] Asociacion Latino Americana de Entomo-logia (AIAE) 129-33

187 Reyes G M 1957 Rice dwarf disease in the Philippines Food and Agriculture Organiiation Plant Prot Bull 617-19

188 Rice S E A A Grigarick and M 0 Way 1982 Effect of leaf and panicle feeding byarmyworm (Lepidoptera Noctuidae) larvae on rice grain yield Environ Entomol 7593-595

189 Risch S J 1) Andow and M A Altieri 1983 Agioccosystem diversity arnd pest control data tentative conclusions and nizw research dirctions Environ Entomol 12625-629

190 Rivera C T 1956 Three organic insecticides in thecontrol of insects affecting upland rice Philipp Agric 39465-472

191 Roffrey 1 1972 Locusta outbreaks in the Philippines Acrida 1177-188

192 Rojas R 1) 1941 The effect of pruning at varying ages upon the yield of Ramairio Phipp Agric 29851-860

193 Roonwal M i 1979 Ternite life and termite control in tropical South Asia Scientilic Pnbl odhpr India 139 p

194 Rosetto C1 F da Silva Martins N C Schmidt and L EAzzini 1978 Damage caused by pasturespittle bugs 4Dvoi)lavopictaandDJ schach) on rice lBragantia Nota 6 (Matcli)25-27

195 Rosztto C1- S S Neto I- Amantc I) Iink 1Garcia-Vieira D M de SoLza N V Bansa t and A M de Olivcira 1971 Rice pests of Brazil [in Portuguese] Pages 149-238 inrFA()-International Rice Commission 2nd meeting of the Rice Committee for the Americas 6-11 l)ecember 1971 Pelotas Rio Grande do Sul Brazil

196 Rothschild G H L 1969 Obseratiois on the armyworm Spodoptera matritia acronrctoiiles Gn (Lep Noctuidae) Bull Entomol Res 59143-160

197 Rothschild G 1- L 1974 Parasites of the armworm Spolopiera mauritiaacroncitoihes in Malaysia Fntornopliaga 19293-299

198 Sachan i N S K Gangwar and 1N Katujar 1980 Field crickets damaging cages in North Eastern IHill Region Indian J Entomol 42526-527

199 Sands 1) P A 1977 The biology and ecology of Lepocorisa (Hemiptera Alydidac) in Papua New Guinea Res Bull 18 Departshyment of Primary Industry Port Moresby Papua New Guinea 104 p

200 Sarawak 1968 titomology division Pages 97-106 in Report of the research branch of the Department of Altriculture 1966 Kuching (Rev AIppl Eintomol A56603)

201 Satpathy JM M S Das and K Naik 1977 Effect of multiple and mixed cropping on the incidence of scine important pestsI Entomol Res 178-85

202 Sauer IHF G 1939 Notes Ott Elasmpnlpualtsligniosellths Zeller (Lep Pyr) a serious pest ofcereals in Sao Paolo [in Portuguese] Arq Inst Biol 10199-206 (Rev Appl Entotnol A28378)

203 Satinders 1L A 1 SKing and C L Vargas S 1983 Crop pests of Central Anerica with t bibliography [in Spanish] Centro Agroshynotnico Tropical de Investigacion y Ensenanza Turrialba Costa Rica (CATIE) Tec Bol 9 90 p

204 Sauphanor B 1985 Some factors of upland rice tolerttce to stem borers in West Africa Inseu Sci Appl 6429434

205 Sen P atnd S Chakravorty 1976 Stentflics in Haritighata Nadia West Bengal India [it Rice Res Newsl 1(12) 15

206 Senapai B and 1M Satpathy 1983 The seedling fly Atherigona orrzae Malloch is a pest of rice in Orissa (India) J Entomol Res 777-79

207 Shhjahan M S Alatn M Ah nad and BN Islam 1983 A new scarabaeid beetle pest of dryland rice in Bangladesh Int Rice Res Ncwsl 8(I)16-17

208 Sharma G L S C Modgol and R C Gautam 1979 Effect of depth of submergence ott incidence of bacterial blight and white grub infestation in transplanted rice Int Rice Res Newsl 4(6)12-13

209 Sharma S 1985 Scarabaeid beetle outbreak itt upland rice Int Rice Res Newsl 1(1(l)24

210 Shiraki T 1917 The paddy borer Schoenobhits incertulas Walker [achaku Agric Exp Stn Formosa 256 p

211 Silva A de Brito and 11P Magalhaes 1981 Insect pests of rice [in Portuguese] Circular Tecnica No 22 EMBRAPA Centro de Pesquisa t ecuaria de Tropico Unido (CPAJU) Belem Brazil 14 p

212 Singh S A 1979 Emergence of rice leaf butterfly as a rice pest in Manipur India int Rice Res Newsl 4(4)16-17

213 SinghS A 1983 Changing trends of insect pestsof rice in Manipur Int Rice Res Newsl 8(f)12

214 Siwi S S and P H van Doesburg 1984 Lenovorisa Latreille in Indonesia (Heteroptera Coreidae Alydinae) Zool Meded (Leiden) 58117-129

215 Soejitno J 1977 Reaction of some upland rice varieties toseedling fly eltherikonaexigua Stein [in Indonesian] Siri Hana I Penyakit 630-35 Lembga I osat Fenclilian Pertanian Bogor Indonesia

216 Socnardi 1967 Insect pests of rice in Indonesia Pages675-683 in The major inse pests of thu rice plant Proceedings of a symposium at the Internatital Rice Rec Wi itt S pternber 1964 The Johns Hopkins Press Baltimore Maryland USA

217 Soto P E A T Petcz and I W Buddenhagen 1976 Survey of insect pests and diseases of rice in different ecological zones in Nigeria Rice Entomol Ncwsl 435-46

218 Soto 11E and Z Ziddiqi 1976 Screening for resistance to African rice insect pests Varietal improsetnent sCmi n r Bouak Ivory Coast 13-17 Sept 1976 West Africa Rice l)evelopment Association Monrovia Iiberia

219 Squire F A 1972 Entomological problems in Bolivia PANS (PestArtic News Sumr) 18249-268

220 Stapely 1 II 1978 The rice leaf roller in the Solomon Islands Int Rice Res Newsl 3(l)13

221 Suenaga H 1937 On aphids attacking the roots of rice Oyo-I)oubts Zasshi 9150-152 [in Japanese] (Rev Appl Entomol A25745)

222 Suenaga H 1952 On the seasonal abundance of three subterranean aphids on upland rice Bull Kyushu Agric Exp Sti 1249-262

223 Sutynso Yas and Dandi Soekaria 1964 Beberapa hana yang ttenperang tana man padi tanah kering [in Indonesian] Rapat kerja Padi Tanah Kering 6 sd 8 Juily 1964 Kaliurang 9 p

224 Tallamy I) W and R F Denno 1983 Alternative life history patterns in risky environments in example from lace bugs Pages 129-147 in Insect life history patterns R F l)cnio and If Dingle eds Spring-Verlag 225 p

225 Tamura 1 1939 Observations on some insect pests of field crops [in Japanese] 1 Plant Prot 26414-420 (Rev Appl EntomolA27-556)

226 Tamura and Suzuki 1964 Influences of the early-transplanting practice in rice culture upon development of the rice stem borer Chilo siql)ressalisWalker Bull Ilokuriku Natil Agric Exp Stn 761-94

227 Tan J P 1924 The rice root aphid Drowhi ihirsoia Philipp Agric 13277-288

228 Tanaka 1 1957 Taxonomy and distribution of some subterranean aphids injurious to upland rice in Japan with description of a new species [in Japanese English summary] Botyu-Kagaku 22168-176

229 Tanaka T 1961 The rice root aphids their ecology and control Spec [full College of Agriculture Utsunomiya University Japan F3 p

230 Tauber M IC A Tauber R Nechols and 1Obrycki 1983 Seasonal activity of parasitoids control by external internal and genetic factors Pages 87-108 itt I)iapause and life cycle strategies in insects V K Brown and I Ilodek eds )r W Junk The Hague Netherlands

231 Taylor W F 1972 Effects of artificial defoliation (simulating pest damage) on varieties of upland rice Exp Agric 879-83

232 feotia T P S and S Natud 1966 Biotoomics of the rice skipper Parnaraniathiis Fabricius (Lepidoptera Hesperiidac) Indian J Entomol 28181-186

233 Teran F 0 1971 Rice pests of Santa Cruz [in Spanish Ministerio de Agricultura Sita Cruz Bolivia Bull 45 8 p

234 Than Itun 1976 Population dynamics of the yellow rice borer Tryporza itcertulas (Walker) and its damage to the rice plant MS thesis University of the Philippines at Ios Bafios Laguna Philippines 92 p

235 Uichanco L B 1911 Pests Pages 55-74 it Rice Philipp Agric 11-200

IRPS No 123 January 1987 41

236 van der Laan P A 195 1 Life history of the rice seedling fly Contrib Gen Agric Res Stn Boor Indonesia 118 15 p

237 van der Laan 11A 1959 Correlation between rainfall in the dry season and the occurrence of the white rice borer (Scirpolhaga innotala WIk) in lava Entomol Fxp Appl 212-20

238 Van Driesche R G 1983 Meaning of percent parasitismin studies of insect parasitism Fnviron Entomol 121 I1- 122

239 Venugopal M S and A litsingcr 1984 Fffect ofcarbofuran on rice growth Prot Ecol 7313-317

240 Vietnam Directorate of Rural Affairs 1962 Annual work progress report on crop improvement program of rice sugarcane vegetable and field crops (lor the period from July 1961 to June 1962) (11) 299 p (Rev Appl Entomol A51526-527)

241 Wan M T K 1972 Observations on rice leaf- and planthoppers in Sarawak (Malaysian Borneo) Malay Agric 148308-335

242 West Africa Rice I)esclopnient Association 1978 Annual research report Vol 2 Statistics Monrovia Liberia

243 West Africa Rice Development Association 1979 Annual research report p47 Monrovia Liberia

244 Williams ) F and CS lofgren 1981 Fli I illy l-468 anew bait toxicant for control of the red imported lire at Fla Entomol 64472477

245 Williams I) 1 1970 The mealybugs OFlomoptera Coccoidea Iscudococcidac) of sugar-cane rice and sorghum Bull Entomol Res 0109-188

246 Williams D) 1 1982 Pulinaria icervi (Signoret) and its allies ott sugar cane and other grasses Btll Entoinol Res 72111-117

247 Wirjosuiardjo S 1976 Study ofchenmical cont rot of insect pests ott upland rice Imiu Pertanian (Jogjakarta) 1323-329

248 Wirjosuhardjo S and N Wirarta 1976 Chemical control of the white grub tIolorithiatlltriin upland rice in M uneng Irobolinggo Indonesia) [in Indonesian] Imut Pertanian (Jogjakarta) 1330-332

249 Yano K T Miyake and V F Eastop 1983 The biology and economic importance of rice aphids (Henuiptera Aphid idae) a review Bull Entomol Res 73539-566

250 Yelp D and P Rhodes 19(4 lhe success of drilled dryland rice iii the Fiji Islands International Rice Commission Working Party on Rice Production and Protection Manila Philippines 3-10 March1964 Food and Agriculture Organization Rome Italy 13 p

251 Young W R 1981 Fifty-five years of research on the sorghum shootfly Insect Sci Appi 23-9

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Page 6: UPLANDRICE. INSECTl--1i M T -E!R.EOLOU t RTANCE ,AND …delphacid.s3.amazonaws.com › 6958.pdf · The significance of abiotic factors in upland rice insect ecology will be apparent

6 IRPS No 123 January 1987

Table I continued

Order Family

Hemiptera Lygaeidae

Cydnidae

Alydldae

Coreidae Corimelaenidae

Pentatomidae

Thysanoptera Thripidae

Species

Nephotettix virescens (Distant) Nephotettix modulatus Melichar Cofana spectra(Distant)Graphocephala spp Hortensiasimnilis (Walker) Exitianusobscurinervis (Stal)Balclutha spp Draeculacephalaclypeata Osborn Cicadulinabipunctella (Matsumura)Cymoninus turaensis(Paiva) Ninus insigniStal Pachybrachiusnenosus Horvath Caenoblissuspilosus Barber Blissus leucoptens (Say) Blissus spp Dlmnorphoptentscornutus novaeguineae GhauriDimorphopterus similis Slater Paroniuspiratoides (Costa)Pangaeus spp Aethus indicus (Westwood)Geotomnus pygmaeus (Dallas) Stibaropus inolginus Schiodte Cyrornenusbergi FroeschnerCyrtomnenus ciliatus (Palisot de Beauvois) Cyrtonenuscrassus Walker Tominetus spp

Riptortuslinearis (Fabricius) Riptorts spp Leptocorisaacuta (Thunberg)Leptocorisabiguttata (Walker)LeptocorisapalawanensisAhmad Leptocorisaoratorius(Fabricius)Leptocorisachinensis Dallas LeprocorisasolomonensisAhmad Leptocorisaspp Stenocorissouthiwoodi Ahmrad Stenocoris claviformis Ahmad MirperustorridusWestwood aetus trigonus (Thunberg)Alkindus atratusDistant ScaptocorisdivergensFroescher Scaptocoris castaneusPertyScotinopharatarsalis (Vollenhoven) Scotinopharascotti Horvath Eysarcoris (=Stollia) ventralis (Westwood) Dolycorisindicus Stal Macrinajuvenca Burmeister Tanda antiguensis (Westwood) Tantiageifi Schout Tantia perditor (Fabricius) Pygotnenidavaripennis (Westwood) Menida sppNezara viridula (Linnaeus)

Oebaluspoecilus (Dallas) Oebalusypsilon-gnseus (de Geer)Oebalusgrisescens (Sailer) Tibracea limbativentris (Stal)

Aspavia armigera(Fabricius)Diploysfissa Erichson Acrosternum marginatum (Palisot de Beauvois) Acrosternum sppFrankliniellarodeos Pergande Breginatothrips enustos Watson Stenchaetothrips(=Baliothrips)biformis

(Bagnall) (=(Jlloethripsoryzae (Williams)HaplothripspriesnerianusBagnall

Distribution

Asia Africa Asia Latin America Latin America Latin America Latin America Latin America Asia Asia Asia Asia Asia Latin America Asia Asia Africa Asia Latin America Asia Asia Asia Latin America Latin America Latin America Latin America

Asia Africa Asia Asia Asia Asia Asia Asia Asia Africa Africa Africa Asia Latin America Latin America Latin America Asia Asia Asia Asia Africa Latin America Africa Latin America Asia Asia Asia Africa

Latin America Latin America Latin America Latin America

Africa Africa Latin America Latin America Latin America Latin America Asia

Asia

Reference

59 241 217 59 241 6166 61 66 81 150 6166 66 81150 b 178 178 30 83 42 81150 203 30 80 62 178 195 178 178 178 203 203 203 203

55 178 12 72 78 120 214 214 b 59 120178114 178 54 636 82 12 6 12 82 178 81 150 60 is 195 59 120 178 78 36 186 36 186 b 55 12 30 54 55 78 82 120 61 66 81195 61 65 66 195 211 150 195 61 66 81 150 219 233 6 12 82 6 82 186 186 61 66 68 213235

178

7 IRPS No 123 January 1987

Table I continued

Order Family Species Distribution Reference

toleoptera Scarabacidae leteroligus (=Aphonoprocts) 15elcs Billberg Africa 36 Dynastinac (=Ileteroligusmees robustui Prell)

Alissonotum pauper Burnieiser Asia 82 Alissonotuon simile Arrow Asia 209 Dscinetusdubibs (Olivie) Latin America 28 195 233 Devscinetusgagates Burmncster Latin America 20 195 Dyscinctt spp Latin America 60 Eutheola Inimilis Bturmcistr Latin America 20 28 8 1 195 203

233Eutheola identarr(Biurnciter) Latin America 121 203 Heteron)ciwv aw rsonJack Africa 82 Hcreron)chusbtuterculatus Kolbe Africa 82 tleteronichtslicas (Klug) Africa 82 fhteron)chus mosamhicus Peringuey Africa 4 6 48 82 (=Veteronychus orzae Britton)

leteronychts lioderes Redtnbacher Asia 77 82 207 ltcteronchusarator (Fabricius) Africa 82 tleterontchusplebepus (Klug) A frica 57 82Ileteron chus pseuocongonnsis Ferreire Africa 32 Ileteronvchns rogiJrons Fairmaire Africa 82 Heteron chus nrsticus niger (K lug) Africa 82 Heteronvchus spp Africa 17 Lachnosterna longipennis (Blanchard) Asia 82 Lachnosternaspp Asia 120 Ligvros fossator Burmeister Latin America 82 Ligyrus nasutus Burmeister Latin America 203 Ligyrus (=Scarabaeus)ebenus (de Geer) Latin America 20 195

Dynastinac Ligtvros huonilis Burmeister Latin America 60 82 Aaladera castanea (Arrow) Asia 82 Maladera orientalis Motschulsky) Asia 82 Malaaerajaponica Motschulsky Asia 82 Phvlognathusdicynsius (Fabricius) Asia 82

Melolonthinae Exopholis h)ypoletca (Wiedemann) Asia 117 Lerlinosterna[= lolotrichia] consanguinea Asia 122

(Blanchard) Leucopholis rorida (Fabricius) Asia 82 117 Lepidiota blanchardi Dalla Torre Asia 76 82 Leucopholis irrorata (Chevrolat) Asia 76 134 Scrica internpta Walker Asia 76 Holotrichialonipennis (Blanchard) Asia 122 Holotrichia serrata Fabricius Asia 122 Holotrichia leucoph thabna (Wiedemann) Asia 82 Holotrichia seticollis Moser Asia 77 78 Holotrichia pnindanaoana Brenske Asia b Stenoeres spp Latin America 71 195 Atoenius spp Latin America Asia 203 Ph luplg aequata (Bates) Latin America 203 Piyvilophagacaraga Saylor Latin America 203 Phyllophagachiriquina (Bates) Latin America 203 Phyllophagadas)poda (Bates) Latin America 203 Phyllophagaclegans Saylor Latin Ameica 203 Phyllophagahondura Saylor Latin America 203 Phyllophagalatipes (Bates) Latin America 203 Phyllophagamontriesi (Blanchard) Latin America 203 Phyllophaga pamisetis (Bates) Latin America 203 Ph vllophaga sanfosicola Saylor Latin America 203 Phllophagasetifera (Burmeister) Latin America 203 Phyllophaga (=tlolotrichia)helleri Breuske Asia 82 117 159 Phvllophagaspp Latin America 81 150 White grub unspecified Asia 84

Rutclinae Lagochile trigona (Ilost) Latin America 82 Anomala dinidiata var barhata Burmeister Asia 77 78 82 Anomala huteralis(Burmeister) Asia 76 Anoimala lurida (Blanchard) Asia 76 Anomala sulcatula (Burmeister) Asia 76 Anontala varians (Olivier) Asia 82 Anomala antigua (Gyllenhal) Asia 82 Anoinala palida (Fabricius) Asia 82

8 IRPS No 123 January 1987

Table 1 continued

Order Family

Tenebrionidae

Elateridae

Languriidae Chrysomelidae

Chrysomelidae

Coccinellidae Curculionidae

Lepidoptera Lymantriidae

Amatidae

Arctiidae

Limacodidae Eupterotidac Noctuidac

Species

Anomaa volita (Blanchard) Adoretus caliginosus Burmeister Adoretus comprcssus (Weber) Papuana huhneri (Fairmaire) Papuana incrmnis Prell Popillia capricollis Hlope Gonocephalu depressur (Fabricius)Gonocephahm acuiangulum (Fairmaire)Gonocephalum sinphx (Fabricius) Epitragus sallei Champion Anaedus ptnctatissimnts ChampionAeolus spp4griotes spp Agriotes mancus (Say) Conoderts spp Aeoloderma brachmana (Candeze)Wireworni unspecified Anadastus filijf rmis (Fabricius)Dicladispa viridicanca(Kraat z)Trichispasericea (Guerin)Oediopalpastervalis (Weise) Oediopalpa guerini (Italy) Chaetocnema cvlindrica (Baly)Chactocnema denticulata S tephens Chaetocnemabasalis (Baly)Aulacophora sp nr similis Olivier Monolepta caripemne Ialy Monolepta bilasciata (Ilornstedt) Monolepta signata Olivier Cerotoma airojcsciata J acoby Diabrotica adelpha Ilarold Diabrotica balteata Leconte Diabrotica graminLa ltaly Diabrotica speciosa (Germar) Diabrotica limitafa quindecinmunctata(Germar)Diabrotica melanocephala (Fabricius)Diabrotica spp Altica (= Fondia)madagascariensis(Allard)Altica spp Epitrix cuctmeris (Harris) Epitrix spp Oulenjo or pae(Kuwayama) Disonchq (= Donacia) spp Flea beetle unspecified Chnootriba (= htIilachna)sinilis assimilis Mulsant Atactogaster iduriens Walker Hypomeces squtamosus (Fabricius) Tan vinecus discoidalis GyllenhalPhaulosoinusmusculinus (= mnus) Csiki Neobaridia anplitarsis CaseyPsalis pennatula (Fabricius) Laeliasufihsa (Walker) Euproctis irgtncula (Walker)Euproctis minor (Snellen)Euproctis xanthorrhoea (Kollar)Amata sp

Diacrisia obliqua (Walker) Creatonotusgangis Linnacus Latoia (= Parasa) bicolor (Walker)Nisaga simplex Walker Sesamia botanephaga Tams amp Bowden Sesamnia nonagroides botanephagaTfums et Bowden Sesamia calamistis llampson Sesamia inferens (Walker) Spodoptera eridania (Cramer)Spodoptera mauritia acron)ctoides Guenee Spodoptera frugiperda ( ESmith)

Distribution Reference

Asia 82 Asia 82 Asia 82 Asia 82 Asia 82 Asia 78 Asia 82 117 178 Asia 82 117 Africa 82 Latin America 203 Latin America 203 Latin America 195 Latin America 195 Latin America 203 Latin America 195 Asia h Asia Latin America 60 73 117 Asia 200 Africa 36 Africa II Latin America 6166 68 Latin America 6166 68 Asia 225 Latin America 61 66 68 81

Asia 178 Asia 55 Asia 55 Asia h Asia 55 Latin America 203 Latin Amefia 203 Latin America 203 Latin America 203 Latin America 6166 68 Latin Amcrie 195 Latin Amw 195 Latin Aierwca 61 81 Africa 57 Asia 178 Latin Americ 203 Latin America 81 Asia 82 Latin America 195 Asia 120 Africa 6 11 Asia 185 Asia 120 Africa 82 Africa 82 Latin America 66 68 195 233 Asia 117 Asia 117 Asia 117 Asia 117 Asia 117 Asia A S Pradhan

unpubl dataAsia 82 Asia 82 Asia 82 Asia 182 Africa 12 Africa 6 36 Africa 6 8 11 12 36 Asia 59 82 84240 Latin America 150 Asia -0 59 196 Latin America 28 61 66 68 81 150

152195203219233

9 IRPS No 123 January 1987

Table I continued

Order Family Species Distribution Reference

Spodoptera exigua (Hubner) Spodoptera exem pta (Walker) Spodoptera ornithogalli (Guenee)

Africa Africa Latin America

32 217 32 33 36217 150

Mythimna loreyi (Duponchel) Asia 30 Mythimna separata (Walker) Mythimna (- Pseudaletia)latifascia [= adultera

Asia Latin America

178 195

(Schaus)] (Walker)Mythimna (=Pseudaletia) sequax (Fabriclus) Latin America 68 Mythimna rosellinea (Walker) Asia b Mythimna yu (Guenee) Asia b Mocisfrugalis (Fabricius) Asia 30 117 Mocis latipes (Guenee) Latin America 66 68 149 150 203

Platysenta (=Spodoptera) compta (Walker) Asia 219233A Barrion unpubl data

Agrotis ipsilon (Hufnagel) Latin America Africa

33 68 81 150 1)5 203

Agrotis spp Latin America 60 Achaeajanata (Linnaeus) EldanasaccharinaWalker

Asia Africa

117 36

Armyworm unspecified Asia 72 84 Pyralidae Cnaphalocrocis medinalis (Guenee)

Marasmia bilinealisHampson Asia Africa

30 4559 72 84233 36

Marasmia trapezalis(Guenee) Africa 32 217 Marasmia (=Sumia) exigua (Butler) Asia 220 Marasnianmralis (Walker) Asia b Marasiniapatnalis Bradley Marasiniaspp

Asia Asia

b b

Leaf roller unspecified M2liarphaseparatellaRagont

Asia Africa

72 12 57

Maliarphasp Asia b Chilo auricilius Dudgeon Asia b Chilo partellus (Swinhoe) Chilo polychrysus (Meyrick)

Africa Asia

12 92 59 72

Qhilo diffusilineus J de Joannis Africa 6 1229 Chilosuppressalis(Walker)Cibozacconius Blezzinski

Asia Africa

59 72 6 11 12

Cravubus spp Latin America 203 Acigona loftini (Dyar) Latin America 203 Acigona chrysographella(Kollar)Scirpophagaincertulas(Walker)

Asia Asia

82 59 72

Scirpophagainnotata (Walker) Asia 59 200 Scirpophaganiuella (Fabricius) Asia b Elasinopalpuslignosellus (Zeller) Latin America 28 60 61 66 81 150

Diatraeasaccharalis(Fabricius) Latin America 162195203219233 28 66 81 111 150 195

Rupela albincllaCramer Latin America 111

Gelechlidae

Stem borer unspecified Undetermined stem borer nr Maliarpha BrachniaarctraeaMayr

Asia Asia Asia

72 b 55

Brachmi spp Africa 32 Hesperiidae Parnaraguttata Bremer et Grey Asia b

Parnaranaso Fabricius Asia 55 Pelopidasagnaagna (Moore) Pelopidasmathias (Fabricius)

Asia Asia Africa

164 36 155217

Pelopidasconjuncta conjuncta (Herrich-Schaffer) Asia b Borbo fanta Evans Africa 6

Satyridac Melanitisleda ismene Cramer Asia 84213 Mycalesis asophisHew Asia 30

Diptera Cecldomyiidae Mycalesis spp Orseoliaoryzae (Wood-Masn)

Asia Asia

b 72 88 131 181

Orseoliaoryzivora Harris et Gagne Africa 61132218 Chloropidae Steleocerellus (=Mepachymens) ensifer (Thomson)

Oscinella spp Asia Asia

205 205

Gaurax sppChloropsoryzae Matsuntura (= kuwonae Aldrich)

Asia Asia

205 82

Mucldae Atherigonaoryzae Malloch Asia 30 82 117130 Atherigona exigua Stein Asia 82 117 178

10 IRPS No 123 January 1987

Table I continued Order Family

Diopsidae

Agronyzidae

lHymenoptera Formicidae

Species

Atherigona oritentalisSchiner Atherigona indica MallochDiopsis longicornis(= thoracica)Macqurt Diopsis spp Pseudonapnyza asiatica Spencer Pseudonapomyza spicata (Malloch) Agronoza orvzae MunakataAcromyren landoltibalzani(Emery) Acromyrmex landoltifracticornis(Forel)Acromyrnex heyeri Forel)Atta bisphaericaFore Atta capiguaraGoncalves Atta laeLigata (F Smi h1) Atta opacicepsBorgmeierSolenopsisgeminata (abricius)Pheidole sp Pheidolegeton diversus Jerdon Monomorium pharaonis (Linnaeus) Ants unspecified

Distribution Reference

Asia 82 Asia Africa 82205Africa 11 1248 Africa 217 Asia 213 Asia h Asia bLatin America 6168 195 Latin America 61 195 Latin America 61 68 195 Latin America 61 68 195 Latin America 14 61 68 195Latin America 61 68 195 Latin America 61 195 Asia b Asia b Asia 1) Asia b Asia 120

aG orlentalls formerly recorded in Asia as G africana thPhilippines reported in this study CSubterranean root feeders

subterranean nests Foraging ants will feed on germinatedseeds only when the supply of ungerminated seed is low However these ant specialists feed on a wide varietyplants and thus help control grassy weeds (39) This dual

of

pest benefit role extends from tending ants which protecthoneydew-excreting pest species including root aphids and mealybugs (workers repel potential predators and even selectively kill parasitized aphids) to predators of leaf- and stem-feeding rice pests such as leaffolders armywormswhite grubs and stem borers (1 17)

The upland rice environment is particularly well suited to ants because of the lack of flooding Frequent tillage perpetuates grasses and granivorous species that are highlyadapted to constructing new nests in recently tilled fieldsAnt nests however are relatively shallow and tilling with a moldboard plow can destroy them Also upland ricefields are uually small and isolated and a given area in a smallfield has more field borders than it has in a larger fieldallowing high rates of infiltration from more sedentaryspecies such as Plieidole that do not colonize tilled fields but nest in soil under shrubs and trees along field bordersPheidole workers forage into fields from borders and store seed in their nests In slash-and-burn or other no-tillage rice cultures nests would be preserved in the field and possiblylead to high rates of seed predation However granivorous species do not prefer forest habitats

Damage is characterized by reduced usually patchy plantstand (120) Seed-thieving ants greatly prefer rice but they can be selective regarding plants they collect and store Theyharvest more seed than they can consume so patches of germip ing surplus seed (such as Rottboellia) will sproutfrom their nests during the rainy season

In Latin America leaf-cutting ants (Aita spp and Acromyrmex spp) occur only in upland fields and defoliate

young rice plants (47 68) They take leaves to fungal gardensin underground nests where fungi predigest the plant material

Termites Subterranean termites of the family Termitidae which lack symbiotic protozoa to help digest plants are the most frequently mentioned termite pests of upland rice (1185 120) This family cultures fungi in special undergroundcells --- fungal combs made of half-digested plant material (117) The fungi inoculated onto the combs break down plant material into food that termits can digest The termites then consume the fungal combs These grasslandtermites build nests below the plow layer in uplandricefields The colonies attack living plants only when dead plant material is not available They will attack a droughtshystressed crop but prefer older plants having greater cellulose content They will damage newly planted crops where clean culture has removed vegetation

Infestations are worse in deep light-textured soils with low moisture content The first sign of damage is yellowingof older leaves (195) Termites feed on roots (the plantyellows then wilts and finally collapses) and germinatingseed (loss of stand) or move above ground at night to cutseedlings at ground level which they cover with soil for later consumption Soft-bodied termites are highly sensitive to desiccation They live within a self-contained system of airtight chambers and build surface tunnels linei with nudand body secretions (carton) to maintain more than 90 relative humidity Termites can be located by their tunnels

Termites apparently can withstand limited submergence (157) and can be a pest of dry lowland seedbeds (117)perhaps by having nests below perched water tables

Termites appear to do more damage in Latin America where large ricefields are plowed with tractors deprivingtermites of preferred vegetation and forcing them to live on

rice plants In Asia upland ricefields are typically small with Iprge field border areas (ecotones) where termites can forage for preferred food other than growing rice plants All grassland termites do not feed on rice In an upland area in Batangas Province Philippines ricefields are commonly infested by Hosiititernesluzonensis (Oshima) however this termite does not damage rice In the same province though Mrroternesgivus(Hagen)feeds on rice seedlings (178)

In Africa termites may damage rice even though fields are small and surrounded by perennial vegetation These termite species perhaps prefer rice over other hosts

Mole crickets Nymphs and adults are nocturnal and burrow through soil feeding on roots of a wide variety of plants or forage above ground as predators of insects Adults live 3-5 mo and are even cannibalistic Mole crickets prefer low-lyin moist upland soils with high organic matter (II 81) In the lowlands they inhabit rice bunds or nonflooded fields Adults are highly mobile and can leave a flooded field to locate a more suitable habitat Losses show as wilted plants (from root feeding) or reduced stands (120) Damage is more common near field borders where mole crickets relocate after tillage (126)

Fiehl crickets Nymphs and adults of field crickets have similar nocturnal habits and damage rice as much as mole crickets do Piles of weeds removed from fields attract them They make subterranean nests and tunnel through the soil to feed on roots Some species prefer seed to roots Others feed at the base of stems causing deadhearts

Tunnel entrances surrounded by excavated soil are easily seen Young plants are cut at ground level and stored in underground cell Field crickets tre highly polyphagous as are all soil pests (117) Sachan et al (198) reported that field crickets prefer maize to upland rice Like mole crickets field crickets cannot survive in standing water and are therefore more prevalent in upland fields

Whitegrubsandhackbeetles Scarab beetles that feed on living roots as larvae but not as adults are called white grubs or chafers (Melolonthinae Rutellinae) and those that feed as adults but not as larvae on plant crowns are called black beetles (Dynastinac) In the tropics chafers or white grubs have a I-yr life cycle starting with adults emerging from the soil about I mo after the first downpour of the rainy season (134 159) They lay eggs at the same time farmers sow upland rice The rice passes its most susceptible stage and damage is mostly avoided when the white grubs are small After several months the long-lived larvae are large enough for two or three to denude the root system of mature rice (223) This intensity of damage is rare but wilting occurs when root loss is combined with drought stress

The larvae need damp (not saturated or dry) soil to survive and pass the unfavorable dry season 05-2 m underground in aestivation in the moist soil The first heavy rains of the season (20-30 amd) stimulate the grubs to resume activity and after several weeks they develop into

IRPS No 123 January 1987 11

pupae and adults and eventually dig their way to the soil surface and fly to nearby trees to seek food and mates The adults of most white grub species feed only on trees but in some species notably Eutheolahumilis the adults defoliate and cut off stems at ground level (28) Putheola hidentata damages sugarcane fields in Venezuela (121) Floods kill white grubs (208) Grasslands can support large populations and therefore white grubs can be more abundant in newly planted upland ricefields that were previously fallow (81)

Black beetles live up to a year adults can attack a crop at any stage by burrowing into the soil (127) and feeding at the base of stems to cause whiteheads (207) Adults emerge with the early rains and are normally more abundant and do more damage on a young rice crop with small root systems (38 81)

The larvae feed only on dead organic matter in dryland fields and do not attack rice Adults arc highly mobile and although sensitive to flooding can invade a field soon after it drains (115)

Root aplhids Several genera of migratory aphids feed on the roots of upland rice (249) Populations build up more in light-textured soils with high percolation rates Damage has also been reported from dry seed heds of lowland rice (52) Tending ants -- Pheidhle Cren aster Tetramoriuw lsius Tapinoma -- are necessary for root aphids to multiply Aiits harbor aphids in their nests over winter or during unfavorable periods and relocate them on rice plants digging tunnels along the root systems to allow these softshybodied insects to penetrate the soil Most intensive studies on aphid life histories have been done in temperate environments (158 222 229) Root aphids fly to rice plants at the beginning of the rice season and pass through several generations continually developing winged forms to relocate on alternate grassy hosts until winter when they move to perennial hosts usually trees Yield loss occurs mainly through reduced tillering but infested plants become yellow and stunted and in extreme cases wilt (227) Aphid populations build up gradually so damage usually begins in the late vegetative and reproductive growth stages

Root-feeding ineal tugs More than half of the mealybug species or rice feed on roots (245) Being soft bodied mealybugs are not adapted to living underground but they survive because tending ants (the same species that tend root aphids) dig burrows for them and move them from plant to plant Well-drained soils ihelp mcalybugs survive Although six species are recorded as subterranean root pests of rice we found no report citing damage Root-feeding mealybugs are probably more prevalent on perennial grasses than on short-lived annual rice

The most common rice mealyb ig Brevennia rehi is a foliar feeder favored by lack ol rain apparently it is responsible for high yield loss in Northeast India and Bangladesh (N Panda pers comm as stated by IRRI [1061) This report needs confirmation Mealybug damage was only briefly mentioned in another report (156)

12 IRPS No 123 January 1987

Root weevils A few species of upland weevils (IHporneces Donacia Atactogaster) attack rice roots as larvae and feed on stems and leaves as adults Damage ispatchy The larvae are not adapted to submergence and are true upland soil weevils isopposed to the so-called water weevils They rarely become sufficiently abundant to cause damage Adults on rice foliage or the soil surface show that the roots are being attacked by the grubs Root weevils are highly polyphagous (185)

Root heeths Soil-inhabiting false wireworms Gono-cephalumt spp are reported in upland ricefields The adults and larvae normally feed on decaying organic matter but land preparation for upland rice removes these sources The larvae feed on seedling roots and nocturnal adults roam on the soil surface to cut off seedlings at ground level (82) This group of ground beetles is highly omnivorous adults even prey on stem borer larvae (183) True wireworms (Elate-ridae) feed on living roots but rice as a host has been reported Infrequently (117) In Latin America the highlypclyphagous larvae of Diabrotica rootworms are also recorded on rice

Root htus Upland rice seedlings can be attacked by the adults anI nlViphs of chinch bugs Blissusand Caenoblissus and bro n bug Scaptocoris which feed on the roots with theirsucking nouthparts (80 83) Three genera of burrower bugs A tms (eotonhus and Stiharopus -are common iil grasslands in the Philippines and may attack the first crop (rice or maize) planted after the dry season Chinch bugs are most abundant after a dry spell These insects are large and can kill seedlings or stunt and reduce tillering on older plants I hey prefer other hosts and rarely damage rice Adults can also feed oii developing rice grains (81)

(utworms Soil-dwelling noctuid larvae hide under-ground during the day to avoid predation by birds and bccone active at night above grour cutting off you ing plants at ground level (hence tile naiie cvtworn) arvaedrag severed plants into hurrows llie larvae have wide host ranges arid pupatc in the soillgrotis ipsilon occurs worldwide and appears to be the most common cutworm pest of upland environments but is more prevalent inlowland rice particularly in dry seedbeds

IFoliar pests Other groups of insects are adapted to upland environments because they pass one growth stage underground either as egg (in heinrietabolous species) or pupa (in holomnetabolous groups)

Rice seedlibig maggots AitJeriota flies occur in non-fhoodcd environments in Asia and Africa (180) Feeding of larvae one per iller causes deadhearts Flies as opposed to moths only secondarily evolved as pliytophagous pestsfroni a saprophagous Origin Athrnonalarvae secondarihv feed onidecaying tillers whereas larvae of stem borer moths actively Iced on living tillers to cause deadlicarts Seedling maggots will die if they cannot sever a tiller on which to feed

as it decays Plants die under heavy attack whereas lighter infestations cause stunting delayed maturity and ragged leaves (236)

Eggs are laid only on actively growing plants (49) aid damage occurs wiihin the first week after crop emergence(170) Larvae pupate more in stems than in soil Seedling maggot attack is highly seasonal (206) normally peaking 2-3 mo after the onset of the rainy season These pests possiblyaestivatc over the dry season Rice isonly one of their many graminaceous crop and weed hosts

The rice stem maggot (hlorops orzae is adapted to temperate upland and lowland rice growing regions of Asia In Japan and China upland rice grows in mountainous regions a habitat of this chloropid fly The larvae tunnel into tilestems The stem maggot is more damaging during the seedling stage larvac hibernate in grasses

File several dipterans reported as stem maggots in dryshyseeded lowland seedlings inIndia (205) could presumably attack upland rio

Leaf urinrs Icaf miners are common on wheat and maize but are rarely reported on upland rice (213) Three leaf miners are more prevalent on upland than lowland rice inthe Philippines Pvudonalomy a asicanh Vficataand lIAgromrza orzae attack at the seedling stage The larvae develop inside the tunnels they construct in the parenchyma tissue P Vpicata and Aorvzac prefer maizeEehtsine and other grasses to rice spicatacan become abundant on wheat

Leaf beetles Chrysomelid and coccinellid eaf bcetles scrape or otherwise remove leaf tissue as larvae or adults or both This damage accelerates desiccation of plants Tile larvae of sonic species are root feeders Flea beetles Epitrix and (lraetocnetna and rootworms l)iahroticaspp feed on rice foliage only as adults Their root-feeding larvae have hosts other than rice The adults and larvae of the hispa-like Oediolmlpa spp beetles of Brazil and (hmootriha (=EPfishyachra) beetle of Africa feed on upland rice leaves Oealioshypapa spp larvae are leaf miners (mootriha is also pshyvalent in lowland environments (11) Leaf beetles prefer vegetative rice (81)

The rice hispa Dicladipaarnigera is not reported on upland rice and prcers more aquatic habitats In Central Africa however I)cladispm viridicyaiea attacks uplandrice in the vegetative stage as well as lowland seedbeds 7richipasericea is reported on upland rice in West Africa and Madagascar (57)

The rice leaf beetle 1d1ea 01izae riiuech like Chioropsor|rzae is adapted to temperate regions of Ashi and is usually a pest on lowland rice but seldom oi inpland rice in the mountains (82) The adult hibernates on grasses

Armmworms Species of grassland-adapted SpoptloeraPlatysemta and A ithimna larvae create widespread epishydemics They can defoliate ricefields generally in patchesfrom early vegetative growth to harvest (41) Armywornms pupate inthe soil and highly favor upland rice (34) Adults

can disperse long distances to colonize even remote upland areas They become more abundant than their natural enemies particularly after favorable rains following a prolonged drought (196) The drought kills the armyworm and natural enemies (mainly parasites) but the armyworm can recolonize rapidly

The natural increase insoil fertility from mineralization of soil N over the drought period promotes luxuriant plant growth to foster a rapid buildup of these highly fecund species Weeding rice increases tilelikelihood of damage The parasites normally return to control the armyworms but not until after serious damage

Outbreaks on upland rice have been reported in Ghana (3) inzibar (38) Central Africa (36) lPanama (165) Brazil (28) Malaysia (196) and India (167) Defoliation may be severe often leaving only the base of stems Armyworm larvae hide during the day under litter or in soil cracks At night they ascend plants to feed Larvae pupate in the soil

Thrips Thrips frequent upland rice but damage is less than in the lowlands Thrips larvae and adults feed on leaf blades by rasping causing leaf rolling and stippling Thrips prefer vegetative stage rice and their survival increases during drought (heavy rain washes them from foliage) although drought stress usually outweighs thrips damage Drought and thrips cause leaf rolling in upland rice (235) but thrips have been overlooked perhaps becaLse of their small size Thrips are most numerous indry season irrigated rice The combination of favorable plant growth and lack of rainfall is ideal for thrips In contrast the lack of rainfall in upland rice hurts the crop and eventually the thrips whose numbers must decline

In [atin America Frankinilla rodeos attacks panicles before they emerge from the boot calusing sterile grains

Stem hug~s Black bugs are the most common group of rice stem-feeding -lemiptera in Asia The most frequently mentioned species ----Scotinophara coarctaia and Shrida - prefer aquatic habitats and therefore could not occur in upland rice as sometimes reported Upland species in the same gerurs - notably S tarsalis and S scotti -- are morphologically similar and remove sap from stems of grasses and upland rice The upland species are notably more abundant in upland rice planted near forests

In Latin America tile Tibraca ima-large pentatomid tiventris causes deadhearts and whitehcads on upland rice Both adults and nymphs remove sap from internodes of plants more than 3 wk old Tillers are killed by mechanical damage and by the entry of secondary microbial infection This damage is often confused with that of lepidopterous stem borers (233) Eggs are laid on the leaves and stems Nymphs are gregarious This species can also be found on irrigated rice Drought reduces population buildup

Stiem borers Lepidopterous and dipterous stem borers are widely recorded on upland rice but cultural practices and alternate hosts determine abtndance Except for Elasmolmll)uS Iinosellts numbers usually build up toward

IRPS No 123 January 1987 13

the end of the crop growth cycle Therefore late-maturing varieties staggered planting and lack offloodingfavorstem borer buildup Many stem borer species can lie dormant in the stUbbleafter harvest and theirsurvival is encouraged by panicle harvesting (leaving tall stems) and lack of tillage after harvest

Stem borers disperse and lay many eggs Many species are polyphagous however the monophagous MaliarpiasepashyratellaDiopsis Ivmgicornis Scirpophaga incertlas and S innotata can also be abundant on upland rice (hilo Suplressalisdevelops fasterand becomes la rger when reared on upland than on lowland rice (226)

The lesser corn stalk borer lasmopal tt Ittoselhs is semisubterranean and is perhaps the stern borer most adapted to upland rice(129) Highly polyphagous it infests maize peanut and cowpea It attacks seedling rice (one larva can kill tip to four plants before maximum tillering) (195) (hiv and Maliarphastem borers prefer older plants But Scirpophagaspp will attack young plants even in the nursery lasn opalls larvae tunnel into the stems at or below the soil level mausing dcadhcarts Larvae are not found in their turnnels inside sterns beca use vhei disturbed they retreat into cases made from soil particles bound by silk The cases areattached to the tunnel entrance (143) This behavior probably evolved as protection from natural enemies Larvae pupate in the soil The lesser corn stalk borer is more abundant during drought damage is often confused with that of drought (233) This species prefers sandy soils arid its distribution within a field may relate to soil texture (143)

Diatraeasaccharalis aside from the lesser corn stalk borer is frequently a stem borer pest in Latin America According to Teran (233) rice and maize are more preferred hosts than sugarcaine its namesake

Atcitonacrvsograpla is a less known upland rice stem borer in Asia with habits similar to Easinopall s (82) because it is not found in lowlands Acutra has grassy weed alternate hosts As a moth it is often confused with the dark-headed stern borer(hi polur(standstriped stern borer (hi sup)lpsrcais

Other common upland rice stem borers are also reported in the lowlands The dominant species in upland rice vary greatly with location In Uttar lPradesh India (78) and in Japan (123) the pink stem borer Sesamia htfirens is more prevalent hIKenya the principal rice stem borer is Chi partelus

(92) The lowland African sten borers Aialiarpha sepashyratellaDiopsis vngicornis and Sesamia calatnisnis are less abundant Oimupland than on lowland rice(217) but perhaps cause greater yield loss because plant injury is greater if combined with drought css (92) The stalk-eyed stern borer D)ivpsis is more prevalent in wet habitats (6 1I217)

In Central and West Africa Maliarpha and Sesamia calanistis are the dominant stern borers of upland rice (8 9 10 II) (lilo dihlfilineui and Chio partellus inhabit the

14 IRPS No 123 January 1987

upland savannas whereas Chilo zacconio prefers the lowlands (II) Aaliarpha is most abundant in the mangrove swamp habitats (91) coastal regions (16) or rain forest zones (II) of Africa and appears to be the homologue ofScirlpophaga incertulas The yellow stem borr can also be prevalent in upland rice (120 167) The sugarcane borer Eldanasaccharinais so far only a potential pest of uplandrice (91) despite Grist and evers (82) warning

Grasshotl)ers andI locitas Short-horned grasshoppers and locusts -- Locvta Patanrga Schistocerca - layeggs in the soil inhabit grasslands and develop into swarms to seriously damage upland rice from time to time (173) Largenumbers can destroy a field leaving only the stubble In low cmuoroer s giassnoppeirs an cut panicles (173) Farmers may not plant if locust swarms are imminent Prolonged dryweather followed by favorable rain favors development of large swarms (173) Usually breeding in di) grasslands theydisperse toattack crops including upland rice(191) Eggsaresusceptible to desiccation d uring drough t and toinundation during seasonal rains With favorable soil moisture over consecutive seasons natural enemies ca nnot prevent a largepopulation increase leading to migratory swarms (64)

Most grasshoppers on rice are adapted to uplands (107)but lowland rice grasshoppers iervgl(rlhusspp and Ora spp occur occasionally on upland rice (2 82) They lay eggson rice foliage Nymphs are scmiaquatic

The gryllid Lsctrtus coucinnus is a pest of lowland and deep water rice in Bangladesh and Thailand In the Philippines it damages upland rice grown in isolated pockets near lakes The gryllid nymphs and ad ults feed on the central portions of folrage leasing only the midrib and leaf nargins

Leal ler Pvralid moths whose larvae fold leaves to make a feeding shelter occur worldwide (217) but arc appareritly most common in Asia Leaffolde rsattacking rice in Asia are a complex of species (26) (Ctalhahcrcisinedinalis the best-known rice Ia flolde r in Asia may prefer grassy weeds over rice (86) It is a late colonizer of uplandrice U1arasmja eviua a lowland species (220) nreflers rice to grass specres (86) and colonizes upland rice during il early growth stages llarasnria patnalis is the most corn-rnonly encountered species in upland rice in the Philippines Perennial grasses are their alternate hosts and they can survive year-round in upland rice environments The leaf-folders 11arasmia ruralisand Alarasmia spp are least aburndant Dormancy is unknown in leaffolders Cnaplha-ocrocis medhnalis however is known as a migrant (90)Leaffolders remove photosynthetic tissue attack during the flag leaf stage is particularly injurious Popurlations are normally held in check by natural enemies but leaffolders become parlicularly abundant in conditions of high plantfertility and shade

Butterflies Skippers PelopidasInathiasand Borhofanta are more prevalent in African lowland than in uplandhabitats (6) but extensive studies in Japan and China

showed that Pelopidas mathias prefers an upland habitat (155) Parnaragutata seasonally migrates between lowland and upland areas and is equally abundant in both environshyments (164)

The green horned caterpillar Melanitis leda ismene is more abundant in lowlands (212) but our experience in the Philippines suggests that Pelopidas niathiasand 11elanifis leda ismene are often more abundant in uplands than in lowlands Bamboo iaunbuiva spp annually sustains P mathiaspopulations in the Philippine uplands Adults feed on nectar from flowers and migrate to upland rice during the late vegetative stage

iPi erlled h i-1 1d dill L tIC i 7-0 l(become numerous probably because of high egg predationEgg and larval parasitization are often low Butterflies can readily disperse long distances to seek renlote uplandricefields However they have low biotic potential and even it predation rates are low the rarely become abundant Upland environments with their diverse microhabitats can provide more favorable sites than lowland rice plains

tutterfly pests of rice have broad host ranges and become dormant Juring urnfavorable times of the year

IolhagrotslaidopwraThe larvae of many polyshyphagous moths are reported to defoliate upland rice tussock moths lsaispelnnatula Iawlia sUifita luproctisirgtwltaE mnorand P vathorrhoeaslug caterpillar

litotiahicolorhairy caterpillars Nisaga sinp vandA mata spp and wooly bears l)iacrisiaohliqtia and Crvatonotts gangis Many of these polyphgous larvac migrate and are transients in rice including HelhWhis arm iera and 1t zea highly pol phagous species (82)

Gall nidges The Asian rice gall midge Orseoliaorrzae (88 131) and its African counterpart 0 ortzivora(6 11 12 32217) prelerlowland to upland rice Gall midges in rainfed areas are prevalent only during tle rainy season and dispersepoorly Their alternate hosts (wild rices LeersiaIschaetnuan and Pasahmt)are mostlyaquatic grasses and are less likelyto be near upland rice areas In an area of mixed uplandfloating and rainfed lowland rice cultures in northern Thailand upland rice which is ilanted a month earlier was attacked firstbut subsequcent populations were much lower than in rainfed lowla id or floating rice Low numbers of gallmidges passed the dry season orr perennial weeds such as Paspahim (88)

Smot (hid(s Reports of aphids attacking upland rice plants are few (249) Without maize the maize leaf aphidRho alosihhunm maidis attacks upland rice in the Philipshypines The plum peach aphid IIrsteroneuraselariae was reported in Sierra Leerne toinfest leavesand tnriperied grain(7) Siphaglrctriaecaused transverse linrea r necrotic striae iir rice leaves in Italy (172) I-lea v rains held populations in check Rice is not a preferred host for shoot aphids

Leafhoper andI )anthol)pers Cicadellids and delshyphacidsdispersecasilyand readilycolonie upland ricefrom nearby lowland areas A study in Sarawak (241) showed

IRPS No 123 January 1987 15

similar species in upland and lowland rice Nephoiettix The principal vectors of rice virus diseases generally have species dominated The most prevalent species accounting narrow host ranges and alternate hosts in the grass family for 90-95 of the sweep net samples wre N virescens N are mostly annuals dying in the dr3 season Some virus umropictus Nilaparvia hgens Soatellafiircifera Cfana vectors such as BPH can disperse lontg distances but few do

sectraand Recilia dorsalisLowlands had higher popula- Th restriction of upland rice and weed hosts to the rainy tions Katanyukul and Chandaratat (120) monitored hopper season the small area planted to upland rice and the hopper numbers in upland and lowland fields of northern Thaiiand vectors having narrow host ranges all work against virus and found similar results A comparative study of plant- diseases in upland rice hoppers in Fiji showed more brown planthoppe BPH) and In Latin America Sogctdhs o(rzicola and S cuhanus whitebacked planthopper (WBPH) in wetland rice than in transmit hoja blanca virus disease but reports on upland lowland rice and hopperburn was prevalent in lowland rice are rare except in the favored upland areas of Colombia areas with standing water (89) Peru and Venezuela (35 42) S rricolaocurs in Brazil

Hopperburn in upland ricefields in Asia is rare (84) In on upland rice but hojaI blanca has not been reported (67) Batangas Philippines isolated fields of upland rice become S orzicola has a wide host ramgc (10) similar to WBPI- in hopperburned infrequently from WBPH A report from Fiji Asia WBPH however is not known to transmit any virus cites hopperburn from BPH and WBPH (250) In these two disease situations upland rice was grown on highly fertic soil and In Africa pale yellow mosaic virus is endemic to swampy N fertilizer was applied Hoppers respond to better nutrition lowland areas where rice ratoons year-round (19) and is Normally upland rice is grown on poor soil and receives no mechanically vectored mainly by hispid beetles A minor fertilizer vector a flea beetle Chaeiocnema spp is an upland rice

In Asia upland rice ras are normally not far from pest and upland ricetields near endemic areas occasionally lowland ricefields-- a source of hoppers Otherwise hopper contract the virus species with greater polyphagy than BPH or Nelhotettiv Spittle bugs Cercopid nymphs producc a protective virescens would be dominant frothy covering that looks like saliva In Latin America

In Africa leathoppers and planthoppers do not become spittle bugs damage upland rice In parts of Brazil where abundant on upland rice (6 II) probably because lowland upland ricefields are surrounded by pastures adults rice is not widely rlanted nearby The only reported migrating from grasses can damage rice severely and cause hopperburn as from Nilapanrata maeander in breeder complete hopperburn in a young rice crop within a week of plots receiving high N rates in Nigeria (98) Hoppers colonization (194 195) Dispersal to rice occurs even if however are a greter problem in Latin America where pastures are verdant and spittle bugs prefer pasture grasses large grassland areas breed polyphagous species (Gra- to rice (168) Nymphs are rarely abundant enough to phocephala spp Hortensia spp lEvitianus ohscurunervis damage rice Regular rainfall favors their development Balclutha spp and Draeculacephala spp) vhich then (233) disperse to cause hopperburn on seedling rice (68) Rosetto et al (195) reported Graphocep7hala spp and Hortensia spp Seed pests to be equally abundant on upland and lowland rice Alydid and pentatomid bugs are seed pests of upland rice

The leafliopper and planthopper epidemics in Asia that worldwide In Asia Leptocorisa sp prevail Their habits caused severe losses from hopperburn and vijus dicases make them particularly suitable for upland rice 1)thev feed over the past several decades occurred principally on on many grasses (and therefore can survive the earl y wet lowland rice Except in a few isolated cases (160) upland season before rice sets its grain) 2)adults are long-ied and rice has been spared from tungro grassy stunt and ragged mobile allowing them to find isolated plantings 0 rice stunt virus diseases vectorcd by green lcafhopper (GLH) (besides seed bugs only birds and rats appear to have shyand BPI] (123) Virus diseases have occurred in upland rice capability) and 3) adults aestivate in forested areas or where lowland rice was nearby Rice dwarf disease however sugarcane fields during the dry sca son when neither rice nor was first recorded on upland rice in the Philippines (187) grassy weeds are present (I199) Rice seed bugs concentrate Virus diseases can be perpetuated only on livir plants or on small-scale upland riceficlds because they can actively insect vectors and no living rice - plans tooned or search them out The prevalence of groves of trees characshyvolunteer - grows year-round in upland arcas excert in a teristic of upland environments ensures nearby aostivation few with 12 mo of rainfall sites keeping the seed bugs close by

The smaller brown planthopper Laodcelphax striatellu Leptocorisae are uncommon in lowland rice plains occurs in upland rice in Japan (18) It spreads black- because 1) their populations are diluted in a sea of rice streaked dwarf and stripe viruses between winter wheat and planted more or less at the same time (they ca n only feed on barley to lowland rice Early planting of lowland rice rice during the milk to hard dough stages) 2) they lack nurseries spreads these viruses but upland ice has remained aestivation sites and 3) weedy fallow areas are limited uninfected The lowands of Asia are dominated by L oratorius

16 IRPS No 123 January 1987

which lays its eggs hign on tile foliage (199) L acuta and L solonlonlesis oviposit at ground level the former on litter and the latter loose on the ground These egg laying habis explain theirtenvironmental prefcrencecs(214) In Japan the postdiapause adults of L chinensis migrate from moun-tainous areas to upland ricefields during floweiing (1 14)

Pentatornid bugs prevail in Latin America (8182 195) and in Africa pentatomid and coreid bugs arc equally cited (6 82) l)amage is often characterstic of the species (5)rcoris ventralis and Mnida spp are the common

pentatomid seed bugs of Philippine upland rice but they prefer lEchinochva spp grass seed to rice Vearaviritda occurs worldwide and hoften more abundant in the uplands(54) We saw few reports of seed bugs on upland rice but see(d bugs apparently opelrate cqually between by land and upland rice Good dispersal is necessary ior species that call feed for only 2-3 xon a crop and must first find those crops at ripening stages

Other pct species feed n flowers In Mato Grosso Brazil (194) and Santa Cruz Bolivia the panicle weevil Aeo-hariuia amllitarsis attacks rice during flowering (68 233)The larva e rice sten horers adutlts feed at the bases of spikelets caIusing empty grains Also inLatin America thecydnid -lkiituttatralusisa seed bug as anad tilt and nymph (194)

A number of beetles ecd on poilen - coccinellids lialrotia Aulac lhora and Monohpl-a (55) - but becatise riCe is self-pollinating and fcrtiliation occurs before pikelets open they pose no threat to yield

IST ECOLOGY

We now look at characteiistics of the life cycles of upland rice insect pests to lea rn hew each plays an adaptive role in upland rice ecology

Life history strategies Orzasativadoes not tolerate drodgh well so upland rice is highly seasonal normally grown in the wettest months of the year Usually rice is present for less than half of the year The rice-fiee fallow poses serious problems of survival to upland rice insects which have evolved att least four mechanisms to ovrcolne tle cvcl ic-l lack of a hostI) polyphagy 2) longevity 3) dormancy and 4) vagility (dispersal)

Poldt)agy lolyphagy is defined as having hosts of at least two botanical families oligophagy of more than one gents and ionophagy of only one ventus (40) - inour case Orvza Most wild rices however are illi tiC and theieiirc are normally far removed from upland rice hahitat othewise more would be alternate hosts for uplanJ iice insects Thte highly mornophagous rice pests shySciriol)haita hiceritlas Nilal)arvaiahiens Iand Nealo--tettiyvircsctws in Asia and Aalirpha secparatella and 1ioj)sis lngicornis inAfrica - occur on upland rice but

are more abundant in lowland culture They can specialize in rice because they are highly vagile and can attack all growth stages By specializing in rice monophagous rice pests can outcompete related but polyphagous speciesIntensive andI extensive lowland rice culture has favored monophagous species Riqwla alhinella (Cramer) the onlymonophagous species in Latin America is rarely reported on upland ice (I1)

Because of its limited temporal and spatial existence upland rice has not favored the evolution of specialized species Clhang(44)shows evidence that upland rice has only recently been cultured by man from the lowlands thus pests have had little time to fully adapt to the crop All but the lowland-adapted insect pest species have alternate planthosts to rice Root pests with limited mobility are highly polyphagous Pests that specialize in one growth stage eg secdling maggots seed bugs are oligophagous or polyphagous

Most alternat hosts of rice pests are annual grasseswhich also are seasonal Upla_rice is also a host but not the preferred one of the most highly polypliagous spcciessoil pests grass hoppers armvwarrs and Vearaviirielda

The d ry season poses igreat obstacle to ani upland riceinsects survival and only sonic perennial plant hosts are suitable food during the off-seasoin Plolyphagy allows upland rice insects to suLrx-ixe during thC rainy season when plants- both annual and perennial -areactivelygrowing and therefore more nutritious

Loiugevitr The ability of aiinsect to live 2-3 mo without undergoing dormancy helps species sich as root-feedingwhite grbs (as larvae) and secd-feeding bugs (as adtilt) to survive unfavorable periods White grubs arc not highly fecund and their life strategy is to improve survivorship of limited progeny Their subterranean habitat hides them from manv natural enemies lBtut root tissue is not highly nutritious so they ltilst eat great quantities and need a largeinsect biomass to digest it Other soil-inhabiting rootshyfeeding insects have short life cycles but are more fecund

Seed pests need greater longcvi vas adults to locate ahost at early grain dcclopment arnd give sedentary offspring time to develop Few groups of upland rice insects live longin active development Polyphagy and longevity only increase their ability to survive during the rainy seasonOther mechanisms are needed to survive the dry season

orantc In the tropics the nonrice season is the dry season while intemperate areas tile off-+season is winter Many upland rice insects undergo dormancy inlsummer (aestivation) or winter (hibernation) Dormancy is simplyinactivity during winters with prolonged temperatures below 10 C insects find shelter arnd cease movement Activity is resu med with warmer t empcera tu res A deep state of irnactivity termed diapauise occurs in sole species that prepare physiologically (havxing low metabolism and convcrting gonad s to fatty fuel reserves) for the unfavorable season (148) They enter diapaUse inanticipation of unshy

IRPS No 123 January 1987 17

favorable weather and resume activity after favorable Armyworms skippers and locusts actively travel hundreds weather begins usually in response to photoperiod or of kilometers in air fronts such as intertropical convergencethrough a delayed response to a stimulus such as heavy zones to descend onto upland rice fields far from their rainfall (237) Not all individuals of a population have the breeding grounds (31) Several generations can build ip on same threshold for environmental responses for entering or alternate hosts in tile grass family before rice is plantedterminating diapause -- this is seen as a mechanism to These large insects are active fliers Smaller insects such as prevent total mortality if a false cue appears such as a thrips aphids and planthoppers passively migrate long sudden warm spell in winter or rain in the middle of the dry distances by the wind season (154) Otherspecies ean readily travel tens of kilometers - stem

Probably more upland rice insects undL go dormancy borers leaffolders leafhoppes cutworms spittle bugs leaf than we realize Candidates are Atherigona spp root beetles seed bugs ants and termites aphids root weevils crickets leaf beetles thrips and seed The least dispersive species are white grabs mealybugSbugs Insects known to entervarious states of aestivation are seedling maggots crickets weevils and gall midgesleptocorisaspp adults (199) Maliarphascparatellalarvae Each species has evolved a unique set of attributes to (9) Scirpophaga innotata larvae (117) Chilo partelhtsv enable it to survive and adapt to the changing uplandlarvae (21) ScirpopThagainccetulas (179) white grub larvae environments Several evolutionary avenues lead to fitness (134) Patangasuccincia adults (6 82) and Schistocerca (Table 2) but there is no reason to believe one set of gregariaeggs (82) attributes is better than another (224)

Insects known to enter hibernation are LodeIphaDstriatellus nymphs (124) Chtio Tvlpressalis larivae (125) DroughtSesaia inferens larvae (146) but not S calavisis(2) Well-drained rainfed upland soils are subject to drought ifIDiaraea saccharalis larvae (75) Scirpophaga- tcerlttas rains do not fall within 2-3 wk Prolonged drought followedlarae(5 lacchrli s lae (75)saSlOhagasshoperseggs (82)and felo pidas athias pOpae (232) by favorable rains stimulates armyworm and locust outshybreaks but mealybtigs root aphids and thrips become

Lcpmotorisa oratorisand L acuta adults aggregate on numerous onlyafter dry spells for two reasons First heavy trees sugarcane or other shady moist sites L paluwanmsis rain normally kills soft-bodied foliage-feeding thrips and confined to Sulu and Palawan Philippines lives on the mealybugs and soil-inhabiting root aphids and root mealyshygrass Brachiariamutica (Forsk) Stapf in the absence of bugs Drought eliminates this source of mortality Second rice The adults can live for 5 mo When aestivating their the rice plant responds to drought by breaking down rate of metabolism drops and they do not feed They will proteins into soluble N compounds which enter the phloem however puncture plant tissues of their aestivation host in and are taken up by these sap-feeding insects (245) Greater search of water This behavior has led to errors of the host nutrition therefore leads to greater survivorship and range of rice bugs (I) Aestivation quickly terminates with fecundity rain probably meaning aestivation is not deep Small rice area

Some stem borers aestivate as mature larvae in rice Except in the highly mechanized large landholdings of stubble or in straw stacked as livestock feed However Latin America upland ricefieldstypically are patchy within moriality increases proportionally with the duration of the a highly diverse flora The small fields result in higher ratios dry season even if the stubble remains undisturbed If the of perimeter to area than with the typically larger lowland stubble is plowed for a crop following rice few stem borers ricefields Upland ricefields themselves may be intercroppedsurvive Rainfall terminates dormancy but it normally with cereals legumes or root crops Mixed intercroppingtakes several weeks for moths to develop and emerge with a wide variety of species also occ in tribal slash-andshy

Last-instar white grub larvae tunnel 1-2 m deep in the soil burn agriculture (50 74) to construct pupal chambers in which they aestivate The Upland ricefields tend to be small because of the labor larvae are very sensitive to dryness so pupal chambers are needed to clear land in slash-and-burn areas and to till land sealed to coierve body moisture Aestivation terminates to minimize weeds Rice requires more tillage than does with the first heavy rains but as in stem borer 3-4 wk pass maize because rice competes less well with weeds Uplandbefore adults develop rice is more a subsistence than a cash crop and small areas

Insects may respond differently in each region to cold (I ha) can feed a family temperature or drought Biotypes or loal populations may Small fields favor some insect pests such as some ant evolve (56) Local populations of Scirpophagaincertulasor species that can forage from more permanent field borders Patangasuccincta may have become adapted to prevailing Trees provide food and mating sites for white grubs (134) as conditions well as aestivation sites for LetTocorisa spp (199) Field

iagiliti The ability to disperse combined with high border grasses plus tIhose within a ricefield provide alternate fecundity orshort life cycles enable some upland rice pests to food for all but the highly monophagous upland rice better exploit the temporarily favorable upland habitat colonizers and a habitat for natural enemies

0 z

Table 2 Life history patterns of upland rice insects or insect groups

Host range Longevity Dormancy Vagility FertilityMonophagy Oligophagy Polyphagy Low Medium High Yes No Low Dispersive Migratory Low Medium High(Oryza spp) (grasses sedges) (angiosperms) (lt3 wk) (gt2 mo) (lt1 kin) (lt50 eggs) (gt500 eggs)

Ants xTermites x x x

Mole crickets x x x xx x xField crickets x xx

White grubs x x x xx xRoot aphids x x xxMealybugs x

x x x xxSeedling maggots x

x x x x xLeaf beetles

x xx xArmyworms x x xx x

Thrips x x xx xChilo spp x x xx xMaliarpha x x x x

xLocusts x x Xx x xGall midges x x xxiVephotettL iirescens x x xx x x xLeprocorisaspp x

x xLeaftolders x x x

Sogatellafurcifera x

xx x x x x x x

00

IRPS No 123 January 1Q87 19

All this appeals to favor insect pest buildup but the notwithstanding the release of stored nutrients in response contrary is true Loevinsohn (138) showed that insect pest to drought that temporarily benefits philoem feeders populations respond exponentially with the proportion of land devoted to rice up to about 75 of the area then the M POR ANCE I I P1A 1) RWI P1St S

rate of response declines In most upland rico areas tie rice crop occupies less than 50i of the area- therefore the Visits to upland ricefields normally reveal 1kw insect pests potential for population buildup is low This relationship Loevinsohn (138) and Locvinsohn ct al (139) compared may not hold for seed pests that can locate small ricefields insect abundance using annual light trap atchcs in upland

Another outcome of the highly diverse flora and phical rainfed lowland and irrigated lowland sites in the Philipshyenvironment of upland rice isthateach upland ricearca has pines Although only one upland site was studied data a unique composition of insect pests showed equal or fewer insects at the upland site than at the

For example in Tanauan Batangas Philippines the rainfcd lowland sites and fewer than at the irrigated sites main upland rice pests are rice leaffolder(RLF) armvworm (Table 3) M Lltiple regression analysis showed that the key Aythntnaseparata WBPH ants Soli psis gentitataand factors explaining insect abundance among a set of cropping white grub Leucopholis irrorata Sixty kilometers away in intensity variables were the number of rice crops grown per Real Quezon the main pests are seedling maggot Athe- year followed by area devoted to rice Table 3 shows that rigona orae flea beetle (Thacitcmnra hasalis thrips the upland rice site had only I rice crop perym rand that rice Stenchaetothrips hiurnmis and rice bug Leptocorisa was planted in only 20Vi of the area Fertiliter and oratorius In a third Philippine upland rice site in Claveria insecticide use had little bea ring on the trend The armters at Misamis Oriental Mindanao the main pests are ants the upland site applied 60 kg N ha higher than at most Solenopsis gemtinata seedling maggot Alieriona orvzae irrigated sites white grubs Holotrichia in)iUaiuoan t and Lucopholiv Insect occurrence at the upland site was omparable to irrorata stem borers Sesamia ifiArelr (hio auriciliusC that at the Cagayan rainfed lowland site parlicularly in 1982stpessallis Scirpophaga incertulas S innoiata and when drought prevented many farmers fron planting(only

4c ona chrrsurapella root aphids Tetraneura niriah- 40(of the rice area eventually was planted that year) M ore dunrinalisand Rhupaosiphunm riahbdminalis lcaffolders favorable weather occurred in 1980 and 1981 which Marasniapataltis and Chaphalocrocis Medinalis army- supported greater rice areas and consequently more insects worm MIthinna separata and rice bug Leltucorisa The most intensive site for insect pests was South Cotabato oratoriusThe great diversity of insect problems is typical Mindanao where farmers plant 23-25 rice crops per year fer upland rice and makes control efforts more difficult with irrigation The Zaragoa Nutva Ecija site also had particularly regarding breeding for insect resistance high pest abundance and at this site the fields were highly

asynchronous Asynchrony was the third most important Low yield potential Iactor in explaining rice insect abundance Most upland rice is grown on low-nutrient and mineral- Rice cropping intensity in upland areas is bound to toxic soils which when combined with eiratic rainfall remain low Only one rainid upland crop is possible per make upland rice a highly riskycrop for the farmerto invest year (Farmers in Claveria Misamis Oriental Mindanao in costly soil amendments and land management Often Philippines tried asecond upland ricecrop i 1984a ycarof upland rice isgrown far from markets making inputs even favorable rains but no harvest was possible b-cause of more expensive Farmers growing a subsistence crop hosi- drought blast and birds) Rice area isaso bound to remain tate to invest in costly inputs to raise yields low because of the high labor and power rcquirelmenlts to

Even having varieties that could double existing yield prepare the land (because of weeds)and because upland rice potential under present management levels would probably wvill not compete with lowland rice as a cash crop Some not prompt many farmers to use purchased insecticide upland rice sites with prolonged rainy seasons howeverare unless tile risk of crop failure were reduced such as it was planted asynchronously such as the Clacra site with irrigation systems in lowland rice The seasonal abundance of Upland rice insect pests

The Tanauan Batangas site is atypical for upland rice attracted toa light trap wasgrapied for crop year 1980-81 in The soils and rainfall pattern are favorable and because of Tanauan FBatangas (Fig I 2) W13P11 was he most its nearness to Manila farmers have cash resources ftm common rice hopper and its population peaked at the sales of vegetables to purchase tertilier They applied an midgrowth stage as it does in lowland rice D-Vac suction average of 60 kg Nha after particle initiation and obtained samples showed an earlier peak than that from a light trap 25-35 tha yields The high fertility isone reason why RLF indicating emigration W13PI usually emigrate after the armyworms and WBPH developed into large numbers vegetative stage Populations in 1980 reached levels that Farmers however used no insecticide on upland rice caused patches of slight yellowing in some fields WBPH although they did on vegetables probably cause hopperburn in Batangas They immigrate

The poor growing conditions fOr rice generally mean year-round as shown by low levels in the light trap BPH poor growth and low fecundity for insect pests fIeding on it however were scarce in the light trap and not collected with

Table 3 Abundance of rice insect pests and factors in rice cropping intensity of Philippine upland rainfed lowland and irrigated lowland locations where light traps were operated daily for atleast I yr -I

0Rainfed Rainfed lowland Irrigated lowlandupland

Pangasinan Cagayan lloilo Santa Maria Nueva Ecija South Cotabato Tanauan Caaringayan Caaringayan Bangag Bangag Bangag Buray

LagunaCale

Batangas Manaoag 1980-81 Santa Motiica 1982-83 Cabanatuan Zaragoza Avancena NaniamanSolana 1981-82 1982-83 Oton

1980-81 1979-80 Oton 1981-82 1981-82 Koronadal Koronadal1980-81 1979-80 1979-80 1983 1983

Insect (kerosene light trapNephotcttixcatch nolyr)virescens 650 1240 500 1800 330 210 3400 1600 160N nigropictus 210 - 360 18400 5500 2900- 140 90 20 - shy 1700 -Reciliadorsalis 710 - 3200 1700180 890 630 140 170 2400 800Nilaparratalugens 130 690 - - 4200 4600300 520 750 180 60 3300 1600 740 310Sogatellafiircifera 1700 31000 13000 8200310 210 3300 610 270 980 1600 6300 -Cyrrorhinus - 12000 6700600 540 3200 290 160 330 120 410Scirpophaga spp 50 410 620 - - 21000 12000150 180 220 270 30 1000Cnaphalocrocismedinalis 70 70 540 610 270 11000 6100200 140 100 6 60 250Chilo suppressalis 0 60 60 1900 300 200- - 30 20 0 6 3 160 -C auricilius - 0 00 - - 20 3 60 20 3 0Sesania inferens 0 - 80 60 3 80 10 51 -Cropping intensity _ -

Rice crops (noyr) 10 10 12 10 10 10 13Rice area (7c) 17 19 20 20 25 2320 85 85 60 70 40 85 85 90Fertilizer (kg Nha) 60 20 85 80 70 7020 0 0 0 30 30Insecticide (no of 40 60 40 300 04 04 0 03 300 13 13 30 50 40 33 33applicationscrop)

__________

IRPS No 123 January 1987 21

Light trapped Insects Drytnd __ 2(notrap-nght) Suctioned Insects (nom )

50 -)o 9 jO40- SOW ffurciWO Light trp

D-Vc suction

20 to 4

20

Egg porasitizahion (M) 75 60

300 SOOampReOftwciffr

45 30shy

15

-insects (notrop-night) 4 Nileporvata Iuen5

Egg parastization()

Light trapped insects (no trop-nght) Suchoned insects (no m 2 )

6 8 Cytorh~nuls hvidipennii 0

4 05 2

50 0 Insects (noltrop-night)40-ui Ipanh pe g

Moohase 1 Seasonal abundance measured by light trap

Rainfall(etod) and D-Vac suction sampler of whitebacked

to ~ hopper Nilaparvata lugens and lea ffolder60)-t Cnaphalocrocis mnedinalis andplthpe g20 L I II I I I parasites and mtrid predator Ortorhjntus livi-Apr May Jun Jul Aug Sep Oct Nov Dec JaIn Feb Mar dipennis set against cropping pattern daily1 9 80o 19 8 1rainfall and moon phase Tanauan Batangas

1980-81

the D- Vac Cyrtorhinus build up on the crop coterminous are collected in light traps during full moon (109)with WBPH buildup Cyrtorhinus disperses during the Because upland rice shares most of the lowland rice pestsrainy season RLF not readily collected in light traps was and those pest species are generally less impotant to uplandprevalent only during the rice crop rice than lowland rice there is a general belief that insect

Nephotettix virescens was more abundant than Nnigro- pests are less important onpicLus in the light trap collections and on the crop GLH upland than lowland rice Arcview of limited data on upland rice shows an averag of population peaked at the reproductive stage as did zigzag 10-21 yield loss from insect pests on a par with that ofleafhopper (ZLH) Crtorhinus tracked the populations of lowland uee determined by the same methodology (132) planthoppers and leafhoppers Both GLH species were (Table 4) McGuire and Crandall (144) made a similarcollected year-round in the light trap indicating immigra- estimate for Central America Cramer (51) howevertion from nearby irrigated rice areas Possibly because of the estimated 35 yieldlow populations loss in South America based on ainsect number did not correlate with similar pest complex - stem borers and seed bugs shymoon phase although reports indicate that more hoppers attacking irrigated rice in North America This estimate did

22 IRPS No 123 January 1987

Light trapped insects D~~nd (no trap-night )

St plht Suctined insects (nom 2

)

D-Voc suction 1520

1o

0 Lih trap 5 ~105 o5

00 Eggparositizotion()5040 NephOtgtha virneens

30 20

I010 A-Insects (notrop -night)

4 Nephatelh nigropccent i

2 o I vV^J - -- ^ VA____ f _

Light trapped insects (no trap-night) Suctioned insects (no rnE

10 -1208 ReCdO doRsol

6 15

4 10 052 iwxALAiamp 0

Light trapped insects (notrap-night) 2Suc oned nsects (nom )

8 CyrtorhmnusI6o nn]10 i0075

4 05

2 0 0

025

2 Seasonal abundance measured by light trap Roinfall (mmd) Moon phase and D-Vac suction sampler of green leafhoppersNephotettix virescens and N ngropicrus and I zigzag leafhopper Recilia dorsalis their egg 60

parasites and micid predator Cyrtorhinus livi-dipennis set against cropping pattern dailyrainfall and moon phase Tanauan Batangas1980-81

40

5 Lf Apr May Jun

1980 Jul Aug Sp Oct Nov I

-aMar

1981

not account for soil pests and was not based on field data in the absence of pests Carbofuran granules are phytotonicAkinsola (I1) cited mainly data from irrigated rice areas in to rice (239) The prevailing hypothesis is that carbofuranAfrica but estimated yield losses of upland rice between 15 prevents soil bacteria from consuming fertilizer The phytoshyand 30 from insects However losses in Thailand over a tonic effect is through greater availability of fertilizer to the4-yr period ranged from I to 13 averaging 5(106) plant Denitrifying bacteria are known to be more active inYield loss data however should be interpreted with aerobic than anaerobic soils therefore it may not be merecaution (184) The Philippine data were all derived from coincidence that the highest recorded yield losses (22-69)trials on farmers fields under farmers normal agronomic occurred in those trials with 12-30 kg basal Nha On themanagement practices except for Bukidnon and Capiz other hand contro with carbofuran granules may not be aswhere researchers used fertilizer Batangas farmers are good as a seed treatment against seed pests and perhapsatypical because ofhigh levels of fertilizer used The trials in yields would have been higher if a systemic seed treatmentBrazil were carried out at an experiment station but insecticide had been used conditions were similar to those of local farmers The 1979 and 1980 trials in Batangas had two full-

Carbofuran granules were used in every trial as a broad- protection treatments one using carbofuran granules andspectrum insecticide applied basally with the seed An the other using bendiocarb WP as a seed treatment Plotsobjection to using insecticides to measure yield losses isthat protected with bendiocarb yielded higher than those proshyinsecticidescandirectlyorindirectlystimulateplantgrowth tected with carbofuran granules The other trials in the

Table 4 Yield losses to insect pests and yield responses to insecticide in upland rice in the Philippines Brazil and Upper Volta

Yield (tha) Yield loss (5) Location Year Cultivar Protected

with Protected

without Check Total Vegetative Reproductive Ripening

Basal N (kgha) Reference

carbofuran carbofuran Protected No with carbofuran

carbofuran

Philippines Pili Camarines Sur Tanauan Batangas

Pangantucan Bukidnon

Dumarao t-apiz

BrazilGoias Minas Ge~ais

Upper Volta6

1975 1974 1974 1975 1975 1976 1977 1978

1979

1980

1979 1980 1979

1977 1978 1979

Bursiging puti Kinanda Dagge Kinanda Dagge Dagge Dagge Dagge UPL Ri5 Dagge UPL Ri5 Dagge UPL RiS UPL Ri5 UPL Ri5 UPL RiS

IAC47 IAC47 IAC47

14 33 11 36 35 31 30 27 37 26 40 30 40 42 46 35

19 21 09

--

-34 24 24 27 --

29 43 31 42 -

47 25

---

10 28 05 27 27 30 26 29 33 28 33 29 43 23 36 11

15 15 06

29 15 55 25 23

3 13 -7 11

-7 18

3 -7 45 22 69

24 28 35

--

21 -11 -20

4 -

-3

23 7

-2 -23 56

-

-

-__0

6 --

-

-23 14

5 0 8 2

22

-

--

0

-

--18 11

3 -10

4 10

7

--7 -

14 -

-1 6

10 -5

8 9 7

-

--

0 0 0 0 0 0 0 0 0 0 0

30 30 30

12 12 12

99 99

100 100 101 101 102 102 102 102 103 103 104 104 104

66 66 66

Farako-Ba

Mean

1977 1978

IRAT10 IRATIO

38 66

--

40 43

-5 35

21

-

10

--

--

--

- 242 243

Z

tJ

-w

24 IRPS No 123 January 1987

Philippines which had two treatments offering protection In the 9 Philippine trials where yield losses were measuredwith and without carbofuran granules relied on further by growth stage 10 of the yield loss (21) occurred at thecontrol with foliar spray and not on seed dressings The vegetative stage and 517 occurred at each of the reproshycarbofuran-protected plots had an 18yield los compared ductive and ripening stages Ibis result is similar to thewith only 10 with foliar sprayS This yield difference is pattern of yield loss in lowland rice measured by successiveprobably due to differences in protection against sown-seed treatments where the crop is unprotected at each growthpests (foliar spray do not control sown-seed pests) rather stage but proticted during the other growth stagesthan phytotonic effects became basal fertilizer wa used in However the insect complexes r sponsible for vegetativeonly 2 of the 10 comparisons stage yield losses in upland rice are entirely diftfrent than in

The recorded yield losses were not related to yield lowland rice The key vegetative stage pests in lowland ricepotential (Fig 3) One might expect higheryielding crops to arc aquatic whereas those in upland rice are sown--seedhave higher percentage of yield losses but this ws not tile (sown in soil) root and seedling pests This dift rence maycase One reason is that insect damage exacerbates plant explain why agricul torists have overlooked tile ilmportanccinjury from drought stress low yielding fields from d ought of upland rice insect pests Many reports of upland rice pestsstress would show disproportionate differences between in Asia ha Ve focused on major lowland species (46 108)insect-protected and -unprotected treatments Shoot and It isalso surprising that tei vegctative stage of upland rice root pests would be more responsible for losses from would record the highest yield losses becatuseseeral studiesdrought stress than sown-scd pests A second reason have shown that upland rice can readily recover from foliageconcerns seed pests which would have a greater impact on reinoval during that stage (117 152 165 188) Upland ricelow tillering low yielding varieties such as )aggc Low poscsses a hitgh obilitV to compensate for ca rly loss oftillering varieties nnot fill in the space created by removal foliage and will even be stinulated by foliage removal toof seeds oweer 1) R i 5 is high yielding because it tillersP1 produce a higher viel than plants ithout any foliageactively which can close the canopy to provide greater weed removed (63 192 231) [ie reason for this apparentcontrol discrepancy hovce is that yield loss at the vegetative

stage does not octUr frol leat arca loss but fron pestsYield loss () removing sown seed and feeding on devcloping tillers and roots -- damage that cannot be readily conpLtnsatCd for

70 - Measuring yield losses in upland rice cnvironrnents is further complicated by a report from Brazil that greater yield losses occurred in response to better insect control60 - =-09 as Insecticide seed treatments resuled in denser plant stands

(n=21) which in turn created a selectively more favorable environshyment for blast (69) This resuIt was parlticurla rlxexacerbated50 when cirbofuran granules were used in seed furrows Aside

ffrom a greater plant stand ca rbofuran iay have auigmented the N lecl and stimulated blast If so ca rbofuran should be40 - replaced with another insecticide or the seeding or fertilizerI rates decreased

Other methods to determine the impact of upland rice30 shy pests wvcrc to interview farners or to develop single speciesbull bull correlations of population levels and yield loss Feneira (66)2 NIM)st (75(1) Golas farmers reporteddid both in Brazil

20- insect losses in ufpland rice and 251 said those lossesS avcragcd 37(1 [his result is consistent with tie yield losses 0 0 measured by field trials in the same statc(Table4) For every

I0 4 of seedlings removC d by leaf-cutting ants a I 7yield loss is predicted Similarly Ferreira (66) found that ai

0 - average of 53 thrips per panicle before panicle initiation meant 3 times more unfillh d grains

-IO0 I 1 1 I I-101 0 2 3 4 5 6 7

Yield (tho) C)N [ROt IM FI IODS 3 Correlation between upland rice yield and yield lols caused by Upland rice insect pests may be controlled by culturalinsects measured in 21 trials in the Philippines Ivory Coast andBrazil genetic biological and chemical methods

IRPS No 123 January 1987 25

Cultural control dry season plowing will have little effect on them Plowing Crop husbandry to reduce upland insect pest populations when the soil is wet from the first monsoon rains will expose can be classified into practices effective in a single field and soil insects that tunnel close to the soil surface to predatory those effective only at the community level Cultural birds chickens dogs and even man (field crickets nole controls should come first in a pest management program crickets and white grubs a re delicacies in man tra ditional They have broad stable effects as pests have little pos- diets) sibility of overcoming them through selection of biotyps The greatest degree i1 inIscct control fron tillage comes (138) On the other hand some cultu ral controls work only from plowing soon after rice ha rvest when most soil insects with high labor or power inputs and connunity-wilt lie close to the soil surface anrd stern borers are in the rice methods require coordination of iany farniters stuble Whitegrubsare mature and tillage exposesthem to

Pantin time The luxury of being able to shift the time of predators Tirnelv control will prtec the crop following pla ntring upland rice is open onl hr la rniers in areas with a rice and lowr the populationIfor succeedinig years Plowingprolonged rainifall pa ttern lI copping intensity or underthe ice stulble helps it decompose killing sten borer rnechanied land preparatiot Gcnerally planlting as soon larvae inside as the rFiinv season begins will lower populatiois of 1110st [requent tillage destroys ait nests bit not termites insect pests Insect populationsarelo aftcra dlvseoll or Tillage and clean culture howes er rcliove food for winter fallow delayed plaiting lets them build tip on termites which then mtay attack a young rice crop Scarab alIternate hosts that gi vigorotsly with the first seasonial beetles also prefer to miposit in eceitly tilled fields Noshyratiris This first flush iof weds cros lItxuttriartlv with iceCult is dibbling faors the survival oftillace Clre Such rineralied N released during the dr season aid made stils i aels tnd insects thtt pttpate or lay eggs- in the soil

available to the plants with the rains It iliso igrtmitcally Zero tilla hio eerl conserves predators ofthe GIH (37) advanttgeous for the rice ciop to tap this trlLItlaI fertility nt it mcttoi lo lessen the impact o0 soil insects and grow vigorotsly to compete with wrus and to tolerate brotdcasting seed or planting in ftltrrows is prefc ired over pest infestation dihlirig in hills Crickets are large enotlgh to eat seedlings in

Larly plantngs terd to escape seedling maggolts Upland a hill atrd tillering normnally ca iriot fill in tlhe missing space rice in Bitangas is planted in Ma and ine itli the Iirst Oftcn dibbll rd is not well covered and iseasily found by rains Fairners are highly mtotiated to plant early tocscape foraging insets ioIdents and birds drlitght and typlotns ricar harct Arn Atrgtst planting Plan uit sii Incrcatsini sced density protects tile cropwould be evcely attacked by It riim orivzo At tti againrst seed and seelling pests especially ants particularly other Philippiie tipland sites Rea l Qteot antd (laelit if the crop will germinate and ermerge quickly M isainis Oriental seedlig mnaggots arnrige ric boe t se hr trftinll itI India rice intcrclopped witi cotton or farnes have a longer growing season and plnit 2 rin or pigeon pci had lower 6IIt and WBIlII poptrlations than mote after the ortset of the rainy season liaaiangas ftrmners rice alone (211) NfailC arid Urplald ir_ 2re coilltori in the plant ca rly becia use ofa shorter raiinfall period arnd the desi re Philippines he chatngC in pest status folii iniercropping is to lliarvet before the typhoor sea son peaks ltighly locatioll-specific (133) cmianarid the net elfc be either

Early plantings cornbind with early-iatiuring varictics nil bcnelficial ortdetriential (189) For example BIatangias provide higher crp tolerartce for white grubs Aestiatior farmers intercrip Itro in their ipland ricefields A liornshyterminates with the first rains but 5-6 moare ineeded for the nrii iotd-Ahis i (I iiiacius) which norlially feeds resting larvae to develop into beetles emerge and lay eggs on taro ill feed or miarbv rice plants Intercropping and for larvae to reach the danriigii g ird-instar stage probably would not afft0 seed pests A crluanion crop (134 In lataigas this normually iseiough tiite for harvest planted ith rice and haing gleater tolerarnce for root

It Brazil howecver delayed planting is a suggested dariage or being toxic nayact isi trapcrop to root-feeditig cultural control for the lesser cornstalk borer and insect pests Acial-fei i ig pests voultId be most affected by pests favored l drought (66 195) Cropping internsity in intrer ropping Shoot aphids leaf beetles sten borers and Brail is low and delayed plraing places the rice crop in the hopplers would be likely ca ntidates to note effcts On the niost stable rainfall period reducing likelihood old rought othlt halnd the corlipalior crop stch~ sitniaie Iiiv be more

In rilniy arca s rinmirsare roi atl to plant other crops benehited than rice from initecropping ( 137) before rice and althNgh it woultl be desirabh to avoid 11echi Weeuling luring the first month after clop infestations by planting rice cuatl lariners lack the labor cstablishent will force pcts such is anyvworris which and time to do so ol a low priority crp prefer grassy cetds ontor tle vorling rice Soil iampsecs will

77age Uplild rice Stils ae light textured and Call be mvnye to the rice crop Cleani citrltirc also lay lore territes plowed duiring the d rv seasor notl only to grin time for to attack I u cetincti inrice ceds i ay he le optinal planting with lie fitst Inilsoon riiains but also to desiccate solition 13) soil insects arid weeds such is ntsedge that foim rhizomes Irtlizr Itneatsing N Iciiliier oin hlwland rice has Htowever becta use most soil insects lie below die plow layer fAtwvored Irigher populains of pliithoppeiv Imlhoppers

26 IRPS No 123 January 1987

and leaffolde rs However because of the need to temper fertilizer use on upland rice o prevent blast fIrtilizer rates will be lower than is comonia r for lowlad rice causing relatixely less hopper and IaIfTfoide r population buildup However inupland rice areas without a history of blast such Is Batangas farmers apply high rates of N and conscqtrently the crop has high infcstation levels of WBPI]and RIF The fhrmrcrs leec insect buildup by splitting the fertili11r in0to two to three applicatiois bit do not applV fertilizer at the vegetati estage because this prodrces a tall crop which illeasily lodge during typhoons Greater crop fertility leads to greater insect survival (better nutrition) larger insects (to cat no re foliage) arid greater fec titd t (to lav nmre eggs) (128 145 171) Not all insect species respondequally hovr Increasing rates (if N l anrid K increased Dialraa s-acchralis and (iaeiurema flea beetle but reduced thrips (66) Zn rcdLuced deadheart density from the lesser cornIstalk borer

Fnldingi Although most Upland rice areas do nIot hawxcaccess to wa ter floodiig the fields is rRrrnriierdcd in areas of Brazil or root bug and white gib control ( 195)

Crop rotatioti Tillage before planting a crop after rc harvest will control stern boreis and other pests renain n the stubble arid will unearth soil pests Planting a non-gratriiaceous crop after rice is recommncrded for termite control (66) Crop rotation is less nttdcd in upland rice culture than in lowlands because atcrop-free period is assured even after harvest of crops following rice

t1ulchilt iryl plastic irli irig designed for weedcoitrol ii upland rice inJapan resulted in ntr highanri liy infestation of (ihilo sulytresvais (93 9-1 95) NIulched Upland rice grew more luxuriantlv and attracted more (Avipositing mothis than u lched rice Also the rice plant and stein borers imatoured 10rC quickly and a partialgeneratiori af vouing ista iNoverwintcred in the stubbleLotili--malitrt varieti Quickly maturing crops reduce the tmbCr of pCst generations that Calt build up Rapid crop establishment olan ca rl v-natu ring vaitvty if)pa 11icLa r will lessen white gib and stem borer damiage White grub larvae intle last iistar are highly destIruCtive so aii early-iiatri ng variety will escape serious root loss Stem borers build 1ip slowlyx in rice to ecormic abundant in a late-Maturing crop anrd caiuse whi tlieads Numibers of somespecies increase exponeitially with each succeeding genera-ti so a xarietyexhat atures I trowh earier thian another will hac less insect damage Early-naturing vaiicties however will not lessen seed pest daiage either ii the soil or oina standing crop

Strwhoi hAIJttam Inrscts tcd irig on thlie aerial portions of rice plats disperse from carly to late plantedfields AS iiiibCirs increase exponentially with eachinsect

gtscnrattm e eiglbrig f rmc~s1(uil upp wh rc generation pests build tIP where neighboring fihrTC e9r1sstagger their plantings II Batangas farmers plant uplandrice within I iro use 120-d s-arieties and plant at the start of the rainy season Ini Misarnis Oriental laricrs stagger their

plantings up to 3 mo apart and use a set of atieties that mature in 4-7 mo Insect pest danage particularly from stem borers is greater in Misarmis Oriental Also seedling maggots damage rice there because rainfall favors frequent planting of maic In Batangas rice escapes seedling maggots bCca usC of distinct drs season and synchron0us early planting with the onset of ains

Therefore combinations of cultural practices ea rlyplanting synchronous planing crop rotation and earlyshymaturing vaneties protect the rice crop against most insect pests

Ilmant resistance Beca tse upland rice has few specialized insect pests and many insect types attack the crop worldwide regional breeding and strong internatiornal cooperation are required

Sown-sced pests arc economically controlled by insectishycide seed treatment and should the relore receive lowxpriori ty inbreeding objectives

Root pests are normally cost ly to control with insecticides but finding resista t sonres mayvbe di fficult Tanaka (229) for exa -ple fi led to find varieties resistant to root aphids Selecting for large root bioniass perhaps should be the strategy for pests sLuch as Whlite grubs that renliove roots Fainter(174) howcvcr cites varietiesof crops tolerant ofor resistant t(i root pests Some sorghun varieties are high in cyanide so rice sorghun might incorporate broadshyspcctruri chemical resistance

Seedling vigor and drought tolerance are high prioritybreeding objectives so tupland rices should have large r root systernsand be able to tolerate ligherlevels of root loss Sonie root loss inolder plants is beneficial if it stimulates new root developnrut and therefore crihances intake of soil nutrients (53) The rice seedling inaggot rtiLotia spp may be

controlled thirough resistant aittics A hi rgc breeding effort on sorghniu has had fair success inideveloping arreties esrstait to Asoccata Rodani (251) A field trial comparing 10 upland ricc varieties showed differences aniong varieties va ring from II to 4(- damaged tilleis (215) Hovver resistarce to 1Iill-0r gtia species may rot cross ox-rti otier

Shiraki (210) reported that upland i-ice stens wcrc harder than lows-land rice stemsperhaps becaIusc (f a higher silicoricontent and thus were more resistant to yellow stern borer hc tortia Iow tillering (f uplaid iice xarneties prodtres a

higher percentage of infested tillers than would occor in higher tillering lowland iice under a siiilar egg density The corollary therefore is to select for high tilleriig upland rices

Stemii bor amcstheh main targes dobrecders inAfrica (571 204 )but nticl oifth e sc reen irig is dotie unde r lo wland co d t osrm s wih A h r~ o l pe i s n acorditions (11 96 218) As with lthrinrnwspecies mayhave to be dealt with independently however initial results show cross resistance between African and Asian species(92)

In Brazil screening Ior insect resistance focttses oil lou I pest problems Varieties tolerant of lasmolwalpus ligi-sellus and resistant to )iatraea saccharalis have been identified (70 153) Current work also emphasies 7iraca limativetntris and Thmialspis (=Iosis) flavouicta (66) These insect pests Ilt onlV citise IIigh yield losses tihey also ire difficult to control chemically ofr by other ilealls

ManV upland rice vaieties aic early maturing and are resistant to pests such as gall midge (I Xl ) Early mIturity is often a highly desirable trait to esuilpe pest buildup but it is not true resista nce bca usc ilthose va reties are plinted late they will be itsdamaged as ain early planted susc ible variety

P~erhaps tile cosiiiopolitall of all upland rice pestsimost are the sced pests (ripenling stagc) which pose a particular problem beta use scuds re ised IsIO for hitnit4ans Ayltoxic substatice introd tcld geticticllyilmay also be toxic to nal iniless it caln he placed old in thfe IdhuffSelectmi fo hairiness or lotng wns kill mIlct sthilrohleiiis dutilig threshing pa rtictfarl maitial threshinfl [hireshers ill complain of itcfimin atid itteascd (fust levels

Attempts itIRRI to fitid tcistatncc to the rice bug plorisa have ilotbeCnl prmitisilg lIIetIattonlids iorc

through the lenilna or palca hillhairs sedstVIor seeds wilh awns that deter bugs promote complaints from threshers Seed bugs liptocorisa and Stcnocoris enter the seed through the opening between tile lemna and palC but varieties with narrow openings are diifficult to mill

Breeding priorities should concentrate ott inotCI gruLtLIps that are most difficult to c nttrol by other meains Steil) horers appear to be the most widespread group that sigiiificantly dalag s rice aind is hard to manage by other Ilnca us

Biological control Very fe stutdic of mitatt~l eneties of rice pests in upland envirollnletits wee in tile wefouiid literature Therefore have not compiled a list of species Ve present a list of natural enemies iOtid in upland ricefields tuic latialln Batangas for some conitno iOldiar pest species ([itble 5)

Muich ofl thc lollowinle itfor ittiol cotlliCs rol Stutdies in the lowlatnds or roi crops otithcr thlin tice

Qu(ailtr o 11ll t11t( Nat trrtf lii hcoeits ight Ile used agaitist ile social insects tilt and termites Sollie predator specice live ill theirf nests n ilnic theirnllt ip-pearanIlCe aMRidbehavior IJtrouMdCing a mote aggressise ant species to displace pest ant codonies is a possibility (169) Patihogens as potential natural encnies have reccived little atteition (I13) Natural ncnuics asa group however hac not been toIl iporta it in Iegulating aintshown be and ternite numbers

The nocturnal behavior aid subteitmien habitat of lole crickets and field crickets protect them from many ntllral enemies There are only limited records of natural cnelics for these groups of pests some pathogens (87) but mainly

IRPS No 123 JattuIny I 271987

parasites - scelionids on eggsand sphecids on ilymphs and adults Sphecid wasps Larra carutmaria (SImitI hI L huzwuensis Rohwer and lsaltiuhta Willi ams speciali in mole crickets pa ralyzing them nd dragging the ninto thcir nests as food lot their voting ()tier sphCcid species Lins aurulenoa (Iabricius) lot maiac (Ashmead ) 1 lhshylssellaus (Smith) and Ml lal)oriosu (Snithf specialie in field clickets

White grUbsO the other hand hae Icti scriuns enuigh to have been extensively studied for cotttroi 1W nati al enemies( 112 140 141) Scoliid parasites C(ajsoMcispp have beetn intrtduced itto tilePhilippites for control of Ielxl ]udisiro la The rcsearchcts w Intttrdced themle during ill era claimed that the white gttionthreak as controlled (142) howevcr it isnormal for pest epidemics to subside and there was no diect evidence to indictte that tfe parasite was responsible A tmore promisitg titctllod might he to identify iso ltcttass prt~f tcartd apply hiactelaln fingal pathogens to tile soil Ihe lalattese beetle in North America has hcci coitrolled hv tite conmnleciall aailable hactcrial prcpa ration itacihlpu pav ofr milksdiscas (71) Spote prtyefott the soil itloted ilrtier reitiain actic fr Veas

Root aphid sardtnctly bugs readilslic tilib to predatots and parlsites once their tetndine ants are controlled The larvac and adtilts of a coccinellid beetle 1Scynms sp prey on llriadlouais in slash and burn upland rice Baits treated with insecticide may be used to control tending ants by killing their ourig (244)

Root-tkcditig blugs false wireoris wireorms and root wccils aic r ofsuch little ifilpoltatice that thenomttll role of liatural Cticniics hias tiot beei asscssed A possibility hoevrl tiglt he tit ItaIIitic niaCIMtodcs lintt1g11en Itese soil-itlhlfbitine mCtitodcs W5uhtfhave to be imiass Produiced

large miltibcr of niatltrll cnemics ircknown ft CltIoriS IC IpblCeii is thtt clttvortnlls ofonlie the crop soott alter land ptel)tritiOti a1tithe bcgit iniig of the raittv

Citsoln hfien tiatllral leniiiy populations aric lo Colltloil WOtuld iii ole mass prodtticing andirclcaslng key spccies

le egg stage is the fpirt (olthe Cedlitgt Ialot life cycle tI hich to f)ctis ftor tittlril elctit cottntrol hplraites o1 pleditors ttsit elleggs highly expoIsYCl I Sgg pailsites or predators stolld probahly e ittass andtbe produced released Littlh i toruialioll is a ailable ott what species to trV o ho to Iniss podLice thiCIi A etiloplhid wasp Icmihlaol sp effctiicly regushy

lates the leatf lcloalapolm2a spioanoiiitr heat andc rice antid should be collsctvcd

leaf beetles whCllose e am] eggs ate laid in the grutindhirO ould be motC difficutlt to control by niatulral enelV Mnll ipuhltiOn But species sUici IS c_gsthispid beetles ith

and larVle oilleascs ate vtilcratble to iparaites and pledators

Natural enemisc of tlirips otl rice are little known

28 IRPS No 123 January 1987

Table S Parasites recorded from upland rice insect pests in Tanauan Batangas Philippines 1977-84

Order Family Species Host stage

I lymttenoptera Braconidae Leaffolders Cnaphalocrocis Marasinia spy

Cardiochilesphilipplinensis Ashmead a Larva

Cotesia angustillasis(Gahan)Cotesia or cypris (Nixon)

Larva Larva

Cotesia or taeniaticornis(Nixon) Larva Cilonusinotakatae Munakata Chelonits SppAlacrocetrts philippinensis (Ashtmead)

Egg to larva Egg to larva Larva

Iclineumonidac Orgilts spp Isclutojppa hireator(Fabricius)

Larva Larva

Itoplectis naraigav(Ashmcad) Larva Tenelhtcha philippiensis (Ashmead) Larva Tenelchha stangli (Ashmead) a

irichomma cnaphalocrosis Uchida Larva Larva

Bethylidac Elasmidac

Goniozms or trianguli r Kieffer lasmtusalhopictus Crawford

Larva Larva

Diptera

Encyrtidac Chalcididac

Tachinidac

Elasmtus spp Copidosomopsisnacohiae (Fidy) Brach nteria evcarinata Gahan Biachyieria lasus (Walker) Ztgohothria ciliata (Wulp)

Larva Egg to larva Pupa Pupa Larva

Argyrophylx nigrotihialis(Blaranov) Larva

l-ymenoptera Braconidac Yellow stern borer Scirpopiaga incertulas

Braco chinensisSzeplegeti Larva CIthelotis spp Chelonus niunakatae Munakata Cotesi flaripes Cameron Stenoracon nicerilli (Bingham) a

Larva Egg to larva Larva Larva

Ichneumonidac Tropobracon schoenobii (Viereck) Amatroinorpiaaccepta metathoracica(Ashmiead)Frihorus sinicus (I lolmgren)

Larva Larva Larva

Ischnojoppaluteator (Fabricius) Larva Isotima nr dammeratiti Rohwer Larva Tenelucha philippinensis (Ashmead) a

Larva Tenehicha stangli (Ashmcad)Trichomma enaphalocrosis Uchida

Larva Lara

Chalcididac Eulophidac Trichogranmmatidae

Xanthopimnpla stemmator (Thunberg) Charops hrachYpterumn Gupta and Malteswary Brachtm eria sppTetrasticttus schoenohii Ferriere Trichorigranm chihunis Ishi

Larva to pupa Larva Pupa Egg Egg

Diptera

Scelionidac Pteromalidae Tachinidae

Tt ichograintnta aponiicuitt (Ashniead) Telenomits rowani (Gahan) a

Trichoinalopsis apattelctena (Crawford) Jeirhaea spp

Egg Fgg Egg pupa Larva

Z4gobothria ciliata (Wulp) Larva

Ilymenoptera lecneumonirlac Gol fringed borer (hilo auricilius

Trichomm enoptalocrosisUchida Larva

Ieioenoptera Braconidae

Aantholimlh steiniator (Thtinberg) a

Pink stet Ioret Sesamia inferens StoInacoil nicrilhi (Bingh am)

Loarva to pupa

larva

Diptera Ilyinenoptera

Armvtnyorms Mythimna separata Spodoptera mauritia acronyetoidesTachinidae Zvgohothriaatropiora( Robinea i -Desvoid y)Eulophidae It-phctrtiscitapadoe (Ashimead)

Larva Larva

(halcididae Braconidac

Brachpin ria spip (oesia spp

Larva larva

IHyntenoptera I)iptera

Rice skipper Ielop ida (halcididae Tachinidae

math ias aidgreen iortedci ciliarNIelan itis eda Brat lhincria sp iii matuqitatt Cameron Irgyrolhirlaxi trotlhiais(marumtv)a1

ismtne Larva Larva

Diptera Ilymenoptera

Tachnidae lelineutionidae

fuown se tilo per Mois frugalisIrgtrophrlax 1igrothiahiv 3 a rat ov) anthopimpa pmnctata (Iabricius)

Larva Larva to pupa

aMost dominant

IRPS No 123 January 1987 29

Normally predatory thrips prey on herbivorous thrips strepsiptera on both Spiders dominate tilenymphaladultNatural enemies of stem bugs --Scolinoplara and Tibraa predators Spittle bugs move to rice from pasture grasses - are in order of importance egg par~isites and predators and biocontrol would be successful only ifcarried out before fungal pathogens and nymphal adult parasites Egg tiley immigrate into ricefields parasites could either be introduced fron other areas or Biological control tactics for hemipterous seed bugs mass produced and released ould follow those outlined for stein bugs

Armyworms are normally lwld in hcck by the activities Coml)arison of environnewtns Natural enemies are of egg and larval parasites When these natural enemies fail perhaps faced with even greaterconstraints than insect pestsusually because of drought armyworns become epidemic to survive ind reproduce in upland environments TheyIt may be worthwhile therefore to release parasites during have adapted ani survived by having wide host ranges the the rainy months following a drought Virus diseases of ability to aestivate or hibernate over unfavorable seasonslarvae would beanotheravnue to exploreforarnyworms and the ability to disperse or by other mechanisms

Parasites have ben traditionallyconsidered for biolegical Comparing the most dominant and therefore the most control attempts against stein borers usually by rekasing adapted species of natural enemies collected in various exotic species( 166) This approach has not met with success environments in the Philippines we can understand how and new avenues should be explored The role of predators different rice environments are arid pathogens is little inderstood and deserves greater Anagqrms olptahiisa mymarid planthopper egg parasiteattention Egg parasitism rates are normally reasonably is cquallyadapted to all Philippine rice environments (Tablehigh but should be supplemented with effective egg 6) It is selective for planthoppers and even parasitizes eggspredators Orthopterans are potential egg predators of the n-iize planthopper Peregrinitsmaidis itt upland areas Metioche vittaticollis (Sta ) Anavipha nifiwnnis (Saus- Alavohis also parasitizes maize planthopper eggs but is sure) and Conocephahs hongipnni (de i-laan) feed on most prevalent on BPH in the irrigated wetlands (onashystem borer eggs etioche and Anaxil)ha specialize in eggs tocerus spp like A optalhilis on BPH and WBPI Iattacks with no hair such as Chilo spp The omnivorous (ono- the eggs of GLH in ell rie environments It is also found cephahsspecializes in eggs cov red with hairs it is a proven parasitizing the eggs ef the white rice licahopper (oimna egg predator of Scirlouphaga spp but has the discouraging spetra Oirosita naias a trichogrammatid egg parasitehabit of eating rice guin (65) attacks BIPH and WBPH in the wetlands and has no other

Grasshoppers have egg (scelionid) and nymphalad ult host A related species 0 aesopi specializes in G1-H in (nemestrinid tachinid and sarcophagid) v irasites Among wetland environments Dryinid strepsipt-an and pipunshypathogens protozoans have been recorded most frequently culid nymphal and adult parasites of planthoppers and Control of locust species by natural enemies could con- leafhoppers also specialize either in hopper species or centrate on their habitual breeding areas environments Among the dryinids Pseuhdgonatoipsflavi-

Leaffolders have rich complexes of natural enemies some feimur prefers the lowlands and BPH while P nuch has no adapted to upland rice (22 27) ranging from egg predators environmental prefirences but is adapted only to WBPH (gryllids coccinellids) to larval parasites (braconids ichncu- The main d ryinid species on GLH aptogonatop)usspp is monids) to larval prcdators (ants and carabids) to larval most dominant in rainfed lowlands and also attacks other pathogeiis (vira and fungal diseases) Efforts to augment leafhoppers within Amnrasca Cicadulinaand Bachaltuathese natural enemies should focus on the egg predators Strepsiptera are more dominant on plarithoppers than (rearing releasing)and viral or fungal pathogens (culturing lealhoppers The Elenchus species in the Philippines attack spraying) all species of planthoppers within the genera Sogawilla

Parasitization of the large conspicuous larvae of Sogatodels NilaparsataHarualia and Opiconsiva Elenshyskippers green horned caterpillars and other polyphagous cusrvasuntaisuiis adapted to the major rice environments Lepidoptera by tachinids and chalcids is normally low ELg but prefers BPH in rainfed arid WBPIH in wetland environshyparasites and predators are perhaps the key to their control ments Ilalictoph(utsmtnroeiand HVpectrus attack G LH

Gall midges are parasitized by pterormalids and platy- and COfimravcctra respectivelygasterids that aestivate within their larval host between Pipunculids are mainly parasitic oii lealhoppers Each seasons Shoot aphids are preyed upon by coccinellids species has a unique environmental preference Tomosshysyrphids and chrysopids varireIa oraetoraprefers Ne7 otetltix nigropictrs and

Planthoppers arid leafhoppers have egg parasites and Pilunwulhtsjavanensisattacks Dehoceplhahls spp which predators as well as nymphialad nlt parasitesand predators are more abundant in upland than irrigated lowland their rich complex of natural enemies includes fungal environments T suhvirescensattacks Nephotettix virescens pathogens The principal egg parasites are myrnarids and and prefers the lowlands Pilpuiulus nttillattls is an trichogramniatids Egg predators are norinally mirid plant upland rice parasite arid attacks only Nephotetix speciesbugs that suck out the yolk Nymphal ad nIt parasites are More than 17 recorded species of leaffolder parasites in dryinids on planthoppers pipunculids on lealhoppers and the Philippines attack species of C(naplwocrocis and

30 IRPS No 123 January 1987

Table 6 Dominant natural enemies of rice pests in upland rainfed lowland and irrigated lowland rice environments of the Philippines

Natural enemy

Host stage Species

Brown planthop-Parasite Egg

MymaridaeAnagrus optabilis(Perkins) Anagrusflaveolus Waterhouse

TrichogrammatidacOligosita naiasGirault

NymphAdult Dryinidac

Pseudogotatopuslaiifemttr Esaki et Hashimoto Elenchidae

Elenchus yasurnatsui Kifune and Hirashima

Environment

Upand Rainfed lowland Irrigated lowland

Ailaparata lugens

x x X x

x

x x

x X X Whitebacked planthoppers Sogatella furcifera and Sogatodes pusanusEgg

MymaridaeAnagrus optabilis(Perkins)

TrichogrammatidacOl~gositanaias Girault

NymphAdult Dryinidae

Pseudogonatopusnudus Perkins Elenchidae

Elenchus yasutatsui Kifune et Hirashima

Parasite Egg

MymaridaeGonatocerus spp

TrichogrammatidaeOligositaaesopi Girault

NymphAdult Pipunculidac

Tomosvaryella subviresccns Loew Tomosvaryella oryzaetora (Koizumi) Pipunclusmutillatus (de Meijere)

DryinidaeIlaplogonatopusspp

Egg Copidosomopsisnacoleiae (Eady)

Larva BraLonidae

Cardiochiles philipphnensisAshnead lchneumonidac

Trichiomnia cnaphalocrosisUchida Temelucha stangli (Ashmead)

Pupa Chalcididae

BrachymeriaexcarinataGahan Brachymerialasus (Walker) Brachymeria nr tachardiaegroup

Parasite Egg

Scelionidae Telenomus rowani (Gahan)

EulophidacTettasticusschoenobhi Ferriere

Larva Braconidae

Stenobraconnicevillei (Bingham) Iclhncurnonidac

Temelucha phiiippinensis(Ashinead)Temelucha stangli (Ashmead)

x x x

x x

X x X

x x Green leafhopper Nephotettix virescens

X x x

x x

X X X X

X

X Rice leaffolders CnaphalocrocisMarasmia

x X x

X

X X

x x x x x X x x x

Yellow stem borer Scirpophaga incertulas

X X x

x

x

x X XAinauromorphaaccepta metathoracica(Ashmead) x

continued on next page

IRPS No 123 January 1987 31

Table 6 cntinued

Natural enemy Environment

Host stage Species Upland Rainfed lowland Irrigated lowland

Larva-pupa Gold fringed borer Chilo attriciiuslchneumonidae

Xanthopimpla stemmator (Thunberg) x aPupa x

Pteromalidae Trichoinalopsis apanteloctena (Crawford) ax

Pink stem borerb Sesamnia inferensParasite

Larva Braconidae

Stenobracon nicevillei (Bingham) x b b Armyworms Mvthinna SpodopteraLarva

Tachinidac Zygobothriaarroptvora (Robineau-Desvoidy) x x

Braconidac Cotesia spp

X Skipper Pelopidas mathias and green horned caterpillar Melanitis leda ismeneF-gg

TrichogramrnatidacTrichogrannaspp x

Scelionidae Undetermined Scelionid

x Larva

Tachinidae Argyroph lax nitrotibialis (Baranov) X x

Pteromalidae Trichomnalopsis apanteloctena (Crawford)

X Predator

CoenagrionidaeAgriocnenis femina femina Brauer x xIschnura senegalensis tambur

Miridae CrtorhinsliidipennisReuter x x

Nabidac Stcnonabis tagalicus (Stal) x

VeliidaeAlicrovelia douglasi atrolineata (Bergroth) x

Coccinellidae Micraspis crocea (Mulsant) X x x

Vespidae Eumenes campaniformis (Fabricius) x

Formicidac Diaecamna spp xOdontoponera transversa (F Smith) x

Carabidae Chaenius spp OphioneaW- aJasciataSchmidt-Goebel

x

Ophionea ishii ishii Habu x x X

Gryllidav Anaxipta longipennis (Saussur-) x xMetioche ittaticollis (Sial) x x x

Tettigoniidae Conocephalus longipennis (de Haan) x x x

aNo rearing made on rainfed lowland bNo rearing made on rainfed lowland or Irrigated lowland

MrasmiaAmong the Laval parasites the braconid Cardio- lowlands and parasitizes oriental maize borer OsIriniachilesphilijpphensis is most adapted to an upland environ- furnacalis (Guenee) larvae The leaffolder pupal parasitesment It also parasitizes Hydelepta indicata a common not only occur equally in all environments but also haveleaffolder of legumes The ichneumonids Trichomma cna- wvide host ranges- parasitizing species of OsIrinia Marucaphalocrosis and Temehcha stangli occur more in the Hydelepta and Honiona - in maize and legumeslowlands T stangli is most adapted to irrigated lowlands The principal egg parasite of yeltow stem borer (YSB)and also parasitizes Chilo and Scirpophaga stem borer Telenomus rowani is widely adapted to all environmentslarvae T cnaphalocrosis is most prevalent in rainfed and has an unusual alternate host -Tabanus eggs (25)

32 IRPS No 123 January 1987

Tetrastictu schoenobii specializes only in YSB and occurs mainly in irrigated lowlands -- further proof that YSB is most adapted to the wetter environments

Larval parasites of YSB are highly environment-specific The braconid Stenohracon nicevillei occurs only in upland rice but also attacks the pink stem borer (PSB) Tieinchwa phiiiplPwnsis an ichneumonid occurs in upland rice and also parasitizes the larvae of (7hiu stqpresaisinatro-morpha accepta melaihoracica like 7 stangi is mnost adapted to the irrigated lowlands

The dark-headed stem borer(l)HSB) (ih pochit3sus was not collected in all rice environments and the gold fringed borer Chilo auricilius nd PSB were collected only in upland rice Xaantiholiplastenrnaloran ichneumonid larval parasite appears to be widely adapted whereas TrihomaslopisapanitI)cteiiaa pteromalid pupal parasite occurs mostly in wetlands and also parasitizes larvae of Chilo suppressais and Pelhuidas mathias

Armyworms Myit hinin and Spodujipera have two main larval parasites - a tachinid Zrgohothriaatropivora in rainfed environments and a braconid Cotesia spp in irrigated lowlands

The butterfly pests Pelopidas and 4elanitishave tricho-grammatid egg parasites found only in irrigated lowland environments

Two larval parasites show environmental preferences --Argrroph av nigrotihialis a tachinid in rainfed habitats and Trichonialopsisapantelkcnela in irrigated ricefields A limgrotibialisalsoattacks the sweet potato hornwormAgrius convolvtdi and T apanteloctena parasitizes DH S1B

Predators as a group show more distinct environmental preferences The aquatic damselflies Coenagrionidae are most abundant in the lowlands Cvriorhinus is also most adapted to the lowlands The nabid Stenonabis tagalicus occurs only in the uplands and isalso prevalent on legumes The coccinellid lady beetles wasps and ants are more prevalent in upland areas AlicrasIpis cit-ea larvae prey ona variety of aphids attacking legumes and maize Etunienes campanonia vespid mud wasp makes nests in trees and is therefore most adapted to the more botanically diverse uplands Soil-dwelling ants cannot tolerate flooding

Upland environments are habitats to arboreal carabid beetles Three species of Chlaeniusbeetles prey on leaffolder larvae and are perhaps more important than parasites (27)

The gryllids and tettigoniids are widely adapted egg predators not only on rice but also on maize and legumes They feed on eggs of most insects that are laid on leaves

The spidercommunity of upland rice environments isrich in species At one site (Batangas Philippines) 31 species have been recorded (23) Of the 176 spider species recorded in Philippine ricefields about one-half (82) occur in upland rice The spiderspecies of upland rice environments overlap more with those of rainfed lowland than with those of irrigated lowland rice (24) The same study showed that of the three environments irrigated rice has (ie greatest spider species diversity foiiowed by rainfed lowland and upland

However of the 10 most prevalent ricefield spiders 9 were abundant in upland rice showing wide adaptation All three spider guilds -- orb-web space-web and huting spiders shywere prevalent in upland ricefields There were however differences in environmental prelerences for some spider species

Of the orb-wcavers Tetragniathamanlihulatawas partishycila rlv abundant 7 japonica was low and Ltucatge dehcorata was absent in upland rice

As in other environments Aiypeiwnaifirtosana was the most dominant space-web spider however its relative numbers were lowerthan for lowland sites Among hunting spiders Ox|opes javanus and Lcosa leatcost fgla were more abundant and Lvcosa lpseudoanmnlata was less abundant in upland compared with lowland rice envishyroniments

Extensive rearing of rice insect pests in upland rainfed lowland and irrigated sites over an 8-yr period in the Philippines provided insight into the effectiveness of parasites as natural enemies (Table 7 8) Most of toe data are from large sample sizes taken over at least one crop which overcomes the pitfalls expressed by Van Driesche (238)

VSB egg parasitization was surprisingly similar across all environments Larval parasitization was highly dynamic among sites and even years within the same site but showed somewhat lowerlevels in uplandsthan in irrigated wetlands This would have been predicted for egg parasites as well since YSB is adapted to the lowlands and so must be its parasites This isevidence that Telenomus rowani is as good at dispersing as its main hosts Trowani clings onto the anal tufts of female YSB and parasitizes eggs as they are laid This phoresy may explain its wide adaptation The more specialized lowland YSB larval parasites therefore appear more effective than their upland counterparts with more alternate hosts

Low collection levels of PSB larvae especially in the wetlands make comparisons between environments difficult Low levels of larval parasifization were recorded in both rainfed environments Except in the Cagayan rainfed wetland site in 1980 low levels ef larval parasitization were evident at irrigated and rainfed sitesalike No differences by environment were evident Higher levels of parasitization occurred in Sarawak during a 1967 outbreak also from tachinids Rothschild (197) indicated that higher parasishytization occurred in upland rice areas

Skipper and green horned caterpillar larval parasitization rates were generally low and showed no environmental effect htighest activity occurred in Cagayan in 1980 (48 parasitization) Although larval parasitization levels of RLF were variable year to year activity appeared greater in the irrigated lowlands than in the uplands Again the highest levels were recorded in Cagayan in 1980 Cagayan has extensive grasslands and fallow around the rice areas and a relatively long rainfall period Rainfall in 1930 was good for crop growth and apparently for increase of larval parasites

IRPS No 123 January 1987 33

Table 7 Parasitization rates of rice planthoppers and green leafhopper in upland rainfed lowland and irrigated lowland environments Philipshypines 1976-840

Environment and site Year Egg parasitization () Nymphaladult parasitization ()

BPH W13P GLH BPII WBPH GLIH

UplandBatangas

Rainfed lowland

1976 1979 1980 1982

15 17 12

2(47)8(39)

4(51)

9(1800)14 (212)

12(304)

8 (101)6 (127)

9(116)

Iloilo

Cagayan

1976 1977 1978 1981 1980 1981 1982

0 (69)

11

44 48 40

-

-

46 55 44

0 (471)

11

52 56 49

0 (200)9 (214)

10 (1259)8(235)6 (900) 3(800)

-7 (462)

11 (270)3 (313)5 (900) 1 (800)

4 (600)10 (725)10 (362)12(870)5 (900) 3 (800)

Pangasinan

Irrigatedlowland

1984 1976 1978 1979 1982

13 (436) 13(379) 14 (457)

6(14)7 (197)

17(198) 11 (486) 18 (275)

3 (81)10 (274)3 (116)

16(871) 19(88)

28(211)9 (764)5 (647)7 (1204) 8(305)

Laguna 1977 11 7 21 1978 31 1979 14 4 4 28 1980 7 8 20 1981 1983

56 7(278)

26 16 (1650)

16 27(1921)

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R I Apostol unpublished 1lo1lo and Pangasinan 136 Cagayan BCanapi unpublished Liuna 175 I C Pantua unpublished 118 234 43 Carino and Shepard unpublished BPH = brown planthopper Nila parvara htgens W131Pi =whitebacked planthopper Sogatelia furcifera GLH = green leafhopper Nephotettrix virescens Figiires in parentheses are sample sizes

Tauher et al (230) reported that internal external and genetic factors influence the seasonal activity of parasitoids

Hopper egg parasites but not nymphaladult parasites were also parlicularly abundant at the Cagayan site possibly the egg parasites have more alternative hopper prey in the weedy areas than nymphaladult paisites

The seasonal dynamics of hopper egg parasites can be seen for upland rice in Batangas(Fig I 2) and for rainfed lowland lice in Pangasinan( 135)and Cagayan( 105) Levels of BPHWBPI-L and GLH egg parasitization were corn-parable in upland and irrigated lowland environments Except for some high rates of paraMsitization early in the season the parasitization rate tended to be steady in all envilonments However hopper nymphaladult parasi-tizution levels were more similar in the rainfed environments and lower than in irrigated rice

Overall parasites do not appear highly effective by themselves against upland or even lowland pests Among natural enemies parasites have attracted the most attention because they can be more readily assesscd

Predators and pathogens lend themselves better to management practices Predators the most important group of natural enemies in riccfields are difficult to quantify On the other hand pathogens appear less im-portant but can be readily cultured and disseminated

The natural enemy commulnity of upland rice is rich in species and differs significantly from that of lowland rice

These beneficial organisms must be conserved by applying insecticides judiciously particularly sprays Seed or soil placement of chemicals minimizes exposure to natural enemies The strategy to derive the greatest benefit from natural enemies isto allow the greatest number of beneficial species to thrive Then perhaps at least one species will be effective against each pest at any tine overcoming the variable abundance of each species within and between years

Programs to introduce exotic species or mass produce indigenous ones are ambitious and expensive because they require trained people on a sustained not ad hoc basis

Chemical control Insecticides offer rapid and efficient control of upland insect pests (190) but-should be used only after other control measures have been considered Insecticides are rarely used on upland rice because of cost (32) Upland rice yields are normally too low to justify the expenditure Also most upland rice is grown as a cash crop Spraying upland rice is more difficult than lowland rice because water is less accessible Government subsidy programs to provide pestishycides to tribal peoples for upland rice production have created problems of toxicity to humans because isolated people have had no experience to be able to handle pesticides safely

However ins-eticide use has beenjustified when a crop is

0Z

Table 8 Parasitization rates of lepidopterous pests in upland rainfed lowland and irrigated lowland environments Philippines 1976-84a

Parasitization (5)

Environment and site Year YSB GFB PSB Earcutting Swarming Skipper Greenborned RLF Brown larva larva caterpillar caterpillar caterpillar caterpillar larva semilooperEgg Larva larva larva larva larva larva

UplandBatangas 1977 - 0 (57) - 0 (63) - 0 (71) - 0 (106) 11(142) 2 (101)1978 43 (4697) 8(451) 5 (38) 3 (40) 6 (61) 2(58) 15 (57) 13 (129) 13 (1286) 4 (30)1979 59(4135) 12(320) 13 (11) 5(28) 9(45) 6(46) 7(23) 8(160) 9(1152) 8(26)

Rainfed lowland Iloilo 1976 - 0 (10) 0(10) 0 (7) 0 (19) 2 13) 15 (180) 5 (76)1978 0 (54) 0 (3) 0 (17) 9 (64) 4 (78) 3 (73) 0 (121) 0 (191)1979 64 (2350) 16 (84) 6 (17) 0(6) 18 (68) 25 (104)

1981 56 (1322) 18(304) 0(5) 12(51) 18(138)Pangasinan 1976 60 (7 7 )b 62(82) 0(111) 5 (39)

1978 58 (53)b 5(51) Cagayan 1979 59(713) 8 (101) 6 (34) 0(7)

1980 0(11) 4 (28) 15 (286)

62 ( 48(177) 61(140)1981 0(5) 12 (2) 13(45) 14 (92)

Irrigatedlowland Laguna 1974 62 8

1974-75 50 41 1975 43 72 1978 96 (623) 21(24) 33 (60)1979 83(2017) 18(110) 8(62) 11 (89) 22(400)1980 78 (1625) 20(75) 0 (16) 4 (72)1983 60

9 (80) 7 (270) 30

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R F Apostol unpublished Iloilo and Pangasnan 136 Cagayan B Canapi unpublished Laguna 175 P CPantua unpublished 118 234 143 Carino and Shepard unpublished YSB = yellow stem borer ScirpophagaincertulasGFB = gold fringed borer Chilo auricilius PSB = pink stem borerSesamia inferens earcutting caterpillar Alythimna separata swarming caterpillar Spcdoptera mauritia acronyctoides skipper Pelopidas mathias reen horned caterpillar Melanitls ledaismene RLF = rice leaffoldersCnaphalocrocismedinalis and Marasmiaspp brown semilooper Mocisfrugalis Figures in parentheses are sample sizes Egg masses

IRPS No 123 January 1987 35

Table 9 Effect of seeding rate and insecticide on plant density and Yield losses from insect pests in upland rice often areyield of Dagge upland rice Batangas Philippines 1976a comparable to those of lowland rainfed or irrigated rice

Seeding rate Insecticide Dosage Plant density Yield (0-Y30o) but usually from different pests The principal (kg aiha) seed treatment (kg aiha) (noIra-row) (tha) groups are soil pests feeding on sown seed and rootsat 14 DE followed by seedling pests These groups have largely gone

50 None - 35 a 20 a unnoticed leading researchers to conclude that stem borers50 Carbofuran F 10 38 ab 23 a and seed bugs are the most important pests (17) Yield loss 10 beCabofuranF100 None - 4551 c 23 a trials that pinpoint major pests need to includc seed

100 Dieldrin WP 10 52 cd 22 a treatments Granules applied in the furrow may not give150 None - 59 d 22 a adequate protection from many seed pests

aAv of 8 fields DE = days after crop emergence In a column means The floristic and pedological diversity of upland rice followed by a common letter are not significantly different at the environments is matched by the diversity of arthropods5level by D)MRT F = flowable Wi = wettable powder both pests and their natural enemies Irrigation has homoshy

genized lowland rice environments Flooded soils tend tothreatened by epidemics of locusts or armyworms New have the same properties and to eliminate soil pests The technology stressing efficiency has minimized insecticide spatial and temporal dominance of cultivated rice has cost benefited specialist pests

Seed treatment is normally inexpensive and can be Upland rice has no specialist arthropod species The economicallyjustified in -nanycases(66) Treating seed with monophagous pests on upland rice prefer the lowlands systemic insecticide (216) but not chlorinated hydrocarbons where rice evolved Rice in the uplands exists only because (247) is most effective in controlling seedling maggots of man and even though lowland rice predates upland riceFoliar sprays have to be repeated to obtain control and specialist species evolved Wheat maize and sorghum do often are washed off by frequent early season rains not have monophagous pests leading us to conclude that Granular insecticides are effective but required dosages are there is no evolutionary advantage for inssects to have too expensive to justify except under heavy infestation narrow host ranges in upland graminaceae(247) When directed against first-instar larvae banding Pests have adapted to the uplands not only by having granules in seed furrows (177) can be effective against white wider plant host ranges but also by having long life cyclesgrubs at low dosages (025 kg aii ha) (134) Seed treatments undergoing dormancy or being dispersive Each pest has a are not promising for white grub control (248) unique life history strategy suited to highly unpredictable

Seed treatment is also effective ainst ants and may be environments In agivenyear someinsect specics an better cheaper than increasing the seeding rate In Batangas exploit upland rice as a food source but no pest is a senious treating 50 kg seed ha was as effective as increasing the one every year on upland rice seeding rate to 100 kgha and using untreated seeds (Table The low yielding potential of upland rice means less 9) In Brazil seed treatments control termites and other soil quantity and lower quality of food for pests to exploit but insects including the lessercornstalk borer(66) Domiciano also less incentive for farmers to undertake control (58) however found seed treatments inconsistent in their The most effective control would be early and synchroshyeffect against high populations of lesser corn stalk borer nous planting of early-maturing upland rice varieties to

Baiting can be an inexpensive way of controlling ants prevent insect pest buildup Breeding for insect resistance seedling maggot flies mole crickets and field crickets Baits should initially focus on stem borers because they are verymade of locally available material and impregnated with difficult to control by other means Biocontrol efforts insecticide can be sparingly distributed taking adantage of should first stress conservation of natural enemies preshythe pests ability to disperse and encounter bait sites dators and pathogens rather than parasites Efficient

The new low-volume hand-held controlled-droplet chemical control includes seed treatment or baiting for seed sprayers may offer an advantage over high-volume knap- seedling and some root pests and using controlled dropletsack sprayes for controlling foliar pests Upland rice sprayers for spot treatments against foliar pests and seed canopies are more open than those in lowland rice and bugs when economic thresholds are reached better droplet penetration should result Controlled droplet applicators would be ideal for seedbugs and defoliators

CONCLUSION

Upland rice is attacked by a wider array of insect pests than is lowland rice mainly beca use of the addition of soil pests Generally population levels of lowland rice pests are lower in upland than lowland rice leading many people to conclude that insect pests are not important on upland rice

36 IRPS No 123 January 1987

REFERENCES CITED

1Abraham C C and K S R Mony 1978 Occurrence ofLeptocorisa acuta Fabr (Coreidae Hemiptera) as a pest of nutmeg trees J Bombay Nat ttistSoc 74553

2 Agyen-Sampong M 1975 HlierogrlphusdiaganensisKrauss (Orthop-tera Acrididae) a new pest of rice in Northern Ghana Ghana J Agric Sci 8249-253

3 Agyen-Samnpong M 1976 Varietal resistance and other inle-ratcd pest control components as factors of rice varietal improvement in West Africa Prcsented at the Second varietal improvement serinar 14-18 Sep 1976 Bouak Ivory Coast West Africa Rice Development Association Monrovia Liberia 18 p

4 Agyen-Sampong M 1977 Outbreak of Ihccrorrdnisorae in rice seedlings near Rokupr Sierra Leone lit Rice Res Newsl 2(6)16-17

5 Agyen-Sampong M 1980 Dirty panicles and rice yield reduction caused by hugs IntRice Res Newsl 5(l)11-12

6 Agyen-Sampong M 1982 The major pest problems of irrigated upland and mangrove swanip rice ecosystems ir the humid tropical Guinea savanna and Salel climate zones Pages 318-328 in Pro-ceedings training course cn concepts techniques and application of irtegrated pest management in rice inWest Africa 10-20 Jan 1982 Fendall Liberia West Africa Rice l)evelopioent Association Monrovia Liberia 505 p

7 Akibo-Betts V I and S A Raymundo 1978 Aphids as rice pests in Sierra Leone lit Rice Res Newsl 3(6)15-16

8 Akinsola F A 1975 Present status of different rice stemborers in parts of Nigeria iicL Entomo Newsl 328

9 Akinsola FA 1979 Ihe biology and ecology of rice stemborers in Nigeria Ph 1) thesis University ol lbadan Nigeria 217 p

10 Akinsola E A 1979 Guidelines of integrated management of insect pests of upland rice in West Africa Presented at the Seminar on integrated management of rice diseases and insect pests at Bobo-Dioulasso Upper Volta 17-22 Sep 1979 West Africa Rice Develop-ment Association Monrovia Liberia

II Akinsola IA 1984 Insect pests of upland rice in Africa Pages 301-305 in An overview of upland rice research Proceedings of the 1982 lIouak Iwsrv Coast upland rice workshop International Rice Research Institute 13O Box 933 Manila Philippines

12 Alam M S V TJohn and Kaung Zan 1985 Insect pests and diseases of rice in Africa Pages 67-82 in Rice improvement in Eastern Central and Southern Africa proceedings of the international rice workshop at Iusaka Zambia 9-19 April 1984

13 Alticri M A A van Schoonhovcn and J D Doll 1977 The ecological role of weeds in insect pest management systems a review illustiated with bean (Phaseohsvulgaris L)cropping systems PANS (Pest Artie News Summ) 23195-205

14 Anmante F 1967 The leaf-cutting ant Atia capiguaraa pest of pastures [in Portuguese] Biol6gico (Sao Paulo) 33113-120 (Rev Appl Entomol A56207)

15Andrade A C and D Puzzi 1951 Preliminary results of experi-merits on the control of Sculptcoris iastantus on sugarcane [in Portuguese] Bioulgico (Sao Paulo) 1744-49 (Rev Appl Entomol A41421422)

16 Appert J 1970 Maliarplia separaelha (rice stem borer) new observations of entomological problems of rice in Madagascar [in French] Agron Trop (Paris) 25329-334

17 Ariadeau Mand Z Harahap 1986 Relevant upland rice breeding objectives Pages 189-197 in Progress in upland rice research Proceedings of the 1985 Jakarta conference International Rice Research Institute 1O Box 933 Manila Philippines

18Asar S and Kisimoto 1975 Seasonal occurrence of the smaller brown planthopper in upland rice [in Japanese] Proc Kanto-Tosan Plant Prot Soc 2228

19 Bakker W 1974 Characterization and ecological aspects of rice yellow mottle virus in Kenya Meded Landbouwhogesh Wage-ningen 152 p

20 Balut F F 1970 White grubs attacking rice Oryza ativa L [in Portuguese] Biol6gico (So Paulo) 36321-322

21 Banerjee S N and L M Pramanik 1967 The lepidopterous stalk borers of rice and their life cycles in the tropics pages 103-124 in The major insect pests ofthe rice plant Proceedings ofa symposium at the International Rice Research Institute September 1964 The Johns Hopkins Press Baltimore Maryland USA

22 Barrion A T and J A Litsinger 1980 Ants a natural enemy of cliaiarac in dryland rice IntRice Res Newsl

5(4)22-23 23 Barrion A T and J A litsinger 1981 The spider fauna of

Philippine rice agroecosystem 1Dryland Philipp Entomnol 5139-166

24 Barrion A T and 1 A Iitsinger 1984 The spider fauna of Philippine rice agroccosystcms II Welland Philipp Eiitomol 611-37

25 Barrion A T1 and J A Litsinger 1984 lihanus (Diptera Tabanidae) eggs an alternative host of rice stei borer (SI3) egg parasite Telenoonis digns (Iliymenoptcra Scelionidae) lit Rice Res Newvsl 9(6)19

26 Barrion A T and J A Litsinger 1985 Identilication of rice leaffolders by wing markings Int Rice Res Newsl 10( 1)24

27 Barrion A T and J A Litsinger 1985 Chctiosspp (Coleoptera Carabidae) a leaffolder ILF)predator lit Rice Res Newsl 10(1)21

28 Bertels A 1970 Insect pests of rice and theircontrol [in Portuguese] Pelotas Brazil Instituto de Expcrimentacao Agropccuararias do Sul Circular 43 24 p

29 Bonzi S M 1982 Ctilo (flisilineus J de Joannis (Lepidoptera Pyralidae) stem borer of irrigated cereals in Upper Volta LioFrench] Agron Trop Riu Rizic Cult Vivrieres Trop 37207-209

30 Bourke T V T L Fenner J N LStihick G Baker EHassan D F OSullivan and C S li 1977 Insect pest survey for the year ending 30th June 1969 Papta New Guinea )epartment of Agrishyculture Stock and Fisheries Port Moresby 57 p

31 Bowden 1 1973 Migration of pests in the tropics Meded Rijksfac Landbouwwet Gent 38785-796

32 Breniere J 1973 The main rice parasites in West Africa and their control Pages 27-42 in Proceedings seminar on plant protection for the rice crop West Africa Rice Development Association Monrovia Liberia

33 Breniere J 1976 The principal insect pests of rice in West Africa and their control West Africa Rice Development Association Monrovia Liberia 52 p

34 Brown ES 1962 The African arinyworn Spoiolptera eXI)Ina (Walker) (Lepidoptera Noctuidae) a review of the literature Cornshymonwealth Institute of Entomology London UK57 p

35 Brown F B 1968 Upland rice in Latin America FAO-International Rice Commission 12th Session of the Working Party on Rice Production and Protection 2-14 Sept Peradeniya Sri lanka Food and Agriculture Organization Rome Italy 6 p

36 Buyckx F J 1 1962 Disease and insect pests of crops inCongo Rwanda and Burundi [iil French] Publications dc LInstilut National pour lestude Agrononiquedu Congo M Weissenbruck S A Brussels Belgium 708 p

37 Cariiio F 0 1981 Role of natural enentics inpopulation suppression and pest management of green rice lealhoppers Ph D thesis University ofthe Philippines at Los Bafitos PhilippinesLaguna 136 -

38 Carpente A J 1983 The development of rice cultivation and extension of vegetable productin in Zanzibar Final Report Vol II1 October 1983 Food and Agricultme Organization Ronte Italy

39 Carroll C P and S1 Risch 1983 Tropical annual cropping systems ant ecology Environ Manage 751-57

4(0 Cates R G 198 1 Host plant predictability and the feeding patterns of monophagous oligophagous and polyphagous insect herbivores Oecologia (Berl) 48319-326

IRPS No 123 January 1987 37

41 Cendcafia S M and F B Caora 1967 Insect pests of rice in the 62 Ene 1 C 1981 I)iiorldhol)ierusihilis Slater (licteroplera Philippines Pages 591-616 in The najor insect pests ofthe rice plant lygacidue Illissinac) a new rice pest front Nigeria lit Rice Res Proceedings of a symposium at the International Rice Research Newsl N3)15 Institute September 1964 The Johns Hopkins Press Baltimore 63 Fugenio T S 1965 rhe effect oi e-vCgetation on rice production Maryland USA 729 p and the possibility of overcoming it by leaf pruning MS thesis

42 Centro Ihternacional de Agricklittira Tropical 1984 Upland rice in University ol the Philippines at Los Baijos ILaguna Ihilippins 69 p Iatin Anerica Pages 93-119 hi An os ervies of uplaind rice research 04 Farrow R A 1975 The Alrican nmigratory locust ii is main Proceedings of the 1982 Boinakamp Ivory Coast upland rice workshop outbreak area of the middle Niger qlinlitatise studies of solitary International Rice Research Institute P0 Blox 933 Manila populations in relation to environmental factors Locusla No II Philippines 198 p

43 Chandra G 1979 Taxonomy ad hn riononlics ofthe imect parasites of 65 Fennal R G 1943 Food crop investigation Windward and rice Icalloppers and planthoppers in the Pihilippines and their Ieeward Islands First report Report onl work dunring the periodiiiportairce iii natural biol)gical cointrol Philipp Fntoltol 4119-139 April 1942-April 1943 Imperial College of Tropical Agriculture

44 Chang T T 1976 The originii colut ion enlIivation dissemination Trinidad 16 p (Rev of Appl Fintornol A3222-25) and diersificition of Asian and African rices -uphytica 25425-44 1 66 Ferreira F 1983 Integrated pest ritaageiret of upland rice [ill

45 Chatierjee P B 1979 Rice leaffolder attacks in India lit Rice Res Portuguese English summary] Pages 323-341 in Upland riceculttre Newsl 4(3)21 factors affecting production E Ferreira T Yamada and F-

46 Cheng Cbien-pan 19(4 Upland rice in Taiwan International Rice Malmolta eds Institutoda Potassae Foslato Institnto Internacional Cnmmission Wiorking Party oit Rice Production ard Protection I()th da Potassa Piracicaha Sao Paulo lraid Meeting 3-10 March 1964 Manila Philippines Food and Agricul- 67 Ferreira E arid F dl Silva Martins 1979 Occurrence of ture Organiaion Rome Italy 5 p Sogatohs orvziola (Muir) inupland rice in Goias Ilrail lit Rice

47 Chcrrett 1 M and 1) 1Peregrine 1976 A review of the status of Rcs Ncwssl 4(4)18 leaf-cutting ants and their control Ann Appl Biol 84124-128 68 Ferreira E and 1F da Silva Martiris 1984 Insect pestof rice ill

48 Chipeta F M 1974 Pests of rice in Malawi Pages 90-95 in Brazil and their control Jill Portugucsej FMIlRAtPA-CNPAF Proceedings Sith eastern Africa cereals research conference 10-15 Goiania Brmiil 67 p March 1974 DRB Mialda ed Ministry of Agriculture lilongwe 69 Ferreira FI F ca Silha Miartins S S Nero and 1 It P de Malawi Carvallio 1982 M tltiplc regression analysis todeterimine the effect of

49 Chu Yau-i 1979 The d atiaging period of the rice seedling fly insect populatiois ott upland rice yield [inIPortuguese Pesqui (Atlterigotiavxigu Stcin) (NItiscide Diptera) oil cori) at Fast Java Agropecu Bras 17671-675 Indonesia Plant Prot Bill 21403413 70 Ferreira F 1F da Silva Martins and F 1PZimmermann 1979

50 Conklin H C 1957 IHaninoo agriculture a report on aln integral Rice varietal resistance to tle lesser corn stalk borer [in Port uguese] systent of shi fti rig cult ivationi in the Pltilippincs Food and Agriculture Pesqui Agropccu Biras 14317-321 Organization Rome Italy 209 p 71 Fleming W F 1968 Biological control of lie hapanese beetle U S

51 Cramer H II 1967 Plant proection and world crop production Dep Agric RLs Sers [ech Bull 1383 78 p Pilaniensclhutz-Nachr 201-524 72 Food and Agriculture Organiation of the United Nations 1982

52 l)ani R C and Majunidar 1978 Prelininary observations on tie Country report surmimary Pages 27-37 in Report on the first incidence of root aphids on differeit rice cultnrcs Sci Cult 4488-89 FAOIUN)IPhail and regional training coiuirse in iniproved ctltural

53 Davidson R I 1979 lffects of root feeding on foliage yield Pages practices for upland rice 5-23 Oct 1981 Northern Region Agrculturd117-120 in T K Ci osb and R B Pottinger eds Proceedings of the Developririt Center Chiang Mai Food and Agricuilture Organiiashy2nd Australian conlerence on grassland invertebrate ecology Govt tion Rome Italy 37 n Printer Wellington New Zealand 73 Franco do Ariaral S 195 1 Residual action of insecticides in turfy

54 l)ean G 1W 1976 Rice insect pests inlaos Int Rice Res Newsl soil--the results of a field experiment [in Portuguese] Biol6gico (Sao 1(2)15 Paulo) 17209-214 (Rev Appl Entomol A41423)

55 I)ean G J W 1978 Insect pests of rice in Laos PANS 24(3)280-289 74 Freeman ) 1970 Report onithe Iban london School of -coroiics 56 )ingle H 1984 Behavior genes and lifc histories conplex adapta- Monographs on Social Anthropology No 41 Athlane Press

tions to uncertain environments Pages 169-194 in A new ecology London UK 317 p novel approaches to interactive systems P W Price C N Slobod- 75 Fuchs T W1 A Harding and 1W Smithlr 1979 Induction and chikoff and W S Gaud cdsl ohn Wiley New York USA termination oldiapause in the sugarcane borer Ann Flitonlol Soc

57 Dobelmann-JP and M Falais 1968 Upland rice cultivation ill tie Am 72271-274 North-western part of Madagascar in 1966-1967 [in French English 76 (hbriel B1) 1975 A review tf the major insect pests of some tpland summary] Trop Agron 2313-21 crops in tlc Philippines Pages 86-111 in Ad Htoc panel of experts on

58 Doniiciano N L 1979 Chemical control of tie lesser corn stalk pest disease arnd weed problems in some rainfed crops Bangkok (lolsonno~paloIgnowslhts) (Zeller 1848) (I-epid(ptera Ihycitid ae) Thailand Food aid Agriculture Organization Roire Italy

on rice in Ponta Gressa Parara 1977-1978 [in Portuguese] Pages 77 Garg D K and N K Shal 1983 White grub outbreak on rainfed 291-303 in Anais 1 Reuniao de tecnicos cmiirizicultura do estado de dryland rice in Uttar Pradeslh Int Rice Res Ncwsl 81(4)16-17 Sao Paulo-Campinas Cati Brazil 78 Garg I) K and IP Tandon 1982 Major insect pests of rice on hilly

59 Dunsmore 1 R 1970 Investigations on tlie varieties pests and tracts of Uttar Pradesh India lilt Rice Res Newsl 7(11)11-12diseases of upland rice (hill padi) in Sarawak Malaysia Int Rice 79 Garrity 1) P 1983 Upland rice research an international bibliograpy Comm Newsl 1929-35 1975-1982 Iternational Rice Research hnstitnte PO Box 933

60 Elias 1R1965 The new decline of rice seedlings is not caused by Manila Philippines 121 p parasites [in Portuguese] Biol6gico (Sao Paulo) 313-6 80 Ghauri M S K 1982 A new subspecies of Iinorh)rone bull

61 Empresa Brasiliera de Pesquisa Agropecuaria 1984 Upland rice itt cornlus Slater (Hemiptera Iygaeidae) front Papua New Guinea on Brazil Pages 121-134 it Av overview of upland rice research rice and carpet grass Ihll lEntoniol Res 72133-137 Proceedings ofthe 1982 Blouak Ivory Coast Upland rice workshop 81 Gonzales F 0 PArregoces R I ihernandef and OT Parada International Rice Research Institute PO Box 933 Manila 1983 Insect and imite pests of rice arid their control in Latin America Philippines [in Spanish] la Federacion Nacional de Arrocerts (Fedearroz)

Bogota Colombia 60 n

38 IRPS No 123 January 1987

82 Grist D H and R J A W Lever 1969 Pests of rice Longmans London UK520 p

83 Hale P R 1979 Two subterranean pests of upland rice in Papua New Guirnca IntRice Res Newsl 4(2)16-17

84 Hale P R and J T Hale 1975 Insect pests of rice in the New Guinea Islands Rice Entomol Newsl 34

85 Harris W V 1969 Termites as pests of crops and trees Common-wealth Institute of Entomology London UK 41 p

86 Hattori 1 1968 A method to distinguish between grass leaf roller (Ctaplha rocis medinalisGuenee) and the rice leaf roller (Susurtia exigua Butler) [in Japanese] Plant Prot (Japan) 23167-170

87 Henry I E and 1A Onsager 1984 Experimental control of the mormon cricket Iahrosshnph by NJsena locustae(Aficrospora Microsporidae) a protozoan parasite of grasshoppers (Ort Acri-didae) Entomophaga 27197-201

88 Hidaka 1 P Vungsilabulr and SKadkao 1974 Studies on ecology and control of the rice gall midge in Thailand Tech Bull 6 TropicalAgricultural Research Center Tokyo Japan 113 p

89 Hincklcy A 1) 1963 Ecology and control of rice planthoppers in Fiji Bull Entomrol Res 53467 481

90 Hirao J 1982 Ecology and chtInical control of the rice leaffolder Jpn Pestic Inf 4114-17

91 Ho DT 1984 Sten borer incifience in rice ecosystems in KenyaEast Africa Pages 307-320 in A overview of upland rice research Proceedings of the 1982 Bouac Ivory Coast upland rice workshop International Rice Research Institute PO Box 933 Manila Philippines

92 Ho )T J 1Njokah and J GKibuka 1983 Studies on rice sten borers in Kenya with emphasis on ifaliarpiaipiraellRag Insect Sci Appl 465-73

93 Ichihara I T Ito A NIatsuo and M Sawada 1971 Chemical control of the rice striped stem borer in upland rice grown in plastic mulch [iii Japanese] Proc Kanto-Tosan Plant Prot Soc 1882

94 IclhiharaIand A Matsuo 1969 Damage caused by rice striped stett borer to upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1686-87

95 Ichhara I atid A MIatsun11970 Conside ration oft lie extraordinary prevalence of rice striped stem borer in upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1778-79

96 Institut de Recherches Agrolnliqes Iropicale et des Cultures Vivrieres 1978 Insect resistance Pages 12-13 in Upland rice 1978 atnual report Bouak Ivory Coast 27 p

97 International Institute for Tropical Agriculture 1983 Aphids itt rice roots I ITA Res Briefs 4(1-3)4-5

98 International Institite fi Tropical Agriculture 1983 Hopperburn in rice caused by brown plathopper IITA Res Briefs 4(l-3)6-7

99 International Rice Research Institute 1975 Annual report for 1974 PO Box 933 Manila Philippines p 216

100 International Rice Restarch Institute 1976 Annual repiort for 1975 PO Box 933 Manila Philippines p 228

101 International Rice Research Institute 1979 Cropping systems program Component technology development and evaluation Page 404 i Annual report for 1978 11O Box 933 Manila Philippines

102 International Rice Research Institute Department of Entomology 1981) Insecticid evaluation 1979 Tables 73 74 PO Box 933Manila Philipes

103 International Rice Research Institute Department of Entonmology 1981 Insecticide evalomtion 1980 Table 81 POBox 933 Manila Philippines

104 International Rice Research Institute 1981 Report of a meeting of Asian rice-based cropping systems entomologists Eleventh cropping systems working group meeting 18-22 May 1981 PO Box 933 Manila Philippines p 132 133 144

105 International Rice Research Institute 1984 Cropping systems program Pages 368-373 in Annual report for 1983 PO Box 933 Manila Philippines

106 International Rice Research Institutc 1984 Upland rice in Asia Pages 45-68 in An overview of upland rice research Proceedings of the 1982 Bouakamp Ivory Coast upland rice workiiop loternational Rice Research Institute 11O Box 933 Manila Philippines

107 Irshad M 1977 Grasshoppers in rice in Pakistan lit Rice Res Newsl 2(3)8-9

108 Ito R and TOno 1964 Upland lice its culture and varieties in Japan lIternational Rice Cormmission loth Meeting of the Working Party on Rice Production and Protection 3-10 March 1964 Manila Philippines Food and Agriculture Organization Rome Italy 12 p

109 Jeffrey M I and V A Dyck 1983 Effect of moonlight on lightshytrap catches of brown planthopper lit Rice Res Newsl 8(6) 13

110 Jennings P R and T Alicia Pineda 1983 Management ofthe rice plnthoppers Sogatodsorivi(coIl in Latin America Presented at the meeting on stable plant resistance FAO and lIndocsian I)inectorate of Food Crops Protection Bali lndonesia 29 Novcniber-2 l)ccemshyber 9 p

IllJimenez Gomez J A 1979 Rice insect pests i lla ios ()riCntales lil Spanish] Pages 99-103 in Rice ciltivation Instituto (olotibiatto Agropecuario Compndio No 29

112 Jongeleen F JJ Soesanto and A Mockidjo 1979 Studies oti pathogens of two species of Scarabacidac froinIndonesia Z Angew Enlomnol 87412-417

113 Jouvenaz 1) PC S lofgren and W A Baiks 1981 Biological control of imported fire antsa review of cirrent knowledge Bull Entomol Soc Am 27203-208

114 Kainoh Y K Shiniiu S MarUani Y Taiaki 1980 lost finding behavior of tie rice bu lplotori+JchinvittjI)allas (liciniptera Corcidac) with special reference to diet pattern ol aggregation and feeding on the rice plant Appl Entomol Zool 15225-233

115 Kakkar K 0 P Sharma and P I Sharma 1977 Note on Ihteronyrchus liodresRedt (Coleopicra I)ynastidae) as asporadic pest of paddy and maie inHimachal Pradesh Corr Agric 156-57

116 Kalode M131 79 Major insect pest problems and their control in upland rice (Unpublishcd mineogriph CWI oJNaaiat lni Chandaratat (198 1)original not seen)

117 Kalshoven L i 98 I The pests of crops in I ldoniesia (translated by P A vatn der Laan) 1 T Ichtiar B3aru-vati Iloeve lakarta Indonesia 701 p

118 Kamal N Q 1981 Suppression of whitebacked planto ner Sogatella fircaifra (Horvath) aLId rice lealfolder (nhal -cis licdb lis (Guenee) populations by natural cnclmie hlI) tiesis Grcgorio Araneta University Foundation Victoncta Park Metto Manila Philippines 179 Ip

119 Karim A N M Reaul 1975 A species ofliealy bug Trioni i70s ceres new oii rice in the Philippines Rice Entoniol Newsl 247

120 Katanyukul Wand C Chandaratat 1981 Insect pestsof uplaid rice and their control Report firstFAOUNlFi Tliailand training course on improved cultural practices for tplad rice Chiang Mai Thailand 5-23 Oct 1981 Food and Agriculture Organiation Rome Italy 22 p

121 Kern F 1955 lhletwioa(I)uninemts) bhhit Biturm(Coleoptera) a new pest of sugarcane inVenezuela [inSpanish with Etnglish sunmary] Agron Trop (Maracay) 527-29 (Rev Appl Eitoniol A4572)

122 Khan K M and S Ghai 1974 White grubs ald their control in India Pesticides (1Bombay) 819-25

123 Kiritani K 1975 Integrated pest management of rice pests Pages 66-70 itt Proceedings platning workshop ott cooperative hIeld research in pest managementP SMotooka ed East-West Center Honololu Hawaii USA

124 Kisimoto R 1967 Ecology of insect vectors forecasting and chemical control Pages 243-255 in The virus diseases of tle rice plant Proceedings of a symposium atthe International Rice Research Institute April 1967 The Johns Hopkins Press altimore Maryshyland USA

125 Kisimoto R and V A Dyck 1976 Climate and rice insects Pages 367-391 in Climate and rice International Rice Research Institute PO Box 933 Manila Philippines

126 Kureha S S Kobayashi SAsari and T Ishihara 1977 Life history and control of the mole cricket Grrllotalpa africana Palisot de Beauvois [in Japanese English translation 7520 1RRI library] Proc Kanto-Tosan Plant Prot Soc 21192-197

127 Kushwaha K S 1981 The black beetle as a pest of rice in Haryana India Int Rice Res Newsl 6(4)14

128 Lawani S M 1982 A review of tile effects of various agronomic practices on cereal stem borers Trop Pest Manage 28266-276

129 Leuck D B 1966 Biology of the lesser cornstalk borer in South Georgia J Econ Entomol 59 797-801

130 lever R J A W 1955 Insect pests of the rice plant in Malaya Int Rice Comm Newsl 1612-21

131 Li Ching-Sing and Shin-Foon Chiu 1951 The study ofthe rice gall midge Iachydiplosis orvza Wood Mason [in Chinese with English summary] 1Taiwan Agric Res 21-13

132 Iitsinger J A 1984 Assesament of the need-based application for rice Proceedings TechnologyTransfer Workshop 15-16 March 1984 International Rice Research Institute 1PO Box 933 Manila Philippines 6 p

133 Litsinger J A 1984 Environmental diversity and insect pest abundance with reference to the Pacific Pages 285-307 in Proceedings sub-regional training course on methods of controlling diseases insects and other pests of plants in the South Pacific 4-20 Oct 1982 Vaini Kingdom of Tonga Nukualofa

134 Litsinger J A RF Apostol and M BObusan 1983 White grub Leucopholis irrorata (Coleoptera Scarabaeidae) pest status popula-tion dynamics and chemical conllol in arice-maize cropping pattern in the Philippines J Econ Entoinol 761133-1138

135 Litsinger J A C de la Cruz F Raymundo A T Barrion M D Lumaban J P Bandong M S Venugopal F Paragna and E Balete 1982 Insect pests and insecticide reponse of dry-seeded bunded rice Pages 179-193 in Report of workshop on cropping systems research in Asia International Rice Research Institute Los Bafos Philippine

136 Litsinger J A M 1) Luinaban J P Bandong A T Barrion W Katanyukul and N Panda 1978 Insect pest management in rainfcd lowland rice agroecosystms of tropical Asia International Rice Research Conference 17-21 April 1978 International Rice Research Institute Los Bafios Philippines

137 Litsinger J A and K Moody 1976 Integrated pest management in multiple cropping systems Pages 293-317 in R I Papendick P A Sanchez and G B Triplett eds Multiple cropping American Society of Agronomy Madison Wisconsin USA

138 Loevinsohn M E 1984 The ecology and control of rice pests in relation to the intensity and synchrony of cultivation Ph D thesis University of London 36) p

139 Loevinsohn M E J A Litsinger N Panda M A Austria R F Apostol C Andrion and J Bandong 1982 A basis for predicting rice insect pest abundance in cropping systems Paper presented at the 1982 International rice research conference on rice and cropping systems 19-23 April 1982 International Rice Research Institute Los Bafios Iaguna Philippines 25 p

140 Lopez A W 1930 Repot of the Entomology Department Pages 145-172 in Annual report of the Philippine Sugar Association 1929-30 Manila

141 Lopez AW 1931 Entomology Department Pages 227-266 in Annual report of the director or research 1930-31 Philippine Sugar Association Manila

142 Lopez A W 1932 Entomology Deparment Pages 252-279 in Annual report of the Philippine Sugar Association 1931-32 Manila

143 Lungibill P and G G Ainslee 1917 The lessercornstalk borer US Dep Agric Agric Res Serv Tech Bull 539 27 p

IRPS No 123 January 1987 39

144 McGuire J U Jr and BS Crandall 1967 Survey of insect pests and plant diseases ofselected food crops of Mexico Central America and Panama U SDepartment of Agriculture International Agriculshytural Development Service Agricultuial Researrh Service October 1967 11 p

145 McNeill Sand T R ESouthwood 1978 The role of nitrogen in the development of insectplant relations Pages 77-9 in lliochemical aspects of plant and animal coevolution J B Ilarbornu ed Academic Press London UK

146 Maebara H 1955 Spatial distribution of lastinstiarlanaeof purplish stem borer (Sesamia iqferens Walker) hibernating in an upland rice field Bull Kagoshima Univ Fac Agric 438-41

147 Mallick E H and N G Hajra 1977 Response of rice plant to defoliation treatments at different developmental stages Indian Agric 2151-56

148 Mansingh A 1971 Physiological classification of dormancies in insects Canl Entomol 103983-10)9

149 Martell C G 1974 Catalogue of plant feeding insects of Mexico [in Spanish] Fitofilo 27(69)1-176

150 Martinez 0 J 1982 Principal rice pests in Ilanos orientalis [in Spanish] P~ages 80-94 in Curso deact nalization de conocimientos en arroz Villaniencis Instituto (olonmbiano Agropecuario

151 Martins J F da Silva F Ferrcira A S Prabhu F 1 P Zimmermann 1980 Use of pesticides for the control of the main subterranean upland rice pests esqui Agropecu Bras 1553-u

152 Martins I F da Silva F Fe eira rod BdamSilveira Pinheiro I Q2 Damage simulation caused by defoliators of upland rice [in Portuguese with English summary] Pesqui Agropecu Bras 171113-1119

153 Martins 1F da Silva N(iyuen van Tan and 1Ida Silveira Pinheiro 1981 Stem borer resistance in upland rice associated with plant morphological characters [in Portuuce Iesqui Agropecu Bras 16187-192

154 Masaki S 1980 Sumnmer diapause Ann Rev Entomiol 251-25 155 Masuzawa T 11 Suwa and F Nakasoji 1983 l)ifkleences of

oviposition preferencc and survival rate of two skipper butterflies Parnara guttata and PIchlpidai iathiais (Iepidoptera HIesperiidac) on rice plant and cogon grass New Entomol 3247-56

156 Maurya D M and C P Vaish 1984 Upland rice in India Pages 69-92 in An overview oftpland rice research Proceedings ofthe 1982 Bouakamp Ivory Coast upland rice workshop International Rice Research Institute IO Box 933 Mnila Philippines

157 Menon M G R R N Katiyai S Kuar and S S Misra 1970 Termites attacking the paddy crop Sci Cult 36294-295

158 Miyahara Y and 1 Nishikubo 1967 Behavior of young nymphs of the red rice root aphid Iholahsipiun ri liabhdontinalisSasaki in relation to movement from the leaf to the roots of upland rice [in Japanese] Pioc Assoc Plant Prot Kyushu 1363-66

159 Mo ljoa Tjien 1952 White grub of upland rice (Holotrichiahelleri Brsk) [in Indonesian with English summary] Contrib Agric Res Stn Bogor No 13134 p

160 Mochida 0 and T Suryana 1979 The occurrence of the brown planthopper and ragged stunt on lowland and upland rice ill Central Java and logyakarta in tile late of January 1979 Entomnolgy Survey Report No 15 Central Research Institute of Agriculture (CRIA) Sukamandi Indonesia I February 1979 9 p

161Monet C 1979 Varietal resistance to upland rice pests [inI French] Pages 35-39 in Annual report 1977 Institut des Savanes Department des Cultures Vivrieres Bouak6 Ivory Coast

162 Monte 0 1942 A caterpillar ofrice-fields[in Portuguese] Biol6gico (Sao Paulo) 8161-163 (Rev Appl Entomol A31414)

163 Munroe I) D 1979 Pest management strategy for hill rice in four resettlement areas in Sarawak Malaysia Pages 89-102 i1 Guidelines for integrated control of rice insect pests Food and Agriculture Organization Plant Production and Protection Paper 14 Rome Italy 115 p

40 IRPS No 123 January 1987

164 Nakasuji F M Ishii and Jen-Zon Ito 1984 Rice skippers in Taiwan and their life histories ZZrC (Taipei) Tech Bull 82 10 p

165 Navas I) 1976 Fall armyworum in rice Pages 99-106 il Ecological animal control by habitat management Proc Tall Timbers Con-ference 28 Feb-I Mar 1974 Fall Timbers Res Stn Misc Publ 6 Tallahassee Horida USA

166 Nickel 1 1 1961 Biological control of rice stem borer A feasibility study Tcch Bull 2 International Rice Rcarch Institute PO Box 933 Manila Philippines IIl p

167 Nigan P M and R A Vernia 1985 Insect pests of uplaild rice in Uttar Pradesh Int Rice Res Newsl 10(6)22

168 Nilakhe S S 1985 -cological observations on spittle bugs with emphasis on their occirence in rice Pcsqtui Agropecu Bras 20407-414

169 Nixon G EJ 195 1 lie association of ats with aphids and coccids Commonwealth Iusti lcte of Entonology Iondon UK 36 p

170 Nurdin F an Ishak Manti 1985 Study oit seedling fly of upland rice Interlational Upland Rice Moitoring Tour Report 26 lebruary-2 MarLI Sukaranti West Sumatra

171 Oka 1 N 1979 Ciltural conttol of the brown planthopper Pages 357-369 in Brown planthopper threat to rice production in Asia International Rice Research Institutc 1p0 Box 933 Manila Philippines

172 Ohni M and A Vilnaii 1975 Rice insect pests of Italy biology of Sipha grcria (Kaltenbach) and its control [in Italian] Riso (Milan) 2459-71

173 Otanes F Q atnd 1 I Sison 194 1 Pests of rice Philipp J Agric 12211-259

174 Painter R I 1951 Insect icsistance in crop plants University Press of Kansits Lawrence Kansas USA 520 p

175 Pantua P C and 1A litsinger 1981 Arthropod abundance in continuous vs biannual rice cropping systeis Int Rice Res Ncwsl 6(2)20-21

176 Pathak M ) and V A l)yck 1975 Sttudics ott insect pests of upland rice Pages 186-191 in Major research in upland rice International Rice Research Institute PO Box 933 Ilanila Philippines

177 Patil B R B M IHasabe and 1) S Ajri 198 I Chemical control of white gritb lIditriihinhorraia Fabr intfesting groundntut paddy atnd sorghum Indian 1Entomol 43232-236

178 Pawar A 1) 1974 (ollection and identification of insect pests of rice rlerminal repirt lInternational Rice Research Institute PO Box 933 Manila Philippines 72 p (tnimco)

179 IerrajU A aid A S Reedy 1965 Observations ott diapausc itt tie steti borer ohrice Rice News Teller (lai-Mar)9-13

180 Pont A C 1972 A review of the Oriental species of theri ona Rondani (I)iptera Muscidac) of econtomic importance Pages 27-102 in Control of sorglium shoot fly M G lomtani and W R Young eds Oxford atd IH1 Ptthl Co New I)clhi India 324 p

181 Prasad S C 1PChand aid M 1PSingh 1983 Incidence of rice gall midge in the upland plateau region of Hihar Oryza 20235-236

182 Rai P S 198 0 Nisaga simpex damage to rice in the hill tracts of South India Int Rice kles News[ 5(I)12

183 Rathorc Y S 1969 Gonocplhahli dcprssunt Fabr as itpredator of (tilo zonwIlhis (Swinhoc) and other moths in field cage at Patnagar Uttar Pradesh Indian 1Entotnol 3127

184 Reed W 1983 Crop losses causcd by insect pists in the developing world Proceedings 10th Internatial Congress of Plant Protection lrighton UK 174-80

185 Reedy M V 1981 Ateactogasterinhens(Walker)a new root-grub pest of tipland paddy and inaize Curr Sci 501076

186 Reveno M A 1968 Catalogue of the insect pests of economic crops in Colombia [in Spaiish] Asociacion Latino Americana de Entomo-logia (AIAE) 129-33

187 Reyes G M 1957 Rice dwarf disease in the Philippines Food and Agriculture Organiiation Plant Prot Bull 617-19

188 Rice S E A A Grigarick and M 0 Way 1982 Effect of leaf and panicle feeding byarmyworm (Lepidoptera Noctuidae) larvae on rice grain yield Environ Entomol 7593-595

189 Risch S J 1) Andow and M A Altieri 1983 Agioccosystem diversity arnd pest control data tentative conclusions and nizw research dirctions Environ Entomol 12625-629

190 Rivera C T 1956 Three organic insecticides in thecontrol of insects affecting upland rice Philipp Agric 39465-472

191 Roffrey 1 1972 Locusta outbreaks in the Philippines Acrida 1177-188

192 Rojas R 1) 1941 The effect of pruning at varying ages upon the yield of Ramairio Phipp Agric 29851-860

193 Roonwal M i 1979 Ternite life and termite control in tropical South Asia Scientilic Pnbl odhpr India 139 p

194 Rosetto C1 F da Silva Martins N C Schmidt and L EAzzini 1978 Damage caused by pasturespittle bugs 4Dvoi)lavopictaandDJ schach) on rice lBragantia Nota 6 (Matcli)25-27

195 Rosztto C1- S S Neto I- Amantc I) Iink 1Garcia-Vieira D M de SoLza N V Bansa t and A M de Olivcira 1971 Rice pests of Brazil [in Portuguese] Pages 149-238 inrFA()-International Rice Commission 2nd meeting of the Rice Committee for the Americas 6-11 l)ecember 1971 Pelotas Rio Grande do Sul Brazil

196 Rothschild G H L 1969 Obseratiois on the armyworm Spodoptera matritia acronrctoiiles Gn (Lep Noctuidae) Bull Entomol Res 59143-160

197 Rothschild G 1- L 1974 Parasites of the armworm Spolopiera mauritiaacroncitoihes in Malaysia Fntornopliaga 19293-299

198 Sachan i N S K Gangwar and 1N Katujar 1980 Field crickets damaging cages in North Eastern IHill Region Indian J Entomol 42526-527

199 Sands 1) P A 1977 The biology and ecology of Lepocorisa (Hemiptera Alydidac) in Papua New Guinea Res Bull 18 Departshyment of Primary Industry Port Moresby Papua New Guinea 104 p

200 Sarawak 1968 titomology division Pages 97-106 in Report of the research branch of the Department of Altriculture 1966 Kuching (Rev AIppl Eintomol A56603)

201 Satpathy JM M S Das and K Naik 1977 Effect of multiple and mixed cropping on the incidence of scine important pestsI Entomol Res 178-85

202 Sauer IHF G 1939 Notes Ott Elasmpnlpualtsligniosellths Zeller (Lep Pyr) a serious pest ofcereals in Sao Paolo [in Portuguese] Arq Inst Biol 10199-206 (Rev Appl Entotnol A28378)

203 Satinders 1L A 1 SKing and C L Vargas S 1983 Crop pests of Central Anerica with t bibliography [in Spanish] Centro Agroshynotnico Tropical de Investigacion y Ensenanza Turrialba Costa Rica (CATIE) Tec Bol 9 90 p

204 Sauphanor B 1985 Some factors of upland rice tolerttce to stem borers in West Africa Inseu Sci Appl 6429434

205 Sen P atnd S Chakravorty 1976 Stentflics in Haritighata Nadia West Bengal India [it Rice Res Newsl 1(12) 15

206 Senapai B and 1M Satpathy 1983 The seedling fly Atherigona orrzae Malloch is a pest of rice in Orissa (India) J Entomol Res 777-79

207 Shhjahan M S Alatn M Ah nad and BN Islam 1983 A new scarabaeid beetle pest of dryland rice in Bangladesh Int Rice Res Ncwsl 8(I)16-17

208 Sharma G L S C Modgol and R C Gautam 1979 Effect of depth of submergence ott incidence of bacterial blight and white grub infestation in transplanted rice Int Rice Res Newsl 4(6)12-13

209 Sharma S 1985 Scarabaeid beetle outbreak itt upland rice Int Rice Res Newsl 1(1(l)24

210 Shiraki T 1917 The paddy borer Schoenobhits incertulas Walker [achaku Agric Exp Stn Formosa 256 p

211 Silva A de Brito and 11P Magalhaes 1981 Insect pests of rice [in Portuguese] Circular Tecnica No 22 EMBRAPA Centro de Pesquisa t ecuaria de Tropico Unido (CPAJU) Belem Brazil 14 p

212 Singh S A 1979 Emergence of rice leaf butterfly as a rice pest in Manipur India int Rice Res Newsl 4(4)16-17

213 SinghS A 1983 Changing trends of insect pestsof rice in Manipur Int Rice Res Newsl 8(f)12

214 Siwi S S and P H van Doesburg 1984 Lenovorisa Latreille in Indonesia (Heteroptera Coreidae Alydinae) Zool Meded (Leiden) 58117-129

215 Soejitno J 1977 Reaction of some upland rice varieties toseedling fly eltherikonaexigua Stein [in Indonesian] Siri Hana I Penyakit 630-35 Lembga I osat Fenclilian Pertanian Bogor Indonesia

216 Socnardi 1967 Insect pests of rice in Indonesia Pages675-683 in The major inse pests of thu rice plant Proceedings of a symposium at the Internatital Rice Rec Wi itt S pternber 1964 The Johns Hopkins Press Baltimore Maryland USA

217 Soto P E A T Petcz and I W Buddenhagen 1976 Survey of insect pests and diseases of rice in different ecological zones in Nigeria Rice Entomol Ncwsl 435-46

218 Soto 11E and Z Ziddiqi 1976 Screening for resistance to African rice insect pests Varietal improsetnent sCmi n r Bouak Ivory Coast 13-17 Sept 1976 West Africa Rice l)evelopment Association Monrovia Iiberia

219 Squire F A 1972 Entomological problems in Bolivia PANS (PestArtic News Sumr) 18249-268

220 Stapely 1 II 1978 The rice leaf roller in the Solomon Islands Int Rice Res Newsl 3(l)13

221 Suenaga H 1937 On aphids attacking the roots of rice Oyo-I)oubts Zasshi 9150-152 [in Japanese] (Rev Appl Entomol A25745)

222 Suenaga H 1952 On the seasonal abundance of three subterranean aphids on upland rice Bull Kyushu Agric Exp Sti 1249-262

223 Sutynso Yas and Dandi Soekaria 1964 Beberapa hana yang ttenperang tana man padi tanah kering [in Indonesian] Rapat kerja Padi Tanah Kering 6 sd 8 Juily 1964 Kaliurang 9 p

224 Tallamy I) W and R F Denno 1983 Alternative life history patterns in risky environments in example from lace bugs Pages 129-147 in Insect life history patterns R F l)cnio and If Dingle eds Spring-Verlag 225 p

225 Tamura 1 1939 Observations on some insect pests of field crops [in Japanese] 1 Plant Prot 26414-420 (Rev Appl EntomolA27-556)

226 Tamura and Suzuki 1964 Influences of the early-transplanting practice in rice culture upon development of the rice stem borer Chilo siql)ressalisWalker Bull Ilokuriku Natil Agric Exp Stn 761-94

227 Tan J P 1924 The rice root aphid Drowhi ihirsoia Philipp Agric 13277-288

228 Tanaka 1 1957 Taxonomy and distribution of some subterranean aphids injurious to upland rice in Japan with description of a new species [in Japanese English summary] Botyu-Kagaku 22168-176

229 Tanaka T 1961 The rice root aphids their ecology and control Spec [full College of Agriculture Utsunomiya University Japan F3 p

230 Tauber M IC A Tauber R Nechols and 1Obrycki 1983 Seasonal activity of parasitoids control by external internal and genetic factors Pages 87-108 itt I)iapause and life cycle strategies in insects V K Brown and I Ilodek eds )r W Junk The Hague Netherlands

231 Taylor W F 1972 Effects of artificial defoliation (simulating pest damage) on varieties of upland rice Exp Agric 879-83

232 feotia T P S and S Natud 1966 Biotoomics of the rice skipper Parnaraniathiis Fabricius (Lepidoptera Hesperiidac) Indian J Entomol 28181-186

233 Teran F 0 1971 Rice pests of Santa Cruz [in Spanish Ministerio de Agricultura Sita Cruz Bolivia Bull 45 8 p

234 Than Itun 1976 Population dynamics of the yellow rice borer Tryporza itcertulas (Walker) and its damage to the rice plant MS thesis University of the Philippines at Ios Bafios Laguna Philippines 92 p

235 Uichanco L B 1911 Pests Pages 55-74 it Rice Philipp Agric 11-200

IRPS No 123 January 1987 41

236 van der Laan P A 195 1 Life history of the rice seedling fly Contrib Gen Agric Res Stn Boor Indonesia 118 15 p

237 van der Laan 11A 1959 Correlation between rainfall in the dry season and the occurrence of the white rice borer (Scirpolhaga innotala WIk) in lava Entomol Fxp Appl 212-20

238 Van Driesche R G 1983 Meaning of percent parasitismin studies of insect parasitism Fnviron Entomol 121 I1- 122

239 Venugopal M S and A litsingcr 1984 Fffect ofcarbofuran on rice growth Prot Ecol 7313-317

240 Vietnam Directorate of Rural Affairs 1962 Annual work progress report on crop improvement program of rice sugarcane vegetable and field crops (lor the period from July 1961 to June 1962) (11) 299 p (Rev Appl Entomol A51526-527)

241 Wan M T K 1972 Observations on rice leaf- and planthoppers in Sarawak (Malaysian Borneo) Malay Agric 148308-335

242 West Africa Rice I)esclopnient Association 1978 Annual research report Vol 2 Statistics Monrovia Liberia

243 West Africa Rice Development Association 1979 Annual research report p47 Monrovia Liberia

244 Williams ) F and CS lofgren 1981 Fli I illy l-468 anew bait toxicant for control of the red imported lire at Fla Entomol 64472477

245 Williams I) 1 1970 The mealybugs OFlomoptera Coccoidea Iscudococcidac) of sugar-cane rice and sorghum Bull Entomol Res 0109-188

246 Williams D) 1 1982 Pulinaria icervi (Signoret) and its allies ott sugar cane and other grasses Btll Entoinol Res 72111-117

247 Wirjosuiardjo S 1976 Study ofchenmical cont rot of insect pests ott upland rice Imiu Pertanian (Jogjakarta) 1323-329

248 Wirjosuhardjo S and N Wirarta 1976 Chemical control of the white grub tIolorithiatlltriin upland rice in M uneng Irobolinggo Indonesia) [in Indonesian] Imut Pertanian (Jogjakarta) 1330-332

249 Yano K T Miyake and V F Eastop 1983 The biology and economic importance of rice aphids (Henuiptera Aphid idae) a review Bull Entomol Res 73539-566

250 Yelp D and P Rhodes 19(4 lhe success of drilled dryland rice iii the Fiji Islands International Rice Commission Working Party on Rice Production and Protection Manila Philippines 3-10 March1964 Food and Agriculture Organization Rome Italy 13 p

251 Young W R 1981 Fifty-five years of research on the sorghum shootfly Insect Sci Appi 23-9

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Page 7: UPLANDRICE. INSECTl--1i M T -E!R.EOLOU t RTANCE ,AND …delphacid.s3.amazonaws.com › 6958.pdf · The significance of abiotic factors in upland rice insect ecology will be apparent

7 IRPS No 123 January 1987

Table I continued

Order Family Species Distribution Reference

toleoptera Scarabacidae leteroligus (=Aphonoprocts) 15elcs Billberg Africa 36 Dynastinac (=Ileteroligusmees robustui Prell)

Alissonotum pauper Burnieiser Asia 82 Alissonotuon simile Arrow Asia 209 Dscinetusdubibs (Olivie) Latin America 28 195 233 Devscinetusgagates Burmncster Latin America 20 195 Dyscinctt spp Latin America 60 Eutheola Inimilis Bturmcistr Latin America 20 28 8 1 195 203

233Eutheola identarr(Biurnciter) Latin America 121 203 Heteron)ciwv aw rsonJack Africa 82 Hcreron)chusbtuterculatus Kolbe Africa 82 tleteronichtslicas (Klug) Africa 82 fhteron)chus mosamhicus Peringuey Africa 4 6 48 82 (=Veteronychus orzae Britton)

leteronychts lioderes Redtnbacher Asia 77 82 207 ltcteronchusarator (Fabricius) Africa 82 tleterontchusplebepus (Klug) A frica 57 82Ileteron chus pseuocongonnsis Ferreire Africa 32 Ileteronvchns rogiJrons Fairmaire Africa 82 Heteron chus nrsticus niger (K lug) Africa 82 Heteronvchus spp Africa 17 Lachnosterna longipennis (Blanchard) Asia 82 Lachnosternaspp Asia 120 Ligvros fossator Burmeister Latin America 82 Ligyrus nasutus Burmeister Latin America 203 Ligyrus (=Scarabaeus)ebenus (de Geer) Latin America 20 195

Dynastinac Ligtvros huonilis Burmeister Latin America 60 82 Aaladera castanea (Arrow) Asia 82 Maladera orientalis Motschulsky) Asia 82 Malaaerajaponica Motschulsky Asia 82 Phvlognathusdicynsius (Fabricius) Asia 82

Melolonthinae Exopholis h)ypoletca (Wiedemann) Asia 117 Lerlinosterna[= lolotrichia] consanguinea Asia 122

(Blanchard) Leucopholis rorida (Fabricius) Asia 82 117 Lepidiota blanchardi Dalla Torre Asia 76 82 Leucopholis irrorata (Chevrolat) Asia 76 134 Scrica internpta Walker Asia 76 Holotrichialonipennis (Blanchard) Asia 122 Holotrichia serrata Fabricius Asia 122 Holotrichia leucoph thabna (Wiedemann) Asia 82 Holotrichia seticollis Moser Asia 77 78 Holotrichia pnindanaoana Brenske Asia b Stenoeres spp Latin America 71 195 Atoenius spp Latin America Asia 203 Ph luplg aequata (Bates) Latin America 203 Piyvilophagacaraga Saylor Latin America 203 Phyllophagachiriquina (Bates) Latin America 203 Phyllophagadas)poda (Bates) Latin America 203 Phyllophagaclegans Saylor Latin Ameica 203 Phyllophagahondura Saylor Latin America 203 Phyllophagalatipes (Bates) Latin America 203 Phyllophagamontriesi (Blanchard) Latin America 203 Phyllophaga pamisetis (Bates) Latin America 203 Ph vllophaga sanfosicola Saylor Latin America 203 Phllophagasetifera (Burmeister) Latin America 203 Phyllophaga (=tlolotrichia)helleri Breuske Asia 82 117 159 Phvllophagaspp Latin America 81 150 White grub unspecified Asia 84

Rutclinae Lagochile trigona (Ilost) Latin America 82 Anomala dinidiata var barhata Burmeister Asia 77 78 82 Anomala huteralis(Burmeister) Asia 76 Anoimala lurida (Blanchard) Asia 76 Anomala sulcatula (Burmeister) Asia 76 Anontala varians (Olivier) Asia 82 Anomala antigua (Gyllenhal) Asia 82 Anoinala palida (Fabricius) Asia 82

8 IRPS No 123 January 1987

Table 1 continued

Order Family

Tenebrionidae

Elateridae

Languriidae Chrysomelidae

Chrysomelidae

Coccinellidae Curculionidae

Lepidoptera Lymantriidae

Amatidae

Arctiidae

Limacodidae Eupterotidac Noctuidac

Species

Anomaa volita (Blanchard) Adoretus caliginosus Burmeister Adoretus comprcssus (Weber) Papuana huhneri (Fairmaire) Papuana incrmnis Prell Popillia capricollis Hlope Gonocephalu depressur (Fabricius)Gonocephahm acuiangulum (Fairmaire)Gonocephalum sinphx (Fabricius) Epitragus sallei Champion Anaedus ptnctatissimnts ChampionAeolus spp4griotes spp Agriotes mancus (Say) Conoderts spp Aeoloderma brachmana (Candeze)Wireworni unspecified Anadastus filijf rmis (Fabricius)Dicladispa viridicanca(Kraat z)Trichispasericea (Guerin)Oediopalpastervalis (Weise) Oediopalpa guerini (Italy) Chaetocnema cvlindrica (Baly)Chactocnema denticulata S tephens Chaetocnemabasalis (Baly)Aulacophora sp nr similis Olivier Monolepta caripemne Ialy Monolepta bilasciata (Ilornstedt) Monolepta signata Olivier Cerotoma airojcsciata J acoby Diabrotica adelpha Ilarold Diabrotica balteata Leconte Diabrotica graminLa ltaly Diabrotica speciosa (Germar) Diabrotica limitafa quindecinmunctata(Germar)Diabrotica melanocephala (Fabricius)Diabrotica spp Altica (= Fondia)madagascariensis(Allard)Altica spp Epitrix cuctmeris (Harris) Epitrix spp Oulenjo or pae(Kuwayama) Disonchq (= Donacia) spp Flea beetle unspecified Chnootriba (= htIilachna)sinilis assimilis Mulsant Atactogaster iduriens Walker Hypomeces squtamosus (Fabricius) Tan vinecus discoidalis GyllenhalPhaulosoinusmusculinus (= mnus) Csiki Neobaridia anplitarsis CaseyPsalis pennatula (Fabricius) Laeliasufihsa (Walker) Euproctis irgtncula (Walker)Euproctis minor (Snellen)Euproctis xanthorrhoea (Kollar)Amata sp

Diacrisia obliqua (Walker) Creatonotusgangis Linnacus Latoia (= Parasa) bicolor (Walker)Nisaga simplex Walker Sesamia botanephaga Tams amp Bowden Sesamnia nonagroides botanephagaTfums et Bowden Sesamia calamistis llampson Sesamia inferens (Walker) Spodoptera eridania (Cramer)Spodoptera mauritia acron)ctoides Guenee Spodoptera frugiperda ( ESmith)

Distribution Reference

Asia 82 Asia 82 Asia 82 Asia 82 Asia 82 Asia 78 Asia 82 117 178 Asia 82 117 Africa 82 Latin America 203 Latin America 203 Latin America 195 Latin America 195 Latin America 203 Latin America 195 Asia h Asia Latin America 60 73 117 Asia 200 Africa 36 Africa II Latin America 6166 68 Latin America 6166 68 Asia 225 Latin America 61 66 68 81

Asia 178 Asia 55 Asia 55 Asia h Asia 55 Latin America 203 Latin Amefia 203 Latin America 203 Latin America 203 Latin America 6166 68 Latin Amcrie 195 Latin Amw 195 Latin Aierwca 61 81 Africa 57 Asia 178 Latin Americ 203 Latin America 81 Asia 82 Latin America 195 Asia 120 Africa 6 11 Asia 185 Asia 120 Africa 82 Africa 82 Latin America 66 68 195 233 Asia 117 Asia 117 Asia 117 Asia 117 Asia 117 Asia A S Pradhan

unpubl dataAsia 82 Asia 82 Asia 82 Asia 182 Africa 12 Africa 6 36 Africa 6 8 11 12 36 Asia 59 82 84240 Latin America 150 Asia -0 59 196 Latin America 28 61 66 68 81 150

152195203219233

9 IRPS No 123 January 1987

Table I continued

Order Family Species Distribution Reference

Spodoptera exigua (Hubner) Spodoptera exem pta (Walker) Spodoptera ornithogalli (Guenee)

Africa Africa Latin America

32 217 32 33 36217 150

Mythimna loreyi (Duponchel) Asia 30 Mythimna separata (Walker) Mythimna (- Pseudaletia)latifascia [= adultera

Asia Latin America

178 195

(Schaus)] (Walker)Mythimna (=Pseudaletia) sequax (Fabriclus) Latin America 68 Mythimna rosellinea (Walker) Asia b Mythimna yu (Guenee) Asia b Mocisfrugalis (Fabricius) Asia 30 117 Mocis latipes (Guenee) Latin America 66 68 149 150 203

Platysenta (=Spodoptera) compta (Walker) Asia 219233A Barrion unpubl data

Agrotis ipsilon (Hufnagel) Latin America Africa

33 68 81 150 1)5 203

Agrotis spp Latin America 60 Achaeajanata (Linnaeus) EldanasaccharinaWalker

Asia Africa

117 36

Armyworm unspecified Asia 72 84 Pyralidae Cnaphalocrocis medinalis (Guenee)

Marasmia bilinealisHampson Asia Africa

30 4559 72 84233 36

Marasmia trapezalis(Guenee) Africa 32 217 Marasmia (=Sumia) exigua (Butler) Asia 220 Marasnianmralis (Walker) Asia b Marasiniapatnalis Bradley Marasiniaspp

Asia Asia

b b

Leaf roller unspecified M2liarphaseparatellaRagont

Asia Africa

72 12 57

Maliarphasp Asia b Chilo auricilius Dudgeon Asia b Chilo partellus (Swinhoe) Chilo polychrysus (Meyrick)

Africa Asia

12 92 59 72

Qhilo diffusilineus J de Joannis Africa 6 1229 Chilosuppressalis(Walker)Cibozacconius Blezzinski

Asia Africa

59 72 6 11 12

Cravubus spp Latin America 203 Acigona loftini (Dyar) Latin America 203 Acigona chrysographella(Kollar)Scirpophagaincertulas(Walker)

Asia Asia

82 59 72

Scirpophagainnotata (Walker) Asia 59 200 Scirpophaganiuella (Fabricius) Asia b Elasinopalpuslignosellus (Zeller) Latin America 28 60 61 66 81 150

Diatraeasaccharalis(Fabricius) Latin America 162195203219233 28 66 81 111 150 195

Rupela albincllaCramer Latin America 111

Gelechlidae

Stem borer unspecified Undetermined stem borer nr Maliarpha BrachniaarctraeaMayr

Asia Asia Asia

72 b 55

Brachmi spp Africa 32 Hesperiidae Parnaraguttata Bremer et Grey Asia b

Parnaranaso Fabricius Asia 55 Pelopidasagnaagna (Moore) Pelopidasmathias (Fabricius)

Asia Asia Africa

164 36 155217

Pelopidasconjuncta conjuncta (Herrich-Schaffer) Asia b Borbo fanta Evans Africa 6

Satyridac Melanitisleda ismene Cramer Asia 84213 Mycalesis asophisHew Asia 30

Diptera Cecldomyiidae Mycalesis spp Orseoliaoryzae (Wood-Masn)

Asia Asia

b 72 88 131 181

Orseoliaoryzivora Harris et Gagne Africa 61132218 Chloropidae Steleocerellus (=Mepachymens) ensifer (Thomson)

Oscinella spp Asia Asia

205 205

Gaurax sppChloropsoryzae Matsuntura (= kuwonae Aldrich)

Asia Asia

205 82

Mucldae Atherigonaoryzae Malloch Asia 30 82 117130 Atherigona exigua Stein Asia 82 117 178

10 IRPS No 123 January 1987

Table I continued Order Family

Diopsidae

Agronyzidae

lHymenoptera Formicidae

Species

Atherigona oritentalisSchiner Atherigona indica MallochDiopsis longicornis(= thoracica)Macqurt Diopsis spp Pseudonapnyza asiatica Spencer Pseudonapomyza spicata (Malloch) Agronoza orvzae MunakataAcromyren landoltibalzani(Emery) Acromyrmex landoltifracticornis(Forel)Acromyrnex heyeri Forel)Atta bisphaericaFore Atta capiguaraGoncalves Atta laeLigata (F Smi h1) Atta opacicepsBorgmeierSolenopsisgeminata (abricius)Pheidole sp Pheidolegeton diversus Jerdon Monomorium pharaonis (Linnaeus) Ants unspecified

Distribution Reference

Asia 82 Asia Africa 82205Africa 11 1248 Africa 217 Asia 213 Asia h Asia bLatin America 6168 195 Latin America 61 195 Latin America 61 68 195 Latin America 61 68 195 Latin America 14 61 68 195Latin America 61 68 195 Latin America 61 195 Asia b Asia b Asia 1) Asia b Asia 120

aG orlentalls formerly recorded in Asia as G africana thPhilippines reported in this study CSubterranean root feeders

subterranean nests Foraging ants will feed on germinatedseeds only when the supply of ungerminated seed is low However these ant specialists feed on a wide varietyplants and thus help control grassy weeds (39) This dual

of

pest benefit role extends from tending ants which protecthoneydew-excreting pest species including root aphids and mealybugs (workers repel potential predators and even selectively kill parasitized aphids) to predators of leaf- and stem-feeding rice pests such as leaffolders armywormswhite grubs and stem borers (1 17)

The upland rice environment is particularly well suited to ants because of the lack of flooding Frequent tillage perpetuates grasses and granivorous species that are highlyadapted to constructing new nests in recently tilled fieldsAnt nests however are relatively shallow and tilling with a moldboard plow can destroy them Also upland ricefields are uually small and isolated and a given area in a smallfield has more field borders than it has in a larger fieldallowing high rates of infiltration from more sedentaryspecies such as Plieidole that do not colonize tilled fields but nest in soil under shrubs and trees along field bordersPheidole workers forage into fields from borders and store seed in their nests In slash-and-burn or other no-tillage rice cultures nests would be preserved in the field and possiblylead to high rates of seed predation However granivorous species do not prefer forest habitats

Damage is characterized by reduced usually patchy plantstand (120) Seed-thieving ants greatly prefer rice but they can be selective regarding plants they collect and store Theyharvest more seed than they can consume so patches of germip ing surplus seed (such as Rottboellia) will sproutfrom their nests during the rainy season

In Latin America leaf-cutting ants (Aita spp and Acromyrmex spp) occur only in upland fields and defoliate

young rice plants (47 68) They take leaves to fungal gardensin underground nests where fungi predigest the plant material

Termites Subterranean termites of the family Termitidae which lack symbiotic protozoa to help digest plants are the most frequently mentioned termite pests of upland rice (1185 120) This family cultures fungi in special undergroundcells --- fungal combs made of half-digested plant material (117) The fungi inoculated onto the combs break down plant material into food that termits can digest The termites then consume the fungal combs These grasslandtermites build nests below the plow layer in uplandricefields The colonies attack living plants only when dead plant material is not available They will attack a droughtshystressed crop but prefer older plants having greater cellulose content They will damage newly planted crops where clean culture has removed vegetation

Infestations are worse in deep light-textured soils with low moisture content The first sign of damage is yellowingof older leaves (195) Termites feed on roots (the plantyellows then wilts and finally collapses) and germinatingseed (loss of stand) or move above ground at night to cutseedlings at ground level which they cover with soil for later consumption Soft-bodied termites are highly sensitive to desiccation They live within a self-contained system of airtight chambers and build surface tunnels linei with nudand body secretions (carton) to maintain more than 90 relative humidity Termites can be located by their tunnels

Termites apparently can withstand limited submergence (157) and can be a pest of dry lowland seedbeds (117)perhaps by having nests below perched water tables

Termites appear to do more damage in Latin America where large ricefields are plowed with tractors deprivingtermites of preferred vegetation and forcing them to live on

rice plants In Asia upland ricefields are typically small with Iprge field border areas (ecotones) where termites can forage for preferred food other than growing rice plants All grassland termites do not feed on rice In an upland area in Batangas Province Philippines ricefields are commonly infested by Hosiititernesluzonensis (Oshima) however this termite does not damage rice In the same province though Mrroternesgivus(Hagen)feeds on rice seedlings (178)

In Africa termites may damage rice even though fields are small and surrounded by perennial vegetation These termite species perhaps prefer rice over other hosts

Mole crickets Nymphs and adults are nocturnal and burrow through soil feeding on roots of a wide variety of plants or forage above ground as predators of insects Adults live 3-5 mo and are even cannibalistic Mole crickets prefer low-lyin moist upland soils with high organic matter (II 81) In the lowlands they inhabit rice bunds or nonflooded fields Adults are highly mobile and can leave a flooded field to locate a more suitable habitat Losses show as wilted plants (from root feeding) or reduced stands (120) Damage is more common near field borders where mole crickets relocate after tillage (126)

Fiehl crickets Nymphs and adults of field crickets have similar nocturnal habits and damage rice as much as mole crickets do Piles of weeds removed from fields attract them They make subterranean nests and tunnel through the soil to feed on roots Some species prefer seed to roots Others feed at the base of stems causing deadhearts

Tunnel entrances surrounded by excavated soil are easily seen Young plants are cut at ground level and stored in underground cell Field crickets tre highly polyphagous as are all soil pests (117) Sachan et al (198) reported that field crickets prefer maize to upland rice Like mole crickets field crickets cannot survive in standing water and are therefore more prevalent in upland fields

Whitegrubsandhackbeetles Scarab beetles that feed on living roots as larvae but not as adults are called white grubs or chafers (Melolonthinae Rutellinae) and those that feed as adults but not as larvae on plant crowns are called black beetles (Dynastinac) In the tropics chafers or white grubs have a I-yr life cycle starting with adults emerging from the soil about I mo after the first downpour of the rainy season (134 159) They lay eggs at the same time farmers sow upland rice The rice passes its most susceptible stage and damage is mostly avoided when the white grubs are small After several months the long-lived larvae are large enough for two or three to denude the root system of mature rice (223) This intensity of damage is rare but wilting occurs when root loss is combined with drought stress

The larvae need damp (not saturated or dry) soil to survive and pass the unfavorable dry season 05-2 m underground in aestivation in the moist soil The first heavy rains of the season (20-30 amd) stimulate the grubs to resume activity and after several weeks they develop into

IRPS No 123 January 1987 11

pupae and adults and eventually dig their way to the soil surface and fly to nearby trees to seek food and mates The adults of most white grub species feed only on trees but in some species notably Eutheolahumilis the adults defoliate and cut off stems at ground level (28) Putheola hidentata damages sugarcane fields in Venezuela (121) Floods kill white grubs (208) Grasslands can support large populations and therefore white grubs can be more abundant in newly planted upland ricefields that were previously fallow (81)

Black beetles live up to a year adults can attack a crop at any stage by burrowing into the soil (127) and feeding at the base of stems to cause whiteheads (207) Adults emerge with the early rains and are normally more abundant and do more damage on a young rice crop with small root systems (38 81)

The larvae feed only on dead organic matter in dryland fields and do not attack rice Adults arc highly mobile and although sensitive to flooding can invade a field soon after it drains (115)

Root aplhids Several genera of migratory aphids feed on the roots of upland rice (249) Populations build up more in light-textured soils with high percolation rates Damage has also been reported from dry seed heds of lowland rice (52) Tending ants -- Pheidhle Cren aster Tetramoriuw lsius Tapinoma -- are necessary for root aphids to multiply Aiits harbor aphids in their nests over winter or during unfavorable periods and relocate them on rice plants digging tunnels along the root systems to allow these softshybodied insects to penetrate the soil Most intensive studies on aphid life histories have been done in temperate environments (158 222 229) Root aphids fly to rice plants at the beginning of the rice season and pass through several generations continually developing winged forms to relocate on alternate grassy hosts until winter when they move to perennial hosts usually trees Yield loss occurs mainly through reduced tillering but infested plants become yellow and stunted and in extreme cases wilt (227) Aphid populations build up gradually so damage usually begins in the late vegetative and reproductive growth stages

Root-feeding ineal tugs More than half of the mealybug species or rice feed on roots (245) Being soft bodied mealybugs are not adapted to living underground but they survive because tending ants (the same species that tend root aphids) dig burrows for them and move them from plant to plant Well-drained soils ihelp mcalybugs survive Although six species are recorded as subterranean root pests of rice we found no report citing damage Root-feeding mealybugs are probably more prevalent on perennial grasses than on short-lived annual rice

The most common rice mealyb ig Brevennia rehi is a foliar feeder favored by lack ol rain apparently it is responsible for high yield loss in Northeast India and Bangladesh (N Panda pers comm as stated by IRRI [1061) This report needs confirmation Mealybug damage was only briefly mentioned in another report (156)

12 IRPS No 123 January 1987

Root weevils A few species of upland weevils (IHporneces Donacia Atactogaster) attack rice roots as larvae and feed on stems and leaves as adults Damage ispatchy The larvae are not adapted to submergence and are true upland soil weevils isopposed to the so-called water weevils They rarely become sufficiently abundant to cause damage Adults on rice foliage or the soil surface show that the roots are being attacked by the grubs Root weevils are highly polyphagous (185)

Root heeths Soil-inhabiting false wireworms Gono-cephalumt spp are reported in upland ricefields The adults and larvae normally feed on decaying organic matter but land preparation for upland rice removes these sources The larvae feed on seedling roots and nocturnal adults roam on the soil surface to cut off seedlings at ground level (82) This group of ground beetles is highly omnivorous adults even prey on stem borer larvae (183) True wireworms (Elate-ridae) feed on living roots but rice as a host has been reported Infrequently (117) In Latin America the highlypclyphagous larvae of Diabrotica rootworms are also recorded on rice

Root htus Upland rice seedlings can be attacked by the adults anI nlViphs of chinch bugs Blissusand Caenoblissus and bro n bug Scaptocoris which feed on the roots with theirsucking nouthparts (80 83) Three genera of burrower bugs A tms (eotonhus and Stiharopus -are common iil grasslands in the Philippines and may attack the first crop (rice or maize) planted after the dry season Chinch bugs are most abundant after a dry spell These insects are large and can kill seedlings or stunt and reduce tillering on older plants I hey prefer other hosts and rarely damage rice Adults can also feed oii developing rice grains (81)

(utworms Soil-dwelling noctuid larvae hide under-ground during the day to avoid predation by birds and bccone active at night above grour cutting off you ing plants at ground level (hence tile naiie cvtworn) arvaedrag severed plants into hurrows llie larvae have wide host ranges arid pupatc in the soillgrotis ipsilon occurs worldwide and appears to be the most common cutworm pest of upland environments but is more prevalent inlowland rice particularly in dry seedbeds

IFoliar pests Other groups of insects are adapted to upland environments because they pass one growth stage underground either as egg (in heinrietabolous species) or pupa (in holomnetabolous groups)

Rice seedlibig maggots AitJeriota flies occur in non-fhoodcd environments in Asia and Africa (180) Feeding of larvae one per iller causes deadhearts Flies as opposed to moths only secondarily evolved as pliytophagous pestsfroni a saprophagous Origin Athrnonalarvae secondarihv feed onidecaying tillers whereas larvae of stem borer moths actively Iced on living tillers to cause deadlicarts Seedling maggots will die if they cannot sever a tiller on which to feed

as it decays Plants die under heavy attack whereas lighter infestations cause stunting delayed maturity and ragged leaves (236)

Eggs are laid only on actively growing plants (49) aid damage occurs wiihin the first week after crop emergence(170) Larvae pupate more in stems than in soil Seedling maggot attack is highly seasonal (206) normally peaking 2-3 mo after the onset of the rainy season These pests possiblyaestivatc over the dry season Rice isonly one of their many graminaceous crop and weed hosts

The rice stem maggot (hlorops orzae is adapted to temperate upland and lowland rice growing regions of Asia In Japan and China upland rice grows in mountainous regions a habitat of this chloropid fly The larvae tunnel into tilestems The stem maggot is more damaging during the seedling stage larvac hibernate in grasses

File several dipterans reported as stem maggots in dryshyseeded lowland seedlings inIndia (205) could presumably attack upland rio

Leaf urinrs Icaf miners are common on wheat and maize but are rarely reported on upland rice (213) Three leaf miners are more prevalent on upland than lowland rice inthe Philippines Pvudonalomy a asicanh Vficataand lIAgromrza orzae attack at the seedling stage The larvae develop inside the tunnels they construct in the parenchyma tissue P Vpicata and Aorvzac prefer maizeEehtsine and other grasses to rice spicatacan become abundant on wheat

Leaf beetles Chrysomelid and coccinellid eaf bcetles scrape or otherwise remove leaf tissue as larvae or adults or both This damage accelerates desiccation of plants Tile larvae of sonic species are root feeders Flea beetles Epitrix and (lraetocnetna and rootworms l)iahroticaspp feed on rice foliage only as adults Their root-feeding larvae have hosts other than rice The adults and larvae of the hispa-like Oediolmlpa spp beetles of Brazil and (hmootriha (=EPfishyachra) beetle of Africa feed on upland rice leaves Oealioshypapa spp larvae are leaf miners (mootriha is also pshyvalent in lowland environments (11) Leaf beetles prefer vegetative rice (81)

The rice hispa Dicladipaarnigera is not reported on upland rice and prcers more aquatic habitats In Central Africa however I)cladispm viridicyaiea attacks uplandrice in the vegetative stage as well as lowland seedbeds 7richipasericea is reported on upland rice in West Africa and Madagascar (57)

The rice leaf beetle 1d1ea 01izae riiuech like Chioropsor|rzae is adapted to temperate regions of Ashi and is usually a pest on lowland rice but seldom oi inpland rice in the mountains (82) The adult hibernates on grasses

Armmworms Species of grassland-adapted SpoptloeraPlatysemta and A ithimna larvae create widespread epishydemics They can defoliate ricefields generally in patchesfrom early vegetative growth to harvest (41) Armywornms pupate inthe soil and highly favor upland rice (34) Adults

can disperse long distances to colonize even remote upland areas They become more abundant than their natural enemies particularly after favorable rains following a prolonged drought (196) The drought kills the armyworm and natural enemies (mainly parasites) but the armyworm can recolonize rapidly

The natural increase insoil fertility from mineralization of soil N over the drought period promotes luxuriant plant growth to foster a rapid buildup of these highly fecund species Weeding rice increases tilelikelihood of damage The parasites normally return to control the armyworms but not until after serious damage

Outbreaks on upland rice have been reported in Ghana (3) inzibar (38) Central Africa (36) lPanama (165) Brazil (28) Malaysia (196) and India (167) Defoliation may be severe often leaving only the base of stems Armyworm larvae hide during the day under litter or in soil cracks At night they ascend plants to feed Larvae pupate in the soil

Thrips Thrips frequent upland rice but damage is less than in the lowlands Thrips larvae and adults feed on leaf blades by rasping causing leaf rolling and stippling Thrips prefer vegetative stage rice and their survival increases during drought (heavy rain washes them from foliage) although drought stress usually outweighs thrips damage Drought and thrips cause leaf rolling in upland rice (235) but thrips have been overlooked perhaps becaLse of their small size Thrips are most numerous indry season irrigated rice The combination of favorable plant growth and lack of rainfall is ideal for thrips In contrast the lack of rainfall in upland rice hurts the crop and eventually the thrips whose numbers must decline

In [atin America Frankinilla rodeos attacks panicles before they emerge from the boot calusing sterile grains

Stem hug~s Black bugs are the most common group of rice stem-feeding -lemiptera in Asia The most frequently mentioned species ----Scotinophara coarctaia and Shrida - prefer aquatic habitats and therefore could not occur in upland rice as sometimes reported Upland species in the same gerurs - notably S tarsalis and S scotti -- are morphologically similar and remove sap from stems of grasses and upland rice The upland species are notably more abundant in upland rice planted near forests

In Latin America tile Tibraca ima-large pentatomid tiventris causes deadhearts and whitehcads on upland rice Both adults and nymphs remove sap from internodes of plants more than 3 wk old Tillers are killed by mechanical damage and by the entry of secondary microbial infection This damage is often confused with that of lepidopterous stem borers (233) Eggs are laid on the leaves and stems Nymphs are gregarious This species can also be found on irrigated rice Drought reduces population buildup

Stiem borers Lepidopterous and dipterous stem borers are widely recorded on upland rice but cultural practices and alternate hosts determine abtndance Except for Elasmolmll)uS Iinosellts numbers usually build up toward

IRPS No 123 January 1987 13

the end of the crop growth cycle Therefore late-maturing varieties staggered planting and lack offloodingfavorstem borer buildup Many stem borer species can lie dormant in the stUbbleafter harvest and theirsurvival is encouraged by panicle harvesting (leaving tall stems) and lack of tillage after harvest

Stem borers disperse and lay many eggs Many species are polyphagous however the monophagous MaliarpiasepashyratellaDiopsis Ivmgicornis Scirpophaga incertlas and S innotata can also be abundant on upland rice (hilo Suplressalisdevelops fasterand becomes la rger when reared on upland than on lowland rice (226)

The lesser corn stalk borer lasmopal tt Ittoselhs is semisubterranean and is perhaps the stern borer most adapted to upland rice(129) Highly polyphagous it infests maize peanut and cowpea It attacks seedling rice (one larva can kill tip to four plants before maximum tillering) (195) (hiv and Maliarphastem borers prefer older plants But Scirpophagaspp will attack young plants even in the nursery lasn opalls larvae tunnel into the stems at or below the soil level mausing dcadhcarts Larvae are not found in their turnnels inside sterns beca use vhei disturbed they retreat into cases made from soil particles bound by silk The cases areattached to the tunnel entrance (143) This behavior probably evolved as protection from natural enemies Larvae pupate in the soil The lesser corn stalk borer is more abundant during drought damage is often confused with that of drought (233) This species prefers sandy soils arid its distribution within a field may relate to soil texture (143)

Diatraeasaccharalis aside from the lesser corn stalk borer is frequently a stem borer pest in Latin America According to Teran (233) rice and maize are more preferred hosts than sugarcaine its namesake

Atcitonacrvsograpla is a less known upland rice stem borer in Asia with habits similar to Easinopall s (82) because it is not found in lowlands Acutra has grassy weed alternate hosts As a moth it is often confused with the dark-headed stern borer(hi polur(standstriped stern borer (hi sup)lpsrcais

Other common upland rice stem borers are also reported in the lowlands The dominant species in upland rice vary greatly with location In Uttar lPradesh India (78) and in Japan (123) the pink stem borer Sesamia htfirens is more prevalent hIKenya the principal rice stem borer is Chi partelus

(92) The lowland African sten borers Aialiarpha sepashyratellaDiopsis vngicornis and Sesamia calatnisnis are less abundant Oimupland than on lowland rice(217) but perhaps cause greater yield loss because plant injury is greater if combined with drought css (92) The stalk-eyed stern borer D)ivpsis is more prevalent in wet habitats (6 1I217)

In Central and West Africa Maliarpha and Sesamia calanistis are the dominant stern borers of upland rice (8 9 10 II) (lilo dihlfilineui and Chio partellus inhabit the

14 IRPS No 123 January 1987

upland savannas whereas Chilo zacconio prefers the lowlands (II) Aaliarpha is most abundant in the mangrove swamp habitats (91) coastal regions (16) or rain forest zones (II) of Africa and appears to be the homologue ofScirlpophaga incertulas The yellow stem borr can also be prevalent in upland rice (120 167) The sugarcane borer Eldanasaccharinais so far only a potential pest of uplandrice (91) despite Grist and evers (82) warning

Grasshotl)ers andI locitas Short-horned grasshoppers and locusts -- Locvta Patanrga Schistocerca - layeggs in the soil inhabit grasslands and develop into swarms to seriously damage upland rice from time to time (173) Largenumbers can destroy a field leaving only the stubble In low cmuoroer s giassnoppeirs an cut panicles (173) Farmers may not plant if locust swarms are imminent Prolonged dryweather followed by favorable rain favors development of large swarms (173) Usually breeding in di) grasslands theydisperse toattack crops including upland rice(191) Eggsaresusceptible to desiccation d uring drough t and toinundation during seasonal rains With favorable soil moisture over consecutive seasons natural enemies ca nnot prevent a largepopulation increase leading to migratory swarms (64)

Most grasshoppers on rice are adapted to uplands (107)but lowland rice grasshoppers iervgl(rlhusspp and Ora spp occur occasionally on upland rice (2 82) They lay eggson rice foliage Nymphs are scmiaquatic

The gryllid Lsctrtus coucinnus is a pest of lowland and deep water rice in Bangladesh and Thailand In the Philippines it damages upland rice grown in isolated pockets near lakes The gryllid nymphs and ad ults feed on the central portions of folrage leasing only the midrib and leaf nargins

Leal ler Pvralid moths whose larvae fold leaves to make a feeding shelter occur worldwide (217) but arc appareritly most common in Asia Leaffolde rsattacking rice in Asia are a complex of species (26) (Ctalhahcrcisinedinalis the best-known rice Ia flolde r in Asia may prefer grassy weeds over rice (86) It is a late colonizer of uplandrice U1arasmja eviua a lowland species (220) nreflers rice to grass specres (86) and colonizes upland rice during il early growth stages llarasnria patnalis is the most corn-rnonly encountered species in upland rice in the Philippines Perennial grasses are their alternate hosts and they can survive year-round in upland rice environments The leaf-folders 11arasmia ruralisand Alarasmia spp are least aburndant Dormancy is unknown in leaffolders Cnaplha-ocrocis medhnalis however is known as a migrant (90)Leaffolders remove photosynthetic tissue attack during the flag leaf stage is particularly injurious Popurlations are normally held in check by natural enemies but leaffolders become parlicularly abundant in conditions of high plantfertility and shade

Butterflies Skippers PelopidasInathiasand Borhofanta are more prevalent in African lowland than in uplandhabitats (6) but extensive studies in Japan and China

showed that Pelopidas mathias prefers an upland habitat (155) Parnaragutata seasonally migrates between lowland and upland areas and is equally abundant in both environshyments (164)

The green horned caterpillar Melanitis leda ismene is more abundant in lowlands (212) but our experience in the Philippines suggests that Pelopidas niathiasand 11elanifis leda ismene are often more abundant in uplands than in lowlands Bamboo iaunbuiva spp annually sustains P mathiaspopulations in the Philippine uplands Adults feed on nectar from flowers and migrate to upland rice during the late vegetative stage

iPi erlled h i-1 1d dill L tIC i 7-0 l(become numerous probably because of high egg predationEgg and larval parasitization are often low Butterflies can readily disperse long distances to seek renlote uplandricefields However they have low biotic potential and even it predation rates are low the rarely become abundant Upland environments with their diverse microhabitats can provide more favorable sites than lowland rice plains

tutterfly pests of rice have broad host ranges and become dormant Juring urnfavorable times of the year

IolhagrotslaidopwraThe larvae of many polyshyphagous moths are reported to defoliate upland rice tussock moths lsaispelnnatula Iawlia sUifita luproctisirgtwltaE mnorand P vathorrhoeaslug caterpillar

litotiahicolorhairy caterpillars Nisaga sinp vandA mata spp and wooly bears l)iacrisiaohliqtia and Crvatonotts gangis Many of these polyphgous larvac migrate and are transients in rice including HelhWhis arm iera and 1t zea highly pol phagous species (82)

Gall nidges The Asian rice gall midge Orseoliaorrzae (88 131) and its African counterpart 0 ortzivora(6 11 12 32217) prelerlowland to upland rice Gall midges in rainfed areas are prevalent only during tle rainy season and dispersepoorly Their alternate hosts (wild rices LeersiaIschaetnuan and Pasahmt)are mostlyaquatic grasses and are less likelyto be near upland rice areas In an area of mixed uplandfloating and rainfed lowland rice cultures in northern Thailand upland rice which is ilanted a month earlier was attacked firstbut subsequcent populations were much lower than in rainfed lowla id or floating rice Low numbers of gallmidges passed the dry season orr perennial weeds such as Paspahim (88)

Smot (hid(s Reports of aphids attacking upland rice plants are few (249) Without maize the maize leaf aphidRho alosihhunm maidis attacks upland rice in the Philipshypines The plum peach aphid IIrsteroneuraselariae was reported in Sierra Leerne toinfest leavesand tnriperied grain(7) Siphaglrctriaecaused transverse linrea r necrotic striae iir rice leaves in Italy (172) I-lea v rains held populations in check Rice is not a preferred host for shoot aphids

Leafhoper andI )anthol)pers Cicadellids and delshyphacidsdispersecasilyand readilycolonie upland ricefrom nearby lowland areas A study in Sarawak (241) showed

IRPS No 123 January 1987 15

similar species in upland and lowland rice Nephoiettix The principal vectors of rice virus diseases generally have species dominated The most prevalent species accounting narrow host ranges and alternate hosts in the grass family for 90-95 of the sweep net samples wre N virescens N are mostly annuals dying in the dr3 season Some virus umropictus Nilaparvia hgens Soatellafiircifera Cfana vectors such as BPH can disperse lontg distances but few do

sectraand Recilia dorsalisLowlands had higher popula- Th restriction of upland rice and weed hosts to the rainy tions Katanyukul and Chandaratat (120) monitored hopper season the small area planted to upland rice and the hopper numbers in upland and lowland fields of northern Thaiiand vectors having narrow host ranges all work against virus and found similar results A comparative study of plant- diseases in upland rice hoppers in Fiji showed more brown planthoppe BPH) and In Latin America Sogctdhs o(rzicola and S cuhanus whitebacked planthopper (WBPH) in wetland rice than in transmit hoja blanca virus disease but reports on upland lowland rice and hopperburn was prevalent in lowland rice are rare except in the favored upland areas of Colombia areas with standing water (89) Peru and Venezuela (35 42) S rricolaocurs in Brazil

Hopperburn in upland ricefields in Asia is rare (84) In on upland rice but hojaI blanca has not been reported (67) Batangas Philippines isolated fields of upland rice become S orzicola has a wide host ramgc (10) similar to WBPI- in hopperburned infrequently from WBPH A report from Fiji Asia WBPH however is not known to transmit any virus cites hopperburn from BPH and WBPH (250) In these two disease situations upland rice was grown on highly fertic soil and In Africa pale yellow mosaic virus is endemic to swampy N fertilizer was applied Hoppers respond to better nutrition lowland areas where rice ratoons year-round (19) and is Normally upland rice is grown on poor soil and receives no mechanically vectored mainly by hispid beetles A minor fertilizer vector a flea beetle Chaeiocnema spp is an upland rice

In Asia upland rice ras are normally not far from pest and upland ricetields near endemic areas occasionally lowland ricefields-- a source of hoppers Otherwise hopper contract the virus species with greater polyphagy than BPH or Nelhotettiv Spittle bugs Cercopid nymphs producc a protective virescens would be dominant frothy covering that looks like saliva In Latin America

In Africa leathoppers and planthoppers do not become spittle bugs damage upland rice In parts of Brazil where abundant on upland rice (6 II) probably because lowland upland ricefields are surrounded by pastures adults rice is not widely rlanted nearby The only reported migrating from grasses can damage rice severely and cause hopperburn as from Nilapanrata maeander in breeder complete hopperburn in a young rice crop within a week of plots receiving high N rates in Nigeria (98) Hoppers colonization (194 195) Dispersal to rice occurs even if however are a greter problem in Latin America where pastures are verdant and spittle bugs prefer pasture grasses large grassland areas breed polyphagous species (Gra- to rice (168) Nymphs are rarely abundant enough to phocephala spp Hortensia spp lEvitianus ohscurunervis damage rice Regular rainfall favors their development Balclutha spp and Draeculacephala spp) vhich then (233) disperse to cause hopperburn on seedling rice (68) Rosetto et al (195) reported Graphocep7hala spp and Hortensia spp Seed pests to be equally abundant on upland and lowland rice Alydid and pentatomid bugs are seed pests of upland rice

The leafliopper and planthopper epidemics in Asia that worldwide In Asia Leptocorisa sp prevail Their habits caused severe losses from hopperburn and vijus dicases make them particularly suitable for upland rice 1)thev feed over the past several decades occurred principally on on many grasses (and therefore can survive the earl y wet lowland rice Except in a few isolated cases (160) upland season before rice sets its grain) 2)adults are long-ied and rice has been spared from tungro grassy stunt and ragged mobile allowing them to find isolated plantings 0 rice stunt virus diseases vectorcd by green lcafhopper (GLH) (besides seed bugs only birds and rats appear to have shyand BPI] (123) Virus diseases have occurred in upland rice capability) and 3) adults aestivate in forested areas or where lowland rice was nearby Rice dwarf disease however sugarcane fields during the dry sca son when neither rice nor was first recorded on upland rice in the Philippines (187) grassy weeds are present (I199) Rice seed bugs concentrate Virus diseases can be perpetuated only on livir plants or on small-scale upland riceficlds because they can actively insect vectors and no living rice - plans tooned or search them out The prevalence of groves of trees characshyvolunteer - grows year-round in upland arcas excert in a teristic of upland environments ensures nearby aostivation few with 12 mo of rainfall sites keeping the seed bugs close by

The smaller brown planthopper Laodcelphax striatellu Leptocorisae are uncommon in lowland rice plains occurs in upland rice in Japan (18) It spreads black- because 1) their populations are diluted in a sea of rice streaked dwarf and stripe viruses between winter wheat and planted more or less at the same time (they ca n only feed on barley to lowland rice Early planting of lowland rice rice during the milk to hard dough stages) 2) they lack nurseries spreads these viruses but upland ice has remained aestivation sites and 3) weedy fallow areas are limited uninfected The lowands of Asia are dominated by L oratorius

16 IRPS No 123 January 1987

which lays its eggs hign on tile foliage (199) L acuta and L solonlonlesis oviposit at ground level the former on litter and the latter loose on the ground These egg laying habis explain theirtenvironmental prefcrencecs(214) In Japan the postdiapause adults of L chinensis migrate from moun-tainous areas to upland ricefields during floweiing (1 14)

Pentatornid bugs prevail in Latin America (8182 195) and in Africa pentatomid and coreid bugs arc equally cited (6 82) l)amage is often characterstic of the species (5)rcoris ventralis and Mnida spp are the common

pentatomid seed bugs of Philippine upland rice but they prefer lEchinochva spp grass seed to rice Vearaviritda occurs worldwide and hoften more abundant in the uplands(54) We saw few reports of seed bugs on upland rice but see(d bugs apparently opelrate cqually between by land and upland rice Good dispersal is necessary ior species that call feed for only 2-3 xon a crop and must first find those crops at ripening stages

Other pct species feed n flowers In Mato Grosso Brazil (194) and Santa Cruz Bolivia the panicle weevil Aeo-hariuia amllitarsis attacks rice during flowering (68 233)The larva e rice sten horers adutlts feed at the bases of spikelets caIusing empty grains Also inLatin America thecydnid -lkiituttatralusisa seed bug as anad tilt and nymph (194)

A number of beetles ecd on poilen - coccinellids lialrotia Aulac lhora and Monohpl-a (55) - but becatise riCe is self-pollinating and fcrtiliation occurs before pikelets open they pose no threat to yield

IST ECOLOGY

We now look at characteiistics of the life cycles of upland rice insect pests to lea rn hew each plays an adaptive role in upland rice ecology

Life history strategies Orzasativadoes not tolerate drodgh well so upland rice is highly seasonal normally grown in the wettest months of the year Usually rice is present for less than half of the year The rice-fiee fallow poses serious problems of survival to upland rice insects which have evolved att least four mechanisms to ovrcolne tle cvcl ic-l lack of a hostI) polyphagy 2) longevity 3) dormancy and 4) vagility (dispersal)

Poldt)agy lolyphagy is defined as having hosts of at least two botanical families oligophagy of more than one gents and ionophagy of only one ventus (40) - inour case Orvza Most wild rices however are illi tiC and theieiirc are normally far removed from upland rice hahitat othewise more would be alternate hosts for uplanJ iice insects Thte highly mornophagous rice pests shySciriol)haita hiceritlas Nilal)arvaiahiens Iand Nealo--tettiyvircsctws in Asia and Aalirpha secparatella and 1ioj)sis lngicornis inAfrica - occur on upland rice but

are more abundant in lowland culture They can specialize in rice because they are highly vagile and can attack all growth stages By specializing in rice monophagous rice pests can outcompete related but polyphagous speciesIntensive andI extensive lowland rice culture has favored monophagous species Riqwla alhinella (Cramer) the onlymonophagous species in Latin America is rarely reported on upland ice (I1)

Because of its limited temporal and spatial existence upland rice has not favored the evolution of specialized species Clhang(44)shows evidence that upland rice has only recently been cultured by man from the lowlands thus pests have had little time to fully adapt to the crop All but the lowland-adapted insect pest species have alternate planthosts to rice Root pests with limited mobility are highly polyphagous Pests that specialize in one growth stage eg secdling maggots seed bugs are oligophagous or polyphagous

Most alternat hosts of rice pests are annual grasseswhich also are seasonal Upla_rice is also a host but not the preferred one of the most highly polypliagous spcciessoil pests grass hoppers armvwarrs and Vearaviirielda

The d ry season poses igreat obstacle to ani upland riceinsects survival and only sonic perennial plant hosts are suitable food during the off-seasoin Plolyphagy allows upland rice insects to suLrx-ixe during thC rainy season when plants- both annual and perennial -areactivelygrowing and therefore more nutritious

Loiugevitr The ability of aiinsect to live 2-3 mo without undergoing dormancy helps species sich as root-feedingwhite grbs (as larvae) and secd-feeding bugs (as adtilt) to survive unfavorable periods White grubs arc not highly fecund and their life strategy is to improve survivorship of limited progeny Their subterranean habitat hides them from manv natural enemies lBtut root tissue is not highly nutritious so they ltilst eat great quantities and need a largeinsect biomass to digest it Other soil-inhabiting rootshyfeeding insects have short life cycles but are more fecund

Seed pests need greater longcvi vas adults to locate ahost at early grain dcclopment arnd give sedentary offspring time to develop Few groups of upland rice insects live longin active development Polyphagy and longevity only increase their ability to survive during the rainy seasonOther mechanisms are needed to survive the dry season

orantc In the tropics the nonrice season is the dry season while intemperate areas tile off-+season is winter Many upland rice insects undergo dormancy inlsummer (aestivation) or winter (hibernation) Dormancy is simplyinactivity during winters with prolonged temperatures below 10 C insects find shelter arnd cease movement Activity is resu med with warmer t empcera tu res A deep state of irnactivity termed diapauise occurs in sole species that prepare physiologically (havxing low metabolism and convcrting gonad s to fatty fuel reserves) for the unfavorable season (148) They enter diapaUse inanticipation of unshy

IRPS No 123 January 1987 17

favorable weather and resume activity after favorable Armyworms skippers and locusts actively travel hundreds weather begins usually in response to photoperiod or of kilometers in air fronts such as intertropical convergencethrough a delayed response to a stimulus such as heavy zones to descend onto upland rice fields far from their rainfall (237) Not all individuals of a population have the breeding grounds (31) Several generations can build ip on same threshold for environmental responses for entering or alternate hosts in tile grass family before rice is plantedterminating diapause -- this is seen as a mechanism to These large insects are active fliers Smaller insects such as prevent total mortality if a false cue appears such as a thrips aphids and planthoppers passively migrate long sudden warm spell in winter or rain in the middle of the dry distances by the wind season (154) Otherspecies ean readily travel tens of kilometers - stem

Probably more upland rice insects undL go dormancy borers leaffolders leafhoppes cutworms spittle bugs leaf than we realize Candidates are Atherigona spp root beetles seed bugs ants and termites aphids root weevils crickets leaf beetles thrips and seed The least dispersive species are white grabs mealybugSbugs Insects known to entervarious states of aestivation are seedling maggots crickets weevils and gall midgesleptocorisaspp adults (199) Maliarphascparatellalarvae Each species has evolved a unique set of attributes to (9) Scirpophaga innotata larvae (117) Chilo partelhtsv enable it to survive and adapt to the changing uplandlarvae (21) ScirpopThagainccetulas (179) white grub larvae environments Several evolutionary avenues lead to fitness (134) Patangasuccincia adults (6 82) and Schistocerca (Table 2) but there is no reason to believe one set of gregariaeggs (82) attributes is better than another (224)

Insects known to enter hibernation are LodeIphaDstriatellus nymphs (124) Chtio Tvlpressalis larivae (125) DroughtSesaia inferens larvae (146) but not S calavisis(2) Well-drained rainfed upland soils are subject to drought ifIDiaraea saccharalis larvae (75) Scirpophaga- tcerlttas rains do not fall within 2-3 wk Prolonged drought followedlarae(5 lacchrli s lae (75)saSlOhagasshoperseggs (82)and felo pidas athias pOpae (232) by favorable rains stimulates armyworm and locust outshybreaks but mealybtigs root aphids and thrips become

Lcpmotorisa oratorisand L acuta adults aggregate on numerous onlyafter dry spells for two reasons First heavy trees sugarcane or other shady moist sites L paluwanmsis rain normally kills soft-bodied foliage-feeding thrips and confined to Sulu and Palawan Philippines lives on the mealybugs and soil-inhabiting root aphids and root mealyshygrass Brachiariamutica (Forsk) Stapf in the absence of bugs Drought eliminates this source of mortality Second rice The adults can live for 5 mo When aestivating their the rice plant responds to drought by breaking down rate of metabolism drops and they do not feed They will proteins into soluble N compounds which enter the phloem however puncture plant tissues of their aestivation host in and are taken up by these sap-feeding insects (245) Greater search of water This behavior has led to errors of the host nutrition therefore leads to greater survivorship and range of rice bugs (I) Aestivation quickly terminates with fecundity rain probably meaning aestivation is not deep Small rice area

Some stem borers aestivate as mature larvae in rice Except in the highly mechanized large landholdings of stubble or in straw stacked as livestock feed However Latin America upland ricefieldstypically are patchy within moriality increases proportionally with the duration of the a highly diverse flora The small fields result in higher ratios dry season even if the stubble remains undisturbed If the of perimeter to area than with the typically larger lowland stubble is plowed for a crop following rice few stem borers ricefields Upland ricefields themselves may be intercroppedsurvive Rainfall terminates dormancy but it normally with cereals legumes or root crops Mixed intercroppingtakes several weeks for moths to develop and emerge with a wide variety of species also occ in tribal slash-andshy

Last-instar white grub larvae tunnel 1-2 m deep in the soil burn agriculture (50 74) to construct pupal chambers in which they aestivate The Upland ricefields tend to be small because of the labor larvae are very sensitive to dryness so pupal chambers are needed to clear land in slash-and-burn areas and to till land sealed to coierve body moisture Aestivation terminates to minimize weeds Rice requires more tillage than does with the first heavy rains but as in stem borer 3-4 wk pass maize because rice competes less well with weeds Uplandbefore adults develop rice is more a subsistence than a cash crop and small areas

Insects may respond differently in each region to cold (I ha) can feed a family temperature or drought Biotypes or loal populations may Small fields favor some insect pests such as some ant evolve (56) Local populations of Scirpophagaincertulasor species that can forage from more permanent field borders Patangasuccincta may have become adapted to prevailing Trees provide food and mating sites for white grubs (134) as conditions well as aestivation sites for LetTocorisa spp (199) Field

iagiliti The ability to disperse combined with high border grasses plus tIhose within a ricefield provide alternate fecundity orshort life cycles enable some upland rice pests to food for all but the highly monophagous upland rice better exploit the temporarily favorable upland habitat colonizers and a habitat for natural enemies

0 z

Table 2 Life history patterns of upland rice insects or insect groups

Host range Longevity Dormancy Vagility FertilityMonophagy Oligophagy Polyphagy Low Medium High Yes No Low Dispersive Migratory Low Medium High(Oryza spp) (grasses sedges) (angiosperms) (lt3 wk) (gt2 mo) (lt1 kin) (lt50 eggs) (gt500 eggs)

Ants xTermites x x x

Mole crickets x x x xx x xField crickets x xx

White grubs x x x xx xRoot aphids x x xxMealybugs x

x x x xxSeedling maggots x

x x x x xLeaf beetles

x xx xArmyworms x x xx x

Thrips x x xx xChilo spp x x xx xMaliarpha x x x x

xLocusts x x Xx x xGall midges x x xxiVephotettL iirescens x x xx x x xLeprocorisaspp x

x xLeaftolders x x x

Sogatellafurcifera x

xx x x x x x x

00

IRPS No 123 January 1Q87 19

All this appeals to favor insect pest buildup but the notwithstanding the release of stored nutrients in response contrary is true Loevinsohn (138) showed that insect pest to drought that temporarily benefits philoem feeders populations respond exponentially with the proportion of land devoted to rice up to about 75 of the area then the M POR ANCE I I P1A 1) RWI P1St S

rate of response declines In most upland rico areas tie rice crop occupies less than 50i of the area- therefore the Visits to upland ricefields normally reveal 1kw insect pests potential for population buildup is low This relationship Loevinsohn (138) and Locvinsohn ct al (139) compared may not hold for seed pests that can locate small ricefields insect abundance using annual light trap atchcs in upland

Another outcome of the highly diverse flora and phical rainfed lowland and irrigated lowland sites in the Philipshyenvironment of upland rice isthateach upland ricearca has pines Although only one upland site was studied data a unique composition of insect pests showed equal or fewer insects at the upland site than at the

For example in Tanauan Batangas Philippines the rainfcd lowland sites and fewer than at the irrigated sites main upland rice pests are rice leaffolder(RLF) armvworm (Table 3) M Lltiple regression analysis showed that the key Aythntnaseparata WBPH ants Soli psis gentitataand factors explaining insect abundance among a set of cropping white grub Leucopholis irrorata Sixty kilometers away in intensity variables were the number of rice crops grown per Real Quezon the main pests are seedling maggot Athe- year followed by area devoted to rice Table 3 shows that rigona orae flea beetle (Thacitcmnra hasalis thrips the upland rice site had only I rice crop perym rand that rice Stenchaetothrips hiurnmis and rice bug Leptocorisa was planted in only 20Vi of the area Fertiliter and oratorius In a third Philippine upland rice site in Claveria insecticide use had little bea ring on the trend The armters at Misamis Oriental Mindanao the main pests are ants the upland site applied 60 kg N ha higher than at most Solenopsis gemtinata seedling maggot Alieriona orvzae irrigated sites white grubs Holotrichia in)iUaiuoan t and Lucopholiv Insect occurrence at the upland site was omparable to irrorata stem borers Sesamia ifiArelr (hio auriciliusC that at the Cagayan rainfed lowland site parlicularly in 1982stpessallis Scirpophaga incertulas S innoiata and when drought prevented many farmers fron planting(only

4c ona chrrsurapella root aphids Tetraneura niriah- 40(of the rice area eventually was planted that year) M ore dunrinalisand Rhupaosiphunm riahbdminalis lcaffolders favorable weather occurred in 1980 and 1981 which Marasniapataltis and Chaphalocrocis Medinalis army- supported greater rice areas and consequently more insects worm MIthinna separata and rice bug Leltucorisa The most intensive site for insect pests was South Cotabato oratoriusThe great diversity of insect problems is typical Mindanao where farmers plant 23-25 rice crops per year fer upland rice and makes control efforts more difficult with irrigation The Zaragoa Nutva Ecija site also had particularly regarding breeding for insect resistance high pest abundance and at this site the fields were highly

asynchronous Asynchrony was the third most important Low yield potential Iactor in explaining rice insect abundance Most upland rice is grown on low-nutrient and mineral- Rice cropping intensity in upland areas is bound to toxic soils which when combined with eiratic rainfall remain low Only one rainid upland crop is possible per make upland rice a highly riskycrop for the farmerto invest year (Farmers in Claveria Misamis Oriental Mindanao in costly soil amendments and land management Often Philippines tried asecond upland ricecrop i 1984a ycarof upland rice isgrown far from markets making inputs even favorable rains but no harvest was possible b-cause of more expensive Farmers growing a subsistence crop hosi- drought blast and birds) Rice area isaso bound to remain tate to invest in costly inputs to raise yields low because of the high labor and power rcquirelmenlts to

Even having varieties that could double existing yield prepare the land (because of weeds)and because upland rice potential under present management levels would probably wvill not compete with lowland rice as a cash crop Some not prompt many farmers to use purchased insecticide upland rice sites with prolonged rainy seasons howeverare unless tile risk of crop failure were reduced such as it was planted asynchronously such as the Clacra site with irrigation systems in lowland rice The seasonal abundance of Upland rice insect pests

The Tanauan Batangas site is atypical for upland rice attracted toa light trap wasgrapied for crop year 1980-81 in The soils and rainfall pattern are favorable and because of Tanauan FBatangas (Fig I 2) W13P11 was he most its nearness to Manila farmers have cash resources ftm common rice hopper and its population peaked at the sales of vegetables to purchase tertilier They applied an midgrowth stage as it does in lowland rice D-Vac suction average of 60 kg Nha after particle initiation and obtained samples showed an earlier peak than that from a light trap 25-35 tha yields The high fertility isone reason why RLF indicating emigration W13PI usually emigrate after the armyworms and WBPH developed into large numbers vegetative stage Populations in 1980 reached levels that Farmers however used no insecticide on upland rice caused patches of slight yellowing in some fields WBPH although they did on vegetables probably cause hopperburn in Batangas They immigrate

The poor growing conditions fOr rice generally mean year-round as shown by low levels in the light trap BPH poor growth and low fecundity for insect pests fIeding on it however were scarce in the light trap and not collected with

Table 3 Abundance of rice insect pests and factors in rice cropping intensity of Philippine upland rainfed lowland and irrigated lowland locations where light traps were operated daily for atleast I yr -I

0Rainfed Rainfed lowland Irrigated lowlandupland

Pangasinan Cagayan lloilo Santa Maria Nueva Ecija South Cotabato Tanauan Caaringayan Caaringayan Bangag Bangag Bangag Buray

LagunaCale

Batangas Manaoag 1980-81 Santa Motiica 1982-83 Cabanatuan Zaragoza Avancena NaniamanSolana 1981-82 1982-83 Oton

1980-81 1979-80 Oton 1981-82 1981-82 Koronadal Koronadal1980-81 1979-80 1979-80 1983 1983

Insect (kerosene light trapNephotcttixcatch nolyr)virescens 650 1240 500 1800 330 210 3400 1600 160N nigropictus 210 - 360 18400 5500 2900- 140 90 20 - shy 1700 -Reciliadorsalis 710 - 3200 1700180 890 630 140 170 2400 800Nilaparratalugens 130 690 - - 4200 4600300 520 750 180 60 3300 1600 740 310Sogatellafiircifera 1700 31000 13000 8200310 210 3300 610 270 980 1600 6300 -Cyrrorhinus - 12000 6700600 540 3200 290 160 330 120 410Scirpophaga spp 50 410 620 - - 21000 12000150 180 220 270 30 1000Cnaphalocrocismedinalis 70 70 540 610 270 11000 6100200 140 100 6 60 250Chilo suppressalis 0 60 60 1900 300 200- - 30 20 0 6 3 160 -C auricilius - 0 00 - - 20 3 60 20 3 0Sesania inferens 0 - 80 60 3 80 10 51 -Cropping intensity _ -

Rice crops (noyr) 10 10 12 10 10 10 13Rice area (7c) 17 19 20 20 25 2320 85 85 60 70 40 85 85 90Fertilizer (kg Nha) 60 20 85 80 70 7020 0 0 0 30 30Insecticide (no of 40 60 40 300 04 04 0 03 300 13 13 30 50 40 33 33applicationscrop)

__________

IRPS No 123 January 1987 21

Light trapped Insects Drytnd __ 2(notrap-nght) Suctioned Insects (nom )

50 -)o 9 jO40- SOW ffurciWO Light trp

D-Vc suction

20 to 4

20

Egg porasitizahion (M) 75 60

300 SOOampReOftwciffr

45 30shy

15

-insects (notrop-night) 4 Nileporvata Iuen5

Egg parastization()

Light trapped insects (no trop-nght) Suchoned insects (no m 2 )

6 8 Cytorh~nuls hvidipennii 0

4 05 2

50 0 Insects (noltrop-night)40-ui Ipanh pe g

Moohase 1 Seasonal abundance measured by light trap

Rainfall(etod) and D-Vac suction sampler of whitebacked

to ~ hopper Nilaparvata lugens and lea ffolder60)-t Cnaphalocrocis mnedinalis andplthpe g20 L I II I I I parasites and mtrid predator Ortorhjntus livi-Apr May Jun Jul Aug Sep Oct Nov Dec JaIn Feb Mar dipennis set against cropping pattern daily1 9 80o 19 8 1rainfall and moon phase Tanauan Batangas

1980-81

the D- Vac Cyrtorhinus build up on the crop coterminous are collected in light traps during full moon (109)with WBPH buildup Cyrtorhinus disperses during the Because upland rice shares most of the lowland rice pestsrainy season RLF not readily collected in light traps was and those pest species are generally less impotant to uplandprevalent only during the rice crop rice than lowland rice there is a general belief that insect

Nephotettix virescens was more abundant than Nnigro- pests are less important onpicLus in the light trap collections and on the crop GLH upland than lowland rice Arcview of limited data on upland rice shows an averag of population peaked at the reproductive stage as did zigzag 10-21 yield loss from insect pests on a par with that ofleafhopper (ZLH) Crtorhinus tracked the populations of lowland uee determined by the same methodology (132) planthoppers and leafhoppers Both GLH species were (Table 4) McGuire and Crandall (144) made a similarcollected year-round in the light trap indicating immigra- estimate for Central America Cramer (51) howevertion from nearby irrigated rice areas Possibly because of the estimated 35 yieldlow populations loss in South America based on ainsect number did not correlate with similar pest complex - stem borers and seed bugs shymoon phase although reports indicate that more hoppers attacking irrigated rice in North America This estimate did

22 IRPS No 123 January 1987

Light trapped insects D~~nd (no trap-night )

St plht Suctined insects (nom 2

)

D-Voc suction 1520

1o

0 Lih trap 5 ~105 o5

00 Eggparositizotion()5040 NephOtgtha virneens

30 20

I010 A-Insects (notrop -night)

4 Nephatelh nigropccent i

2 o I vV^J - -- ^ VA____ f _

Light trapped insects (no trap-night) Suctioned insects (no rnE

10 -1208 ReCdO doRsol

6 15

4 10 052 iwxALAiamp 0

Light trapped insects (notrap-night) 2Suc oned nsects (nom )

8 CyrtorhmnusI6o nn]10 i0075

4 05

2 0 0

025

2 Seasonal abundance measured by light trap Roinfall (mmd) Moon phase and D-Vac suction sampler of green leafhoppersNephotettix virescens and N ngropicrus and I zigzag leafhopper Recilia dorsalis their egg 60

parasites and micid predator Cyrtorhinus livi-dipennis set against cropping pattern dailyrainfall and moon phase Tanauan Batangas1980-81

40

5 Lf Apr May Jun

1980 Jul Aug Sp Oct Nov I

-aMar

1981

not account for soil pests and was not based on field data in the absence of pests Carbofuran granules are phytotonicAkinsola (I1) cited mainly data from irrigated rice areas in to rice (239) The prevailing hypothesis is that carbofuranAfrica but estimated yield losses of upland rice between 15 prevents soil bacteria from consuming fertilizer The phytoshyand 30 from insects However losses in Thailand over a tonic effect is through greater availability of fertilizer to the4-yr period ranged from I to 13 averaging 5(106) plant Denitrifying bacteria are known to be more active inYield loss data however should be interpreted with aerobic than anaerobic soils therefore it may not be merecaution (184) The Philippine data were all derived from coincidence that the highest recorded yield losses (22-69)trials on farmers fields under farmers normal agronomic occurred in those trials with 12-30 kg basal Nha On themanagement practices except for Bukidnon and Capiz other hand contro with carbofuran granules may not be aswhere researchers used fertilizer Batangas farmers are good as a seed treatment against seed pests and perhapsatypical because ofhigh levels of fertilizer used The trials in yields would have been higher if a systemic seed treatmentBrazil were carried out at an experiment station but insecticide had been used conditions were similar to those of local farmers The 1979 and 1980 trials in Batangas had two full-

Carbofuran granules were used in every trial as a broad- protection treatments one using carbofuran granules andspectrum insecticide applied basally with the seed An the other using bendiocarb WP as a seed treatment Plotsobjection to using insecticides to measure yield losses isthat protected with bendiocarb yielded higher than those proshyinsecticidescandirectlyorindirectlystimulateplantgrowth tected with carbofuran granules The other trials in the

Table 4 Yield losses to insect pests and yield responses to insecticide in upland rice in the Philippines Brazil and Upper Volta

Yield (tha) Yield loss (5) Location Year Cultivar Protected

with Protected

without Check Total Vegetative Reproductive Ripening

Basal N (kgha) Reference

carbofuran carbofuran Protected No with carbofuran

carbofuran

Philippines Pili Camarines Sur Tanauan Batangas

Pangantucan Bukidnon

Dumarao t-apiz

BrazilGoias Minas Ge~ais

Upper Volta6

1975 1974 1974 1975 1975 1976 1977 1978

1979

1980

1979 1980 1979

1977 1978 1979

Bursiging puti Kinanda Dagge Kinanda Dagge Dagge Dagge Dagge UPL Ri5 Dagge UPL Ri5 Dagge UPL RiS UPL Ri5 UPL Ri5 UPL RiS

IAC47 IAC47 IAC47

14 33 11 36 35 31 30 27 37 26 40 30 40 42 46 35

19 21 09

--

-34 24 24 27 --

29 43 31 42 -

47 25

---

10 28 05 27 27 30 26 29 33 28 33 29 43 23 36 11

15 15 06

29 15 55 25 23

3 13 -7 11

-7 18

3 -7 45 22 69

24 28 35

--

21 -11 -20

4 -

-3

23 7

-2 -23 56

-

-

-__0

6 --

-

-23 14

5 0 8 2

22

-

--

0

-

--18 11

3 -10

4 10

7

--7 -

14 -

-1 6

10 -5

8 9 7

-

--

0 0 0 0 0 0 0 0 0 0 0

30 30 30

12 12 12

99 99

100 100 101 101 102 102 102 102 103 103 104 104 104

66 66 66

Farako-Ba

Mean

1977 1978

IRAT10 IRATIO

38 66

--

40 43

-5 35

21

-

10

--

--

--

- 242 243

Z

tJ

-w

24 IRPS No 123 January 1987

Philippines which had two treatments offering protection In the 9 Philippine trials where yield losses were measuredwith and without carbofuran granules relied on further by growth stage 10 of the yield loss (21) occurred at thecontrol with foliar spray and not on seed dressings The vegetative stage and 517 occurred at each of the reproshycarbofuran-protected plots had an 18yield los compared ductive and ripening stages Ibis result is similar to thewith only 10 with foliar sprayS This yield difference is pattern of yield loss in lowland rice measured by successiveprobably due to differences in protection against sown-seed treatments where the crop is unprotected at each growthpests (foliar spray do not control sown-seed pests) rather stage but proticted during the other growth stagesthan phytotonic effects became basal fertilizer wa used in However the insect complexes r sponsible for vegetativeonly 2 of the 10 comparisons stage yield losses in upland rice are entirely diftfrent than in

The recorded yield losses were not related to yield lowland rice The key vegetative stage pests in lowland ricepotential (Fig 3) One might expect higheryielding crops to arc aquatic whereas those in upland rice are sown--seedhave higher percentage of yield losses but this ws not tile (sown in soil) root and seedling pests This dift rence maycase One reason is that insect damage exacerbates plant explain why agricul torists have overlooked tile ilmportanccinjury from drought stress low yielding fields from d ought of upland rice insect pests Many reports of upland rice pestsstress would show disproportionate differences between in Asia ha Ve focused on major lowland species (46 108)insect-protected and -unprotected treatments Shoot and It isalso surprising that tei vegctative stage of upland rice root pests would be more responsible for losses from would record the highest yield losses becatuseseeral studiesdrought stress than sown-scd pests A second reason have shown that upland rice can readily recover from foliageconcerns seed pests which would have a greater impact on reinoval during that stage (117 152 165 188) Upland ricelow tillering low yielding varieties such as )aggc Low poscsses a hitgh obilitV to compensate for ca rly loss oftillering varieties nnot fill in the space created by removal foliage and will even be stinulated by foliage removal toof seeds oweer 1) R i 5 is high yielding because it tillersP1 produce a higher viel than plants ithout any foliageactively which can close the canopy to provide greater weed removed (63 192 231) [ie reason for this apparentcontrol discrepancy hovce is that yield loss at the vegetative

stage does not octUr frol leat arca loss but fron pestsYield loss () removing sown seed and feeding on devcloping tillers and roots -- damage that cannot be readily conpLtnsatCd for

70 - Measuring yield losses in upland rice cnvironrnents is further complicated by a report from Brazil that greater yield losses occurred in response to better insect control60 - =-09 as Insecticide seed treatments resuled in denser plant stands

(n=21) which in turn created a selectively more favorable environshyment for blast (69) This resuIt was parlticurla rlxexacerbated50 when cirbofuran granules were used in seed furrows Aside

ffrom a greater plant stand ca rbofuran iay have auigmented the N lecl and stimulated blast If so ca rbofuran should be40 - replaced with another insecticide or the seeding or fertilizerI rates decreased

Other methods to determine the impact of upland rice30 shy pests wvcrc to interview farners or to develop single speciesbull bull correlations of population levels and yield loss Feneira (66)2 NIM)st (75(1) Golas farmers reporteddid both in Brazil

20- insect losses in ufpland rice and 251 said those lossesS avcragcd 37(1 [his result is consistent with tie yield losses 0 0 measured by field trials in the same statc(Table4) For every

I0 4 of seedlings removC d by leaf-cutting ants a I 7yield loss is predicted Similarly Ferreira (66) found that ai

0 - average of 53 thrips per panicle before panicle initiation meant 3 times more unfillh d grains

-IO0 I 1 1 I I-101 0 2 3 4 5 6 7

Yield (tho) C)N [ROt IM FI IODS 3 Correlation between upland rice yield and yield lols caused by Upland rice insect pests may be controlled by culturalinsects measured in 21 trials in the Philippines Ivory Coast andBrazil genetic biological and chemical methods

IRPS No 123 January 1987 25

Cultural control dry season plowing will have little effect on them Plowing Crop husbandry to reduce upland insect pest populations when the soil is wet from the first monsoon rains will expose can be classified into practices effective in a single field and soil insects that tunnel close to the soil surface to predatory those effective only at the community level Cultural birds chickens dogs and even man (field crickets nole controls should come first in a pest management program crickets and white grubs a re delicacies in man tra ditional They have broad stable effects as pests have little pos- diets) sibility of overcoming them through selection of biotyps The greatest degree i1 inIscct control fron tillage comes (138) On the other hand some cultu ral controls work only from plowing soon after rice ha rvest when most soil insects with high labor or power inputs and connunity-wilt lie close to the soil surface anrd stern borers are in the rice methods require coordination of iany farniters stuble Whitegrubsare mature and tillage exposesthem to

Pantin time The luxury of being able to shift the time of predators Tirnelv control will prtec the crop following pla ntring upland rice is open onl hr la rniers in areas with a rice and lowr the populationIfor succeedinig years Plowingprolonged rainifall pa ttern lI copping intensity or underthe ice stulble helps it decompose killing sten borer rnechanied land preparatiot Gcnerally planlting as soon larvae inside as the rFiinv season begins will lower populatiois of 1110st [requent tillage destroys ait nests bit not termites insect pests Insect populationsarelo aftcra dlvseoll or Tillage and clean culture howes er rcliove food for winter fallow delayed plaiting lets them build tip on termites which then mtay attack a young rice crop Scarab alIternate hosts that gi vigorotsly with the first seasonial beetles also prefer to miposit in eceitly tilled fields Noshyratiris This first flush iof weds cros lItxuttriartlv with iceCult is dibbling faors the survival oftillace Clre Such rineralied N released during the dr season aid made stils i aels tnd insects thtt pttpate or lay eggs- in the soil

available to the plants with the rains It iliso igrtmitcally Zero tilla hio eerl conserves predators ofthe GIH (37) advanttgeous for the rice ciop to tap this trlLItlaI fertility nt it mcttoi lo lessen the impact o0 soil insects and grow vigorotsly to compete with wrus and to tolerate brotdcasting seed or planting in ftltrrows is prefc ired over pest infestation dihlirig in hills Crickets are large enotlgh to eat seedlings in

Larly plantngs terd to escape seedling maggolts Upland a hill atrd tillering normnally ca iriot fill in tlhe missing space rice in Bitangas is planted in Ma and ine itli the Iirst Oftcn dibbll rd is not well covered and iseasily found by rains Fairners are highly mtotiated to plant early tocscape foraging insets ioIdents and birds drlitght and typlotns ricar harct Arn Atrgtst planting Plan uit sii Incrcatsini sced density protects tile cropwould be evcely attacked by It riim orivzo At tti againrst seed and seelling pests especially ants particularly other Philippiie tipland sites Rea l Qteot antd (laelit if the crop will germinate and ermerge quickly M isainis Oriental seedlig mnaggots arnrige ric boe t se hr trftinll itI India rice intcrclopped witi cotton or farnes have a longer growing season and plnit 2 rin or pigeon pci had lower 6IIt and WBIlII poptrlations than mote after the ortset of the rainy season liaaiangas ftrmners rice alone (211) NfailC arid Urplald ir_ 2re coilltori in the plant ca rly becia use ofa shorter raiinfall period arnd the desi re Philippines he chatngC in pest status folii iniercropping is to lliarvet before the typhoor sea son peaks ltighly locatioll-specific (133) cmianarid the net elfc be either

Early plantings cornbind with early-iatiuring varictics nil bcnelficial ortdetriential (189) For example BIatangias provide higher crp tolerartce for white grubs Aestiatior farmers intercrip Itro in their ipland ricefields A liornshyterminates with the first rains but 5-6 moare ineeded for the nrii iotd-Ahis i (I iiiacius) which norlially feeds resting larvae to develop into beetles emerge and lay eggs on taro ill feed or miarbv rice plants Intercropping and for larvae to reach the danriigii g ird-instar stage probably would not afft0 seed pests A crluanion crop (134 In lataigas this normually iseiough tiite for harvest planted ith rice and haing gleater tolerarnce for root

It Brazil howecver delayed planting is a suggested dariage or being toxic nayact isi trapcrop to root-feeditig cultural control for the lesser cornstalk borer and insect pests Acial-fei i ig pests voultId be most affected by pests favored l drought (66 195) Cropping internsity in intrer ropping Shoot aphids leaf beetles sten borers and Brail is low and delayed plraing places the rice crop in the hopplers would be likely ca ntidates to note effcts On the niost stable rainfall period reducing likelihood old rought othlt halnd the corlipalior crop stch~ sitniaie Iiiv be more

In rilniy arca s rinmirsare roi atl to plant other crops benehited than rice from initecropping ( 137) before rice and althNgh it woultl be desirabh to avoid 11echi Weeuling luring the first month after clop infestations by planting rice cuatl lariners lack the labor cstablishent will force pcts such is anyvworris which and time to do so ol a low priority crp prefer grassy cetds ontor tle vorling rice Soil iampsecs will

77age Uplild rice Stils ae light textured and Call be mvnye to the rice crop Cleani citrltirc also lay lore territes plowed duiring the d rv seasor notl only to grin time for to attack I u cetincti inrice ceds i ay he le optinal planting with lie fitst Inilsoon riiains but also to desiccate solition 13) soil insects arid weeds such is ntsedge that foim rhizomes Irtlizr Itneatsing N Iciiliier oin hlwland rice has Htowever becta use most soil insects lie below die plow layer fAtwvored Irigher populains of pliithoppeiv Imlhoppers

26 IRPS No 123 January 1987

and leaffolde rs However because of the need to temper fertilizer use on upland rice o prevent blast fIrtilizer rates will be lower than is comonia r for lowlad rice causing relatixely less hopper and IaIfTfoide r population buildup However inupland rice areas without a history of blast such Is Batangas farmers apply high rates of N and conscqtrently the crop has high infcstation levels of WBPI]and RIF The fhrmrcrs leec insect buildup by splitting the fertili11r in0to two to three applicatiois bit do not applV fertilizer at the vegetati estage because this prodrces a tall crop which illeasily lodge during typhoons Greater crop fertility leads to greater insect survival (better nutrition) larger insects (to cat no re foliage) arid greater fec titd t (to lav nmre eggs) (128 145 171) Not all insect species respondequally hovr Increasing rates (if N l anrid K increased Dialraa s-acchralis and (iaeiurema flea beetle but reduced thrips (66) Zn rcdLuced deadheart density from the lesser cornIstalk borer

Fnldingi Although most Upland rice areas do nIot hawxcaccess to wa ter floodiig the fields is rRrrnriierdcd in areas of Brazil or root bug and white gib control ( 195)

Crop rotatioti Tillage before planting a crop after rc harvest will control stern boreis and other pests renain n the stubble arid will unearth soil pests Planting a non-gratriiaceous crop after rice is recommncrded for termite control (66) Crop rotation is less nttdcd in upland rice culture than in lowlands because atcrop-free period is assured even after harvest of crops following rice

t1ulchilt iryl plastic irli irig designed for weedcoitrol ii upland rice inJapan resulted in ntr highanri liy infestation of (ihilo sulytresvais (93 9-1 95) NIulched Upland rice grew more luxuriantlv and attracted more (Avipositing mothis than u lched rice Also the rice plant and stein borers imatoured 10rC quickly and a partialgeneratiori af vouing ista iNoverwintcred in the stubbleLotili--malitrt varieti Quickly maturing crops reduce the tmbCr of pCst generations that Calt build up Rapid crop establishment olan ca rl v-natu ring vaitvty if)pa 11icLa r will lessen white gib and stem borer damiage White grub larvae intle last iistar are highly destIruCtive so aii early-iiatri ng variety will escape serious root loss Stem borers build 1ip slowlyx in rice to ecormic abundant in a late-Maturing crop anrd caiuse whi tlieads Numibers of somespecies increase exponeitially with each succeeding genera-ti so a xarietyexhat atures I trowh earier thian another will hac less insect damage Early-naturing vaiicties however will not lessen seed pest daiage either ii the soil or oina standing crop

Strwhoi hAIJttam Inrscts tcd irig on thlie aerial portions of rice plats disperse from carly to late plantedfields AS iiiibCirs increase exponentially with eachinsect

gtscnrattm e eiglbrig f rmc~s1(uil upp wh rc generation pests build tIP where neighboring fihrTC e9r1sstagger their plantings II Batangas farmers plant uplandrice within I iro use 120-d s-arieties and plant at the start of the rainy season Ini Misarnis Oriental laricrs stagger their

plantings up to 3 mo apart and use a set of atieties that mature in 4-7 mo Insect pest danage particularly from stem borers is greater in Misarmis Oriental Also seedling maggots damage rice there because rainfall favors frequent planting of maic In Batangas rice escapes seedling maggots bCca usC of distinct drs season and synchron0us early planting with the onset of ains

Therefore combinations of cultural practices ea rlyplanting synchronous planing crop rotation and earlyshymaturing vaneties protect the rice crop against most insect pests

Ilmant resistance Beca tse upland rice has few specialized insect pests and many insect types attack the crop worldwide regional breeding and strong internatiornal cooperation are required

Sown-sced pests arc economically controlled by insectishycide seed treatment and should the relore receive lowxpriori ty inbreeding objectives

Root pests are normally cost ly to control with insecticides but finding resista t sonres mayvbe di fficult Tanaka (229) for exa -ple fi led to find varieties resistant to root aphids Selecting for large root bioniass perhaps should be the strategy for pests sLuch as Whlite grubs that renliove roots Fainter(174) howcvcr cites varietiesof crops tolerant ofor resistant t(i root pests Some sorghun varieties are high in cyanide so rice sorghun might incorporate broadshyspcctruri chemical resistance

Seedling vigor and drought tolerance are high prioritybreeding objectives so tupland rices should have large r root systernsand be able to tolerate ligherlevels of root loss Sonie root loss inolder plants is beneficial if it stimulates new root developnrut and therefore crihances intake of soil nutrients (53) The rice seedling inaggot rtiLotia spp may be

controlled thirough resistant aittics A hi rgc breeding effort on sorghniu has had fair success inideveloping arreties esrstait to Asoccata Rodani (251) A field trial comparing 10 upland ricc varieties showed differences aniong varieties va ring from II to 4(- damaged tilleis (215) Hovver resistarce to 1Iill-0r gtia species may rot cross ox-rti otier

Shiraki (210) reported that upland i-ice stens wcrc harder than lows-land rice stemsperhaps becaIusc (f a higher silicoricontent and thus were more resistant to yellow stern borer hc tortia Iow tillering (f uplaid iice xarneties prodtres a

higher percentage of infested tillers than would occor in higher tillering lowland iice under a siiilar egg density The corollary therefore is to select for high tilleriig upland rices

Stemii bor amcstheh main targes dobrecders inAfrica (571 204 )but nticl oifth e sc reen irig is dotie unde r lo wland co d t osrm s wih A h r~ o l pe i s n acorditions (11 96 218) As with lthrinrnwspecies mayhave to be dealt with independently however initial results show cross resistance between African and Asian species(92)

In Brazil screening Ior insect resistance focttses oil lou I pest problems Varieties tolerant of lasmolwalpus ligi-sellus and resistant to )iatraea saccharalis have been identified (70 153) Current work also emphasies 7iraca limativetntris and Thmialspis (=Iosis) flavouicta (66) These insect pests Ilt onlV citise IIigh yield losses tihey also ire difficult to control chemically ofr by other ilealls

ManV upland rice vaieties aic early maturing and are resistant to pests such as gall midge (I Xl ) Early mIturity is often a highly desirable trait to esuilpe pest buildup but it is not true resista nce bca usc ilthose va reties are plinted late they will be itsdamaged as ain early planted susc ible variety

P~erhaps tile cosiiiopolitall of all upland rice pestsimost are the sced pests (ripenling stagc) which pose a particular problem beta use scuds re ised IsIO for hitnit4ans Ayltoxic substatice introd tcld geticticllyilmay also be toxic to nal iniless it caln he placed old in thfe IdhuffSelectmi fo hairiness or lotng wns kill mIlct sthilrohleiiis dutilig threshing pa rtictfarl maitial threshinfl [hireshers ill complain of itcfimin atid itteascd (fust levels

Attempts itIRRI to fitid tcistatncc to the rice bug plorisa have ilotbeCnl prmitisilg lIIetIattonlids iorc

through the lenilna or palca hillhairs sedstVIor seeds wilh awns that deter bugs promote complaints from threshers Seed bugs liptocorisa and Stcnocoris enter the seed through the opening between tile lemna and palC but varieties with narrow openings are diifficult to mill

Breeding priorities should concentrate ott inotCI gruLtLIps that are most difficult to c nttrol by other meains Steil) horers appear to be the most widespread group that sigiiificantly dalag s rice aind is hard to manage by other Ilnca us

Biological control Very fe stutdic of mitatt~l eneties of rice pests in upland envirollnletits wee in tile wefouiid literature Therefore have not compiled a list of species Ve present a list of natural enemies iOtid in upland ricefields tuic latialln Batangas for some conitno iOldiar pest species ([itble 5)

Muich ofl thc lollowinle itfor ittiol cotlliCs rol Stutdies in the lowlatnds or roi crops otithcr thlin tice

Qu(ailtr o 11ll t11t( Nat trrtf lii hcoeits ight Ile used agaitist ile social insects tilt and termites Sollie predator specice live ill theirf nests n ilnic theirnllt ip-pearanIlCe aMRidbehavior IJtrouMdCing a mote aggressise ant species to displace pest ant codonies is a possibility (169) Patihogens as potential natural encnies have reccived little atteition (I13) Natural ncnuics asa group however hac not been toIl iporta it in Iegulating aintshown be and ternite numbers

The nocturnal behavior aid subteitmien habitat of lole crickets and field crickets protect them from many ntllral enemies There are only limited records of natural cnelics for these groups of pests some pathogens (87) but mainly

IRPS No 123 JattuIny I 271987

parasites - scelionids on eggsand sphecids on ilymphs and adults Sphecid wasps Larra carutmaria (SImitI hI L huzwuensis Rohwer and lsaltiuhta Willi ams speciali in mole crickets pa ralyzing them nd dragging the ninto thcir nests as food lot their voting ()tier sphCcid species Lins aurulenoa (Iabricius) lot maiac (Ashmead ) 1 lhshylssellaus (Smith) and Ml lal)oriosu (Snithf specialie in field clickets

White grUbsO the other hand hae Icti scriuns enuigh to have been extensively studied for cotttroi 1W nati al enemies( 112 140 141) Scoliid parasites C(ajsoMcispp have beetn intrtduced itto tilePhilippites for control of Ielxl ]udisiro la The rcsearchcts w Intttrdced themle during ill era claimed that the white gttionthreak as controlled (142) howevcr it isnormal for pest epidemics to subside and there was no diect evidence to indictte that tfe parasite was responsible A tmore promisitg titctllod might he to identify iso ltcttass prt~f tcartd apply hiactelaln fingal pathogens to tile soil Ihe lalattese beetle in North America has hcci coitrolled hv tite conmnleciall aailable hactcrial prcpa ration itacihlpu pav ofr milksdiscas (71) Spote prtyefott the soil itloted ilrtier reitiain actic fr Veas

Root aphid sardtnctly bugs readilslic tilib to predatots and parlsites once their tetndine ants are controlled The larvac and adtilts of a coccinellid beetle 1Scynms sp prey on llriadlouais in slash and burn upland rice Baits treated with insecticide may be used to control tending ants by killing their ourig (244)

Root-tkcditig blugs false wireoris wireorms and root wccils aic r ofsuch little ifilpoltatice that thenomttll role of liatural Cticniics hias tiot beei asscssed A possibility hoevrl tiglt he tit ItaIIitic niaCIMtodcs lintt1g11en Itese soil-itlhlfbitine mCtitodcs W5uhtfhave to be imiass Produiced

large miltibcr of niatltrll cnemics ircknown ft CltIoriS IC IpblCeii is thtt clttvortnlls ofonlie the crop soott alter land ptel)tritiOti a1tithe bcgit iniig of the raittv

Citsoln hfien tiatllral leniiiy populations aric lo Colltloil WOtuld iii ole mass prodtticing andirclcaslng key spccies

le egg stage is the fpirt (olthe Cedlitgt Ialot life cycle tI hich to f)ctis ftor tittlril elctit cottntrol hplraites o1 pleditors ttsit elleggs highly expoIsYCl I Sgg pailsites or predators stolld probahly e ittass andtbe produced released Littlh i toruialioll is a ailable ott what species to trV o ho to Iniss podLice thiCIi A etiloplhid wasp Icmihlaol sp effctiicly regushy

lates the leatf lcloalapolm2a spioanoiiitr heat andc rice antid should be collsctvcd

leaf beetles whCllose e am] eggs ate laid in the grutindhirO ould be motC difficutlt to control by niatulral enelV Mnll ipuhltiOn But species sUici IS c_gsthispid beetles ith

and larVle oilleascs ate vtilcratble to iparaites and pledators

Natural enemisc of tlirips otl rice are little known

28 IRPS No 123 January 1987

Table S Parasites recorded from upland rice insect pests in Tanauan Batangas Philippines 1977-84

Order Family Species Host stage

I lymttenoptera Braconidae Leaffolders Cnaphalocrocis Marasinia spy

Cardiochilesphilipplinensis Ashmead a Larva

Cotesia angustillasis(Gahan)Cotesia or cypris (Nixon)

Larva Larva

Cotesia or taeniaticornis(Nixon) Larva Cilonusinotakatae Munakata Chelonits SppAlacrocetrts philippinensis (Ashtmead)

Egg to larva Egg to larva Larva

Iclineumonidac Orgilts spp Isclutojppa hireator(Fabricius)

Larva Larva

Itoplectis naraigav(Ashmcad) Larva Tenelhtcha philippiensis (Ashmead) Larva Tenelchha stangli (Ashmead) a

irichomma cnaphalocrosis Uchida Larva Larva

Bethylidac Elasmidac

Goniozms or trianguli r Kieffer lasmtusalhopictus Crawford

Larva Larva

Diptera

Encyrtidac Chalcididac

Tachinidac

Elasmtus spp Copidosomopsisnacohiae (Fidy) Brach nteria evcarinata Gahan Biachyieria lasus (Walker) Ztgohothria ciliata (Wulp)

Larva Egg to larva Pupa Pupa Larva

Argyrophylx nigrotihialis(Blaranov) Larva

l-ymenoptera Braconidac Yellow stern borer Scirpopiaga incertulas

Braco chinensisSzeplegeti Larva CIthelotis spp Chelonus niunakatae Munakata Cotesi flaripes Cameron Stenoracon nicerilli (Bingham) a

Larva Egg to larva Larva Larva

Ichneumonidac Tropobracon schoenobii (Viereck) Amatroinorpiaaccepta metathoracica(Ashmiead)Frihorus sinicus (I lolmgren)

Larva Larva Larva

Ischnojoppaluteator (Fabricius) Larva Isotima nr dammeratiti Rohwer Larva Tenelucha philippinensis (Ashmead) a

Larva Tenehicha stangli (Ashmcad)Trichomma enaphalocrosis Uchida

Larva Lara

Chalcididac Eulophidac Trichogranmmatidae

Xanthopimnpla stemmator (Thunberg) Charops hrachYpterumn Gupta and Malteswary Brachtm eria sppTetrasticttus schoenohii Ferriere Trichorigranm chihunis Ishi

Larva to pupa Larva Pupa Egg Egg

Diptera

Scelionidac Pteromalidae Tachinidae

Tt ichograintnta aponiicuitt (Ashniead) Telenomits rowani (Gahan) a

Trichoinalopsis apattelctena (Crawford) Jeirhaea spp

Egg Fgg Egg pupa Larva

Z4gobothria ciliata (Wulp) Larva

Ilymenoptera lecneumonirlac Gol fringed borer (hilo auricilius

Trichomm enoptalocrosisUchida Larva

Ieioenoptera Braconidae

Aantholimlh steiniator (Thtinberg) a

Pink stet Ioret Sesamia inferens StoInacoil nicrilhi (Bingh am)

Loarva to pupa

larva

Diptera Ilyinenoptera

Armvtnyorms Mythimna separata Spodoptera mauritia acronyetoidesTachinidae Zvgohothriaatropiora( Robinea i -Desvoid y)Eulophidae It-phctrtiscitapadoe (Ashimead)

Larva Larva

(halcididae Braconidac

Brachpin ria spip (oesia spp

Larva larva

IHyntenoptera I)iptera

Rice skipper Ielop ida (halcididae Tachinidae

math ias aidgreen iortedci ciliarNIelan itis eda Brat lhincria sp iii matuqitatt Cameron Irgyrolhirlaxi trotlhiais(marumtv)a1

ismtne Larva Larva

Diptera Ilymenoptera

Tachnidae lelineutionidae

fuown se tilo per Mois frugalisIrgtrophrlax 1igrothiahiv 3 a rat ov) anthopimpa pmnctata (Iabricius)

Larva Larva to pupa

aMost dominant

IRPS No 123 January 1987 29

Normally predatory thrips prey on herbivorous thrips strepsiptera on both Spiders dominate tilenymphaladultNatural enemies of stem bugs --Scolinoplara and Tibraa predators Spittle bugs move to rice from pasture grasses - are in order of importance egg par~isites and predators and biocontrol would be successful only ifcarried out before fungal pathogens and nymphal adult parasites Egg tiley immigrate into ricefields parasites could either be introduced fron other areas or Biological control tactics for hemipterous seed bugs mass produced and released ould follow those outlined for stein bugs

Armyworms are normally lwld in hcck by the activities Coml)arison of environnewtns Natural enemies are of egg and larval parasites When these natural enemies fail perhaps faced with even greaterconstraints than insect pestsusually because of drought armyworns become epidemic to survive ind reproduce in upland environments TheyIt may be worthwhile therefore to release parasites during have adapted ani survived by having wide host ranges the the rainy months following a drought Virus diseases of ability to aestivate or hibernate over unfavorable seasonslarvae would beanotheravnue to exploreforarnyworms and the ability to disperse or by other mechanisms

Parasites have ben traditionallyconsidered for biolegical Comparing the most dominant and therefore the most control attempts against stein borers usually by rekasing adapted species of natural enemies collected in various exotic species( 166) This approach has not met with success environments in the Philippines we can understand how and new avenues should be explored The role of predators different rice environments are arid pathogens is little inderstood and deserves greater Anagqrms olptahiisa mymarid planthopper egg parasiteattention Egg parasitism rates are normally reasonably is cquallyadapted to all Philippine rice environments (Tablehigh but should be supplemented with effective egg 6) It is selective for planthoppers and even parasitizes eggspredators Orthopterans are potential egg predators of the n-iize planthopper Peregrinitsmaidis itt upland areas Metioche vittaticollis (Sta ) Anavipha nifiwnnis (Saus- Alavohis also parasitizes maize planthopper eggs but is sure) and Conocephahs hongipnni (de i-laan) feed on most prevalent on BPH in the irrigated wetlands (onashystem borer eggs etioche and Anaxil)ha specialize in eggs tocerus spp like A optalhilis on BPH and WBPI Iattacks with no hair such as Chilo spp The omnivorous (ono- the eggs of GLH in ell rie environments It is also found cephahsspecializes in eggs cov red with hairs it is a proven parasitizing the eggs ef the white rice licahopper (oimna egg predator of Scirlouphaga spp but has the discouraging spetra Oirosita naias a trichogrammatid egg parasitehabit of eating rice guin (65) attacks BIPH and WBPH in the wetlands and has no other

Grasshoppers have egg (scelionid) and nymphalad ult host A related species 0 aesopi specializes in G1-H in (nemestrinid tachinid and sarcophagid) v irasites Among wetland environments Dryinid strepsipt-an and pipunshypathogens protozoans have been recorded most frequently culid nymphal and adult parasites of planthoppers and Control of locust species by natural enemies could con- leafhoppers also specialize either in hopper species or centrate on their habitual breeding areas environments Among the dryinids Pseuhdgonatoipsflavi-

Leaffolders have rich complexes of natural enemies some feimur prefers the lowlands and BPH while P nuch has no adapted to upland rice (22 27) ranging from egg predators environmental prefirences but is adapted only to WBPH (gryllids coccinellids) to larval parasites (braconids ichncu- The main d ryinid species on GLH aptogonatop)usspp is monids) to larval prcdators (ants and carabids) to larval most dominant in rainfed lowlands and also attacks other pathogeiis (vira and fungal diseases) Efforts to augment leafhoppers within Amnrasca Cicadulinaand Bachaltuathese natural enemies should focus on the egg predators Strepsiptera are more dominant on plarithoppers than (rearing releasing)and viral or fungal pathogens (culturing lealhoppers The Elenchus species in the Philippines attack spraying) all species of planthoppers within the genera Sogawilla

Parasitization of the large conspicuous larvae of Sogatodels NilaparsataHarualia and Opiconsiva Elenshyskippers green horned caterpillars and other polyphagous cusrvasuntaisuiis adapted to the major rice environments Lepidoptera by tachinids and chalcids is normally low ELg but prefers BPH in rainfed arid WBPIH in wetland environshyparasites and predators are perhaps the key to their control ments Ilalictoph(utsmtnroeiand HVpectrus attack G LH

Gall midges are parasitized by pterormalids and platy- and COfimravcctra respectivelygasterids that aestivate within their larval host between Pipunculids are mainly parasitic oii lealhoppers Each seasons Shoot aphids are preyed upon by coccinellids species has a unique environmental preference Tomosshysyrphids and chrysopids varireIa oraetoraprefers Ne7 otetltix nigropictrs and

Planthoppers arid leafhoppers have egg parasites and Pilunwulhtsjavanensisattacks Dehoceplhahls spp which predators as well as nymphialad nlt parasitesand predators are more abundant in upland than irrigated lowland their rich complex of natural enemies includes fungal environments T suhvirescensattacks Nephotettix virescens pathogens The principal egg parasites are myrnarids and and prefers the lowlands Pilpuiulus nttillattls is an trichogramniatids Egg predators are norinally mirid plant upland rice parasite arid attacks only Nephotetix speciesbugs that suck out the yolk Nymphal ad nIt parasites are More than 17 recorded species of leaffolder parasites in dryinids on planthoppers pipunculids on lealhoppers and the Philippines attack species of C(naplwocrocis and

30 IRPS No 123 January 1987

Table 6 Dominant natural enemies of rice pests in upland rainfed lowland and irrigated lowland rice environments of the Philippines

Natural enemy

Host stage Species

Brown planthop-Parasite Egg

MymaridaeAnagrus optabilis(Perkins) Anagrusflaveolus Waterhouse

TrichogrammatidacOligosita naiasGirault

NymphAdult Dryinidac

Pseudogotatopuslaiifemttr Esaki et Hashimoto Elenchidae

Elenchus yasurnatsui Kifune and Hirashima

Environment

Upand Rainfed lowland Irrigated lowland

Ailaparata lugens

x x X x

x

x x

x X X Whitebacked planthoppers Sogatella furcifera and Sogatodes pusanusEgg

MymaridaeAnagrus optabilis(Perkins)

TrichogrammatidacOl~gositanaias Girault

NymphAdult Dryinidae

Pseudogonatopusnudus Perkins Elenchidae

Elenchus yasutatsui Kifune et Hirashima

Parasite Egg

MymaridaeGonatocerus spp

TrichogrammatidaeOligositaaesopi Girault

NymphAdult Pipunculidac

Tomosvaryella subviresccns Loew Tomosvaryella oryzaetora (Koizumi) Pipunclusmutillatus (de Meijere)

DryinidaeIlaplogonatopusspp

Egg Copidosomopsisnacoleiae (Eady)

Larva BraLonidae

Cardiochiles philipphnensisAshnead lchneumonidac

Trichiomnia cnaphalocrosisUchida Temelucha stangli (Ashmead)

Pupa Chalcididae

BrachymeriaexcarinataGahan Brachymerialasus (Walker) Brachymeria nr tachardiaegroup

Parasite Egg

Scelionidae Telenomus rowani (Gahan)

EulophidacTettasticusschoenobhi Ferriere

Larva Braconidae

Stenobraconnicevillei (Bingham) Iclhncurnonidac

Temelucha phiiippinensis(Ashinead)Temelucha stangli (Ashmead)

x x x

x x

X x X

x x Green leafhopper Nephotettix virescens

X x x

x x

X X X X

X

X Rice leaffolders CnaphalocrocisMarasmia

x X x

X

X X

x x x x x X x x x

Yellow stem borer Scirpophaga incertulas

X X x

x

x

x X XAinauromorphaaccepta metathoracica(Ashmead) x

continued on next page

IRPS No 123 January 1987 31

Table 6 cntinued

Natural enemy Environment

Host stage Species Upland Rainfed lowland Irrigated lowland

Larva-pupa Gold fringed borer Chilo attriciiuslchneumonidae

Xanthopimpla stemmator (Thunberg) x aPupa x

Pteromalidae Trichoinalopsis apanteloctena (Crawford) ax

Pink stem borerb Sesamnia inferensParasite

Larva Braconidae

Stenobracon nicevillei (Bingham) x b b Armyworms Mvthinna SpodopteraLarva

Tachinidac Zygobothriaarroptvora (Robineau-Desvoidy) x x

Braconidac Cotesia spp

X Skipper Pelopidas mathias and green horned caterpillar Melanitis leda ismeneF-gg

TrichogramrnatidacTrichogrannaspp x

Scelionidae Undetermined Scelionid

x Larva

Tachinidae Argyroph lax nitrotibialis (Baranov) X x

Pteromalidae Trichomnalopsis apanteloctena (Crawford)

X Predator

CoenagrionidaeAgriocnenis femina femina Brauer x xIschnura senegalensis tambur

Miridae CrtorhinsliidipennisReuter x x

Nabidac Stcnonabis tagalicus (Stal) x

VeliidaeAlicrovelia douglasi atrolineata (Bergroth) x

Coccinellidae Micraspis crocea (Mulsant) X x x

Vespidae Eumenes campaniformis (Fabricius) x

Formicidac Diaecamna spp xOdontoponera transversa (F Smith) x

Carabidae Chaenius spp OphioneaW- aJasciataSchmidt-Goebel

x

Ophionea ishii ishii Habu x x X

Gryllidav Anaxipta longipennis (Saussur-) x xMetioche ittaticollis (Sial) x x x

Tettigoniidae Conocephalus longipennis (de Haan) x x x

aNo rearing made on rainfed lowland bNo rearing made on rainfed lowland or Irrigated lowland

MrasmiaAmong the Laval parasites the braconid Cardio- lowlands and parasitizes oriental maize borer OsIriniachilesphilijpphensis is most adapted to an upland environ- furnacalis (Guenee) larvae The leaffolder pupal parasitesment It also parasitizes Hydelepta indicata a common not only occur equally in all environments but also haveleaffolder of legumes The ichneumonids Trichomma cna- wvide host ranges- parasitizing species of OsIrinia Marucaphalocrosis and Temehcha stangli occur more in the Hydelepta and Honiona - in maize and legumeslowlands T stangli is most adapted to irrigated lowlands The principal egg parasite of yeltow stem borer (YSB)and also parasitizes Chilo and Scirpophaga stem borer Telenomus rowani is widely adapted to all environmentslarvae T cnaphalocrosis is most prevalent in rainfed and has an unusual alternate host -Tabanus eggs (25)

32 IRPS No 123 January 1987

Tetrastictu schoenobii specializes only in YSB and occurs mainly in irrigated lowlands -- further proof that YSB is most adapted to the wetter environments

Larval parasites of YSB are highly environment-specific The braconid Stenohracon nicevillei occurs only in upland rice but also attacks the pink stem borer (PSB) Tieinchwa phiiiplPwnsis an ichneumonid occurs in upland rice and also parasitizes the larvae of (7hiu stqpresaisinatro-morpha accepta melaihoracica like 7 stangi is mnost adapted to the irrigated lowlands

The dark-headed stem borer(l)HSB) (ih pochit3sus was not collected in all rice environments and the gold fringed borer Chilo auricilius nd PSB were collected only in upland rice Xaantiholiplastenrnaloran ichneumonid larval parasite appears to be widely adapted whereas TrihomaslopisapanitI)cteiiaa pteromalid pupal parasite occurs mostly in wetlands and also parasitizes larvae of Chilo suppressais and Pelhuidas mathias

Armyworms Myit hinin and Spodujipera have two main larval parasites - a tachinid Zrgohothriaatropivora in rainfed environments and a braconid Cotesia spp in irrigated lowlands

The butterfly pests Pelopidas and 4elanitishave tricho-grammatid egg parasites found only in irrigated lowland environments

Two larval parasites show environmental preferences --Argrroph av nigrotihialis a tachinid in rainfed habitats and Trichonialopsisapantelkcnela in irrigated ricefields A limgrotibialisalsoattacks the sweet potato hornwormAgrius convolvtdi and T apanteloctena parasitizes DH S1B

Predators as a group show more distinct environmental preferences The aquatic damselflies Coenagrionidae are most abundant in the lowlands Cvriorhinus is also most adapted to the lowlands The nabid Stenonabis tagalicus occurs only in the uplands and isalso prevalent on legumes The coccinellid lady beetles wasps and ants are more prevalent in upland areas AlicrasIpis cit-ea larvae prey ona variety of aphids attacking legumes and maize Etunienes campanonia vespid mud wasp makes nests in trees and is therefore most adapted to the more botanically diverse uplands Soil-dwelling ants cannot tolerate flooding

Upland environments are habitats to arboreal carabid beetles Three species of Chlaeniusbeetles prey on leaffolder larvae and are perhaps more important than parasites (27)

The gryllids and tettigoniids are widely adapted egg predators not only on rice but also on maize and legumes They feed on eggs of most insects that are laid on leaves

The spidercommunity of upland rice environments isrich in species At one site (Batangas Philippines) 31 species have been recorded (23) Of the 176 spider species recorded in Philippine ricefields about one-half (82) occur in upland rice The spiderspecies of upland rice environments overlap more with those of rainfed lowland than with those of irrigated lowland rice (24) The same study showed that of the three environments irrigated rice has (ie greatest spider species diversity foiiowed by rainfed lowland and upland

However of the 10 most prevalent ricefield spiders 9 were abundant in upland rice showing wide adaptation All three spider guilds -- orb-web space-web and huting spiders shywere prevalent in upland ricefields There were however differences in environmental prelerences for some spider species

Of the orb-wcavers Tetragniathamanlihulatawas partishycila rlv abundant 7 japonica was low and Ltucatge dehcorata was absent in upland rice

As in other environments Aiypeiwnaifirtosana was the most dominant space-web spider however its relative numbers were lowerthan for lowland sites Among hunting spiders Ox|opes javanus and Lcosa leatcost fgla were more abundant and Lvcosa lpseudoanmnlata was less abundant in upland compared with lowland rice envishyroniments

Extensive rearing of rice insect pests in upland rainfed lowland and irrigated sites over an 8-yr period in the Philippines provided insight into the effectiveness of parasites as natural enemies (Table 7 8) Most of toe data are from large sample sizes taken over at least one crop which overcomes the pitfalls expressed by Van Driesche (238)

VSB egg parasitization was surprisingly similar across all environments Larval parasitization was highly dynamic among sites and even years within the same site but showed somewhat lowerlevels in uplandsthan in irrigated wetlands This would have been predicted for egg parasites as well since YSB is adapted to the lowlands and so must be its parasites This isevidence that Telenomus rowani is as good at dispersing as its main hosts Trowani clings onto the anal tufts of female YSB and parasitizes eggs as they are laid This phoresy may explain its wide adaptation The more specialized lowland YSB larval parasites therefore appear more effective than their upland counterparts with more alternate hosts

Low collection levels of PSB larvae especially in the wetlands make comparisons between environments difficult Low levels of larval parasifization were recorded in both rainfed environments Except in the Cagayan rainfed wetland site in 1980 low levels ef larval parasitization were evident at irrigated and rainfed sitesalike No differences by environment were evident Higher levels of parasitization occurred in Sarawak during a 1967 outbreak also from tachinids Rothschild (197) indicated that higher parasishytization occurred in upland rice areas

Skipper and green horned caterpillar larval parasitization rates were generally low and showed no environmental effect htighest activity occurred in Cagayan in 1980 (48 parasitization) Although larval parasitization levels of RLF were variable year to year activity appeared greater in the irrigated lowlands than in the uplands Again the highest levels were recorded in Cagayan in 1980 Cagayan has extensive grasslands and fallow around the rice areas and a relatively long rainfall period Rainfall in 1930 was good for crop growth and apparently for increase of larval parasites

IRPS No 123 January 1987 33

Table 7 Parasitization rates of rice planthoppers and green leafhopper in upland rainfed lowland and irrigated lowland environments Philipshypines 1976-840

Environment and site Year Egg parasitization () Nymphaladult parasitization ()

BPH W13P GLH BPII WBPH GLIH

UplandBatangas

Rainfed lowland

1976 1979 1980 1982

15 17 12

2(47)8(39)

4(51)

9(1800)14 (212)

12(304)

8 (101)6 (127)

9(116)

Iloilo

Cagayan

1976 1977 1978 1981 1980 1981 1982

0 (69)

11

44 48 40

-

-

46 55 44

0 (471)

11

52 56 49

0 (200)9 (214)

10 (1259)8(235)6 (900) 3(800)

-7 (462)

11 (270)3 (313)5 (900) 1 (800)

4 (600)10 (725)10 (362)12(870)5 (900) 3 (800)

Pangasinan

Irrigatedlowland

1984 1976 1978 1979 1982

13 (436) 13(379) 14 (457)

6(14)7 (197)

17(198) 11 (486) 18 (275)

3 (81)10 (274)3 (116)

16(871) 19(88)

28(211)9 (764)5 (647)7 (1204) 8(305)

Laguna 1977 11 7 21 1978 31 1979 14 4 4 28 1980 7 8 20 1981 1983

56 7(278)

26 16 (1650)

16 27(1921)

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R I Apostol unpublished 1lo1lo and Pangasinan 136 Cagayan BCanapi unpublished Liuna 175 I C Pantua unpublished 118 234 43 Carino and Shepard unpublished BPH = brown planthopper Nila parvara htgens W131Pi =whitebacked planthopper Sogatelia furcifera GLH = green leafhopper Nephotettrix virescens Figiires in parentheses are sample sizes

Tauher et al (230) reported that internal external and genetic factors influence the seasonal activity of parasitoids

Hopper egg parasites but not nymphaladult parasites were also parlicularly abundant at the Cagayan site possibly the egg parasites have more alternative hopper prey in the weedy areas than nymphaladult paisites

The seasonal dynamics of hopper egg parasites can be seen for upland rice in Batangas(Fig I 2) and for rainfed lowland lice in Pangasinan( 135)and Cagayan( 105) Levels of BPHWBPI-L and GLH egg parasitization were corn-parable in upland and irrigated lowland environments Except for some high rates of paraMsitization early in the season the parasitization rate tended to be steady in all envilonments However hopper nymphaladult parasi-tizution levels were more similar in the rainfed environments and lower than in irrigated rice

Overall parasites do not appear highly effective by themselves against upland or even lowland pests Among natural enemies parasites have attracted the most attention because they can be more readily assesscd

Predators and pathogens lend themselves better to management practices Predators the most important group of natural enemies in riccfields are difficult to quantify On the other hand pathogens appear less im-portant but can be readily cultured and disseminated

The natural enemy commulnity of upland rice is rich in species and differs significantly from that of lowland rice

These beneficial organisms must be conserved by applying insecticides judiciously particularly sprays Seed or soil placement of chemicals minimizes exposure to natural enemies The strategy to derive the greatest benefit from natural enemies isto allow the greatest number of beneficial species to thrive Then perhaps at least one species will be effective against each pest at any tine overcoming the variable abundance of each species within and between years

Programs to introduce exotic species or mass produce indigenous ones are ambitious and expensive because they require trained people on a sustained not ad hoc basis

Chemical control Insecticides offer rapid and efficient control of upland insect pests (190) but-should be used only after other control measures have been considered Insecticides are rarely used on upland rice because of cost (32) Upland rice yields are normally too low to justify the expenditure Also most upland rice is grown as a cash crop Spraying upland rice is more difficult than lowland rice because water is less accessible Government subsidy programs to provide pestishycides to tribal peoples for upland rice production have created problems of toxicity to humans because isolated people have had no experience to be able to handle pesticides safely

However ins-eticide use has beenjustified when a crop is

0Z

Table 8 Parasitization rates of lepidopterous pests in upland rainfed lowland and irrigated lowland environments Philippines 1976-84a

Parasitization (5)

Environment and site Year YSB GFB PSB Earcutting Swarming Skipper Greenborned RLF Brown larva larva caterpillar caterpillar caterpillar caterpillar larva semilooperEgg Larva larva larva larva larva larva

UplandBatangas 1977 - 0 (57) - 0 (63) - 0 (71) - 0 (106) 11(142) 2 (101)1978 43 (4697) 8(451) 5 (38) 3 (40) 6 (61) 2(58) 15 (57) 13 (129) 13 (1286) 4 (30)1979 59(4135) 12(320) 13 (11) 5(28) 9(45) 6(46) 7(23) 8(160) 9(1152) 8(26)

Rainfed lowland Iloilo 1976 - 0 (10) 0(10) 0 (7) 0 (19) 2 13) 15 (180) 5 (76)1978 0 (54) 0 (3) 0 (17) 9 (64) 4 (78) 3 (73) 0 (121) 0 (191)1979 64 (2350) 16 (84) 6 (17) 0(6) 18 (68) 25 (104)

1981 56 (1322) 18(304) 0(5) 12(51) 18(138)Pangasinan 1976 60 (7 7 )b 62(82) 0(111) 5 (39)

1978 58 (53)b 5(51) Cagayan 1979 59(713) 8 (101) 6 (34) 0(7)

1980 0(11) 4 (28) 15 (286)

62 ( 48(177) 61(140)1981 0(5) 12 (2) 13(45) 14 (92)

Irrigatedlowland Laguna 1974 62 8

1974-75 50 41 1975 43 72 1978 96 (623) 21(24) 33 (60)1979 83(2017) 18(110) 8(62) 11 (89) 22(400)1980 78 (1625) 20(75) 0 (16) 4 (72)1983 60

9 (80) 7 (270) 30

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R F Apostol unpublished Iloilo and Pangasnan 136 Cagayan B Canapi unpublished Laguna 175 P CPantua unpublished 118 234 143 Carino and Shepard unpublished YSB = yellow stem borer ScirpophagaincertulasGFB = gold fringed borer Chilo auricilius PSB = pink stem borerSesamia inferens earcutting caterpillar Alythimna separata swarming caterpillar Spcdoptera mauritia acronyctoides skipper Pelopidas mathias reen horned caterpillar Melanitls ledaismene RLF = rice leaffoldersCnaphalocrocismedinalis and Marasmiaspp brown semilooper Mocisfrugalis Figures in parentheses are sample sizes Egg masses

IRPS No 123 January 1987 35

Table 9 Effect of seeding rate and insecticide on plant density and Yield losses from insect pests in upland rice often areyield of Dagge upland rice Batangas Philippines 1976a comparable to those of lowland rainfed or irrigated rice

Seeding rate Insecticide Dosage Plant density Yield (0-Y30o) but usually from different pests The principal (kg aiha) seed treatment (kg aiha) (noIra-row) (tha) groups are soil pests feeding on sown seed and rootsat 14 DE followed by seedling pests These groups have largely gone

50 None - 35 a 20 a unnoticed leading researchers to conclude that stem borers50 Carbofuran F 10 38 ab 23 a and seed bugs are the most important pests (17) Yield loss 10 beCabofuranF100 None - 4551 c 23 a trials that pinpoint major pests need to includc seed

100 Dieldrin WP 10 52 cd 22 a treatments Granules applied in the furrow may not give150 None - 59 d 22 a adequate protection from many seed pests

aAv of 8 fields DE = days after crop emergence In a column means The floristic and pedological diversity of upland rice followed by a common letter are not significantly different at the environments is matched by the diversity of arthropods5level by D)MRT F = flowable Wi = wettable powder both pests and their natural enemies Irrigation has homoshy

genized lowland rice environments Flooded soils tend tothreatened by epidemics of locusts or armyworms New have the same properties and to eliminate soil pests The technology stressing efficiency has minimized insecticide spatial and temporal dominance of cultivated rice has cost benefited specialist pests

Seed treatment is normally inexpensive and can be Upland rice has no specialist arthropod species The economicallyjustified in -nanycases(66) Treating seed with monophagous pests on upland rice prefer the lowlands systemic insecticide (216) but not chlorinated hydrocarbons where rice evolved Rice in the uplands exists only because (247) is most effective in controlling seedling maggots of man and even though lowland rice predates upland riceFoliar sprays have to be repeated to obtain control and specialist species evolved Wheat maize and sorghum do often are washed off by frequent early season rains not have monophagous pests leading us to conclude that Granular insecticides are effective but required dosages are there is no evolutionary advantage for inssects to have too expensive to justify except under heavy infestation narrow host ranges in upland graminaceae(247) When directed against first-instar larvae banding Pests have adapted to the uplands not only by having granules in seed furrows (177) can be effective against white wider plant host ranges but also by having long life cyclesgrubs at low dosages (025 kg aii ha) (134) Seed treatments undergoing dormancy or being dispersive Each pest has a are not promising for white grub control (248) unique life history strategy suited to highly unpredictable

Seed treatment is also effective ainst ants and may be environments In agivenyear someinsect specics an better cheaper than increasing the seeding rate In Batangas exploit upland rice as a food source but no pest is a senious treating 50 kg seed ha was as effective as increasing the one every year on upland rice seeding rate to 100 kgha and using untreated seeds (Table The low yielding potential of upland rice means less 9) In Brazil seed treatments control termites and other soil quantity and lower quality of food for pests to exploit but insects including the lessercornstalk borer(66) Domiciano also less incentive for farmers to undertake control (58) however found seed treatments inconsistent in their The most effective control would be early and synchroshyeffect against high populations of lesser corn stalk borer nous planting of early-maturing upland rice varieties to

Baiting can be an inexpensive way of controlling ants prevent insect pest buildup Breeding for insect resistance seedling maggot flies mole crickets and field crickets Baits should initially focus on stem borers because they are verymade of locally available material and impregnated with difficult to control by other means Biocontrol efforts insecticide can be sparingly distributed taking adantage of should first stress conservation of natural enemies preshythe pests ability to disperse and encounter bait sites dators and pathogens rather than parasites Efficient

The new low-volume hand-held controlled-droplet chemical control includes seed treatment or baiting for seed sprayers may offer an advantage over high-volume knap- seedling and some root pests and using controlled dropletsack sprayes for controlling foliar pests Upland rice sprayers for spot treatments against foliar pests and seed canopies are more open than those in lowland rice and bugs when economic thresholds are reached better droplet penetration should result Controlled droplet applicators would be ideal for seedbugs and defoliators

CONCLUSION

Upland rice is attacked by a wider array of insect pests than is lowland rice mainly beca use of the addition of soil pests Generally population levels of lowland rice pests are lower in upland than lowland rice leading many people to conclude that insect pests are not important on upland rice

36 IRPS No 123 January 1987

REFERENCES CITED

1Abraham C C and K S R Mony 1978 Occurrence ofLeptocorisa acuta Fabr (Coreidae Hemiptera) as a pest of nutmeg trees J Bombay Nat ttistSoc 74553

2 Agyen-Sampong M 1975 HlierogrlphusdiaganensisKrauss (Orthop-tera Acrididae) a new pest of rice in Northern Ghana Ghana J Agric Sci 8249-253

3 Agyen-Samnpong M 1976 Varietal resistance and other inle-ratcd pest control components as factors of rice varietal improvement in West Africa Prcsented at the Second varietal improvement serinar 14-18 Sep 1976 Bouak Ivory Coast West Africa Rice Development Association Monrovia Liberia 18 p

4 Agyen-Sampong M 1977 Outbreak of Ihccrorrdnisorae in rice seedlings near Rokupr Sierra Leone lit Rice Res Newsl 2(6)16-17

5 Agyen-Sampong M 1980 Dirty panicles and rice yield reduction caused by hugs IntRice Res Newsl 5(l)11-12

6 Agyen-Sampong M 1982 The major pest problems of irrigated upland and mangrove swanip rice ecosystems ir the humid tropical Guinea savanna and Salel climate zones Pages 318-328 in Pro-ceedings training course cn concepts techniques and application of irtegrated pest management in rice inWest Africa 10-20 Jan 1982 Fendall Liberia West Africa Rice l)evelopioent Association Monrovia Liberia 505 p

7 Akibo-Betts V I and S A Raymundo 1978 Aphids as rice pests in Sierra Leone lit Rice Res Newsl 3(6)15-16

8 Akinsola F A 1975 Present status of different rice stemborers in parts of Nigeria iicL Entomo Newsl 328

9 Akinsola FA 1979 Ihe biology and ecology of rice stemborers in Nigeria Ph 1) thesis University ol lbadan Nigeria 217 p

10 Akinsola E A 1979 Guidelines of integrated management of insect pests of upland rice in West Africa Presented at the Seminar on integrated management of rice diseases and insect pests at Bobo-Dioulasso Upper Volta 17-22 Sep 1979 West Africa Rice Develop-ment Association Monrovia Liberia

II Akinsola IA 1984 Insect pests of upland rice in Africa Pages 301-305 in An overview of upland rice research Proceedings of the 1982 lIouak Iwsrv Coast upland rice workshop International Rice Research Institute 13O Box 933 Manila Philippines

12 Alam M S V TJohn and Kaung Zan 1985 Insect pests and diseases of rice in Africa Pages 67-82 in Rice improvement in Eastern Central and Southern Africa proceedings of the international rice workshop at Iusaka Zambia 9-19 April 1984

13 Alticri M A A van Schoonhovcn and J D Doll 1977 The ecological role of weeds in insect pest management systems a review illustiated with bean (Phaseohsvulgaris L)cropping systems PANS (Pest Artie News Summ) 23195-205

14 Anmante F 1967 The leaf-cutting ant Atia capiguaraa pest of pastures [in Portuguese] Biol6gico (Sao Paulo) 33113-120 (Rev Appl Entomol A56207)

15Andrade A C and D Puzzi 1951 Preliminary results of experi-merits on the control of Sculptcoris iastantus on sugarcane [in Portuguese] Bioulgico (Sao Paulo) 1744-49 (Rev Appl Entomol A41421422)

16 Appert J 1970 Maliarplia separaelha (rice stem borer) new observations of entomological problems of rice in Madagascar [in French] Agron Trop (Paris) 25329-334

17 Ariadeau Mand Z Harahap 1986 Relevant upland rice breeding objectives Pages 189-197 in Progress in upland rice research Proceedings of the 1985 Jakarta conference International Rice Research Institute 1O Box 933 Manila Philippines

18Asar S and Kisimoto 1975 Seasonal occurrence of the smaller brown planthopper in upland rice [in Japanese] Proc Kanto-Tosan Plant Prot Soc 2228

19 Bakker W 1974 Characterization and ecological aspects of rice yellow mottle virus in Kenya Meded Landbouwhogesh Wage-ningen 152 p

20 Balut F F 1970 White grubs attacking rice Oryza ativa L [in Portuguese] Biol6gico (So Paulo) 36321-322

21 Banerjee S N and L M Pramanik 1967 The lepidopterous stalk borers of rice and their life cycles in the tropics pages 103-124 in The major insect pests ofthe rice plant Proceedings ofa symposium at the International Rice Research Institute September 1964 The Johns Hopkins Press Baltimore Maryland USA

22 Barrion A T and J A Litsinger 1980 Ants a natural enemy of cliaiarac in dryland rice IntRice Res Newsl

5(4)22-23 23 Barrion A T and J A litsinger 1981 The spider fauna of

Philippine rice agroecosystem 1Dryland Philipp Entomnol 5139-166

24 Barrion A T and 1 A Iitsinger 1984 The spider fauna of Philippine rice agroccosystcms II Welland Philipp Eiitomol 611-37

25 Barrion A T1 and J A Litsinger 1984 lihanus (Diptera Tabanidae) eggs an alternative host of rice stei borer (SI3) egg parasite Telenoonis digns (Iliymenoptcra Scelionidae) lit Rice Res Newvsl 9(6)19

26 Barrion A T and J A Litsinger 1985 Identilication of rice leaffolders by wing markings Int Rice Res Newsl 10( 1)24

27 Barrion A T and J A Litsinger 1985 Chctiosspp (Coleoptera Carabidae) a leaffolder ILF)predator lit Rice Res Newsl 10(1)21

28 Bertels A 1970 Insect pests of rice and theircontrol [in Portuguese] Pelotas Brazil Instituto de Expcrimentacao Agropccuararias do Sul Circular 43 24 p

29 Bonzi S M 1982 Ctilo (flisilineus J de Joannis (Lepidoptera Pyralidae) stem borer of irrigated cereals in Upper Volta LioFrench] Agron Trop Riu Rizic Cult Vivrieres Trop 37207-209

30 Bourke T V T L Fenner J N LStihick G Baker EHassan D F OSullivan and C S li 1977 Insect pest survey for the year ending 30th June 1969 Papta New Guinea )epartment of Agrishyculture Stock and Fisheries Port Moresby 57 p

31 Bowden 1 1973 Migration of pests in the tropics Meded Rijksfac Landbouwwet Gent 38785-796

32 Breniere J 1973 The main rice parasites in West Africa and their control Pages 27-42 in Proceedings seminar on plant protection for the rice crop West Africa Rice Development Association Monrovia Liberia

33 Breniere J 1976 The principal insect pests of rice in West Africa and their control West Africa Rice Development Association Monrovia Liberia 52 p

34 Brown ES 1962 The African arinyworn Spoiolptera eXI)Ina (Walker) (Lepidoptera Noctuidae) a review of the literature Cornshymonwealth Institute of Entomology London UK57 p

35 Brown F B 1968 Upland rice in Latin America FAO-International Rice Commission 12th Session of the Working Party on Rice Production and Protection 2-14 Sept Peradeniya Sri lanka Food and Agriculture Organization Rome Italy 6 p

36 Buyckx F J 1 1962 Disease and insect pests of crops inCongo Rwanda and Burundi [iil French] Publications dc LInstilut National pour lestude Agrononiquedu Congo M Weissenbruck S A Brussels Belgium 708 p

37 Cariiio F 0 1981 Role of natural enentics inpopulation suppression and pest management of green rice lealhoppers Ph D thesis University ofthe Philippines at Los Bafitos PhilippinesLaguna 136 -

38 Carpente A J 1983 The development of rice cultivation and extension of vegetable productin in Zanzibar Final Report Vol II1 October 1983 Food and Agricultme Organization Ronte Italy

39 Carroll C P and S1 Risch 1983 Tropical annual cropping systems ant ecology Environ Manage 751-57

4(0 Cates R G 198 1 Host plant predictability and the feeding patterns of monophagous oligophagous and polyphagous insect herbivores Oecologia (Berl) 48319-326

IRPS No 123 January 1987 37

41 Cendcafia S M and F B Caora 1967 Insect pests of rice in the 62 Ene 1 C 1981 I)iiorldhol)ierusihilis Slater (licteroplera Philippines Pages 591-616 in The najor insect pests ofthe rice plant lygacidue Illissinac) a new rice pest front Nigeria lit Rice Res Proceedings of a symposium at the International Rice Research Newsl N3)15 Institute September 1964 The Johns Hopkins Press Baltimore 63 Fugenio T S 1965 rhe effect oi e-vCgetation on rice production Maryland USA 729 p and the possibility of overcoming it by leaf pruning MS thesis

42 Centro Ihternacional de Agricklittira Tropical 1984 Upland rice in University ol the Philippines at Los Baijos ILaguna Ihilippins 69 p Iatin Anerica Pages 93-119 hi An os ervies of uplaind rice research 04 Farrow R A 1975 The Alrican nmigratory locust ii is main Proceedings of the 1982 Boinakamp Ivory Coast upland rice workshop outbreak area of the middle Niger qlinlitatise studies of solitary International Rice Research Institute P0 Blox 933 Manila populations in relation to environmental factors Locusla No II Philippines 198 p

43 Chandra G 1979 Taxonomy ad hn riononlics ofthe imect parasites of 65 Fennal R G 1943 Food crop investigation Windward and rice Icalloppers and planthoppers in the Pihilippines and their Ieeward Islands First report Report onl work dunring the periodiiiportairce iii natural biol)gical cointrol Philipp Fntoltol 4119-139 April 1942-April 1943 Imperial College of Tropical Agriculture

44 Chang T T 1976 The originii colut ion enlIivation dissemination Trinidad 16 p (Rev of Appl Fintornol A3222-25) and diersificition of Asian and African rices -uphytica 25425-44 1 66 Ferreira F 1983 Integrated pest ritaageiret of upland rice [ill

45 Chatierjee P B 1979 Rice leaffolder attacks in India lit Rice Res Portuguese English summary] Pages 323-341 in Upland riceculttre Newsl 4(3)21 factors affecting production E Ferreira T Yamada and F-

46 Cheng Cbien-pan 19(4 Upland rice in Taiwan International Rice Malmolta eds Institutoda Potassae Foslato Institnto Internacional Cnmmission Wiorking Party oit Rice Production ard Protection I()th da Potassa Piracicaha Sao Paulo lraid Meeting 3-10 March 1964 Manila Philippines Food and Agricul- 67 Ferreira E arid F dl Silva Martins 1979 Occurrence of ture Organiaion Rome Italy 5 p Sogatohs orvziola (Muir) inupland rice in Goias Ilrail lit Rice

47 Chcrrett 1 M and 1) 1Peregrine 1976 A review of the status of Rcs Ncwssl 4(4)18 leaf-cutting ants and their control Ann Appl Biol 84124-128 68 Ferreira E and 1F da Silva Martiris 1984 Insect pestof rice ill

48 Chipeta F M 1974 Pests of rice in Malawi Pages 90-95 in Brazil and their control Jill Portugucsej FMIlRAtPA-CNPAF Proceedings Sith eastern Africa cereals research conference 10-15 Goiania Brmiil 67 p March 1974 DRB Mialda ed Ministry of Agriculture lilongwe 69 Ferreira FI F ca Silha Miartins S S Nero and 1 It P de Malawi Carvallio 1982 M tltiplc regression analysis todeterimine the effect of

49 Chu Yau-i 1979 The d atiaging period of the rice seedling fly insect populatiois ott upland rice yield [inIPortuguese Pesqui (Atlterigotiavxigu Stcin) (NItiscide Diptera) oil cori) at Fast Java Agropecu Bras 17671-675 Indonesia Plant Prot Bill 21403413 70 Ferreira F 1F da Silva Martins and F 1PZimmermann 1979

50 Conklin H C 1957 IHaninoo agriculture a report on aln integral Rice varietal resistance to tle lesser corn stalk borer [in Port uguese] systent of shi fti rig cult ivationi in the Pltilippincs Food and Agriculture Pesqui Agropccu Biras 14317-321 Organization Rome Italy 209 p 71 Fleming W F 1968 Biological control of lie hapanese beetle U S

51 Cramer H II 1967 Plant proection and world crop production Dep Agric RLs Sers [ech Bull 1383 78 p Pilaniensclhutz-Nachr 201-524 72 Food and Agriculture Organiation of the United Nations 1982

52 l)ani R C and Majunidar 1978 Prelininary observations on tie Country report surmimary Pages 27-37 in Report on the first incidence of root aphids on differeit rice cultnrcs Sci Cult 4488-89 FAOIUN)IPhail and regional training coiuirse in iniproved ctltural

53 Davidson R I 1979 lffects of root feeding on foliage yield Pages practices for upland rice 5-23 Oct 1981 Northern Region Agrculturd117-120 in T K Ci osb and R B Pottinger eds Proceedings of the Developririt Center Chiang Mai Food and Agricuilture Organiiashy2nd Australian conlerence on grassland invertebrate ecology Govt tion Rome Italy 37 n Printer Wellington New Zealand 73 Franco do Ariaral S 195 1 Residual action of insecticides in turfy

54 l)ean G 1W 1976 Rice insect pests inlaos Int Rice Res Newsl soil--the results of a field experiment [in Portuguese] Biol6gico (Sao 1(2)15 Paulo) 17209-214 (Rev Appl Entomol A41423)

55 I)ean G J W 1978 Insect pests of rice in Laos PANS 24(3)280-289 74 Freeman ) 1970 Report onithe Iban london School of -coroiics 56 )ingle H 1984 Behavior genes and lifc histories conplex adapta- Monographs on Social Anthropology No 41 Athlane Press

tions to uncertain environments Pages 169-194 in A new ecology London UK 317 p novel approaches to interactive systems P W Price C N Slobod- 75 Fuchs T W1 A Harding and 1W Smithlr 1979 Induction and chikoff and W S Gaud cdsl ohn Wiley New York USA termination oldiapause in the sugarcane borer Ann Flitonlol Soc

57 Dobelmann-JP and M Falais 1968 Upland rice cultivation ill tie Am 72271-274 North-western part of Madagascar in 1966-1967 [in French English 76 (hbriel B1) 1975 A review tf the major insect pests of some tpland summary] Trop Agron 2313-21 crops in tlc Philippines Pages 86-111 in Ad Htoc panel of experts on

58 Doniiciano N L 1979 Chemical control of tie lesser corn stalk pest disease arnd weed problems in some rainfed crops Bangkok (lolsonno~paloIgnowslhts) (Zeller 1848) (I-epid(ptera Ihycitid ae) Thailand Food aid Agriculture Organization Roire Italy

on rice in Ponta Gressa Parara 1977-1978 [in Portuguese] Pages 77 Garg D K and N K Shal 1983 White grub outbreak on rainfed 291-303 in Anais 1 Reuniao de tecnicos cmiirizicultura do estado de dryland rice in Uttar Pradeslh Int Rice Res Ncwsl 81(4)16-17 Sao Paulo-Campinas Cati Brazil 78 Garg I) K and IP Tandon 1982 Major insect pests of rice on hilly

59 Dunsmore 1 R 1970 Investigations on tlie varieties pests and tracts of Uttar Pradesh India lilt Rice Res Newsl 7(11)11-12diseases of upland rice (hill padi) in Sarawak Malaysia Int Rice 79 Garrity 1) P 1983 Upland rice research an international bibliograpy Comm Newsl 1929-35 1975-1982 Iternational Rice Research hnstitnte PO Box 933

60 Elias 1R1965 The new decline of rice seedlings is not caused by Manila Philippines 121 p parasites [in Portuguese] Biol6gico (Sao Paulo) 313-6 80 Ghauri M S K 1982 A new subspecies of Iinorh)rone bull

61 Empresa Brasiliera de Pesquisa Agropecuaria 1984 Upland rice itt cornlus Slater (Hemiptera Iygaeidae) front Papua New Guinea on Brazil Pages 121-134 it Av overview of upland rice research rice and carpet grass Ihll lEntoniol Res 72133-137 Proceedings ofthe 1982 Blouak Ivory Coast Upland rice workshop 81 Gonzales F 0 PArregoces R I ihernandef and OT Parada International Rice Research Institute PO Box 933 Manila 1983 Insect and imite pests of rice arid their control in Latin America Philippines [in Spanish] la Federacion Nacional de Arrocerts (Fedearroz)

Bogota Colombia 60 n

38 IRPS No 123 January 1987

82 Grist D H and R J A W Lever 1969 Pests of rice Longmans London UK520 p

83 Hale P R 1979 Two subterranean pests of upland rice in Papua New Guirnca IntRice Res Newsl 4(2)16-17

84 Hale P R and J T Hale 1975 Insect pests of rice in the New Guinea Islands Rice Entomol Newsl 34

85 Harris W V 1969 Termites as pests of crops and trees Common-wealth Institute of Entomology London UK 41 p

86 Hattori 1 1968 A method to distinguish between grass leaf roller (Ctaplha rocis medinalisGuenee) and the rice leaf roller (Susurtia exigua Butler) [in Japanese] Plant Prot (Japan) 23167-170

87 Henry I E and 1A Onsager 1984 Experimental control of the mormon cricket Iahrosshnph by NJsena locustae(Aficrospora Microsporidae) a protozoan parasite of grasshoppers (Ort Acri-didae) Entomophaga 27197-201

88 Hidaka 1 P Vungsilabulr and SKadkao 1974 Studies on ecology and control of the rice gall midge in Thailand Tech Bull 6 TropicalAgricultural Research Center Tokyo Japan 113 p

89 Hincklcy A 1) 1963 Ecology and control of rice planthoppers in Fiji Bull Entomrol Res 53467 481

90 Hirao J 1982 Ecology and chtInical control of the rice leaffolder Jpn Pestic Inf 4114-17

91 Ho DT 1984 Sten borer incifience in rice ecosystems in KenyaEast Africa Pages 307-320 in A overview of upland rice research Proceedings of the 1982 Bouac Ivory Coast upland rice workshop International Rice Research Institute PO Box 933 Manila Philippines

92 Ho )T J 1Njokah and J GKibuka 1983 Studies on rice sten borers in Kenya with emphasis on ifaliarpiaipiraellRag Insect Sci Appl 465-73

93 Ichihara I T Ito A NIatsuo and M Sawada 1971 Chemical control of the rice striped stem borer in upland rice grown in plastic mulch [iii Japanese] Proc Kanto-Tosan Plant Prot Soc 1882

94 IclhiharaIand A Matsuo 1969 Damage caused by rice striped stett borer to upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1686-87

95 Ichhara I atid A MIatsun11970 Conside ration oft lie extraordinary prevalence of rice striped stem borer in upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1778-79

96 Institut de Recherches Agrolnliqes Iropicale et des Cultures Vivrieres 1978 Insect resistance Pages 12-13 in Upland rice 1978 atnual report Bouak Ivory Coast 27 p

97 International Institute for Tropical Agriculture 1983 Aphids itt rice roots I ITA Res Briefs 4(1-3)4-5

98 International Institite fi Tropical Agriculture 1983 Hopperburn in rice caused by brown plathopper IITA Res Briefs 4(l-3)6-7

99 International Rice Research Institute 1975 Annual report for 1974 PO Box 933 Manila Philippines p 216

100 International Rice Restarch Institute 1976 Annual repiort for 1975 PO Box 933 Manila Philippines p 228

101 International Rice Research Institute 1979 Cropping systems program Component technology development and evaluation Page 404 i Annual report for 1978 11O Box 933 Manila Philippines

102 International Rice Research Institute Department of Entomology 1981) Insecticid evaluation 1979 Tables 73 74 PO Box 933Manila Philipes

103 International Rice Research Institute Department of Entonmology 1981 Insecticide evalomtion 1980 Table 81 POBox 933 Manila Philippines

104 International Rice Research Institute 1981 Report of a meeting of Asian rice-based cropping systems entomologists Eleventh cropping systems working group meeting 18-22 May 1981 PO Box 933 Manila Philippines p 132 133 144

105 International Rice Research Institute 1984 Cropping systems program Pages 368-373 in Annual report for 1983 PO Box 933 Manila Philippines

106 International Rice Research Institutc 1984 Upland rice in Asia Pages 45-68 in An overview of upland rice research Proceedings of the 1982 Bouakamp Ivory Coast upland rice workiiop loternational Rice Research Institute 11O Box 933 Manila Philippines

107 Irshad M 1977 Grasshoppers in rice in Pakistan lit Rice Res Newsl 2(3)8-9

108 Ito R and TOno 1964 Upland lice its culture and varieties in Japan lIternational Rice Cormmission loth Meeting of the Working Party on Rice Production and Protection 3-10 March 1964 Manila Philippines Food and Agriculture Organization Rome Italy 12 p

109 Jeffrey M I and V A Dyck 1983 Effect of moonlight on lightshytrap catches of brown planthopper lit Rice Res Newsl 8(6) 13

110 Jennings P R and T Alicia Pineda 1983 Management ofthe rice plnthoppers Sogatodsorivi(coIl in Latin America Presented at the meeting on stable plant resistance FAO and lIndocsian I)inectorate of Food Crops Protection Bali lndonesia 29 Novcniber-2 l)ccemshyber 9 p

IllJimenez Gomez J A 1979 Rice insect pests i lla ios ()riCntales lil Spanish] Pages 99-103 in Rice ciltivation Instituto (olotibiatto Agropecuario Compndio No 29

112 Jongeleen F JJ Soesanto and A Mockidjo 1979 Studies oti pathogens of two species of Scarabacidac froinIndonesia Z Angew Enlomnol 87412-417

113 Jouvenaz 1) PC S lofgren and W A Baiks 1981 Biological control of imported fire antsa review of cirrent knowledge Bull Entomol Soc Am 27203-208

114 Kainoh Y K Shiniiu S MarUani Y Taiaki 1980 lost finding behavior of tie rice bu lplotori+JchinvittjI)allas (liciniptera Corcidac) with special reference to diet pattern ol aggregation and feeding on the rice plant Appl Entomol Zool 15225-233

115 Kakkar K 0 P Sharma and P I Sharma 1977 Note on Ihteronyrchus liodresRedt (Coleopicra I)ynastidae) as asporadic pest of paddy and maie inHimachal Pradesh Corr Agric 156-57

116 Kalode M131 79 Major insect pest problems and their control in upland rice (Unpublishcd mineogriph CWI oJNaaiat lni Chandaratat (198 1)original not seen)

117 Kalshoven L i 98 I The pests of crops in I ldoniesia (translated by P A vatn der Laan) 1 T Ichtiar B3aru-vati Iloeve lakarta Indonesia 701 p

118 Kamal N Q 1981 Suppression of whitebacked planto ner Sogatella fircaifra (Horvath) aLId rice lealfolder (nhal -cis licdb lis (Guenee) populations by natural cnclmie hlI) tiesis Grcgorio Araneta University Foundation Victoncta Park Metto Manila Philippines 179 Ip

119 Karim A N M Reaul 1975 A species ofliealy bug Trioni i70s ceres new oii rice in the Philippines Rice Entoniol Newsl 247

120 Katanyukul Wand C Chandaratat 1981 Insect pestsof uplaid rice and their control Report firstFAOUNlFi Tliailand training course on improved cultural practices for tplad rice Chiang Mai Thailand 5-23 Oct 1981 Food and Agriculture Organiation Rome Italy 22 p

121 Kern F 1955 lhletwioa(I)uninemts) bhhit Biturm(Coleoptera) a new pest of sugarcane inVenezuela [inSpanish with Etnglish sunmary] Agron Trop (Maracay) 527-29 (Rev Appl Eitoniol A4572)

122 Khan K M and S Ghai 1974 White grubs ald their control in India Pesticides (1Bombay) 819-25

123 Kiritani K 1975 Integrated pest management of rice pests Pages 66-70 itt Proceedings platning workshop ott cooperative hIeld research in pest managementP SMotooka ed East-West Center Honololu Hawaii USA

124 Kisimoto R 1967 Ecology of insect vectors forecasting and chemical control Pages 243-255 in The virus diseases of tle rice plant Proceedings of a symposium atthe International Rice Research Institute April 1967 The Johns Hopkins Press altimore Maryshyland USA

125 Kisimoto R and V A Dyck 1976 Climate and rice insects Pages 367-391 in Climate and rice International Rice Research Institute PO Box 933 Manila Philippines

126 Kureha S S Kobayashi SAsari and T Ishihara 1977 Life history and control of the mole cricket Grrllotalpa africana Palisot de Beauvois [in Japanese English translation 7520 1RRI library] Proc Kanto-Tosan Plant Prot Soc 21192-197

127 Kushwaha K S 1981 The black beetle as a pest of rice in Haryana India Int Rice Res Newsl 6(4)14

128 Lawani S M 1982 A review of tile effects of various agronomic practices on cereal stem borers Trop Pest Manage 28266-276

129 Leuck D B 1966 Biology of the lesser cornstalk borer in South Georgia J Econ Entomol 59 797-801

130 lever R J A W 1955 Insect pests of the rice plant in Malaya Int Rice Comm Newsl 1612-21

131 Li Ching-Sing and Shin-Foon Chiu 1951 The study ofthe rice gall midge Iachydiplosis orvza Wood Mason [in Chinese with English summary] 1Taiwan Agric Res 21-13

132 Iitsinger J A 1984 Assesament of the need-based application for rice Proceedings TechnologyTransfer Workshop 15-16 March 1984 International Rice Research Institute 1PO Box 933 Manila Philippines 6 p

133 Litsinger J A 1984 Environmental diversity and insect pest abundance with reference to the Pacific Pages 285-307 in Proceedings sub-regional training course on methods of controlling diseases insects and other pests of plants in the South Pacific 4-20 Oct 1982 Vaini Kingdom of Tonga Nukualofa

134 Litsinger J A RF Apostol and M BObusan 1983 White grub Leucopholis irrorata (Coleoptera Scarabaeidae) pest status popula-tion dynamics and chemical conllol in arice-maize cropping pattern in the Philippines J Econ Entoinol 761133-1138

135 Litsinger J A C de la Cruz F Raymundo A T Barrion M D Lumaban J P Bandong M S Venugopal F Paragna and E Balete 1982 Insect pests and insecticide reponse of dry-seeded bunded rice Pages 179-193 in Report of workshop on cropping systems research in Asia International Rice Research Institute Los Bafos Philippine

136 Litsinger J A M 1) Luinaban J P Bandong A T Barrion W Katanyukul and N Panda 1978 Insect pest management in rainfcd lowland rice agroecosystms of tropical Asia International Rice Research Conference 17-21 April 1978 International Rice Research Institute Los Bafios Philippines

137 Litsinger J A and K Moody 1976 Integrated pest management in multiple cropping systems Pages 293-317 in R I Papendick P A Sanchez and G B Triplett eds Multiple cropping American Society of Agronomy Madison Wisconsin USA

138 Loevinsohn M E 1984 The ecology and control of rice pests in relation to the intensity and synchrony of cultivation Ph D thesis University of London 36) p

139 Loevinsohn M E J A Litsinger N Panda M A Austria R F Apostol C Andrion and J Bandong 1982 A basis for predicting rice insect pest abundance in cropping systems Paper presented at the 1982 International rice research conference on rice and cropping systems 19-23 April 1982 International Rice Research Institute Los Bafios Iaguna Philippines 25 p

140 Lopez A W 1930 Repot of the Entomology Department Pages 145-172 in Annual report of the Philippine Sugar Association 1929-30 Manila

141 Lopez AW 1931 Entomology Department Pages 227-266 in Annual report of the director or research 1930-31 Philippine Sugar Association Manila

142 Lopez A W 1932 Entomology Deparment Pages 252-279 in Annual report of the Philippine Sugar Association 1931-32 Manila

143 Lungibill P and G G Ainslee 1917 The lessercornstalk borer US Dep Agric Agric Res Serv Tech Bull 539 27 p

IRPS No 123 January 1987 39

144 McGuire J U Jr and BS Crandall 1967 Survey of insect pests and plant diseases ofselected food crops of Mexico Central America and Panama U SDepartment of Agriculture International Agriculshytural Development Service Agricultuial Researrh Service October 1967 11 p

145 McNeill Sand T R ESouthwood 1978 The role of nitrogen in the development of insectplant relations Pages 77-9 in lliochemical aspects of plant and animal coevolution J B Ilarbornu ed Academic Press London UK

146 Maebara H 1955 Spatial distribution of lastinstiarlanaeof purplish stem borer (Sesamia iqferens Walker) hibernating in an upland rice field Bull Kagoshima Univ Fac Agric 438-41

147 Mallick E H and N G Hajra 1977 Response of rice plant to defoliation treatments at different developmental stages Indian Agric 2151-56

148 Mansingh A 1971 Physiological classification of dormancies in insects Canl Entomol 103983-10)9

149 Martell C G 1974 Catalogue of plant feeding insects of Mexico [in Spanish] Fitofilo 27(69)1-176

150 Martinez 0 J 1982 Principal rice pests in Ilanos orientalis [in Spanish] P~ages 80-94 in Curso deact nalization de conocimientos en arroz Villaniencis Instituto (olonmbiano Agropecuario

151 Martins J F da Silva F Ferrcira A S Prabhu F 1 P Zimmermann 1980 Use of pesticides for the control of the main subterranean upland rice pests esqui Agropecu Bras 1553-u

152 Martins I F da Silva F Fe eira rod BdamSilveira Pinheiro I Q2 Damage simulation caused by defoliators of upland rice [in Portuguese with English summary] Pesqui Agropecu Bras 171113-1119

153 Martins 1F da Silva N(iyuen van Tan and 1Ida Silveira Pinheiro 1981 Stem borer resistance in upland rice associated with plant morphological characters [in Portuuce Iesqui Agropecu Bras 16187-192

154 Masaki S 1980 Sumnmer diapause Ann Rev Entomiol 251-25 155 Masuzawa T 11 Suwa and F Nakasoji 1983 l)ifkleences of

oviposition preferencc and survival rate of two skipper butterflies Parnara guttata and PIchlpidai iathiais (Iepidoptera HIesperiidac) on rice plant and cogon grass New Entomol 3247-56

156 Maurya D M and C P Vaish 1984 Upland rice in India Pages 69-92 in An overview oftpland rice research Proceedings ofthe 1982 Bouakamp Ivory Coast upland rice workshop International Rice Research Institute IO Box 933 Mnila Philippines

157 Menon M G R R N Katiyai S Kuar and S S Misra 1970 Termites attacking the paddy crop Sci Cult 36294-295

158 Miyahara Y and 1 Nishikubo 1967 Behavior of young nymphs of the red rice root aphid Iholahsipiun ri liabhdontinalisSasaki in relation to movement from the leaf to the roots of upland rice [in Japanese] Pioc Assoc Plant Prot Kyushu 1363-66

159 Mo ljoa Tjien 1952 White grub of upland rice (Holotrichiahelleri Brsk) [in Indonesian with English summary] Contrib Agric Res Stn Bogor No 13134 p

160 Mochida 0 and T Suryana 1979 The occurrence of the brown planthopper and ragged stunt on lowland and upland rice ill Central Java and logyakarta in tile late of January 1979 Entomnolgy Survey Report No 15 Central Research Institute of Agriculture (CRIA) Sukamandi Indonesia I February 1979 9 p

161Monet C 1979 Varietal resistance to upland rice pests [inI French] Pages 35-39 in Annual report 1977 Institut des Savanes Department des Cultures Vivrieres Bouak6 Ivory Coast

162 Monte 0 1942 A caterpillar ofrice-fields[in Portuguese] Biol6gico (Sao Paulo) 8161-163 (Rev Appl Entomol A31414)

163 Munroe I) D 1979 Pest management strategy for hill rice in four resettlement areas in Sarawak Malaysia Pages 89-102 i1 Guidelines for integrated control of rice insect pests Food and Agriculture Organization Plant Production and Protection Paper 14 Rome Italy 115 p

40 IRPS No 123 January 1987

164 Nakasuji F M Ishii and Jen-Zon Ito 1984 Rice skippers in Taiwan and their life histories ZZrC (Taipei) Tech Bull 82 10 p

165 Navas I) 1976 Fall armyworum in rice Pages 99-106 il Ecological animal control by habitat management Proc Tall Timbers Con-ference 28 Feb-I Mar 1974 Fall Timbers Res Stn Misc Publ 6 Tallahassee Horida USA

166 Nickel 1 1 1961 Biological control of rice stem borer A feasibility study Tcch Bull 2 International Rice Rcarch Institute PO Box 933 Manila Philippines IIl p

167 Nigan P M and R A Vernia 1985 Insect pests of uplaild rice in Uttar Pradesh Int Rice Res Newsl 10(6)22

168 Nilakhe S S 1985 -cological observations on spittle bugs with emphasis on their occirence in rice Pcsqtui Agropecu Bras 20407-414

169 Nixon G EJ 195 1 lie association of ats with aphids and coccids Commonwealth Iusti lcte of Entonology Iondon UK 36 p

170 Nurdin F an Ishak Manti 1985 Study oit seedling fly of upland rice Interlational Upland Rice Moitoring Tour Report 26 lebruary-2 MarLI Sukaranti West Sumatra

171 Oka 1 N 1979 Ciltural conttol of the brown planthopper Pages 357-369 in Brown planthopper threat to rice production in Asia International Rice Research Institutc 1p0 Box 933 Manila Philippines

172 Ohni M and A Vilnaii 1975 Rice insect pests of Italy biology of Sipha grcria (Kaltenbach) and its control [in Italian] Riso (Milan) 2459-71

173 Otanes F Q atnd 1 I Sison 194 1 Pests of rice Philipp J Agric 12211-259

174 Painter R I 1951 Insect icsistance in crop plants University Press of Kansits Lawrence Kansas USA 520 p

175 Pantua P C and 1A litsinger 1981 Arthropod abundance in continuous vs biannual rice cropping systeis Int Rice Res Ncwsl 6(2)20-21

176 Pathak M ) and V A l)yck 1975 Sttudics ott insect pests of upland rice Pages 186-191 in Major research in upland rice International Rice Research Institute PO Box 933 Ilanila Philippines

177 Patil B R B M IHasabe and 1) S Ajri 198 I Chemical control of white gritb lIditriihinhorraia Fabr intfesting groundntut paddy atnd sorghum Indian 1Entomol 43232-236

178 Pawar A 1) 1974 (ollection and identification of insect pests of rice rlerminal repirt lInternational Rice Research Institute PO Box 933 Manila Philippines 72 p (tnimco)

179 IerrajU A aid A S Reedy 1965 Observations ott diapausc itt tie steti borer ohrice Rice News Teller (lai-Mar)9-13

180 Pont A C 1972 A review of the Oriental species of theri ona Rondani (I)iptera Muscidac) of econtomic importance Pages 27-102 in Control of sorglium shoot fly M G lomtani and W R Young eds Oxford atd IH1 Ptthl Co New I)clhi India 324 p

181 Prasad S C 1PChand aid M 1PSingh 1983 Incidence of rice gall midge in the upland plateau region of Hihar Oryza 20235-236

182 Rai P S 198 0 Nisaga simpex damage to rice in the hill tracts of South India Int Rice kles News[ 5(I)12

183 Rathorc Y S 1969 Gonocplhahli dcprssunt Fabr as itpredator of (tilo zonwIlhis (Swinhoc) and other moths in field cage at Patnagar Uttar Pradesh Indian 1Entotnol 3127

184 Reed W 1983 Crop losses causcd by insect pists in the developing world Proceedings 10th Internatial Congress of Plant Protection lrighton UK 174-80

185 Reedy M V 1981 Ateactogasterinhens(Walker)a new root-grub pest of tipland paddy and inaize Curr Sci 501076

186 Reveno M A 1968 Catalogue of the insect pests of economic crops in Colombia [in Spaiish] Asociacion Latino Americana de Entomo-logia (AIAE) 129-33

187 Reyes G M 1957 Rice dwarf disease in the Philippines Food and Agriculture Organiiation Plant Prot Bull 617-19

188 Rice S E A A Grigarick and M 0 Way 1982 Effect of leaf and panicle feeding byarmyworm (Lepidoptera Noctuidae) larvae on rice grain yield Environ Entomol 7593-595

189 Risch S J 1) Andow and M A Altieri 1983 Agioccosystem diversity arnd pest control data tentative conclusions and nizw research dirctions Environ Entomol 12625-629

190 Rivera C T 1956 Three organic insecticides in thecontrol of insects affecting upland rice Philipp Agric 39465-472

191 Roffrey 1 1972 Locusta outbreaks in the Philippines Acrida 1177-188

192 Rojas R 1) 1941 The effect of pruning at varying ages upon the yield of Ramairio Phipp Agric 29851-860

193 Roonwal M i 1979 Ternite life and termite control in tropical South Asia Scientilic Pnbl odhpr India 139 p

194 Rosetto C1 F da Silva Martins N C Schmidt and L EAzzini 1978 Damage caused by pasturespittle bugs 4Dvoi)lavopictaandDJ schach) on rice lBragantia Nota 6 (Matcli)25-27

195 Rosztto C1- S S Neto I- Amantc I) Iink 1Garcia-Vieira D M de SoLza N V Bansa t and A M de Olivcira 1971 Rice pests of Brazil [in Portuguese] Pages 149-238 inrFA()-International Rice Commission 2nd meeting of the Rice Committee for the Americas 6-11 l)ecember 1971 Pelotas Rio Grande do Sul Brazil

196 Rothschild G H L 1969 Obseratiois on the armyworm Spodoptera matritia acronrctoiiles Gn (Lep Noctuidae) Bull Entomol Res 59143-160

197 Rothschild G 1- L 1974 Parasites of the armworm Spolopiera mauritiaacroncitoihes in Malaysia Fntornopliaga 19293-299

198 Sachan i N S K Gangwar and 1N Katujar 1980 Field crickets damaging cages in North Eastern IHill Region Indian J Entomol 42526-527

199 Sands 1) P A 1977 The biology and ecology of Lepocorisa (Hemiptera Alydidac) in Papua New Guinea Res Bull 18 Departshyment of Primary Industry Port Moresby Papua New Guinea 104 p

200 Sarawak 1968 titomology division Pages 97-106 in Report of the research branch of the Department of Altriculture 1966 Kuching (Rev AIppl Eintomol A56603)

201 Satpathy JM M S Das and K Naik 1977 Effect of multiple and mixed cropping on the incidence of scine important pestsI Entomol Res 178-85

202 Sauer IHF G 1939 Notes Ott Elasmpnlpualtsligniosellths Zeller (Lep Pyr) a serious pest ofcereals in Sao Paolo [in Portuguese] Arq Inst Biol 10199-206 (Rev Appl Entotnol A28378)

203 Satinders 1L A 1 SKing and C L Vargas S 1983 Crop pests of Central Anerica with t bibliography [in Spanish] Centro Agroshynotnico Tropical de Investigacion y Ensenanza Turrialba Costa Rica (CATIE) Tec Bol 9 90 p

204 Sauphanor B 1985 Some factors of upland rice tolerttce to stem borers in West Africa Inseu Sci Appl 6429434

205 Sen P atnd S Chakravorty 1976 Stentflics in Haritighata Nadia West Bengal India [it Rice Res Newsl 1(12) 15

206 Senapai B and 1M Satpathy 1983 The seedling fly Atherigona orrzae Malloch is a pest of rice in Orissa (India) J Entomol Res 777-79

207 Shhjahan M S Alatn M Ah nad and BN Islam 1983 A new scarabaeid beetle pest of dryland rice in Bangladesh Int Rice Res Ncwsl 8(I)16-17

208 Sharma G L S C Modgol and R C Gautam 1979 Effect of depth of submergence ott incidence of bacterial blight and white grub infestation in transplanted rice Int Rice Res Newsl 4(6)12-13

209 Sharma S 1985 Scarabaeid beetle outbreak itt upland rice Int Rice Res Newsl 1(1(l)24

210 Shiraki T 1917 The paddy borer Schoenobhits incertulas Walker [achaku Agric Exp Stn Formosa 256 p

211 Silva A de Brito and 11P Magalhaes 1981 Insect pests of rice [in Portuguese] Circular Tecnica No 22 EMBRAPA Centro de Pesquisa t ecuaria de Tropico Unido (CPAJU) Belem Brazil 14 p

212 Singh S A 1979 Emergence of rice leaf butterfly as a rice pest in Manipur India int Rice Res Newsl 4(4)16-17

213 SinghS A 1983 Changing trends of insect pestsof rice in Manipur Int Rice Res Newsl 8(f)12

214 Siwi S S and P H van Doesburg 1984 Lenovorisa Latreille in Indonesia (Heteroptera Coreidae Alydinae) Zool Meded (Leiden) 58117-129

215 Soejitno J 1977 Reaction of some upland rice varieties toseedling fly eltherikonaexigua Stein [in Indonesian] Siri Hana I Penyakit 630-35 Lembga I osat Fenclilian Pertanian Bogor Indonesia

216 Socnardi 1967 Insect pests of rice in Indonesia Pages675-683 in The major inse pests of thu rice plant Proceedings of a symposium at the Internatital Rice Rec Wi itt S pternber 1964 The Johns Hopkins Press Baltimore Maryland USA

217 Soto P E A T Petcz and I W Buddenhagen 1976 Survey of insect pests and diseases of rice in different ecological zones in Nigeria Rice Entomol Ncwsl 435-46

218 Soto 11E and Z Ziddiqi 1976 Screening for resistance to African rice insect pests Varietal improsetnent sCmi n r Bouak Ivory Coast 13-17 Sept 1976 West Africa Rice l)evelopment Association Monrovia Iiberia

219 Squire F A 1972 Entomological problems in Bolivia PANS (PestArtic News Sumr) 18249-268

220 Stapely 1 II 1978 The rice leaf roller in the Solomon Islands Int Rice Res Newsl 3(l)13

221 Suenaga H 1937 On aphids attacking the roots of rice Oyo-I)oubts Zasshi 9150-152 [in Japanese] (Rev Appl Entomol A25745)

222 Suenaga H 1952 On the seasonal abundance of three subterranean aphids on upland rice Bull Kyushu Agric Exp Sti 1249-262

223 Sutynso Yas and Dandi Soekaria 1964 Beberapa hana yang ttenperang tana man padi tanah kering [in Indonesian] Rapat kerja Padi Tanah Kering 6 sd 8 Juily 1964 Kaliurang 9 p

224 Tallamy I) W and R F Denno 1983 Alternative life history patterns in risky environments in example from lace bugs Pages 129-147 in Insect life history patterns R F l)cnio and If Dingle eds Spring-Verlag 225 p

225 Tamura 1 1939 Observations on some insect pests of field crops [in Japanese] 1 Plant Prot 26414-420 (Rev Appl EntomolA27-556)

226 Tamura and Suzuki 1964 Influences of the early-transplanting practice in rice culture upon development of the rice stem borer Chilo siql)ressalisWalker Bull Ilokuriku Natil Agric Exp Stn 761-94

227 Tan J P 1924 The rice root aphid Drowhi ihirsoia Philipp Agric 13277-288

228 Tanaka 1 1957 Taxonomy and distribution of some subterranean aphids injurious to upland rice in Japan with description of a new species [in Japanese English summary] Botyu-Kagaku 22168-176

229 Tanaka T 1961 The rice root aphids their ecology and control Spec [full College of Agriculture Utsunomiya University Japan F3 p

230 Tauber M IC A Tauber R Nechols and 1Obrycki 1983 Seasonal activity of parasitoids control by external internal and genetic factors Pages 87-108 itt I)iapause and life cycle strategies in insects V K Brown and I Ilodek eds )r W Junk The Hague Netherlands

231 Taylor W F 1972 Effects of artificial defoliation (simulating pest damage) on varieties of upland rice Exp Agric 879-83

232 feotia T P S and S Natud 1966 Biotoomics of the rice skipper Parnaraniathiis Fabricius (Lepidoptera Hesperiidac) Indian J Entomol 28181-186

233 Teran F 0 1971 Rice pests of Santa Cruz [in Spanish Ministerio de Agricultura Sita Cruz Bolivia Bull 45 8 p

234 Than Itun 1976 Population dynamics of the yellow rice borer Tryporza itcertulas (Walker) and its damage to the rice plant MS thesis University of the Philippines at Ios Bafios Laguna Philippines 92 p

235 Uichanco L B 1911 Pests Pages 55-74 it Rice Philipp Agric 11-200

IRPS No 123 January 1987 41

236 van der Laan P A 195 1 Life history of the rice seedling fly Contrib Gen Agric Res Stn Boor Indonesia 118 15 p

237 van der Laan 11A 1959 Correlation between rainfall in the dry season and the occurrence of the white rice borer (Scirpolhaga innotala WIk) in lava Entomol Fxp Appl 212-20

238 Van Driesche R G 1983 Meaning of percent parasitismin studies of insect parasitism Fnviron Entomol 121 I1- 122

239 Venugopal M S and A litsingcr 1984 Fffect ofcarbofuran on rice growth Prot Ecol 7313-317

240 Vietnam Directorate of Rural Affairs 1962 Annual work progress report on crop improvement program of rice sugarcane vegetable and field crops (lor the period from July 1961 to June 1962) (11) 299 p (Rev Appl Entomol A51526-527)

241 Wan M T K 1972 Observations on rice leaf- and planthoppers in Sarawak (Malaysian Borneo) Malay Agric 148308-335

242 West Africa Rice I)esclopnient Association 1978 Annual research report Vol 2 Statistics Monrovia Liberia

243 West Africa Rice Development Association 1979 Annual research report p47 Monrovia Liberia

244 Williams ) F and CS lofgren 1981 Fli I illy l-468 anew bait toxicant for control of the red imported lire at Fla Entomol 64472477

245 Williams I) 1 1970 The mealybugs OFlomoptera Coccoidea Iscudococcidac) of sugar-cane rice and sorghum Bull Entomol Res 0109-188

246 Williams D) 1 1982 Pulinaria icervi (Signoret) and its allies ott sugar cane and other grasses Btll Entoinol Res 72111-117

247 Wirjosuiardjo S 1976 Study ofchenmical cont rot of insect pests ott upland rice Imiu Pertanian (Jogjakarta) 1323-329

248 Wirjosuhardjo S and N Wirarta 1976 Chemical control of the white grub tIolorithiatlltriin upland rice in M uneng Irobolinggo Indonesia) [in Indonesian] Imut Pertanian (Jogjakarta) 1330-332

249 Yano K T Miyake and V F Eastop 1983 The biology and economic importance of rice aphids (Henuiptera Aphid idae) a review Bull Entomol Res 73539-566

250 Yelp D and P Rhodes 19(4 lhe success of drilled dryland rice iii the Fiji Islands International Rice Commission Working Party on Rice Production and Protection Manila Philippines 3-10 March1964 Food and Agriculture Organization Rome Italy 13 p

251 Young W R 1981 Fifty-five years of research on the sorghum shootfly Insect Sci Appi 23-9

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No 101The economics of hybrid rice production in (hina No 113 Boron toxicity in rice No 102 Rice ratooning No 114 Energy analysis rice production systems and rice research No 103 Growth and development of the (Jeep water rice plant No 115 Production risk and optimal fertilizer rates an application of the

No 104 Faridpur acomputer-assisted instruction nmodel forrainfed lowland random coefficient model No 116 Consumer demand for rice grain quality in Thailand Indonesia andrice

No 105 A readingand listening comprehcnsion test inEnglish for non native the Philippines speakers applying for tiraining at IRIRI No 117 Morphological changes in rice panicle development a review of

No 106 Rice grassy stunt virus 2 a new strain of rice grassy stunt in tire literature

Philippines No 118 IRRI-Korca collaborative project for the development of cold-No 107 Physical losses and quality deterioration in rice plstprodnCtion tolerant lines through anthei culture

systemis No 119 Problen soils as potenial areas for adverse soils-tolerant rice No 108 (olptblication ofIRRI materials a survey of translators and varieties in South and Ssuthcasi Asia

publishers No 120 Changes isinall-farm iLtireliing techii)hgy in Thailand and [he No 109 Classification of Philippine rainfall patterns Philippines No 110 Contributions of modern rice varieties to rutrition in Asia No 121 Landforms and modern rice varieties No Ill Changes in rice breeding in 10 Asian countries 1965-84 No 122 Yield stability and modern rice technology No 112 Design parameters affecting the performance of the IRRI-designed

axial-flow pump

Page 8: UPLANDRICE. INSECTl--1i M T -E!R.EOLOU t RTANCE ,AND …delphacid.s3.amazonaws.com › 6958.pdf · The significance of abiotic factors in upland rice insect ecology will be apparent

8 IRPS No 123 January 1987

Table 1 continued

Order Family

Tenebrionidae

Elateridae

Languriidae Chrysomelidae

Chrysomelidae

Coccinellidae Curculionidae

Lepidoptera Lymantriidae

Amatidae

Arctiidae

Limacodidae Eupterotidac Noctuidac

Species

Anomaa volita (Blanchard) Adoretus caliginosus Burmeister Adoretus comprcssus (Weber) Papuana huhneri (Fairmaire) Papuana incrmnis Prell Popillia capricollis Hlope Gonocephalu depressur (Fabricius)Gonocephahm acuiangulum (Fairmaire)Gonocephalum sinphx (Fabricius) Epitragus sallei Champion Anaedus ptnctatissimnts ChampionAeolus spp4griotes spp Agriotes mancus (Say) Conoderts spp Aeoloderma brachmana (Candeze)Wireworni unspecified Anadastus filijf rmis (Fabricius)Dicladispa viridicanca(Kraat z)Trichispasericea (Guerin)Oediopalpastervalis (Weise) Oediopalpa guerini (Italy) Chaetocnema cvlindrica (Baly)Chactocnema denticulata S tephens Chaetocnemabasalis (Baly)Aulacophora sp nr similis Olivier Monolepta caripemne Ialy Monolepta bilasciata (Ilornstedt) Monolepta signata Olivier Cerotoma airojcsciata J acoby Diabrotica adelpha Ilarold Diabrotica balteata Leconte Diabrotica graminLa ltaly Diabrotica speciosa (Germar) Diabrotica limitafa quindecinmunctata(Germar)Diabrotica melanocephala (Fabricius)Diabrotica spp Altica (= Fondia)madagascariensis(Allard)Altica spp Epitrix cuctmeris (Harris) Epitrix spp Oulenjo or pae(Kuwayama) Disonchq (= Donacia) spp Flea beetle unspecified Chnootriba (= htIilachna)sinilis assimilis Mulsant Atactogaster iduriens Walker Hypomeces squtamosus (Fabricius) Tan vinecus discoidalis GyllenhalPhaulosoinusmusculinus (= mnus) Csiki Neobaridia anplitarsis CaseyPsalis pennatula (Fabricius) Laeliasufihsa (Walker) Euproctis irgtncula (Walker)Euproctis minor (Snellen)Euproctis xanthorrhoea (Kollar)Amata sp

Diacrisia obliqua (Walker) Creatonotusgangis Linnacus Latoia (= Parasa) bicolor (Walker)Nisaga simplex Walker Sesamia botanephaga Tams amp Bowden Sesamnia nonagroides botanephagaTfums et Bowden Sesamia calamistis llampson Sesamia inferens (Walker) Spodoptera eridania (Cramer)Spodoptera mauritia acron)ctoides Guenee Spodoptera frugiperda ( ESmith)

Distribution Reference

Asia 82 Asia 82 Asia 82 Asia 82 Asia 82 Asia 78 Asia 82 117 178 Asia 82 117 Africa 82 Latin America 203 Latin America 203 Latin America 195 Latin America 195 Latin America 203 Latin America 195 Asia h Asia Latin America 60 73 117 Asia 200 Africa 36 Africa II Latin America 6166 68 Latin America 6166 68 Asia 225 Latin America 61 66 68 81

Asia 178 Asia 55 Asia 55 Asia h Asia 55 Latin America 203 Latin Amefia 203 Latin America 203 Latin America 203 Latin America 6166 68 Latin Amcrie 195 Latin Amw 195 Latin Aierwca 61 81 Africa 57 Asia 178 Latin Americ 203 Latin America 81 Asia 82 Latin America 195 Asia 120 Africa 6 11 Asia 185 Asia 120 Africa 82 Africa 82 Latin America 66 68 195 233 Asia 117 Asia 117 Asia 117 Asia 117 Asia 117 Asia A S Pradhan

unpubl dataAsia 82 Asia 82 Asia 82 Asia 182 Africa 12 Africa 6 36 Africa 6 8 11 12 36 Asia 59 82 84240 Latin America 150 Asia -0 59 196 Latin America 28 61 66 68 81 150

152195203219233

9 IRPS No 123 January 1987

Table I continued

Order Family Species Distribution Reference

Spodoptera exigua (Hubner) Spodoptera exem pta (Walker) Spodoptera ornithogalli (Guenee)

Africa Africa Latin America

32 217 32 33 36217 150

Mythimna loreyi (Duponchel) Asia 30 Mythimna separata (Walker) Mythimna (- Pseudaletia)latifascia [= adultera

Asia Latin America

178 195

(Schaus)] (Walker)Mythimna (=Pseudaletia) sequax (Fabriclus) Latin America 68 Mythimna rosellinea (Walker) Asia b Mythimna yu (Guenee) Asia b Mocisfrugalis (Fabricius) Asia 30 117 Mocis latipes (Guenee) Latin America 66 68 149 150 203

Platysenta (=Spodoptera) compta (Walker) Asia 219233A Barrion unpubl data

Agrotis ipsilon (Hufnagel) Latin America Africa

33 68 81 150 1)5 203

Agrotis spp Latin America 60 Achaeajanata (Linnaeus) EldanasaccharinaWalker

Asia Africa

117 36

Armyworm unspecified Asia 72 84 Pyralidae Cnaphalocrocis medinalis (Guenee)

Marasmia bilinealisHampson Asia Africa

30 4559 72 84233 36

Marasmia trapezalis(Guenee) Africa 32 217 Marasmia (=Sumia) exigua (Butler) Asia 220 Marasnianmralis (Walker) Asia b Marasiniapatnalis Bradley Marasiniaspp

Asia Asia

b b

Leaf roller unspecified M2liarphaseparatellaRagont

Asia Africa

72 12 57

Maliarphasp Asia b Chilo auricilius Dudgeon Asia b Chilo partellus (Swinhoe) Chilo polychrysus (Meyrick)

Africa Asia

12 92 59 72

Qhilo diffusilineus J de Joannis Africa 6 1229 Chilosuppressalis(Walker)Cibozacconius Blezzinski

Asia Africa

59 72 6 11 12

Cravubus spp Latin America 203 Acigona loftini (Dyar) Latin America 203 Acigona chrysographella(Kollar)Scirpophagaincertulas(Walker)

Asia Asia

82 59 72

Scirpophagainnotata (Walker) Asia 59 200 Scirpophaganiuella (Fabricius) Asia b Elasinopalpuslignosellus (Zeller) Latin America 28 60 61 66 81 150

Diatraeasaccharalis(Fabricius) Latin America 162195203219233 28 66 81 111 150 195

Rupela albincllaCramer Latin America 111

Gelechlidae

Stem borer unspecified Undetermined stem borer nr Maliarpha BrachniaarctraeaMayr

Asia Asia Asia

72 b 55

Brachmi spp Africa 32 Hesperiidae Parnaraguttata Bremer et Grey Asia b

Parnaranaso Fabricius Asia 55 Pelopidasagnaagna (Moore) Pelopidasmathias (Fabricius)

Asia Asia Africa

164 36 155217

Pelopidasconjuncta conjuncta (Herrich-Schaffer) Asia b Borbo fanta Evans Africa 6

Satyridac Melanitisleda ismene Cramer Asia 84213 Mycalesis asophisHew Asia 30

Diptera Cecldomyiidae Mycalesis spp Orseoliaoryzae (Wood-Masn)

Asia Asia

b 72 88 131 181

Orseoliaoryzivora Harris et Gagne Africa 61132218 Chloropidae Steleocerellus (=Mepachymens) ensifer (Thomson)

Oscinella spp Asia Asia

205 205

Gaurax sppChloropsoryzae Matsuntura (= kuwonae Aldrich)

Asia Asia

205 82

Mucldae Atherigonaoryzae Malloch Asia 30 82 117130 Atherigona exigua Stein Asia 82 117 178

10 IRPS No 123 January 1987

Table I continued Order Family

Diopsidae

Agronyzidae

lHymenoptera Formicidae

Species

Atherigona oritentalisSchiner Atherigona indica MallochDiopsis longicornis(= thoracica)Macqurt Diopsis spp Pseudonapnyza asiatica Spencer Pseudonapomyza spicata (Malloch) Agronoza orvzae MunakataAcromyren landoltibalzani(Emery) Acromyrmex landoltifracticornis(Forel)Acromyrnex heyeri Forel)Atta bisphaericaFore Atta capiguaraGoncalves Atta laeLigata (F Smi h1) Atta opacicepsBorgmeierSolenopsisgeminata (abricius)Pheidole sp Pheidolegeton diversus Jerdon Monomorium pharaonis (Linnaeus) Ants unspecified

Distribution Reference

Asia 82 Asia Africa 82205Africa 11 1248 Africa 217 Asia 213 Asia h Asia bLatin America 6168 195 Latin America 61 195 Latin America 61 68 195 Latin America 61 68 195 Latin America 14 61 68 195Latin America 61 68 195 Latin America 61 195 Asia b Asia b Asia 1) Asia b Asia 120

aG orlentalls formerly recorded in Asia as G africana thPhilippines reported in this study CSubterranean root feeders

subterranean nests Foraging ants will feed on germinatedseeds only when the supply of ungerminated seed is low However these ant specialists feed on a wide varietyplants and thus help control grassy weeds (39) This dual

of

pest benefit role extends from tending ants which protecthoneydew-excreting pest species including root aphids and mealybugs (workers repel potential predators and even selectively kill parasitized aphids) to predators of leaf- and stem-feeding rice pests such as leaffolders armywormswhite grubs and stem borers (1 17)

The upland rice environment is particularly well suited to ants because of the lack of flooding Frequent tillage perpetuates grasses and granivorous species that are highlyadapted to constructing new nests in recently tilled fieldsAnt nests however are relatively shallow and tilling with a moldboard plow can destroy them Also upland ricefields are uually small and isolated and a given area in a smallfield has more field borders than it has in a larger fieldallowing high rates of infiltration from more sedentaryspecies such as Plieidole that do not colonize tilled fields but nest in soil under shrubs and trees along field bordersPheidole workers forage into fields from borders and store seed in their nests In slash-and-burn or other no-tillage rice cultures nests would be preserved in the field and possiblylead to high rates of seed predation However granivorous species do not prefer forest habitats

Damage is characterized by reduced usually patchy plantstand (120) Seed-thieving ants greatly prefer rice but they can be selective regarding plants they collect and store Theyharvest more seed than they can consume so patches of germip ing surplus seed (such as Rottboellia) will sproutfrom their nests during the rainy season

In Latin America leaf-cutting ants (Aita spp and Acromyrmex spp) occur only in upland fields and defoliate

young rice plants (47 68) They take leaves to fungal gardensin underground nests where fungi predigest the plant material

Termites Subterranean termites of the family Termitidae which lack symbiotic protozoa to help digest plants are the most frequently mentioned termite pests of upland rice (1185 120) This family cultures fungi in special undergroundcells --- fungal combs made of half-digested plant material (117) The fungi inoculated onto the combs break down plant material into food that termits can digest The termites then consume the fungal combs These grasslandtermites build nests below the plow layer in uplandricefields The colonies attack living plants only when dead plant material is not available They will attack a droughtshystressed crop but prefer older plants having greater cellulose content They will damage newly planted crops where clean culture has removed vegetation

Infestations are worse in deep light-textured soils with low moisture content The first sign of damage is yellowingof older leaves (195) Termites feed on roots (the plantyellows then wilts and finally collapses) and germinatingseed (loss of stand) or move above ground at night to cutseedlings at ground level which they cover with soil for later consumption Soft-bodied termites are highly sensitive to desiccation They live within a self-contained system of airtight chambers and build surface tunnels linei with nudand body secretions (carton) to maintain more than 90 relative humidity Termites can be located by their tunnels

Termites apparently can withstand limited submergence (157) and can be a pest of dry lowland seedbeds (117)perhaps by having nests below perched water tables

Termites appear to do more damage in Latin America where large ricefields are plowed with tractors deprivingtermites of preferred vegetation and forcing them to live on

rice plants In Asia upland ricefields are typically small with Iprge field border areas (ecotones) where termites can forage for preferred food other than growing rice plants All grassland termites do not feed on rice In an upland area in Batangas Province Philippines ricefields are commonly infested by Hosiititernesluzonensis (Oshima) however this termite does not damage rice In the same province though Mrroternesgivus(Hagen)feeds on rice seedlings (178)

In Africa termites may damage rice even though fields are small and surrounded by perennial vegetation These termite species perhaps prefer rice over other hosts

Mole crickets Nymphs and adults are nocturnal and burrow through soil feeding on roots of a wide variety of plants or forage above ground as predators of insects Adults live 3-5 mo and are even cannibalistic Mole crickets prefer low-lyin moist upland soils with high organic matter (II 81) In the lowlands they inhabit rice bunds or nonflooded fields Adults are highly mobile and can leave a flooded field to locate a more suitable habitat Losses show as wilted plants (from root feeding) or reduced stands (120) Damage is more common near field borders where mole crickets relocate after tillage (126)

Fiehl crickets Nymphs and adults of field crickets have similar nocturnal habits and damage rice as much as mole crickets do Piles of weeds removed from fields attract them They make subterranean nests and tunnel through the soil to feed on roots Some species prefer seed to roots Others feed at the base of stems causing deadhearts

Tunnel entrances surrounded by excavated soil are easily seen Young plants are cut at ground level and stored in underground cell Field crickets tre highly polyphagous as are all soil pests (117) Sachan et al (198) reported that field crickets prefer maize to upland rice Like mole crickets field crickets cannot survive in standing water and are therefore more prevalent in upland fields

Whitegrubsandhackbeetles Scarab beetles that feed on living roots as larvae but not as adults are called white grubs or chafers (Melolonthinae Rutellinae) and those that feed as adults but not as larvae on plant crowns are called black beetles (Dynastinac) In the tropics chafers or white grubs have a I-yr life cycle starting with adults emerging from the soil about I mo after the first downpour of the rainy season (134 159) They lay eggs at the same time farmers sow upland rice The rice passes its most susceptible stage and damage is mostly avoided when the white grubs are small After several months the long-lived larvae are large enough for two or three to denude the root system of mature rice (223) This intensity of damage is rare but wilting occurs when root loss is combined with drought stress

The larvae need damp (not saturated or dry) soil to survive and pass the unfavorable dry season 05-2 m underground in aestivation in the moist soil The first heavy rains of the season (20-30 amd) stimulate the grubs to resume activity and after several weeks they develop into

IRPS No 123 January 1987 11

pupae and adults and eventually dig their way to the soil surface and fly to nearby trees to seek food and mates The adults of most white grub species feed only on trees but in some species notably Eutheolahumilis the adults defoliate and cut off stems at ground level (28) Putheola hidentata damages sugarcane fields in Venezuela (121) Floods kill white grubs (208) Grasslands can support large populations and therefore white grubs can be more abundant in newly planted upland ricefields that were previously fallow (81)

Black beetles live up to a year adults can attack a crop at any stage by burrowing into the soil (127) and feeding at the base of stems to cause whiteheads (207) Adults emerge with the early rains and are normally more abundant and do more damage on a young rice crop with small root systems (38 81)

The larvae feed only on dead organic matter in dryland fields and do not attack rice Adults arc highly mobile and although sensitive to flooding can invade a field soon after it drains (115)

Root aplhids Several genera of migratory aphids feed on the roots of upland rice (249) Populations build up more in light-textured soils with high percolation rates Damage has also been reported from dry seed heds of lowland rice (52) Tending ants -- Pheidhle Cren aster Tetramoriuw lsius Tapinoma -- are necessary for root aphids to multiply Aiits harbor aphids in their nests over winter or during unfavorable periods and relocate them on rice plants digging tunnels along the root systems to allow these softshybodied insects to penetrate the soil Most intensive studies on aphid life histories have been done in temperate environments (158 222 229) Root aphids fly to rice plants at the beginning of the rice season and pass through several generations continually developing winged forms to relocate on alternate grassy hosts until winter when they move to perennial hosts usually trees Yield loss occurs mainly through reduced tillering but infested plants become yellow and stunted and in extreme cases wilt (227) Aphid populations build up gradually so damage usually begins in the late vegetative and reproductive growth stages

Root-feeding ineal tugs More than half of the mealybug species or rice feed on roots (245) Being soft bodied mealybugs are not adapted to living underground but they survive because tending ants (the same species that tend root aphids) dig burrows for them and move them from plant to plant Well-drained soils ihelp mcalybugs survive Although six species are recorded as subterranean root pests of rice we found no report citing damage Root-feeding mealybugs are probably more prevalent on perennial grasses than on short-lived annual rice

The most common rice mealyb ig Brevennia rehi is a foliar feeder favored by lack ol rain apparently it is responsible for high yield loss in Northeast India and Bangladesh (N Panda pers comm as stated by IRRI [1061) This report needs confirmation Mealybug damage was only briefly mentioned in another report (156)

12 IRPS No 123 January 1987

Root weevils A few species of upland weevils (IHporneces Donacia Atactogaster) attack rice roots as larvae and feed on stems and leaves as adults Damage ispatchy The larvae are not adapted to submergence and are true upland soil weevils isopposed to the so-called water weevils They rarely become sufficiently abundant to cause damage Adults on rice foliage or the soil surface show that the roots are being attacked by the grubs Root weevils are highly polyphagous (185)

Root heeths Soil-inhabiting false wireworms Gono-cephalumt spp are reported in upland ricefields The adults and larvae normally feed on decaying organic matter but land preparation for upland rice removes these sources The larvae feed on seedling roots and nocturnal adults roam on the soil surface to cut off seedlings at ground level (82) This group of ground beetles is highly omnivorous adults even prey on stem borer larvae (183) True wireworms (Elate-ridae) feed on living roots but rice as a host has been reported Infrequently (117) In Latin America the highlypclyphagous larvae of Diabrotica rootworms are also recorded on rice

Root htus Upland rice seedlings can be attacked by the adults anI nlViphs of chinch bugs Blissusand Caenoblissus and bro n bug Scaptocoris which feed on the roots with theirsucking nouthparts (80 83) Three genera of burrower bugs A tms (eotonhus and Stiharopus -are common iil grasslands in the Philippines and may attack the first crop (rice or maize) planted after the dry season Chinch bugs are most abundant after a dry spell These insects are large and can kill seedlings or stunt and reduce tillering on older plants I hey prefer other hosts and rarely damage rice Adults can also feed oii developing rice grains (81)

(utworms Soil-dwelling noctuid larvae hide under-ground during the day to avoid predation by birds and bccone active at night above grour cutting off you ing plants at ground level (hence tile naiie cvtworn) arvaedrag severed plants into hurrows llie larvae have wide host ranges arid pupatc in the soillgrotis ipsilon occurs worldwide and appears to be the most common cutworm pest of upland environments but is more prevalent inlowland rice particularly in dry seedbeds

IFoliar pests Other groups of insects are adapted to upland environments because they pass one growth stage underground either as egg (in heinrietabolous species) or pupa (in holomnetabolous groups)

Rice seedlibig maggots AitJeriota flies occur in non-fhoodcd environments in Asia and Africa (180) Feeding of larvae one per iller causes deadhearts Flies as opposed to moths only secondarily evolved as pliytophagous pestsfroni a saprophagous Origin Athrnonalarvae secondarihv feed onidecaying tillers whereas larvae of stem borer moths actively Iced on living tillers to cause deadlicarts Seedling maggots will die if they cannot sever a tiller on which to feed

as it decays Plants die under heavy attack whereas lighter infestations cause stunting delayed maturity and ragged leaves (236)

Eggs are laid only on actively growing plants (49) aid damage occurs wiihin the first week after crop emergence(170) Larvae pupate more in stems than in soil Seedling maggot attack is highly seasonal (206) normally peaking 2-3 mo after the onset of the rainy season These pests possiblyaestivatc over the dry season Rice isonly one of their many graminaceous crop and weed hosts

The rice stem maggot (hlorops orzae is adapted to temperate upland and lowland rice growing regions of Asia In Japan and China upland rice grows in mountainous regions a habitat of this chloropid fly The larvae tunnel into tilestems The stem maggot is more damaging during the seedling stage larvac hibernate in grasses

File several dipterans reported as stem maggots in dryshyseeded lowland seedlings inIndia (205) could presumably attack upland rio

Leaf urinrs Icaf miners are common on wheat and maize but are rarely reported on upland rice (213) Three leaf miners are more prevalent on upland than lowland rice inthe Philippines Pvudonalomy a asicanh Vficataand lIAgromrza orzae attack at the seedling stage The larvae develop inside the tunnels they construct in the parenchyma tissue P Vpicata and Aorvzac prefer maizeEehtsine and other grasses to rice spicatacan become abundant on wheat

Leaf beetles Chrysomelid and coccinellid eaf bcetles scrape or otherwise remove leaf tissue as larvae or adults or both This damage accelerates desiccation of plants Tile larvae of sonic species are root feeders Flea beetles Epitrix and (lraetocnetna and rootworms l)iahroticaspp feed on rice foliage only as adults Their root-feeding larvae have hosts other than rice The adults and larvae of the hispa-like Oediolmlpa spp beetles of Brazil and (hmootriha (=EPfishyachra) beetle of Africa feed on upland rice leaves Oealioshypapa spp larvae are leaf miners (mootriha is also pshyvalent in lowland environments (11) Leaf beetles prefer vegetative rice (81)

The rice hispa Dicladipaarnigera is not reported on upland rice and prcers more aquatic habitats In Central Africa however I)cladispm viridicyaiea attacks uplandrice in the vegetative stage as well as lowland seedbeds 7richipasericea is reported on upland rice in West Africa and Madagascar (57)

The rice leaf beetle 1d1ea 01izae riiuech like Chioropsor|rzae is adapted to temperate regions of Ashi and is usually a pest on lowland rice but seldom oi inpland rice in the mountains (82) The adult hibernates on grasses

Armmworms Species of grassland-adapted SpoptloeraPlatysemta and A ithimna larvae create widespread epishydemics They can defoliate ricefields generally in patchesfrom early vegetative growth to harvest (41) Armywornms pupate inthe soil and highly favor upland rice (34) Adults

can disperse long distances to colonize even remote upland areas They become more abundant than their natural enemies particularly after favorable rains following a prolonged drought (196) The drought kills the armyworm and natural enemies (mainly parasites) but the armyworm can recolonize rapidly

The natural increase insoil fertility from mineralization of soil N over the drought period promotes luxuriant plant growth to foster a rapid buildup of these highly fecund species Weeding rice increases tilelikelihood of damage The parasites normally return to control the armyworms but not until after serious damage

Outbreaks on upland rice have been reported in Ghana (3) inzibar (38) Central Africa (36) lPanama (165) Brazil (28) Malaysia (196) and India (167) Defoliation may be severe often leaving only the base of stems Armyworm larvae hide during the day under litter or in soil cracks At night they ascend plants to feed Larvae pupate in the soil

Thrips Thrips frequent upland rice but damage is less than in the lowlands Thrips larvae and adults feed on leaf blades by rasping causing leaf rolling and stippling Thrips prefer vegetative stage rice and their survival increases during drought (heavy rain washes them from foliage) although drought stress usually outweighs thrips damage Drought and thrips cause leaf rolling in upland rice (235) but thrips have been overlooked perhaps becaLse of their small size Thrips are most numerous indry season irrigated rice The combination of favorable plant growth and lack of rainfall is ideal for thrips In contrast the lack of rainfall in upland rice hurts the crop and eventually the thrips whose numbers must decline

In [atin America Frankinilla rodeos attacks panicles before they emerge from the boot calusing sterile grains

Stem hug~s Black bugs are the most common group of rice stem-feeding -lemiptera in Asia The most frequently mentioned species ----Scotinophara coarctaia and Shrida - prefer aquatic habitats and therefore could not occur in upland rice as sometimes reported Upland species in the same gerurs - notably S tarsalis and S scotti -- are morphologically similar and remove sap from stems of grasses and upland rice The upland species are notably more abundant in upland rice planted near forests

In Latin America tile Tibraca ima-large pentatomid tiventris causes deadhearts and whitehcads on upland rice Both adults and nymphs remove sap from internodes of plants more than 3 wk old Tillers are killed by mechanical damage and by the entry of secondary microbial infection This damage is often confused with that of lepidopterous stem borers (233) Eggs are laid on the leaves and stems Nymphs are gregarious This species can also be found on irrigated rice Drought reduces population buildup

Stiem borers Lepidopterous and dipterous stem borers are widely recorded on upland rice but cultural practices and alternate hosts determine abtndance Except for Elasmolmll)uS Iinosellts numbers usually build up toward

IRPS No 123 January 1987 13

the end of the crop growth cycle Therefore late-maturing varieties staggered planting and lack offloodingfavorstem borer buildup Many stem borer species can lie dormant in the stUbbleafter harvest and theirsurvival is encouraged by panicle harvesting (leaving tall stems) and lack of tillage after harvest

Stem borers disperse and lay many eggs Many species are polyphagous however the monophagous MaliarpiasepashyratellaDiopsis Ivmgicornis Scirpophaga incertlas and S innotata can also be abundant on upland rice (hilo Suplressalisdevelops fasterand becomes la rger when reared on upland than on lowland rice (226)

The lesser corn stalk borer lasmopal tt Ittoselhs is semisubterranean and is perhaps the stern borer most adapted to upland rice(129) Highly polyphagous it infests maize peanut and cowpea It attacks seedling rice (one larva can kill tip to four plants before maximum tillering) (195) (hiv and Maliarphastem borers prefer older plants But Scirpophagaspp will attack young plants even in the nursery lasn opalls larvae tunnel into the stems at or below the soil level mausing dcadhcarts Larvae are not found in their turnnels inside sterns beca use vhei disturbed they retreat into cases made from soil particles bound by silk The cases areattached to the tunnel entrance (143) This behavior probably evolved as protection from natural enemies Larvae pupate in the soil The lesser corn stalk borer is more abundant during drought damage is often confused with that of drought (233) This species prefers sandy soils arid its distribution within a field may relate to soil texture (143)

Diatraeasaccharalis aside from the lesser corn stalk borer is frequently a stem borer pest in Latin America According to Teran (233) rice and maize are more preferred hosts than sugarcaine its namesake

Atcitonacrvsograpla is a less known upland rice stem borer in Asia with habits similar to Easinopall s (82) because it is not found in lowlands Acutra has grassy weed alternate hosts As a moth it is often confused with the dark-headed stern borer(hi polur(standstriped stern borer (hi sup)lpsrcais

Other common upland rice stem borers are also reported in the lowlands The dominant species in upland rice vary greatly with location In Uttar lPradesh India (78) and in Japan (123) the pink stem borer Sesamia htfirens is more prevalent hIKenya the principal rice stem borer is Chi partelus

(92) The lowland African sten borers Aialiarpha sepashyratellaDiopsis vngicornis and Sesamia calatnisnis are less abundant Oimupland than on lowland rice(217) but perhaps cause greater yield loss because plant injury is greater if combined with drought css (92) The stalk-eyed stern borer D)ivpsis is more prevalent in wet habitats (6 1I217)

In Central and West Africa Maliarpha and Sesamia calanistis are the dominant stern borers of upland rice (8 9 10 II) (lilo dihlfilineui and Chio partellus inhabit the

14 IRPS No 123 January 1987

upland savannas whereas Chilo zacconio prefers the lowlands (II) Aaliarpha is most abundant in the mangrove swamp habitats (91) coastal regions (16) or rain forest zones (II) of Africa and appears to be the homologue ofScirlpophaga incertulas The yellow stem borr can also be prevalent in upland rice (120 167) The sugarcane borer Eldanasaccharinais so far only a potential pest of uplandrice (91) despite Grist and evers (82) warning

Grasshotl)ers andI locitas Short-horned grasshoppers and locusts -- Locvta Patanrga Schistocerca - layeggs in the soil inhabit grasslands and develop into swarms to seriously damage upland rice from time to time (173) Largenumbers can destroy a field leaving only the stubble In low cmuoroer s giassnoppeirs an cut panicles (173) Farmers may not plant if locust swarms are imminent Prolonged dryweather followed by favorable rain favors development of large swarms (173) Usually breeding in di) grasslands theydisperse toattack crops including upland rice(191) Eggsaresusceptible to desiccation d uring drough t and toinundation during seasonal rains With favorable soil moisture over consecutive seasons natural enemies ca nnot prevent a largepopulation increase leading to migratory swarms (64)

Most grasshoppers on rice are adapted to uplands (107)but lowland rice grasshoppers iervgl(rlhusspp and Ora spp occur occasionally on upland rice (2 82) They lay eggson rice foliage Nymphs are scmiaquatic

The gryllid Lsctrtus coucinnus is a pest of lowland and deep water rice in Bangladesh and Thailand In the Philippines it damages upland rice grown in isolated pockets near lakes The gryllid nymphs and ad ults feed on the central portions of folrage leasing only the midrib and leaf nargins

Leal ler Pvralid moths whose larvae fold leaves to make a feeding shelter occur worldwide (217) but arc appareritly most common in Asia Leaffolde rsattacking rice in Asia are a complex of species (26) (Ctalhahcrcisinedinalis the best-known rice Ia flolde r in Asia may prefer grassy weeds over rice (86) It is a late colonizer of uplandrice U1arasmja eviua a lowland species (220) nreflers rice to grass specres (86) and colonizes upland rice during il early growth stages llarasnria patnalis is the most corn-rnonly encountered species in upland rice in the Philippines Perennial grasses are their alternate hosts and they can survive year-round in upland rice environments The leaf-folders 11arasmia ruralisand Alarasmia spp are least aburndant Dormancy is unknown in leaffolders Cnaplha-ocrocis medhnalis however is known as a migrant (90)Leaffolders remove photosynthetic tissue attack during the flag leaf stage is particularly injurious Popurlations are normally held in check by natural enemies but leaffolders become parlicularly abundant in conditions of high plantfertility and shade

Butterflies Skippers PelopidasInathiasand Borhofanta are more prevalent in African lowland than in uplandhabitats (6) but extensive studies in Japan and China

showed that Pelopidas mathias prefers an upland habitat (155) Parnaragutata seasonally migrates between lowland and upland areas and is equally abundant in both environshyments (164)

The green horned caterpillar Melanitis leda ismene is more abundant in lowlands (212) but our experience in the Philippines suggests that Pelopidas niathiasand 11elanifis leda ismene are often more abundant in uplands than in lowlands Bamboo iaunbuiva spp annually sustains P mathiaspopulations in the Philippine uplands Adults feed on nectar from flowers and migrate to upland rice during the late vegetative stage

iPi erlled h i-1 1d dill L tIC i 7-0 l(become numerous probably because of high egg predationEgg and larval parasitization are often low Butterflies can readily disperse long distances to seek renlote uplandricefields However they have low biotic potential and even it predation rates are low the rarely become abundant Upland environments with their diverse microhabitats can provide more favorable sites than lowland rice plains

tutterfly pests of rice have broad host ranges and become dormant Juring urnfavorable times of the year

IolhagrotslaidopwraThe larvae of many polyshyphagous moths are reported to defoliate upland rice tussock moths lsaispelnnatula Iawlia sUifita luproctisirgtwltaE mnorand P vathorrhoeaslug caterpillar

litotiahicolorhairy caterpillars Nisaga sinp vandA mata spp and wooly bears l)iacrisiaohliqtia and Crvatonotts gangis Many of these polyphgous larvac migrate and are transients in rice including HelhWhis arm iera and 1t zea highly pol phagous species (82)

Gall nidges The Asian rice gall midge Orseoliaorrzae (88 131) and its African counterpart 0 ortzivora(6 11 12 32217) prelerlowland to upland rice Gall midges in rainfed areas are prevalent only during tle rainy season and dispersepoorly Their alternate hosts (wild rices LeersiaIschaetnuan and Pasahmt)are mostlyaquatic grasses and are less likelyto be near upland rice areas In an area of mixed uplandfloating and rainfed lowland rice cultures in northern Thailand upland rice which is ilanted a month earlier was attacked firstbut subsequcent populations were much lower than in rainfed lowla id or floating rice Low numbers of gallmidges passed the dry season orr perennial weeds such as Paspahim (88)

Smot (hid(s Reports of aphids attacking upland rice plants are few (249) Without maize the maize leaf aphidRho alosihhunm maidis attacks upland rice in the Philipshypines The plum peach aphid IIrsteroneuraselariae was reported in Sierra Leerne toinfest leavesand tnriperied grain(7) Siphaglrctriaecaused transverse linrea r necrotic striae iir rice leaves in Italy (172) I-lea v rains held populations in check Rice is not a preferred host for shoot aphids

Leafhoper andI )anthol)pers Cicadellids and delshyphacidsdispersecasilyand readilycolonie upland ricefrom nearby lowland areas A study in Sarawak (241) showed

IRPS No 123 January 1987 15

similar species in upland and lowland rice Nephoiettix The principal vectors of rice virus diseases generally have species dominated The most prevalent species accounting narrow host ranges and alternate hosts in the grass family for 90-95 of the sweep net samples wre N virescens N are mostly annuals dying in the dr3 season Some virus umropictus Nilaparvia hgens Soatellafiircifera Cfana vectors such as BPH can disperse lontg distances but few do

sectraand Recilia dorsalisLowlands had higher popula- Th restriction of upland rice and weed hosts to the rainy tions Katanyukul and Chandaratat (120) monitored hopper season the small area planted to upland rice and the hopper numbers in upland and lowland fields of northern Thaiiand vectors having narrow host ranges all work against virus and found similar results A comparative study of plant- diseases in upland rice hoppers in Fiji showed more brown planthoppe BPH) and In Latin America Sogctdhs o(rzicola and S cuhanus whitebacked planthopper (WBPH) in wetland rice than in transmit hoja blanca virus disease but reports on upland lowland rice and hopperburn was prevalent in lowland rice are rare except in the favored upland areas of Colombia areas with standing water (89) Peru and Venezuela (35 42) S rricolaocurs in Brazil

Hopperburn in upland ricefields in Asia is rare (84) In on upland rice but hojaI blanca has not been reported (67) Batangas Philippines isolated fields of upland rice become S orzicola has a wide host ramgc (10) similar to WBPI- in hopperburned infrequently from WBPH A report from Fiji Asia WBPH however is not known to transmit any virus cites hopperburn from BPH and WBPH (250) In these two disease situations upland rice was grown on highly fertic soil and In Africa pale yellow mosaic virus is endemic to swampy N fertilizer was applied Hoppers respond to better nutrition lowland areas where rice ratoons year-round (19) and is Normally upland rice is grown on poor soil and receives no mechanically vectored mainly by hispid beetles A minor fertilizer vector a flea beetle Chaeiocnema spp is an upland rice

In Asia upland rice ras are normally not far from pest and upland ricetields near endemic areas occasionally lowland ricefields-- a source of hoppers Otherwise hopper contract the virus species with greater polyphagy than BPH or Nelhotettiv Spittle bugs Cercopid nymphs producc a protective virescens would be dominant frothy covering that looks like saliva In Latin America

In Africa leathoppers and planthoppers do not become spittle bugs damage upland rice In parts of Brazil where abundant on upland rice (6 II) probably because lowland upland ricefields are surrounded by pastures adults rice is not widely rlanted nearby The only reported migrating from grasses can damage rice severely and cause hopperburn as from Nilapanrata maeander in breeder complete hopperburn in a young rice crop within a week of plots receiving high N rates in Nigeria (98) Hoppers colonization (194 195) Dispersal to rice occurs even if however are a greter problem in Latin America where pastures are verdant and spittle bugs prefer pasture grasses large grassland areas breed polyphagous species (Gra- to rice (168) Nymphs are rarely abundant enough to phocephala spp Hortensia spp lEvitianus ohscurunervis damage rice Regular rainfall favors their development Balclutha spp and Draeculacephala spp) vhich then (233) disperse to cause hopperburn on seedling rice (68) Rosetto et al (195) reported Graphocep7hala spp and Hortensia spp Seed pests to be equally abundant on upland and lowland rice Alydid and pentatomid bugs are seed pests of upland rice

The leafliopper and planthopper epidemics in Asia that worldwide In Asia Leptocorisa sp prevail Their habits caused severe losses from hopperburn and vijus dicases make them particularly suitable for upland rice 1)thev feed over the past several decades occurred principally on on many grasses (and therefore can survive the earl y wet lowland rice Except in a few isolated cases (160) upland season before rice sets its grain) 2)adults are long-ied and rice has been spared from tungro grassy stunt and ragged mobile allowing them to find isolated plantings 0 rice stunt virus diseases vectorcd by green lcafhopper (GLH) (besides seed bugs only birds and rats appear to have shyand BPI] (123) Virus diseases have occurred in upland rice capability) and 3) adults aestivate in forested areas or where lowland rice was nearby Rice dwarf disease however sugarcane fields during the dry sca son when neither rice nor was first recorded on upland rice in the Philippines (187) grassy weeds are present (I199) Rice seed bugs concentrate Virus diseases can be perpetuated only on livir plants or on small-scale upland riceficlds because they can actively insect vectors and no living rice - plans tooned or search them out The prevalence of groves of trees characshyvolunteer - grows year-round in upland arcas excert in a teristic of upland environments ensures nearby aostivation few with 12 mo of rainfall sites keeping the seed bugs close by

The smaller brown planthopper Laodcelphax striatellu Leptocorisae are uncommon in lowland rice plains occurs in upland rice in Japan (18) It spreads black- because 1) their populations are diluted in a sea of rice streaked dwarf and stripe viruses between winter wheat and planted more or less at the same time (they ca n only feed on barley to lowland rice Early planting of lowland rice rice during the milk to hard dough stages) 2) they lack nurseries spreads these viruses but upland ice has remained aestivation sites and 3) weedy fallow areas are limited uninfected The lowands of Asia are dominated by L oratorius

16 IRPS No 123 January 1987

which lays its eggs hign on tile foliage (199) L acuta and L solonlonlesis oviposit at ground level the former on litter and the latter loose on the ground These egg laying habis explain theirtenvironmental prefcrencecs(214) In Japan the postdiapause adults of L chinensis migrate from moun-tainous areas to upland ricefields during floweiing (1 14)

Pentatornid bugs prevail in Latin America (8182 195) and in Africa pentatomid and coreid bugs arc equally cited (6 82) l)amage is often characterstic of the species (5)rcoris ventralis and Mnida spp are the common

pentatomid seed bugs of Philippine upland rice but they prefer lEchinochva spp grass seed to rice Vearaviritda occurs worldwide and hoften more abundant in the uplands(54) We saw few reports of seed bugs on upland rice but see(d bugs apparently opelrate cqually between by land and upland rice Good dispersal is necessary ior species that call feed for only 2-3 xon a crop and must first find those crops at ripening stages

Other pct species feed n flowers In Mato Grosso Brazil (194) and Santa Cruz Bolivia the panicle weevil Aeo-hariuia amllitarsis attacks rice during flowering (68 233)The larva e rice sten horers adutlts feed at the bases of spikelets caIusing empty grains Also inLatin America thecydnid -lkiituttatralusisa seed bug as anad tilt and nymph (194)

A number of beetles ecd on poilen - coccinellids lialrotia Aulac lhora and Monohpl-a (55) - but becatise riCe is self-pollinating and fcrtiliation occurs before pikelets open they pose no threat to yield

IST ECOLOGY

We now look at characteiistics of the life cycles of upland rice insect pests to lea rn hew each plays an adaptive role in upland rice ecology

Life history strategies Orzasativadoes not tolerate drodgh well so upland rice is highly seasonal normally grown in the wettest months of the year Usually rice is present for less than half of the year The rice-fiee fallow poses serious problems of survival to upland rice insects which have evolved att least four mechanisms to ovrcolne tle cvcl ic-l lack of a hostI) polyphagy 2) longevity 3) dormancy and 4) vagility (dispersal)

Poldt)agy lolyphagy is defined as having hosts of at least two botanical families oligophagy of more than one gents and ionophagy of only one ventus (40) - inour case Orvza Most wild rices however are illi tiC and theieiirc are normally far removed from upland rice hahitat othewise more would be alternate hosts for uplanJ iice insects Thte highly mornophagous rice pests shySciriol)haita hiceritlas Nilal)arvaiahiens Iand Nealo--tettiyvircsctws in Asia and Aalirpha secparatella and 1ioj)sis lngicornis inAfrica - occur on upland rice but

are more abundant in lowland culture They can specialize in rice because they are highly vagile and can attack all growth stages By specializing in rice monophagous rice pests can outcompete related but polyphagous speciesIntensive andI extensive lowland rice culture has favored monophagous species Riqwla alhinella (Cramer) the onlymonophagous species in Latin America is rarely reported on upland ice (I1)

Because of its limited temporal and spatial existence upland rice has not favored the evolution of specialized species Clhang(44)shows evidence that upland rice has only recently been cultured by man from the lowlands thus pests have had little time to fully adapt to the crop All but the lowland-adapted insect pest species have alternate planthosts to rice Root pests with limited mobility are highly polyphagous Pests that specialize in one growth stage eg secdling maggots seed bugs are oligophagous or polyphagous

Most alternat hosts of rice pests are annual grasseswhich also are seasonal Upla_rice is also a host but not the preferred one of the most highly polypliagous spcciessoil pests grass hoppers armvwarrs and Vearaviirielda

The d ry season poses igreat obstacle to ani upland riceinsects survival and only sonic perennial plant hosts are suitable food during the off-seasoin Plolyphagy allows upland rice insects to suLrx-ixe during thC rainy season when plants- both annual and perennial -areactivelygrowing and therefore more nutritious

Loiugevitr The ability of aiinsect to live 2-3 mo without undergoing dormancy helps species sich as root-feedingwhite grbs (as larvae) and secd-feeding bugs (as adtilt) to survive unfavorable periods White grubs arc not highly fecund and their life strategy is to improve survivorship of limited progeny Their subterranean habitat hides them from manv natural enemies lBtut root tissue is not highly nutritious so they ltilst eat great quantities and need a largeinsect biomass to digest it Other soil-inhabiting rootshyfeeding insects have short life cycles but are more fecund

Seed pests need greater longcvi vas adults to locate ahost at early grain dcclopment arnd give sedentary offspring time to develop Few groups of upland rice insects live longin active development Polyphagy and longevity only increase their ability to survive during the rainy seasonOther mechanisms are needed to survive the dry season

orantc In the tropics the nonrice season is the dry season while intemperate areas tile off-+season is winter Many upland rice insects undergo dormancy inlsummer (aestivation) or winter (hibernation) Dormancy is simplyinactivity during winters with prolonged temperatures below 10 C insects find shelter arnd cease movement Activity is resu med with warmer t empcera tu res A deep state of irnactivity termed diapauise occurs in sole species that prepare physiologically (havxing low metabolism and convcrting gonad s to fatty fuel reserves) for the unfavorable season (148) They enter diapaUse inanticipation of unshy

IRPS No 123 January 1987 17

favorable weather and resume activity after favorable Armyworms skippers and locusts actively travel hundreds weather begins usually in response to photoperiod or of kilometers in air fronts such as intertropical convergencethrough a delayed response to a stimulus such as heavy zones to descend onto upland rice fields far from their rainfall (237) Not all individuals of a population have the breeding grounds (31) Several generations can build ip on same threshold for environmental responses for entering or alternate hosts in tile grass family before rice is plantedterminating diapause -- this is seen as a mechanism to These large insects are active fliers Smaller insects such as prevent total mortality if a false cue appears such as a thrips aphids and planthoppers passively migrate long sudden warm spell in winter or rain in the middle of the dry distances by the wind season (154) Otherspecies ean readily travel tens of kilometers - stem

Probably more upland rice insects undL go dormancy borers leaffolders leafhoppes cutworms spittle bugs leaf than we realize Candidates are Atherigona spp root beetles seed bugs ants and termites aphids root weevils crickets leaf beetles thrips and seed The least dispersive species are white grabs mealybugSbugs Insects known to entervarious states of aestivation are seedling maggots crickets weevils and gall midgesleptocorisaspp adults (199) Maliarphascparatellalarvae Each species has evolved a unique set of attributes to (9) Scirpophaga innotata larvae (117) Chilo partelhtsv enable it to survive and adapt to the changing uplandlarvae (21) ScirpopThagainccetulas (179) white grub larvae environments Several evolutionary avenues lead to fitness (134) Patangasuccincia adults (6 82) and Schistocerca (Table 2) but there is no reason to believe one set of gregariaeggs (82) attributes is better than another (224)

Insects known to enter hibernation are LodeIphaDstriatellus nymphs (124) Chtio Tvlpressalis larivae (125) DroughtSesaia inferens larvae (146) but not S calavisis(2) Well-drained rainfed upland soils are subject to drought ifIDiaraea saccharalis larvae (75) Scirpophaga- tcerlttas rains do not fall within 2-3 wk Prolonged drought followedlarae(5 lacchrli s lae (75)saSlOhagasshoperseggs (82)and felo pidas athias pOpae (232) by favorable rains stimulates armyworm and locust outshybreaks but mealybtigs root aphids and thrips become

Lcpmotorisa oratorisand L acuta adults aggregate on numerous onlyafter dry spells for two reasons First heavy trees sugarcane or other shady moist sites L paluwanmsis rain normally kills soft-bodied foliage-feeding thrips and confined to Sulu and Palawan Philippines lives on the mealybugs and soil-inhabiting root aphids and root mealyshygrass Brachiariamutica (Forsk) Stapf in the absence of bugs Drought eliminates this source of mortality Second rice The adults can live for 5 mo When aestivating their the rice plant responds to drought by breaking down rate of metabolism drops and they do not feed They will proteins into soluble N compounds which enter the phloem however puncture plant tissues of their aestivation host in and are taken up by these sap-feeding insects (245) Greater search of water This behavior has led to errors of the host nutrition therefore leads to greater survivorship and range of rice bugs (I) Aestivation quickly terminates with fecundity rain probably meaning aestivation is not deep Small rice area

Some stem borers aestivate as mature larvae in rice Except in the highly mechanized large landholdings of stubble or in straw stacked as livestock feed However Latin America upland ricefieldstypically are patchy within moriality increases proportionally with the duration of the a highly diverse flora The small fields result in higher ratios dry season even if the stubble remains undisturbed If the of perimeter to area than with the typically larger lowland stubble is plowed for a crop following rice few stem borers ricefields Upland ricefields themselves may be intercroppedsurvive Rainfall terminates dormancy but it normally with cereals legumes or root crops Mixed intercroppingtakes several weeks for moths to develop and emerge with a wide variety of species also occ in tribal slash-andshy

Last-instar white grub larvae tunnel 1-2 m deep in the soil burn agriculture (50 74) to construct pupal chambers in which they aestivate The Upland ricefields tend to be small because of the labor larvae are very sensitive to dryness so pupal chambers are needed to clear land in slash-and-burn areas and to till land sealed to coierve body moisture Aestivation terminates to minimize weeds Rice requires more tillage than does with the first heavy rains but as in stem borer 3-4 wk pass maize because rice competes less well with weeds Uplandbefore adults develop rice is more a subsistence than a cash crop and small areas

Insects may respond differently in each region to cold (I ha) can feed a family temperature or drought Biotypes or loal populations may Small fields favor some insect pests such as some ant evolve (56) Local populations of Scirpophagaincertulasor species that can forage from more permanent field borders Patangasuccincta may have become adapted to prevailing Trees provide food and mating sites for white grubs (134) as conditions well as aestivation sites for LetTocorisa spp (199) Field

iagiliti The ability to disperse combined with high border grasses plus tIhose within a ricefield provide alternate fecundity orshort life cycles enable some upland rice pests to food for all but the highly monophagous upland rice better exploit the temporarily favorable upland habitat colonizers and a habitat for natural enemies

0 z

Table 2 Life history patterns of upland rice insects or insect groups

Host range Longevity Dormancy Vagility FertilityMonophagy Oligophagy Polyphagy Low Medium High Yes No Low Dispersive Migratory Low Medium High(Oryza spp) (grasses sedges) (angiosperms) (lt3 wk) (gt2 mo) (lt1 kin) (lt50 eggs) (gt500 eggs)

Ants xTermites x x x

Mole crickets x x x xx x xField crickets x xx

White grubs x x x xx xRoot aphids x x xxMealybugs x

x x x xxSeedling maggots x

x x x x xLeaf beetles

x xx xArmyworms x x xx x

Thrips x x xx xChilo spp x x xx xMaliarpha x x x x

xLocusts x x Xx x xGall midges x x xxiVephotettL iirescens x x xx x x xLeprocorisaspp x

x xLeaftolders x x x

Sogatellafurcifera x

xx x x x x x x

00

IRPS No 123 January 1Q87 19

All this appeals to favor insect pest buildup but the notwithstanding the release of stored nutrients in response contrary is true Loevinsohn (138) showed that insect pest to drought that temporarily benefits philoem feeders populations respond exponentially with the proportion of land devoted to rice up to about 75 of the area then the M POR ANCE I I P1A 1) RWI P1St S

rate of response declines In most upland rico areas tie rice crop occupies less than 50i of the area- therefore the Visits to upland ricefields normally reveal 1kw insect pests potential for population buildup is low This relationship Loevinsohn (138) and Locvinsohn ct al (139) compared may not hold for seed pests that can locate small ricefields insect abundance using annual light trap atchcs in upland

Another outcome of the highly diverse flora and phical rainfed lowland and irrigated lowland sites in the Philipshyenvironment of upland rice isthateach upland ricearca has pines Although only one upland site was studied data a unique composition of insect pests showed equal or fewer insects at the upland site than at the

For example in Tanauan Batangas Philippines the rainfcd lowland sites and fewer than at the irrigated sites main upland rice pests are rice leaffolder(RLF) armvworm (Table 3) M Lltiple regression analysis showed that the key Aythntnaseparata WBPH ants Soli psis gentitataand factors explaining insect abundance among a set of cropping white grub Leucopholis irrorata Sixty kilometers away in intensity variables were the number of rice crops grown per Real Quezon the main pests are seedling maggot Athe- year followed by area devoted to rice Table 3 shows that rigona orae flea beetle (Thacitcmnra hasalis thrips the upland rice site had only I rice crop perym rand that rice Stenchaetothrips hiurnmis and rice bug Leptocorisa was planted in only 20Vi of the area Fertiliter and oratorius In a third Philippine upland rice site in Claveria insecticide use had little bea ring on the trend The armters at Misamis Oriental Mindanao the main pests are ants the upland site applied 60 kg N ha higher than at most Solenopsis gemtinata seedling maggot Alieriona orvzae irrigated sites white grubs Holotrichia in)iUaiuoan t and Lucopholiv Insect occurrence at the upland site was omparable to irrorata stem borers Sesamia ifiArelr (hio auriciliusC that at the Cagayan rainfed lowland site parlicularly in 1982stpessallis Scirpophaga incertulas S innoiata and when drought prevented many farmers fron planting(only

4c ona chrrsurapella root aphids Tetraneura niriah- 40(of the rice area eventually was planted that year) M ore dunrinalisand Rhupaosiphunm riahbdminalis lcaffolders favorable weather occurred in 1980 and 1981 which Marasniapataltis and Chaphalocrocis Medinalis army- supported greater rice areas and consequently more insects worm MIthinna separata and rice bug Leltucorisa The most intensive site for insect pests was South Cotabato oratoriusThe great diversity of insect problems is typical Mindanao where farmers plant 23-25 rice crops per year fer upland rice and makes control efforts more difficult with irrigation The Zaragoa Nutva Ecija site also had particularly regarding breeding for insect resistance high pest abundance and at this site the fields were highly

asynchronous Asynchrony was the third most important Low yield potential Iactor in explaining rice insect abundance Most upland rice is grown on low-nutrient and mineral- Rice cropping intensity in upland areas is bound to toxic soils which when combined with eiratic rainfall remain low Only one rainid upland crop is possible per make upland rice a highly riskycrop for the farmerto invest year (Farmers in Claveria Misamis Oriental Mindanao in costly soil amendments and land management Often Philippines tried asecond upland ricecrop i 1984a ycarof upland rice isgrown far from markets making inputs even favorable rains but no harvest was possible b-cause of more expensive Farmers growing a subsistence crop hosi- drought blast and birds) Rice area isaso bound to remain tate to invest in costly inputs to raise yields low because of the high labor and power rcquirelmenlts to

Even having varieties that could double existing yield prepare the land (because of weeds)and because upland rice potential under present management levels would probably wvill not compete with lowland rice as a cash crop Some not prompt many farmers to use purchased insecticide upland rice sites with prolonged rainy seasons howeverare unless tile risk of crop failure were reduced such as it was planted asynchronously such as the Clacra site with irrigation systems in lowland rice The seasonal abundance of Upland rice insect pests

The Tanauan Batangas site is atypical for upland rice attracted toa light trap wasgrapied for crop year 1980-81 in The soils and rainfall pattern are favorable and because of Tanauan FBatangas (Fig I 2) W13P11 was he most its nearness to Manila farmers have cash resources ftm common rice hopper and its population peaked at the sales of vegetables to purchase tertilier They applied an midgrowth stage as it does in lowland rice D-Vac suction average of 60 kg Nha after particle initiation and obtained samples showed an earlier peak than that from a light trap 25-35 tha yields The high fertility isone reason why RLF indicating emigration W13PI usually emigrate after the armyworms and WBPH developed into large numbers vegetative stage Populations in 1980 reached levels that Farmers however used no insecticide on upland rice caused patches of slight yellowing in some fields WBPH although they did on vegetables probably cause hopperburn in Batangas They immigrate

The poor growing conditions fOr rice generally mean year-round as shown by low levels in the light trap BPH poor growth and low fecundity for insect pests fIeding on it however were scarce in the light trap and not collected with

Table 3 Abundance of rice insect pests and factors in rice cropping intensity of Philippine upland rainfed lowland and irrigated lowland locations where light traps were operated daily for atleast I yr -I

0Rainfed Rainfed lowland Irrigated lowlandupland

Pangasinan Cagayan lloilo Santa Maria Nueva Ecija South Cotabato Tanauan Caaringayan Caaringayan Bangag Bangag Bangag Buray

LagunaCale

Batangas Manaoag 1980-81 Santa Motiica 1982-83 Cabanatuan Zaragoza Avancena NaniamanSolana 1981-82 1982-83 Oton

1980-81 1979-80 Oton 1981-82 1981-82 Koronadal Koronadal1980-81 1979-80 1979-80 1983 1983

Insect (kerosene light trapNephotcttixcatch nolyr)virescens 650 1240 500 1800 330 210 3400 1600 160N nigropictus 210 - 360 18400 5500 2900- 140 90 20 - shy 1700 -Reciliadorsalis 710 - 3200 1700180 890 630 140 170 2400 800Nilaparratalugens 130 690 - - 4200 4600300 520 750 180 60 3300 1600 740 310Sogatellafiircifera 1700 31000 13000 8200310 210 3300 610 270 980 1600 6300 -Cyrrorhinus - 12000 6700600 540 3200 290 160 330 120 410Scirpophaga spp 50 410 620 - - 21000 12000150 180 220 270 30 1000Cnaphalocrocismedinalis 70 70 540 610 270 11000 6100200 140 100 6 60 250Chilo suppressalis 0 60 60 1900 300 200- - 30 20 0 6 3 160 -C auricilius - 0 00 - - 20 3 60 20 3 0Sesania inferens 0 - 80 60 3 80 10 51 -Cropping intensity _ -

Rice crops (noyr) 10 10 12 10 10 10 13Rice area (7c) 17 19 20 20 25 2320 85 85 60 70 40 85 85 90Fertilizer (kg Nha) 60 20 85 80 70 7020 0 0 0 30 30Insecticide (no of 40 60 40 300 04 04 0 03 300 13 13 30 50 40 33 33applicationscrop)

__________

IRPS No 123 January 1987 21

Light trapped Insects Drytnd __ 2(notrap-nght) Suctioned Insects (nom )

50 -)o 9 jO40- SOW ffurciWO Light trp

D-Vc suction

20 to 4

20

Egg porasitizahion (M) 75 60

300 SOOampReOftwciffr

45 30shy

15

-insects (notrop-night) 4 Nileporvata Iuen5

Egg parastization()

Light trapped insects (no trop-nght) Suchoned insects (no m 2 )

6 8 Cytorh~nuls hvidipennii 0

4 05 2

50 0 Insects (noltrop-night)40-ui Ipanh pe g

Moohase 1 Seasonal abundance measured by light trap

Rainfall(etod) and D-Vac suction sampler of whitebacked

to ~ hopper Nilaparvata lugens and lea ffolder60)-t Cnaphalocrocis mnedinalis andplthpe g20 L I II I I I parasites and mtrid predator Ortorhjntus livi-Apr May Jun Jul Aug Sep Oct Nov Dec JaIn Feb Mar dipennis set against cropping pattern daily1 9 80o 19 8 1rainfall and moon phase Tanauan Batangas

1980-81

the D- Vac Cyrtorhinus build up on the crop coterminous are collected in light traps during full moon (109)with WBPH buildup Cyrtorhinus disperses during the Because upland rice shares most of the lowland rice pestsrainy season RLF not readily collected in light traps was and those pest species are generally less impotant to uplandprevalent only during the rice crop rice than lowland rice there is a general belief that insect

Nephotettix virescens was more abundant than Nnigro- pests are less important onpicLus in the light trap collections and on the crop GLH upland than lowland rice Arcview of limited data on upland rice shows an averag of population peaked at the reproductive stage as did zigzag 10-21 yield loss from insect pests on a par with that ofleafhopper (ZLH) Crtorhinus tracked the populations of lowland uee determined by the same methodology (132) planthoppers and leafhoppers Both GLH species were (Table 4) McGuire and Crandall (144) made a similarcollected year-round in the light trap indicating immigra- estimate for Central America Cramer (51) howevertion from nearby irrigated rice areas Possibly because of the estimated 35 yieldlow populations loss in South America based on ainsect number did not correlate with similar pest complex - stem borers and seed bugs shymoon phase although reports indicate that more hoppers attacking irrigated rice in North America This estimate did

22 IRPS No 123 January 1987

Light trapped insects D~~nd (no trap-night )

St plht Suctined insects (nom 2

)

D-Voc suction 1520

1o

0 Lih trap 5 ~105 o5

00 Eggparositizotion()5040 NephOtgtha virneens

30 20

I010 A-Insects (notrop -night)

4 Nephatelh nigropccent i

2 o I vV^J - -- ^ VA____ f _

Light trapped insects (no trap-night) Suctioned insects (no rnE

10 -1208 ReCdO doRsol

6 15

4 10 052 iwxALAiamp 0

Light trapped insects (notrap-night) 2Suc oned nsects (nom )

8 CyrtorhmnusI6o nn]10 i0075

4 05

2 0 0

025

2 Seasonal abundance measured by light trap Roinfall (mmd) Moon phase and D-Vac suction sampler of green leafhoppersNephotettix virescens and N ngropicrus and I zigzag leafhopper Recilia dorsalis their egg 60

parasites and micid predator Cyrtorhinus livi-dipennis set against cropping pattern dailyrainfall and moon phase Tanauan Batangas1980-81

40

5 Lf Apr May Jun

1980 Jul Aug Sp Oct Nov I

-aMar

1981

not account for soil pests and was not based on field data in the absence of pests Carbofuran granules are phytotonicAkinsola (I1) cited mainly data from irrigated rice areas in to rice (239) The prevailing hypothesis is that carbofuranAfrica but estimated yield losses of upland rice between 15 prevents soil bacteria from consuming fertilizer The phytoshyand 30 from insects However losses in Thailand over a tonic effect is through greater availability of fertilizer to the4-yr period ranged from I to 13 averaging 5(106) plant Denitrifying bacteria are known to be more active inYield loss data however should be interpreted with aerobic than anaerobic soils therefore it may not be merecaution (184) The Philippine data were all derived from coincidence that the highest recorded yield losses (22-69)trials on farmers fields under farmers normal agronomic occurred in those trials with 12-30 kg basal Nha On themanagement practices except for Bukidnon and Capiz other hand contro with carbofuran granules may not be aswhere researchers used fertilizer Batangas farmers are good as a seed treatment against seed pests and perhapsatypical because ofhigh levels of fertilizer used The trials in yields would have been higher if a systemic seed treatmentBrazil were carried out at an experiment station but insecticide had been used conditions were similar to those of local farmers The 1979 and 1980 trials in Batangas had two full-

Carbofuran granules were used in every trial as a broad- protection treatments one using carbofuran granules andspectrum insecticide applied basally with the seed An the other using bendiocarb WP as a seed treatment Plotsobjection to using insecticides to measure yield losses isthat protected with bendiocarb yielded higher than those proshyinsecticidescandirectlyorindirectlystimulateplantgrowth tected with carbofuran granules The other trials in the

Table 4 Yield losses to insect pests and yield responses to insecticide in upland rice in the Philippines Brazil and Upper Volta

Yield (tha) Yield loss (5) Location Year Cultivar Protected

with Protected

without Check Total Vegetative Reproductive Ripening

Basal N (kgha) Reference

carbofuran carbofuran Protected No with carbofuran

carbofuran

Philippines Pili Camarines Sur Tanauan Batangas

Pangantucan Bukidnon

Dumarao t-apiz

BrazilGoias Minas Ge~ais

Upper Volta6

1975 1974 1974 1975 1975 1976 1977 1978

1979

1980

1979 1980 1979

1977 1978 1979

Bursiging puti Kinanda Dagge Kinanda Dagge Dagge Dagge Dagge UPL Ri5 Dagge UPL Ri5 Dagge UPL RiS UPL Ri5 UPL Ri5 UPL RiS

IAC47 IAC47 IAC47

14 33 11 36 35 31 30 27 37 26 40 30 40 42 46 35

19 21 09

--

-34 24 24 27 --

29 43 31 42 -

47 25

---

10 28 05 27 27 30 26 29 33 28 33 29 43 23 36 11

15 15 06

29 15 55 25 23

3 13 -7 11

-7 18

3 -7 45 22 69

24 28 35

--

21 -11 -20

4 -

-3

23 7

-2 -23 56

-

-

-__0

6 --

-

-23 14

5 0 8 2

22

-

--

0

-

--18 11

3 -10

4 10

7

--7 -

14 -

-1 6

10 -5

8 9 7

-

--

0 0 0 0 0 0 0 0 0 0 0

30 30 30

12 12 12

99 99

100 100 101 101 102 102 102 102 103 103 104 104 104

66 66 66

Farako-Ba

Mean

1977 1978

IRAT10 IRATIO

38 66

--

40 43

-5 35

21

-

10

--

--

--

- 242 243

Z

tJ

-w

24 IRPS No 123 January 1987

Philippines which had two treatments offering protection In the 9 Philippine trials where yield losses were measuredwith and without carbofuran granules relied on further by growth stage 10 of the yield loss (21) occurred at thecontrol with foliar spray and not on seed dressings The vegetative stage and 517 occurred at each of the reproshycarbofuran-protected plots had an 18yield los compared ductive and ripening stages Ibis result is similar to thewith only 10 with foliar sprayS This yield difference is pattern of yield loss in lowland rice measured by successiveprobably due to differences in protection against sown-seed treatments where the crop is unprotected at each growthpests (foliar spray do not control sown-seed pests) rather stage but proticted during the other growth stagesthan phytotonic effects became basal fertilizer wa used in However the insect complexes r sponsible for vegetativeonly 2 of the 10 comparisons stage yield losses in upland rice are entirely diftfrent than in

The recorded yield losses were not related to yield lowland rice The key vegetative stage pests in lowland ricepotential (Fig 3) One might expect higheryielding crops to arc aquatic whereas those in upland rice are sown--seedhave higher percentage of yield losses but this ws not tile (sown in soil) root and seedling pests This dift rence maycase One reason is that insect damage exacerbates plant explain why agricul torists have overlooked tile ilmportanccinjury from drought stress low yielding fields from d ought of upland rice insect pests Many reports of upland rice pestsstress would show disproportionate differences between in Asia ha Ve focused on major lowland species (46 108)insect-protected and -unprotected treatments Shoot and It isalso surprising that tei vegctative stage of upland rice root pests would be more responsible for losses from would record the highest yield losses becatuseseeral studiesdrought stress than sown-scd pests A second reason have shown that upland rice can readily recover from foliageconcerns seed pests which would have a greater impact on reinoval during that stage (117 152 165 188) Upland ricelow tillering low yielding varieties such as )aggc Low poscsses a hitgh obilitV to compensate for ca rly loss oftillering varieties nnot fill in the space created by removal foliage and will even be stinulated by foliage removal toof seeds oweer 1) R i 5 is high yielding because it tillersP1 produce a higher viel than plants ithout any foliageactively which can close the canopy to provide greater weed removed (63 192 231) [ie reason for this apparentcontrol discrepancy hovce is that yield loss at the vegetative

stage does not octUr frol leat arca loss but fron pestsYield loss () removing sown seed and feeding on devcloping tillers and roots -- damage that cannot be readily conpLtnsatCd for

70 - Measuring yield losses in upland rice cnvironrnents is further complicated by a report from Brazil that greater yield losses occurred in response to better insect control60 - =-09 as Insecticide seed treatments resuled in denser plant stands

(n=21) which in turn created a selectively more favorable environshyment for blast (69) This resuIt was parlticurla rlxexacerbated50 when cirbofuran granules were used in seed furrows Aside

ffrom a greater plant stand ca rbofuran iay have auigmented the N lecl and stimulated blast If so ca rbofuran should be40 - replaced with another insecticide or the seeding or fertilizerI rates decreased

Other methods to determine the impact of upland rice30 shy pests wvcrc to interview farners or to develop single speciesbull bull correlations of population levels and yield loss Feneira (66)2 NIM)st (75(1) Golas farmers reporteddid both in Brazil

20- insect losses in ufpland rice and 251 said those lossesS avcragcd 37(1 [his result is consistent with tie yield losses 0 0 measured by field trials in the same statc(Table4) For every

I0 4 of seedlings removC d by leaf-cutting ants a I 7yield loss is predicted Similarly Ferreira (66) found that ai

0 - average of 53 thrips per panicle before panicle initiation meant 3 times more unfillh d grains

-IO0 I 1 1 I I-101 0 2 3 4 5 6 7

Yield (tho) C)N [ROt IM FI IODS 3 Correlation between upland rice yield and yield lols caused by Upland rice insect pests may be controlled by culturalinsects measured in 21 trials in the Philippines Ivory Coast andBrazil genetic biological and chemical methods

IRPS No 123 January 1987 25

Cultural control dry season plowing will have little effect on them Plowing Crop husbandry to reduce upland insect pest populations when the soil is wet from the first monsoon rains will expose can be classified into practices effective in a single field and soil insects that tunnel close to the soil surface to predatory those effective only at the community level Cultural birds chickens dogs and even man (field crickets nole controls should come first in a pest management program crickets and white grubs a re delicacies in man tra ditional They have broad stable effects as pests have little pos- diets) sibility of overcoming them through selection of biotyps The greatest degree i1 inIscct control fron tillage comes (138) On the other hand some cultu ral controls work only from plowing soon after rice ha rvest when most soil insects with high labor or power inputs and connunity-wilt lie close to the soil surface anrd stern borers are in the rice methods require coordination of iany farniters stuble Whitegrubsare mature and tillage exposesthem to

Pantin time The luxury of being able to shift the time of predators Tirnelv control will prtec the crop following pla ntring upland rice is open onl hr la rniers in areas with a rice and lowr the populationIfor succeedinig years Plowingprolonged rainifall pa ttern lI copping intensity or underthe ice stulble helps it decompose killing sten borer rnechanied land preparatiot Gcnerally planlting as soon larvae inside as the rFiinv season begins will lower populatiois of 1110st [requent tillage destroys ait nests bit not termites insect pests Insect populationsarelo aftcra dlvseoll or Tillage and clean culture howes er rcliove food for winter fallow delayed plaiting lets them build tip on termites which then mtay attack a young rice crop Scarab alIternate hosts that gi vigorotsly with the first seasonial beetles also prefer to miposit in eceitly tilled fields Noshyratiris This first flush iof weds cros lItxuttriartlv with iceCult is dibbling faors the survival oftillace Clre Such rineralied N released during the dr season aid made stils i aels tnd insects thtt pttpate or lay eggs- in the soil

available to the plants with the rains It iliso igrtmitcally Zero tilla hio eerl conserves predators ofthe GIH (37) advanttgeous for the rice ciop to tap this trlLItlaI fertility nt it mcttoi lo lessen the impact o0 soil insects and grow vigorotsly to compete with wrus and to tolerate brotdcasting seed or planting in ftltrrows is prefc ired over pest infestation dihlirig in hills Crickets are large enotlgh to eat seedlings in

Larly plantngs terd to escape seedling maggolts Upland a hill atrd tillering normnally ca iriot fill in tlhe missing space rice in Bitangas is planted in Ma and ine itli the Iirst Oftcn dibbll rd is not well covered and iseasily found by rains Fairners are highly mtotiated to plant early tocscape foraging insets ioIdents and birds drlitght and typlotns ricar harct Arn Atrgtst planting Plan uit sii Incrcatsini sced density protects tile cropwould be evcely attacked by It riim orivzo At tti againrst seed and seelling pests especially ants particularly other Philippiie tipland sites Rea l Qteot antd (laelit if the crop will germinate and ermerge quickly M isainis Oriental seedlig mnaggots arnrige ric boe t se hr trftinll itI India rice intcrclopped witi cotton or farnes have a longer growing season and plnit 2 rin or pigeon pci had lower 6IIt and WBIlII poptrlations than mote after the ortset of the rainy season liaaiangas ftrmners rice alone (211) NfailC arid Urplald ir_ 2re coilltori in the plant ca rly becia use ofa shorter raiinfall period arnd the desi re Philippines he chatngC in pest status folii iniercropping is to lliarvet before the typhoor sea son peaks ltighly locatioll-specific (133) cmianarid the net elfc be either

Early plantings cornbind with early-iatiuring varictics nil bcnelficial ortdetriential (189) For example BIatangias provide higher crp tolerartce for white grubs Aestiatior farmers intercrip Itro in their ipland ricefields A liornshyterminates with the first rains but 5-6 moare ineeded for the nrii iotd-Ahis i (I iiiacius) which norlially feeds resting larvae to develop into beetles emerge and lay eggs on taro ill feed or miarbv rice plants Intercropping and for larvae to reach the danriigii g ird-instar stage probably would not afft0 seed pests A crluanion crop (134 In lataigas this normually iseiough tiite for harvest planted ith rice and haing gleater tolerarnce for root

It Brazil howecver delayed planting is a suggested dariage or being toxic nayact isi trapcrop to root-feeditig cultural control for the lesser cornstalk borer and insect pests Acial-fei i ig pests voultId be most affected by pests favored l drought (66 195) Cropping internsity in intrer ropping Shoot aphids leaf beetles sten borers and Brail is low and delayed plraing places the rice crop in the hopplers would be likely ca ntidates to note effcts On the niost stable rainfall period reducing likelihood old rought othlt halnd the corlipalior crop stch~ sitniaie Iiiv be more

In rilniy arca s rinmirsare roi atl to plant other crops benehited than rice from initecropping ( 137) before rice and althNgh it woultl be desirabh to avoid 11echi Weeuling luring the first month after clop infestations by planting rice cuatl lariners lack the labor cstablishent will force pcts such is anyvworris which and time to do so ol a low priority crp prefer grassy cetds ontor tle vorling rice Soil iampsecs will

77age Uplild rice Stils ae light textured and Call be mvnye to the rice crop Cleani citrltirc also lay lore territes plowed duiring the d rv seasor notl only to grin time for to attack I u cetincti inrice ceds i ay he le optinal planting with lie fitst Inilsoon riiains but also to desiccate solition 13) soil insects arid weeds such is ntsedge that foim rhizomes Irtlizr Itneatsing N Iciiliier oin hlwland rice has Htowever becta use most soil insects lie below die plow layer fAtwvored Irigher populains of pliithoppeiv Imlhoppers

26 IRPS No 123 January 1987

and leaffolde rs However because of the need to temper fertilizer use on upland rice o prevent blast fIrtilizer rates will be lower than is comonia r for lowlad rice causing relatixely less hopper and IaIfTfoide r population buildup However inupland rice areas without a history of blast such Is Batangas farmers apply high rates of N and conscqtrently the crop has high infcstation levels of WBPI]and RIF The fhrmrcrs leec insect buildup by splitting the fertili11r in0to two to three applicatiois bit do not applV fertilizer at the vegetati estage because this prodrces a tall crop which illeasily lodge during typhoons Greater crop fertility leads to greater insect survival (better nutrition) larger insects (to cat no re foliage) arid greater fec titd t (to lav nmre eggs) (128 145 171) Not all insect species respondequally hovr Increasing rates (if N l anrid K increased Dialraa s-acchralis and (iaeiurema flea beetle but reduced thrips (66) Zn rcdLuced deadheart density from the lesser cornIstalk borer

Fnldingi Although most Upland rice areas do nIot hawxcaccess to wa ter floodiig the fields is rRrrnriierdcd in areas of Brazil or root bug and white gib control ( 195)

Crop rotatioti Tillage before planting a crop after rc harvest will control stern boreis and other pests renain n the stubble arid will unearth soil pests Planting a non-gratriiaceous crop after rice is recommncrded for termite control (66) Crop rotation is less nttdcd in upland rice culture than in lowlands because atcrop-free period is assured even after harvest of crops following rice

t1ulchilt iryl plastic irli irig designed for weedcoitrol ii upland rice inJapan resulted in ntr highanri liy infestation of (ihilo sulytresvais (93 9-1 95) NIulched Upland rice grew more luxuriantlv and attracted more (Avipositing mothis than u lched rice Also the rice plant and stein borers imatoured 10rC quickly and a partialgeneratiori af vouing ista iNoverwintcred in the stubbleLotili--malitrt varieti Quickly maturing crops reduce the tmbCr of pCst generations that Calt build up Rapid crop establishment olan ca rl v-natu ring vaitvty if)pa 11icLa r will lessen white gib and stem borer damiage White grub larvae intle last iistar are highly destIruCtive so aii early-iiatri ng variety will escape serious root loss Stem borers build 1ip slowlyx in rice to ecormic abundant in a late-Maturing crop anrd caiuse whi tlieads Numibers of somespecies increase exponeitially with each succeeding genera-ti so a xarietyexhat atures I trowh earier thian another will hac less insect damage Early-naturing vaiicties however will not lessen seed pest daiage either ii the soil or oina standing crop

Strwhoi hAIJttam Inrscts tcd irig on thlie aerial portions of rice plats disperse from carly to late plantedfields AS iiiibCirs increase exponentially with eachinsect

gtscnrattm e eiglbrig f rmc~s1(uil upp wh rc generation pests build tIP where neighboring fihrTC e9r1sstagger their plantings II Batangas farmers plant uplandrice within I iro use 120-d s-arieties and plant at the start of the rainy season Ini Misarnis Oriental laricrs stagger their

plantings up to 3 mo apart and use a set of atieties that mature in 4-7 mo Insect pest danage particularly from stem borers is greater in Misarmis Oriental Also seedling maggots damage rice there because rainfall favors frequent planting of maic In Batangas rice escapes seedling maggots bCca usC of distinct drs season and synchron0us early planting with the onset of ains

Therefore combinations of cultural practices ea rlyplanting synchronous planing crop rotation and earlyshymaturing vaneties protect the rice crop against most insect pests

Ilmant resistance Beca tse upland rice has few specialized insect pests and many insect types attack the crop worldwide regional breeding and strong internatiornal cooperation are required

Sown-sced pests arc economically controlled by insectishycide seed treatment and should the relore receive lowxpriori ty inbreeding objectives

Root pests are normally cost ly to control with insecticides but finding resista t sonres mayvbe di fficult Tanaka (229) for exa -ple fi led to find varieties resistant to root aphids Selecting for large root bioniass perhaps should be the strategy for pests sLuch as Whlite grubs that renliove roots Fainter(174) howcvcr cites varietiesof crops tolerant ofor resistant t(i root pests Some sorghun varieties are high in cyanide so rice sorghun might incorporate broadshyspcctruri chemical resistance

Seedling vigor and drought tolerance are high prioritybreeding objectives so tupland rices should have large r root systernsand be able to tolerate ligherlevels of root loss Sonie root loss inolder plants is beneficial if it stimulates new root developnrut and therefore crihances intake of soil nutrients (53) The rice seedling inaggot rtiLotia spp may be

controlled thirough resistant aittics A hi rgc breeding effort on sorghniu has had fair success inideveloping arreties esrstait to Asoccata Rodani (251) A field trial comparing 10 upland ricc varieties showed differences aniong varieties va ring from II to 4(- damaged tilleis (215) Hovver resistarce to 1Iill-0r gtia species may rot cross ox-rti otier

Shiraki (210) reported that upland i-ice stens wcrc harder than lows-land rice stemsperhaps becaIusc (f a higher silicoricontent and thus were more resistant to yellow stern borer hc tortia Iow tillering (f uplaid iice xarneties prodtres a

higher percentage of infested tillers than would occor in higher tillering lowland iice under a siiilar egg density The corollary therefore is to select for high tilleriig upland rices

Stemii bor amcstheh main targes dobrecders inAfrica (571 204 )but nticl oifth e sc reen irig is dotie unde r lo wland co d t osrm s wih A h r~ o l pe i s n acorditions (11 96 218) As with lthrinrnwspecies mayhave to be dealt with independently however initial results show cross resistance between African and Asian species(92)

In Brazil screening Ior insect resistance focttses oil lou I pest problems Varieties tolerant of lasmolwalpus ligi-sellus and resistant to )iatraea saccharalis have been identified (70 153) Current work also emphasies 7iraca limativetntris and Thmialspis (=Iosis) flavouicta (66) These insect pests Ilt onlV citise IIigh yield losses tihey also ire difficult to control chemically ofr by other ilealls

ManV upland rice vaieties aic early maturing and are resistant to pests such as gall midge (I Xl ) Early mIturity is often a highly desirable trait to esuilpe pest buildup but it is not true resista nce bca usc ilthose va reties are plinted late they will be itsdamaged as ain early planted susc ible variety

P~erhaps tile cosiiiopolitall of all upland rice pestsimost are the sced pests (ripenling stagc) which pose a particular problem beta use scuds re ised IsIO for hitnit4ans Ayltoxic substatice introd tcld geticticllyilmay also be toxic to nal iniless it caln he placed old in thfe IdhuffSelectmi fo hairiness or lotng wns kill mIlct sthilrohleiiis dutilig threshing pa rtictfarl maitial threshinfl [hireshers ill complain of itcfimin atid itteascd (fust levels

Attempts itIRRI to fitid tcistatncc to the rice bug plorisa have ilotbeCnl prmitisilg lIIetIattonlids iorc

through the lenilna or palca hillhairs sedstVIor seeds wilh awns that deter bugs promote complaints from threshers Seed bugs liptocorisa and Stcnocoris enter the seed through the opening between tile lemna and palC but varieties with narrow openings are diifficult to mill

Breeding priorities should concentrate ott inotCI gruLtLIps that are most difficult to c nttrol by other meains Steil) horers appear to be the most widespread group that sigiiificantly dalag s rice aind is hard to manage by other Ilnca us

Biological control Very fe stutdic of mitatt~l eneties of rice pests in upland envirollnletits wee in tile wefouiid literature Therefore have not compiled a list of species Ve present a list of natural enemies iOtid in upland ricefields tuic latialln Batangas for some conitno iOldiar pest species ([itble 5)

Muich ofl thc lollowinle itfor ittiol cotlliCs rol Stutdies in the lowlatnds or roi crops otithcr thlin tice

Qu(ailtr o 11ll t11t( Nat trrtf lii hcoeits ight Ile used agaitist ile social insects tilt and termites Sollie predator specice live ill theirf nests n ilnic theirnllt ip-pearanIlCe aMRidbehavior IJtrouMdCing a mote aggressise ant species to displace pest ant codonies is a possibility (169) Patihogens as potential natural encnies have reccived little atteition (I13) Natural ncnuics asa group however hac not been toIl iporta it in Iegulating aintshown be and ternite numbers

The nocturnal behavior aid subteitmien habitat of lole crickets and field crickets protect them from many ntllral enemies There are only limited records of natural cnelics for these groups of pests some pathogens (87) but mainly

IRPS No 123 JattuIny I 271987

parasites - scelionids on eggsand sphecids on ilymphs and adults Sphecid wasps Larra carutmaria (SImitI hI L huzwuensis Rohwer and lsaltiuhta Willi ams speciali in mole crickets pa ralyzing them nd dragging the ninto thcir nests as food lot their voting ()tier sphCcid species Lins aurulenoa (Iabricius) lot maiac (Ashmead ) 1 lhshylssellaus (Smith) and Ml lal)oriosu (Snithf specialie in field clickets

White grUbsO the other hand hae Icti scriuns enuigh to have been extensively studied for cotttroi 1W nati al enemies( 112 140 141) Scoliid parasites C(ajsoMcispp have beetn intrtduced itto tilePhilippites for control of Ielxl ]udisiro la The rcsearchcts w Intttrdced themle during ill era claimed that the white gttionthreak as controlled (142) howevcr it isnormal for pest epidemics to subside and there was no diect evidence to indictte that tfe parasite was responsible A tmore promisitg titctllod might he to identify iso ltcttass prt~f tcartd apply hiactelaln fingal pathogens to tile soil Ihe lalattese beetle in North America has hcci coitrolled hv tite conmnleciall aailable hactcrial prcpa ration itacihlpu pav ofr milksdiscas (71) Spote prtyefott the soil itloted ilrtier reitiain actic fr Veas

Root aphid sardtnctly bugs readilslic tilib to predatots and parlsites once their tetndine ants are controlled The larvac and adtilts of a coccinellid beetle 1Scynms sp prey on llriadlouais in slash and burn upland rice Baits treated with insecticide may be used to control tending ants by killing their ourig (244)

Root-tkcditig blugs false wireoris wireorms and root wccils aic r ofsuch little ifilpoltatice that thenomttll role of liatural Cticniics hias tiot beei asscssed A possibility hoevrl tiglt he tit ItaIIitic niaCIMtodcs lintt1g11en Itese soil-itlhlfbitine mCtitodcs W5uhtfhave to be imiass Produiced

large miltibcr of niatltrll cnemics ircknown ft CltIoriS IC IpblCeii is thtt clttvortnlls ofonlie the crop soott alter land ptel)tritiOti a1tithe bcgit iniig of the raittv

Citsoln hfien tiatllral leniiiy populations aric lo Colltloil WOtuld iii ole mass prodtticing andirclcaslng key spccies

le egg stage is the fpirt (olthe Cedlitgt Ialot life cycle tI hich to f)ctis ftor tittlril elctit cottntrol hplraites o1 pleditors ttsit elleggs highly expoIsYCl I Sgg pailsites or predators stolld probahly e ittass andtbe produced released Littlh i toruialioll is a ailable ott what species to trV o ho to Iniss podLice thiCIi A etiloplhid wasp Icmihlaol sp effctiicly regushy

lates the leatf lcloalapolm2a spioanoiiitr heat andc rice antid should be collsctvcd

leaf beetles whCllose e am] eggs ate laid in the grutindhirO ould be motC difficutlt to control by niatulral enelV Mnll ipuhltiOn But species sUici IS c_gsthispid beetles ith

and larVle oilleascs ate vtilcratble to iparaites and pledators

Natural enemisc of tlirips otl rice are little known

28 IRPS No 123 January 1987

Table S Parasites recorded from upland rice insect pests in Tanauan Batangas Philippines 1977-84

Order Family Species Host stage

I lymttenoptera Braconidae Leaffolders Cnaphalocrocis Marasinia spy

Cardiochilesphilipplinensis Ashmead a Larva

Cotesia angustillasis(Gahan)Cotesia or cypris (Nixon)

Larva Larva

Cotesia or taeniaticornis(Nixon) Larva Cilonusinotakatae Munakata Chelonits SppAlacrocetrts philippinensis (Ashtmead)

Egg to larva Egg to larva Larva

Iclineumonidac Orgilts spp Isclutojppa hireator(Fabricius)

Larva Larva

Itoplectis naraigav(Ashmcad) Larva Tenelhtcha philippiensis (Ashmead) Larva Tenelchha stangli (Ashmead) a

irichomma cnaphalocrosis Uchida Larva Larva

Bethylidac Elasmidac

Goniozms or trianguli r Kieffer lasmtusalhopictus Crawford

Larva Larva

Diptera

Encyrtidac Chalcididac

Tachinidac

Elasmtus spp Copidosomopsisnacohiae (Fidy) Brach nteria evcarinata Gahan Biachyieria lasus (Walker) Ztgohothria ciliata (Wulp)

Larva Egg to larva Pupa Pupa Larva

Argyrophylx nigrotihialis(Blaranov) Larva

l-ymenoptera Braconidac Yellow stern borer Scirpopiaga incertulas

Braco chinensisSzeplegeti Larva CIthelotis spp Chelonus niunakatae Munakata Cotesi flaripes Cameron Stenoracon nicerilli (Bingham) a

Larva Egg to larva Larva Larva

Ichneumonidac Tropobracon schoenobii (Viereck) Amatroinorpiaaccepta metathoracica(Ashmiead)Frihorus sinicus (I lolmgren)

Larva Larva Larva

Ischnojoppaluteator (Fabricius) Larva Isotima nr dammeratiti Rohwer Larva Tenelucha philippinensis (Ashmead) a

Larva Tenehicha stangli (Ashmcad)Trichomma enaphalocrosis Uchida

Larva Lara

Chalcididac Eulophidac Trichogranmmatidae

Xanthopimnpla stemmator (Thunberg) Charops hrachYpterumn Gupta and Malteswary Brachtm eria sppTetrasticttus schoenohii Ferriere Trichorigranm chihunis Ishi

Larva to pupa Larva Pupa Egg Egg

Diptera

Scelionidac Pteromalidae Tachinidae

Tt ichograintnta aponiicuitt (Ashniead) Telenomits rowani (Gahan) a

Trichoinalopsis apattelctena (Crawford) Jeirhaea spp

Egg Fgg Egg pupa Larva

Z4gobothria ciliata (Wulp) Larva

Ilymenoptera lecneumonirlac Gol fringed borer (hilo auricilius

Trichomm enoptalocrosisUchida Larva

Ieioenoptera Braconidae

Aantholimlh steiniator (Thtinberg) a

Pink stet Ioret Sesamia inferens StoInacoil nicrilhi (Bingh am)

Loarva to pupa

larva

Diptera Ilyinenoptera

Armvtnyorms Mythimna separata Spodoptera mauritia acronyetoidesTachinidae Zvgohothriaatropiora( Robinea i -Desvoid y)Eulophidae It-phctrtiscitapadoe (Ashimead)

Larva Larva

(halcididae Braconidac

Brachpin ria spip (oesia spp

Larva larva

IHyntenoptera I)iptera

Rice skipper Ielop ida (halcididae Tachinidae

math ias aidgreen iortedci ciliarNIelan itis eda Brat lhincria sp iii matuqitatt Cameron Irgyrolhirlaxi trotlhiais(marumtv)a1

ismtne Larva Larva

Diptera Ilymenoptera

Tachnidae lelineutionidae

fuown se tilo per Mois frugalisIrgtrophrlax 1igrothiahiv 3 a rat ov) anthopimpa pmnctata (Iabricius)

Larva Larva to pupa

aMost dominant

IRPS No 123 January 1987 29

Normally predatory thrips prey on herbivorous thrips strepsiptera on both Spiders dominate tilenymphaladultNatural enemies of stem bugs --Scolinoplara and Tibraa predators Spittle bugs move to rice from pasture grasses - are in order of importance egg par~isites and predators and biocontrol would be successful only ifcarried out before fungal pathogens and nymphal adult parasites Egg tiley immigrate into ricefields parasites could either be introduced fron other areas or Biological control tactics for hemipterous seed bugs mass produced and released ould follow those outlined for stein bugs

Armyworms are normally lwld in hcck by the activities Coml)arison of environnewtns Natural enemies are of egg and larval parasites When these natural enemies fail perhaps faced with even greaterconstraints than insect pestsusually because of drought armyworns become epidemic to survive ind reproduce in upland environments TheyIt may be worthwhile therefore to release parasites during have adapted ani survived by having wide host ranges the the rainy months following a drought Virus diseases of ability to aestivate or hibernate over unfavorable seasonslarvae would beanotheravnue to exploreforarnyworms and the ability to disperse or by other mechanisms

Parasites have ben traditionallyconsidered for biolegical Comparing the most dominant and therefore the most control attempts against stein borers usually by rekasing adapted species of natural enemies collected in various exotic species( 166) This approach has not met with success environments in the Philippines we can understand how and new avenues should be explored The role of predators different rice environments are arid pathogens is little inderstood and deserves greater Anagqrms olptahiisa mymarid planthopper egg parasiteattention Egg parasitism rates are normally reasonably is cquallyadapted to all Philippine rice environments (Tablehigh but should be supplemented with effective egg 6) It is selective for planthoppers and even parasitizes eggspredators Orthopterans are potential egg predators of the n-iize planthopper Peregrinitsmaidis itt upland areas Metioche vittaticollis (Sta ) Anavipha nifiwnnis (Saus- Alavohis also parasitizes maize planthopper eggs but is sure) and Conocephahs hongipnni (de i-laan) feed on most prevalent on BPH in the irrigated wetlands (onashystem borer eggs etioche and Anaxil)ha specialize in eggs tocerus spp like A optalhilis on BPH and WBPI Iattacks with no hair such as Chilo spp The omnivorous (ono- the eggs of GLH in ell rie environments It is also found cephahsspecializes in eggs cov red with hairs it is a proven parasitizing the eggs ef the white rice licahopper (oimna egg predator of Scirlouphaga spp but has the discouraging spetra Oirosita naias a trichogrammatid egg parasitehabit of eating rice guin (65) attacks BIPH and WBPH in the wetlands and has no other

Grasshoppers have egg (scelionid) and nymphalad ult host A related species 0 aesopi specializes in G1-H in (nemestrinid tachinid and sarcophagid) v irasites Among wetland environments Dryinid strepsipt-an and pipunshypathogens protozoans have been recorded most frequently culid nymphal and adult parasites of planthoppers and Control of locust species by natural enemies could con- leafhoppers also specialize either in hopper species or centrate on their habitual breeding areas environments Among the dryinids Pseuhdgonatoipsflavi-

Leaffolders have rich complexes of natural enemies some feimur prefers the lowlands and BPH while P nuch has no adapted to upland rice (22 27) ranging from egg predators environmental prefirences but is adapted only to WBPH (gryllids coccinellids) to larval parasites (braconids ichncu- The main d ryinid species on GLH aptogonatop)usspp is monids) to larval prcdators (ants and carabids) to larval most dominant in rainfed lowlands and also attacks other pathogeiis (vira and fungal diseases) Efforts to augment leafhoppers within Amnrasca Cicadulinaand Bachaltuathese natural enemies should focus on the egg predators Strepsiptera are more dominant on plarithoppers than (rearing releasing)and viral or fungal pathogens (culturing lealhoppers The Elenchus species in the Philippines attack spraying) all species of planthoppers within the genera Sogawilla

Parasitization of the large conspicuous larvae of Sogatodels NilaparsataHarualia and Opiconsiva Elenshyskippers green horned caterpillars and other polyphagous cusrvasuntaisuiis adapted to the major rice environments Lepidoptera by tachinids and chalcids is normally low ELg but prefers BPH in rainfed arid WBPIH in wetland environshyparasites and predators are perhaps the key to their control ments Ilalictoph(utsmtnroeiand HVpectrus attack G LH

Gall midges are parasitized by pterormalids and platy- and COfimravcctra respectivelygasterids that aestivate within their larval host between Pipunculids are mainly parasitic oii lealhoppers Each seasons Shoot aphids are preyed upon by coccinellids species has a unique environmental preference Tomosshysyrphids and chrysopids varireIa oraetoraprefers Ne7 otetltix nigropictrs and

Planthoppers arid leafhoppers have egg parasites and Pilunwulhtsjavanensisattacks Dehoceplhahls spp which predators as well as nymphialad nlt parasitesand predators are more abundant in upland than irrigated lowland their rich complex of natural enemies includes fungal environments T suhvirescensattacks Nephotettix virescens pathogens The principal egg parasites are myrnarids and and prefers the lowlands Pilpuiulus nttillattls is an trichogramniatids Egg predators are norinally mirid plant upland rice parasite arid attacks only Nephotetix speciesbugs that suck out the yolk Nymphal ad nIt parasites are More than 17 recorded species of leaffolder parasites in dryinids on planthoppers pipunculids on lealhoppers and the Philippines attack species of C(naplwocrocis and

30 IRPS No 123 January 1987

Table 6 Dominant natural enemies of rice pests in upland rainfed lowland and irrigated lowland rice environments of the Philippines

Natural enemy

Host stage Species

Brown planthop-Parasite Egg

MymaridaeAnagrus optabilis(Perkins) Anagrusflaveolus Waterhouse

TrichogrammatidacOligosita naiasGirault

NymphAdult Dryinidac

Pseudogotatopuslaiifemttr Esaki et Hashimoto Elenchidae

Elenchus yasurnatsui Kifune and Hirashima

Environment

Upand Rainfed lowland Irrigated lowland

Ailaparata lugens

x x X x

x

x x

x X X Whitebacked planthoppers Sogatella furcifera and Sogatodes pusanusEgg

MymaridaeAnagrus optabilis(Perkins)

TrichogrammatidacOl~gositanaias Girault

NymphAdult Dryinidae

Pseudogonatopusnudus Perkins Elenchidae

Elenchus yasutatsui Kifune et Hirashima

Parasite Egg

MymaridaeGonatocerus spp

TrichogrammatidaeOligositaaesopi Girault

NymphAdult Pipunculidac

Tomosvaryella subviresccns Loew Tomosvaryella oryzaetora (Koizumi) Pipunclusmutillatus (de Meijere)

DryinidaeIlaplogonatopusspp

Egg Copidosomopsisnacoleiae (Eady)

Larva BraLonidae

Cardiochiles philipphnensisAshnead lchneumonidac

Trichiomnia cnaphalocrosisUchida Temelucha stangli (Ashmead)

Pupa Chalcididae

BrachymeriaexcarinataGahan Brachymerialasus (Walker) Brachymeria nr tachardiaegroup

Parasite Egg

Scelionidae Telenomus rowani (Gahan)

EulophidacTettasticusschoenobhi Ferriere

Larva Braconidae

Stenobraconnicevillei (Bingham) Iclhncurnonidac

Temelucha phiiippinensis(Ashinead)Temelucha stangli (Ashmead)

x x x

x x

X x X

x x Green leafhopper Nephotettix virescens

X x x

x x

X X X X

X

X Rice leaffolders CnaphalocrocisMarasmia

x X x

X

X X

x x x x x X x x x

Yellow stem borer Scirpophaga incertulas

X X x

x

x

x X XAinauromorphaaccepta metathoracica(Ashmead) x

continued on next page

IRPS No 123 January 1987 31

Table 6 cntinued

Natural enemy Environment

Host stage Species Upland Rainfed lowland Irrigated lowland

Larva-pupa Gold fringed borer Chilo attriciiuslchneumonidae

Xanthopimpla stemmator (Thunberg) x aPupa x

Pteromalidae Trichoinalopsis apanteloctena (Crawford) ax

Pink stem borerb Sesamnia inferensParasite

Larva Braconidae

Stenobracon nicevillei (Bingham) x b b Armyworms Mvthinna SpodopteraLarva

Tachinidac Zygobothriaarroptvora (Robineau-Desvoidy) x x

Braconidac Cotesia spp

X Skipper Pelopidas mathias and green horned caterpillar Melanitis leda ismeneF-gg

TrichogramrnatidacTrichogrannaspp x

Scelionidae Undetermined Scelionid

x Larva

Tachinidae Argyroph lax nitrotibialis (Baranov) X x

Pteromalidae Trichomnalopsis apanteloctena (Crawford)

X Predator

CoenagrionidaeAgriocnenis femina femina Brauer x xIschnura senegalensis tambur

Miridae CrtorhinsliidipennisReuter x x

Nabidac Stcnonabis tagalicus (Stal) x

VeliidaeAlicrovelia douglasi atrolineata (Bergroth) x

Coccinellidae Micraspis crocea (Mulsant) X x x

Vespidae Eumenes campaniformis (Fabricius) x

Formicidac Diaecamna spp xOdontoponera transversa (F Smith) x

Carabidae Chaenius spp OphioneaW- aJasciataSchmidt-Goebel

x

Ophionea ishii ishii Habu x x X

Gryllidav Anaxipta longipennis (Saussur-) x xMetioche ittaticollis (Sial) x x x

Tettigoniidae Conocephalus longipennis (de Haan) x x x

aNo rearing made on rainfed lowland bNo rearing made on rainfed lowland or Irrigated lowland

MrasmiaAmong the Laval parasites the braconid Cardio- lowlands and parasitizes oriental maize borer OsIriniachilesphilijpphensis is most adapted to an upland environ- furnacalis (Guenee) larvae The leaffolder pupal parasitesment It also parasitizes Hydelepta indicata a common not only occur equally in all environments but also haveleaffolder of legumes The ichneumonids Trichomma cna- wvide host ranges- parasitizing species of OsIrinia Marucaphalocrosis and Temehcha stangli occur more in the Hydelepta and Honiona - in maize and legumeslowlands T stangli is most adapted to irrigated lowlands The principal egg parasite of yeltow stem borer (YSB)and also parasitizes Chilo and Scirpophaga stem borer Telenomus rowani is widely adapted to all environmentslarvae T cnaphalocrosis is most prevalent in rainfed and has an unusual alternate host -Tabanus eggs (25)

32 IRPS No 123 January 1987

Tetrastictu schoenobii specializes only in YSB and occurs mainly in irrigated lowlands -- further proof that YSB is most adapted to the wetter environments

Larval parasites of YSB are highly environment-specific The braconid Stenohracon nicevillei occurs only in upland rice but also attacks the pink stem borer (PSB) Tieinchwa phiiiplPwnsis an ichneumonid occurs in upland rice and also parasitizes the larvae of (7hiu stqpresaisinatro-morpha accepta melaihoracica like 7 stangi is mnost adapted to the irrigated lowlands

The dark-headed stem borer(l)HSB) (ih pochit3sus was not collected in all rice environments and the gold fringed borer Chilo auricilius nd PSB were collected only in upland rice Xaantiholiplastenrnaloran ichneumonid larval parasite appears to be widely adapted whereas TrihomaslopisapanitI)cteiiaa pteromalid pupal parasite occurs mostly in wetlands and also parasitizes larvae of Chilo suppressais and Pelhuidas mathias

Armyworms Myit hinin and Spodujipera have two main larval parasites - a tachinid Zrgohothriaatropivora in rainfed environments and a braconid Cotesia spp in irrigated lowlands

The butterfly pests Pelopidas and 4elanitishave tricho-grammatid egg parasites found only in irrigated lowland environments

Two larval parasites show environmental preferences --Argrroph av nigrotihialis a tachinid in rainfed habitats and Trichonialopsisapantelkcnela in irrigated ricefields A limgrotibialisalsoattacks the sweet potato hornwormAgrius convolvtdi and T apanteloctena parasitizes DH S1B

Predators as a group show more distinct environmental preferences The aquatic damselflies Coenagrionidae are most abundant in the lowlands Cvriorhinus is also most adapted to the lowlands The nabid Stenonabis tagalicus occurs only in the uplands and isalso prevalent on legumes The coccinellid lady beetles wasps and ants are more prevalent in upland areas AlicrasIpis cit-ea larvae prey ona variety of aphids attacking legumes and maize Etunienes campanonia vespid mud wasp makes nests in trees and is therefore most adapted to the more botanically diverse uplands Soil-dwelling ants cannot tolerate flooding

Upland environments are habitats to arboreal carabid beetles Three species of Chlaeniusbeetles prey on leaffolder larvae and are perhaps more important than parasites (27)

The gryllids and tettigoniids are widely adapted egg predators not only on rice but also on maize and legumes They feed on eggs of most insects that are laid on leaves

The spidercommunity of upland rice environments isrich in species At one site (Batangas Philippines) 31 species have been recorded (23) Of the 176 spider species recorded in Philippine ricefields about one-half (82) occur in upland rice The spiderspecies of upland rice environments overlap more with those of rainfed lowland than with those of irrigated lowland rice (24) The same study showed that of the three environments irrigated rice has (ie greatest spider species diversity foiiowed by rainfed lowland and upland

However of the 10 most prevalent ricefield spiders 9 were abundant in upland rice showing wide adaptation All three spider guilds -- orb-web space-web and huting spiders shywere prevalent in upland ricefields There were however differences in environmental prelerences for some spider species

Of the orb-wcavers Tetragniathamanlihulatawas partishycila rlv abundant 7 japonica was low and Ltucatge dehcorata was absent in upland rice

As in other environments Aiypeiwnaifirtosana was the most dominant space-web spider however its relative numbers were lowerthan for lowland sites Among hunting spiders Ox|opes javanus and Lcosa leatcost fgla were more abundant and Lvcosa lpseudoanmnlata was less abundant in upland compared with lowland rice envishyroniments

Extensive rearing of rice insect pests in upland rainfed lowland and irrigated sites over an 8-yr period in the Philippines provided insight into the effectiveness of parasites as natural enemies (Table 7 8) Most of toe data are from large sample sizes taken over at least one crop which overcomes the pitfalls expressed by Van Driesche (238)

VSB egg parasitization was surprisingly similar across all environments Larval parasitization was highly dynamic among sites and even years within the same site but showed somewhat lowerlevels in uplandsthan in irrigated wetlands This would have been predicted for egg parasites as well since YSB is adapted to the lowlands and so must be its parasites This isevidence that Telenomus rowani is as good at dispersing as its main hosts Trowani clings onto the anal tufts of female YSB and parasitizes eggs as they are laid This phoresy may explain its wide adaptation The more specialized lowland YSB larval parasites therefore appear more effective than their upland counterparts with more alternate hosts

Low collection levels of PSB larvae especially in the wetlands make comparisons between environments difficult Low levels of larval parasifization were recorded in both rainfed environments Except in the Cagayan rainfed wetland site in 1980 low levels ef larval parasitization were evident at irrigated and rainfed sitesalike No differences by environment were evident Higher levels of parasitization occurred in Sarawak during a 1967 outbreak also from tachinids Rothschild (197) indicated that higher parasishytization occurred in upland rice areas

Skipper and green horned caterpillar larval parasitization rates were generally low and showed no environmental effect htighest activity occurred in Cagayan in 1980 (48 parasitization) Although larval parasitization levels of RLF were variable year to year activity appeared greater in the irrigated lowlands than in the uplands Again the highest levels were recorded in Cagayan in 1980 Cagayan has extensive grasslands and fallow around the rice areas and a relatively long rainfall period Rainfall in 1930 was good for crop growth and apparently for increase of larval parasites

IRPS No 123 January 1987 33

Table 7 Parasitization rates of rice planthoppers and green leafhopper in upland rainfed lowland and irrigated lowland environments Philipshypines 1976-840

Environment and site Year Egg parasitization () Nymphaladult parasitization ()

BPH W13P GLH BPII WBPH GLIH

UplandBatangas

Rainfed lowland

1976 1979 1980 1982

15 17 12

2(47)8(39)

4(51)

9(1800)14 (212)

12(304)

8 (101)6 (127)

9(116)

Iloilo

Cagayan

1976 1977 1978 1981 1980 1981 1982

0 (69)

11

44 48 40

-

-

46 55 44

0 (471)

11

52 56 49

0 (200)9 (214)

10 (1259)8(235)6 (900) 3(800)

-7 (462)

11 (270)3 (313)5 (900) 1 (800)

4 (600)10 (725)10 (362)12(870)5 (900) 3 (800)

Pangasinan

Irrigatedlowland

1984 1976 1978 1979 1982

13 (436) 13(379) 14 (457)

6(14)7 (197)

17(198) 11 (486) 18 (275)

3 (81)10 (274)3 (116)

16(871) 19(88)

28(211)9 (764)5 (647)7 (1204) 8(305)

Laguna 1977 11 7 21 1978 31 1979 14 4 4 28 1980 7 8 20 1981 1983

56 7(278)

26 16 (1650)

16 27(1921)

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R I Apostol unpublished 1lo1lo and Pangasinan 136 Cagayan BCanapi unpublished Liuna 175 I C Pantua unpublished 118 234 43 Carino and Shepard unpublished BPH = brown planthopper Nila parvara htgens W131Pi =whitebacked planthopper Sogatelia furcifera GLH = green leafhopper Nephotettrix virescens Figiires in parentheses are sample sizes

Tauher et al (230) reported that internal external and genetic factors influence the seasonal activity of parasitoids

Hopper egg parasites but not nymphaladult parasites were also parlicularly abundant at the Cagayan site possibly the egg parasites have more alternative hopper prey in the weedy areas than nymphaladult paisites

The seasonal dynamics of hopper egg parasites can be seen for upland rice in Batangas(Fig I 2) and for rainfed lowland lice in Pangasinan( 135)and Cagayan( 105) Levels of BPHWBPI-L and GLH egg parasitization were corn-parable in upland and irrigated lowland environments Except for some high rates of paraMsitization early in the season the parasitization rate tended to be steady in all envilonments However hopper nymphaladult parasi-tizution levels were more similar in the rainfed environments and lower than in irrigated rice

Overall parasites do not appear highly effective by themselves against upland or even lowland pests Among natural enemies parasites have attracted the most attention because they can be more readily assesscd

Predators and pathogens lend themselves better to management practices Predators the most important group of natural enemies in riccfields are difficult to quantify On the other hand pathogens appear less im-portant but can be readily cultured and disseminated

The natural enemy commulnity of upland rice is rich in species and differs significantly from that of lowland rice

These beneficial organisms must be conserved by applying insecticides judiciously particularly sprays Seed or soil placement of chemicals minimizes exposure to natural enemies The strategy to derive the greatest benefit from natural enemies isto allow the greatest number of beneficial species to thrive Then perhaps at least one species will be effective against each pest at any tine overcoming the variable abundance of each species within and between years

Programs to introduce exotic species or mass produce indigenous ones are ambitious and expensive because they require trained people on a sustained not ad hoc basis

Chemical control Insecticides offer rapid and efficient control of upland insect pests (190) but-should be used only after other control measures have been considered Insecticides are rarely used on upland rice because of cost (32) Upland rice yields are normally too low to justify the expenditure Also most upland rice is grown as a cash crop Spraying upland rice is more difficult than lowland rice because water is less accessible Government subsidy programs to provide pestishycides to tribal peoples for upland rice production have created problems of toxicity to humans because isolated people have had no experience to be able to handle pesticides safely

However ins-eticide use has beenjustified when a crop is

0Z

Table 8 Parasitization rates of lepidopterous pests in upland rainfed lowland and irrigated lowland environments Philippines 1976-84a

Parasitization (5)

Environment and site Year YSB GFB PSB Earcutting Swarming Skipper Greenborned RLF Brown larva larva caterpillar caterpillar caterpillar caterpillar larva semilooperEgg Larva larva larva larva larva larva

UplandBatangas 1977 - 0 (57) - 0 (63) - 0 (71) - 0 (106) 11(142) 2 (101)1978 43 (4697) 8(451) 5 (38) 3 (40) 6 (61) 2(58) 15 (57) 13 (129) 13 (1286) 4 (30)1979 59(4135) 12(320) 13 (11) 5(28) 9(45) 6(46) 7(23) 8(160) 9(1152) 8(26)

Rainfed lowland Iloilo 1976 - 0 (10) 0(10) 0 (7) 0 (19) 2 13) 15 (180) 5 (76)1978 0 (54) 0 (3) 0 (17) 9 (64) 4 (78) 3 (73) 0 (121) 0 (191)1979 64 (2350) 16 (84) 6 (17) 0(6) 18 (68) 25 (104)

1981 56 (1322) 18(304) 0(5) 12(51) 18(138)Pangasinan 1976 60 (7 7 )b 62(82) 0(111) 5 (39)

1978 58 (53)b 5(51) Cagayan 1979 59(713) 8 (101) 6 (34) 0(7)

1980 0(11) 4 (28) 15 (286)

62 ( 48(177) 61(140)1981 0(5) 12 (2) 13(45) 14 (92)

Irrigatedlowland Laguna 1974 62 8

1974-75 50 41 1975 43 72 1978 96 (623) 21(24) 33 (60)1979 83(2017) 18(110) 8(62) 11 (89) 22(400)1980 78 (1625) 20(75) 0 (16) 4 (72)1983 60

9 (80) 7 (270) 30

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R F Apostol unpublished Iloilo and Pangasnan 136 Cagayan B Canapi unpublished Laguna 175 P CPantua unpublished 118 234 143 Carino and Shepard unpublished YSB = yellow stem borer ScirpophagaincertulasGFB = gold fringed borer Chilo auricilius PSB = pink stem borerSesamia inferens earcutting caterpillar Alythimna separata swarming caterpillar Spcdoptera mauritia acronyctoides skipper Pelopidas mathias reen horned caterpillar Melanitls ledaismene RLF = rice leaffoldersCnaphalocrocismedinalis and Marasmiaspp brown semilooper Mocisfrugalis Figures in parentheses are sample sizes Egg masses

IRPS No 123 January 1987 35

Table 9 Effect of seeding rate and insecticide on plant density and Yield losses from insect pests in upland rice often areyield of Dagge upland rice Batangas Philippines 1976a comparable to those of lowland rainfed or irrigated rice

Seeding rate Insecticide Dosage Plant density Yield (0-Y30o) but usually from different pests The principal (kg aiha) seed treatment (kg aiha) (noIra-row) (tha) groups are soil pests feeding on sown seed and rootsat 14 DE followed by seedling pests These groups have largely gone

50 None - 35 a 20 a unnoticed leading researchers to conclude that stem borers50 Carbofuran F 10 38 ab 23 a and seed bugs are the most important pests (17) Yield loss 10 beCabofuranF100 None - 4551 c 23 a trials that pinpoint major pests need to includc seed

100 Dieldrin WP 10 52 cd 22 a treatments Granules applied in the furrow may not give150 None - 59 d 22 a adequate protection from many seed pests

aAv of 8 fields DE = days after crop emergence In a column means The floristic and pedological diversity of upland rice followed by a common letter are not significantly different at the environments is matched by the diversity of arthropods5level by D)MRT F = flowable Wi = wettable powder both pests and their natural enemies Irrigation has homoshy

genized lowland rice environments Flooded soils tend tothreatened by epidemics of locusts or armyworms New have the same properties and to eliminate soil pests The technology stressing efficiency has minimized insecticide spatial and temporal dominance of cultivated rice has cost benefited specialist pests

Seed treatment is normally inexpensive and can be Upland rice has no specialist arthropod species The economicallyjustified in -nanycases(66) Treating seed with monophagous pests on upland rice prefer the lowlands systemic insecticide (216) but not chlorinated hydrocarbons where rice evolved Rice in the uplands exists only because (247) is most effective in controlling seedling maggots of man and even though lowland rice predates upland riceFoliar sprays have to be repeated to obtain control and specialist species evolved Wheat maize and sorghum do often are washed off by frequent early season rains not have monophagous pests leading us to conclude that Granular insecticides are effective but required dosages are there is no evolutionary advantage for inssects to have too expensive to justify except under heavy infestation narrow host ranges in upland graminaceae(247) When directed against first-instar larvae banding Pests have adapted to the uplands not only by having granules in seed furrows (177) can be effective against white wider plant host ranges but also by having long life cyclesgrubs at low dosages (025 kg aii ha) (134) Seed treatments undergoing dormancy or being dispersive Each pest has a are not promising for white grub control (248) unique life history strategy suited to highly unpredictable

Seed treatment is also effective ainst ants and may be environments In agivenyear someinsect specics an better cheaper than increasing the seeding rate In Batangas exploit upland rice as a food source but no pest is a senious treating 50 kg seed ha was as effective as increasing the one every year on upland rice seeding rate to 100 kgha and using untreated seeds (Table The low yielding potential of upland rice means less 9) In Brazil seed treatments control termites and other soil quantity and lower quality of food for pests to exploit but insects including the lessercornstalk borer(66) Domiciano also less incentive for farmers to undertake control (58) however found seed treatments inconsistent in their The most effective control would be early and synchroshyeffect against high populations of lesser corn stalk borer nous planting of early-maturing upland rice varieties to

Baiting can be an inexpensive way of controlling ants prevent insect pest buildup Breeding for insect resistance seedling maggot flies mole crickets and field crickets Baits should initially focus on stem borers because they are verymade of locally available material and impregnated with difficult to control by other means Biocontrol efforts insecticide can be sparingly distributed taking adantage of should first stress conservation of natural enemies preshythe pests ability to disperse and encounter bait sites dators and pathogens rather than parasites Efficient

The new low-volume hand-held controlled-droplet chemical control includes seed treatment or baiting for seed sprayers may offer an advantage over high-volume knap- seedling and some root pests and using controlled dropletsack sprayes for controlling foliar pests Upland rice sprayers for spot treatments against foliar pests and seed canopies are more open than those in lowland rice and bugs when economic thresholds are reached better droplet penetration should result Controlled droplet applicators would be ideal for seedbugs and defoliators

CONCLUSION

Upland rice is attacked by a wider array of insect pests than is lowland rice mainly beca use of the addition of soil pests Generally population levels of lowland rice pests are lower in upland than lowland rice leading many people to conclude that insect pests are not important on upland rice

36 IRPS No 123 January 1987

REFERENCES CITED

1Abraham C C and K S R Mony 1978 Occurrence ofLeptocorisa acuta Fabr (Coreidae Hemiptera) as a pest of nutmeg trees J Bombay Nat ttistSoc 74553

2 Agyen-Sampong M 1975 HlierogrlphusdiaganensisKrauss (Orthop-tera Acrididae) a new pest of rice in Northern Ghana Ghana J Agric Sci 8249-253

3 Agyen-Samnpong M 1976 Varietal resistance and other inle-ratcd pest control components as factors of rice varietal improvement in West Africa Prcsented at the Second varietal improvement serinar 14-18 Sep 1976 Bouak Ivory Coast West Africa Rice Development Association Monrovia Liberia 18 p

4 Agyen-Sampong M 1977 Outbreak of Ihccrorrdnisorae in rice seedlings near Rokupr Sierra Leone lit Rice Res Newsl 2(6)16-17

5 Agyen-Sampong M 1980 Dirty panicles and rice yield reduction caused by hugs IntRice Res Newsl 5(l)11-12

6 Agyen-Sampong M 1982 The major pest problems of irrigated upland and mangrove swanip rice ecosystems ir the humid tropical Guinea savanna and Salel climate zones Pages 318-328 in Pro-ceedings training course cn concepts techniques and application of irtegrated pest management in rice inWest Africa 10-20 Jan 1982 Fendall Liberia West Africa Rice l)evelopioent Association Monrovia Liberia 505 p

7 Akibo-Betts V I and S A Raymundo 1978 Aphids as rice pests in Sierra Leone lit Rice Res Newsl 3(6)15-16

8 Akinsola F A 1975 Present status of different rice stemborers in parts of Nigeria iicL Entomo Newsl 328

9 Akinsola FA 1979 Ihe biology and ecology of rice stemborers in Nigeria Ph 1) thesis University ol lbadan Nigeria 217 p

10 Akinsola E A 1979 Guidelines of integrated management of insect pests of upland rice in West Africa Presented at the Seminar on integrated management of rice diseases and insect pests at Bobo-Dioulasso Upper Volta 17-22 Sep 1979 West Africa Rice Develop-ment Association Monrovia Liberia

II Akinsola IA 1984 Insect pests of upland rice in Africa Pages 301-305 in An overview of upland rice research Proceedings of the 1982 lIouak Iwsrv Coast upland rice workshop International Rice Research Institute 13O Box 933 Manila Philippines

12 Alam M S V TJohn and Kaung Zan 1985 Insect pests and diseases of rice in Africa Pages 67-82 in Rice improvement in Eastern Central and Southern Africa proceedings of the international rice workshop at Iusaka Zambia 9-19 April 1984

13 Alticri M A A van Schoonhovcn and J D Doll 1977 The ecological role of weeds in insect pest management systems a review illustiated with bean (Phaseohsvulgaris L)cropping systems PANS (Pest Artie News Summ) 23195-205

14 Anmante F 1967 The leaf-cutting ant Atia capiguaraa pest of pastures [in Portuguese] Biol6gico (Sao Paulo) 33113-120 (Rev Appl Entomol A56207)

15Andrade A C and D Puzzi 1951 Preliminary results of experi-merits on the control of Sculptcoris iastantus on sugarcane [in Portuguese] Bioulgico (Sao Paulo) 1744-49 (Rev Appl Entomol A41421422)

16 Appert J 1970 Maliarplia separaelha (rice stem borer) new observations of entomological problems of rice in Madagascar [in French] Agron Trop (Paris) 25329-334

17 Ariadeau Mand Z Harahap 1986 Relevant upland rice breeding objectives Pages 189-197 in Progress in upland rice research Proceedings of the 1985 Jakarta conference International Rice Research Institute 1O Box 933 Manila Philippines

18Asar S and Kisimoto 1975 Seasonal occurrence of the smaller brown planthopper in upland rice [in Japanese] Proc Kanto-Tosan Plant Prot Soc 2228

19 Bakker W 1974 Characterization and ecological aspects of rice yellow mottle virus in Kenya Meded Landbouwhogesh Wage-ningen 152 p

20 Balut F F 1970 White grubs attacking rice Oryza ativa L [in Portuguese] Biol6gico (So Paulo) 36321-322

21 Banerjee S N and L M Pramanik 1967 The lepidopterous stalk borers of rice and their life cycles in the tropics pages 103-124 in The major insect pests ofthe rice plant Proceedings ofa symposium at the International Rice Research Institute September 1964 The Johns Hopkins Press Baltimore Maryland USA

22 Barrion A T and J A Litsinger 1980 Ants a natural enemy of cliaiarac in dryland rice IntRice Res Newsl

5(4)22-23 23 Barrion A T and J A litsinger 1981 The spider fauna of

Philippine rice agroecosystem 1Dryland Philipp Entomnol 5139-166

24 Barrion A T and 1 A Iitsinger 1984 The spider fauna of Philippine rice agroccosystcms II Welland Philipp Eiitomol 611-37

25 Barrion A T1 and J A Litsinger 1984 lihanus (Diptera Tabanidae) eggs an alternative host of rice stei borer (SI3) egg parasite Telenoonis digns (Iliymenoptcra Scelionidae) lit Rice Res Newvsl 9(6)19

26 Barrion A T and J A Litsinger 1985 Identilication of rice leaffolders by wing markings Int Rice Res Newsl 10( 1)24

27 Barrion A T and J A Litsinger 1985 Chctiosspp (Coleoptera Carabidae) a leaffolder ILF)predator lit Rice Res Newsl 10(1)21

28 Bertels A 1970 Insect pests of rice and theircontrol [in Portuguese] Pelotas Brazil Instituto de Expcrimentacao Agropccuararias do Sul Circular 43 24 p

29 Bonzi S M 1982 Ctilo (flisilineus J de Joannis (Lepidoptera Pyralidae) stem borer of irrigated cereals in Upper Volta LioFrench] Agron Trop Riu Rizic Cult Vivrieres Trop 37207-209

30 Bourke T V T L Fenner J N LStihick G Baker EHassan D F OSullivan and C S li 1977 Insect pest survey for the year ending 30th June 1969 Papta New Guinea )epartment of Agrishyculture Stock and Fisheries Port Moresby 57 p

31 Bowden 1 1973 Migration of pests in the tropics Meded Rijksfac Landbouwwet Gent 38785-796

32 Breniere J 1973 The main rice parasites in West Africa and their control Pages 27-42 in Proceedings seminar on plant protection for the rice crop West Africa Rice Development Association Monrovia Liberia

33 Breniere J 1976 The principal insect pests of rice in West Africa and their control West Africa Rice Development Association Monrovia Liberia 52 p

34 Brown ES 1962 The African arinyworn Spoiolptera eXI)Ina (Walker) (Lepidoptera Noctuidae) a review of the literature Cornshymonwealth Institute of Entomology London UK57 p

35 Brown F B 1968 Upland rice in Latin America FAO-International Rice Commission 12th Session of the Working Party on Rice Production and Protection 2-14 Sept Peradeniya Sri lanka Food and Agriculture Organization Rome Italy 6 p

36 Buyckx F J 1 1962 Disease and insect pests of crops inCongo Rwanda and Burundi [iil French] Publications dc LInstilut National pour lestude Agrononiquedu Congo M Weissenbruck S A Brussels Belgium 708 p

37 Cariiio F 0 1981 Role of natural enentics inpopulation suppression and pest management of green rice lealhoppers Ph D thesis University ofthe Philippines at Los Bafitos PhilippinesLaguna 136 -

38 Carpente A J 1983 The development of rice cultivation and extension of vegetable productin in Zanzibar Final Report Vol II1 October 1983 Food and Agricultme Organization Ronte Italy

39 Carroll C P and S1 Risch 1983 Tropical annual cropping systems ant ecology Environ Manage 751-57

4(0 Cates R G 198 1 Host plant predictability and the feeding patterns of monophagous oligophagous and polyphagous insect herbivores Oecologia (Berl) 48319-326

IRPS No 123 January 1987 37

41 Cendcafia S M and F B Caora 1967 Insect pests of rice in the 62 Ene 1 C 1981 I)iiorldhol)ierusihilis Slater (licteroplera Philippines Pages 591-616 in The najor insect pests ofthe rice plant lygacidue Illissinac) a new rice pest front Nigeria lit Rice Res Proceedings of a symposium at the International Rice Research Newsl N3)15 Institute September 1964 The Johns Hopkins Press Baltimore 63 Fugenio T S 1965 rhe effect oi e-vCgetation on rice production Maryland USA 729 p and the possibility of overcoming it by leaf pruning MS thesis

42 Centro Ihternacional de Agricklittira Tropical 1984 Upland rice in University ol the Philippines at Los Baijos ILaguna Ihilippins 69 p Iatin Anerica Pages 93-119 hi An os ervies of uplaind rice research 04 Farrow R A 1975 The Alrican nmigratory locust ii is main Proceedings of the 1982 Boinakamp Ivory Coast upland rice workshop outbreak area of the middle Niger qlinlitatise studies of solitary International Rice Research Institute P0 Blox 933 Manila populations in relation to environmental factors Locusla No II Philippines 198 p

43 Chandra G 1979 Taxonomy ad hn riononlics ofthe imect parasites of 65 Fennal R G 1943 Food crop investigation Windward and rice Icalloppers and planthoppers in the Pihilippines and their Ieeward Islands First report Report onl work dunring the periodiiiportairce iii natural biol)gical cointrol Philipp Fntoltol 4119-139 April 1942-April 1943 Imperial College of Tropical Agriculture

44 Chang T T 1976 The originii colut ion enlIivation dissemination Trinidad 16 p (Rev of Appl Fintornol A3222-25) and diersificition of Asian and African rices -uphytica 25425-44 1 66 Ferreira F 1983 Integrated pest ritaageiret of upland rice [ill

45 Chatierjee P B 1979 Rice leaffolder attacks in India lit Rice Res Portuguese English summary] Pages 323-341 in Upland riceculttre Newsl 4(3)21 factors affecting production E Ferreira T Yamada and F-

46 Cheng Cbien-pan 19(4 Upland rice in Taiwan International Rice Malmolta eds Institutoda Potassae Foslato Institnto Internacional Cnmmission Wiorking Party oit Rice Production ard Protection I()th da Potassa Piracicaha Sao Paulo lraid Meeting 3-10 March 1964 Manila Philippines Food and Agricul- 67 Ferreira E arid F dl Silva Martins 1979 Occurrence of ture Organiaion Rome Italy 5 p Sogatohs orvziola (Muir) inupland rice in Goias Ilrail lit Rice

47 Chcrrett 1 M and 1) 1Peregrine 1976 A review of the status of Rcs Ncwssl 4(4)18 leaf-cutting ants and their control Ann Appl Biol 84124-128 68 Ferreira E and 1F da Silva Martiris 1984 Insect pestof rice ill

48 Chipeta F M 1974 Pests of rice in Malawi Pages 90-95 in Brazil and their control Jill Portugucsej FMIlRAtPA-CNPAF Proceedings Sith eastern Africa cereals research conference 10-15 Goiania Brmiil 67 p March 1974 DRB Mialda ed Ministry of Agriculture lilongwe 69 Ferreira FI F ca Silha Miartins S S Nero and 1 It P de Malawi Carvallio 1982 M tltiplc regression analysis todeterimine the effect of

49 Chu Yau-i 1979 The d atiaging period of the rice seedling fly insect populatiois ott upland rice yield [inIPortuguese Pesqui (Atlterigotiavxigu Stcin) (NItiscide Diptera) oil cori) at Fast Java Agropecu Bras 17671-675 Indonesia Plant Prot Bill 21403413 70 Ferreira F 1F da Silva Martins and F 1PZimmermann 1979

50 Conklin H C 1957 IHaninoo agriculture a report on aln integral Rice varietal resistance to tle lesser corn stalk borer [in Port uguese] systent of shi fti rig cult ivationi in the Pltilippincs Food and Agriculture Pesqui Agropccu Biras 14317-321 Organization Rome Italy 209 p 71 Fleming W F 1968 Biological control of lie hapanese beetle U S

51 Cramer H II 1967 Plant proection and world crop production Dep Agric RLs Sers [ech Bull 1383 78 p Pilaniensclhutz-Nachr 201-524 72 Food and Agriculture Organiation of the United Nations 1982

52 l)ani R C and Majunidar 1978 Prelininary observations on tie Country report surmimary Pages 27-37 in Report on the first incidence of root aphids on differeit rice cultnrcs Sci Cult 4488-89 FAOIUN)IPhail and regional training coiuirse in iniproved ctltural

53 Davidson R I 1979 lffects of root feeding on foliage yield Pages practices for upland rice 5-23 Oct 1981 Northern Region Agrculturd117-120 in T K Ci osb and R B Pottinger eds Proceedings of the Developririt Center Chiang Mai Food and Agricuilture Organiiashy2nd Australian conlerence on grassland invertebrate ecology Govt tion Rome Italy 37 n Printer Wellington New Zealand 73 Franco do Ariaral S 195 1 Residual action of insecticides in turfy

54 l)ean G 1W 1976 Rice insect pests inlaos Int Rice Res Newsl soil--the results of a field experiment [in Portuguese] Biol6gico (Sao 1(2)15 Paulo) 17209-214 (Rev Appl Entomol A41423)

55 I)ean G J W 1978 Insect pests of rice in Laos PANS 24(3)280-289 74 Freeman ) 1970 Report onithe Iban london School of -coroiics 56 )ingle H 1984 Behavior genes and lifc histories conplex adapta- Monographs on Social Anthropology No 41 Athlane Press

tions to uncertain environments Pages 169-194 in A new ecology London UK 317 p novel approaches to interactive systems P W Price C N Slobod- 75 Fuchs T W1 A Harding and 1W Smithlr 1979 Induction and chikoff and W S Gaud cdsl ohn Wiley New York USA termination oldiapause in the sugarcane borer Ann Flitonlol Soc

57 Dobelmann-JP and M Falais 1968 Upland rice cultivation ill tie Am 72271-274 North-western part of Madagascar in 1966-1967 [in French English 76 (hbriel B1) 1975 A review tf the major insect pests of some tpland summary] Trop Agron 2313-21 crops in tlc Philippines Pages 86-111 in Ad Htoc panel of experts on

58 Doniiciano N L 1979 Chemical control of tie lesser corn stalk pest disease arnd weed problems in some rainfed crops Bangkok (lolsonno~paloIgnowslhts) (Zeller 1848) (I-epid(ptera Ihycitid ae) Thailand Food aid Agriculture Organization Roire Italy

on rice in Ponta Gressa Parara 1977-1978 [in Portuguese] Pages 77 Garg D K and N K Shal 1983 White grub outbreak on rainfed 291-303 in Anais 1 Reuniao de tecnicos cmiirizicultura do estado de dryland rice in Uttar Pradeslh Int Rice Res Ncwsl 81(4)16-17 Sao Paulo-Campinas Cati Brazil 78 Garg I) K and IP Tandon 1982 Major insect pests of rice on hilly

59 Dunsmore 1 R 1970 Investigations on tlie varieties pests and tracts of Uttar Pradesh India lilt Rice Res Newsl 7(11)11-12diseases of upland rice (hill padi) in Sarawak Malaysia Int Rice 79 Garrity 1) P 1983 Upland rice research an international bibliograpy Comm Newsl 1929-35 1975-1982 Iternational Rice Research hnstitnte PO Box 933

60 Elias 1R1965 The new decline of rice seedlings is not caused by Manila Philippines 121 p parasites [in Portuguese] Biol6gico (Sao Paulo) 313-6 80 Ghauri M S K 1982 A new subspecies of Iinorh)rone bull

61 Empresa Brasiliera de Pesquisa Agropecuaria 1984 Upland rice itt cornlus Slater (Hemiptera Iygaeidae) front Papua New Guinea on Brazil Pages 121-134 it Av overview of upland rice research rice and carpet grass Ihll lEntoniol Res 72133-137 Proceedings ofthe 1982 Blouak Ivory Coast Upland rice workshop 81 Gonzales F 0 PArregoces R I ihernandef and OT Parada International Rice Research Institute PO Box 933 Manila 1983 Insect and imite pests of rice arid their control in Latin America Philippines [in Spanish] la Federacion Nacional de Arrocerts (Fedearroz)

Bogota Colombia 60 n

38 IRPS No 123 January 1987

82 Grist D H and R J A W Lever 1969 Pests of rice Longmans London UK520 p

83 Hale P R 1979 Two subterranean pests of upland rice in Papua New Guirnca IntRice Res Newsl 4(2)16-17

84 Hale P R and J T Hale 1975 Insect pests of rice in the New Guinea Islands Rice Entomol Newsl 34

85 Harris W V 1969 Termites as pests of crops and trees Common-wealth Institute of Entomology London UK 41 p

86 Hattori 1 1968 A method to distinguish between grass leaf roller (Ctaplha rocis medinalisGuenee) and the rice leaf roller (Susurtia exigua Butler) [in Japanese] Plant Prot (Japan) 23167-170

87 Henry I E and 1A Onsager 1984 Experimental control of the mormon cricket Iahrosshnph by NJsena locustae(Aficrospora Microsporidae) a protozoan parasite of grasshoppers (Ort Acri-didae) Entomophaga 27197-201

88 Hidaka 1 P Vungsilabulr and SKadkao 1974 Studies on ecology and control of the rice gall midge in Thailand Tech Bull 6 TropicalAgricultural Research Center Tokyo Japan 113 p

89 Hincklcy A 1) 1963 Ecology and control of rice planthoppers in Fiji Bull Entomrol Res 53467 481

90 Hirao J 1982 Ecology and chtInical control of the rice leaffolder Jpn Pestic Inf 4114-17

91 Ho DT 1984 Sten borer incifience in rice ecosystems in KenyaEast Africa Pages 307-320 in A overview of upland rice research Proceedings of the 1982 Bouac Ivory Coast upland rice workshop International Rice Research Institute PO Box 933 Manila Philippines

92 Ho )T J 1Njokah and J GKibuka 1983 Studies on rice sten borers in Kenya with emphasis on ifaliarpiaipiraellRag Insect Sci Appl 465-73

93 Ichihara I T Ito A NIatsuo and M Sawada 1971 Chemical control of the rice striped stem borer in upland rice grown in plastic mulch [iii Japanese] Proc Kanto-Tosan Plant Prot Soc 1882

94 IclhiharaIand A Matsuo 1969 Damage caused by rice striped stett borer to upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1686-87

95 Ichhara I atid A MIatsun11970 Conside ration oft lie extraordinary prevalence of rice striped stem borer in upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1778-79

96 Institut de Recherches Agrolnliqes Iropicale et des Cultures Vivrieres 1978 Insect resistance Pages 12-13 in Upland rice 1978 atnual report Bouak Ivory Coast 27 p

97 International Institute for Tropical Agriculture 1983 Aphids itt rice roots I ITA Res Briefs 4(1-3)4-5

98 International Institite fi Tropical Agriculture 1983 Hopperburn in rice caused by brown plathopper IITA Res Briefs 4(l-3)6-7

99 International Rice Research Institute 1975 Annual report for 1974 PO Box 933 Manila Philippines p 216

100 International Rice Restarch Institute 1976 Annual repiort for 1975 PO Box 933 Manila Philippines p 228

101 International Rice Research Institute 1979 Cropping systems program Component technology development and evaluation Page 404 i Annual report for 1978 11O Box 933 Manila Philippines

102 International Rice Research Institute Department of Entomology 1981) Insecticid evaluation 1979 Tables 73 74 PO Box 933Manila Philipes

103 International Rice Research Institute Department of Entonmology 1981 Insecticide evalomtion 1980 Table 81 POBox 933 Manila Philippines

104 International Rice Research Institute 1981 Report of a meeting of Asian rice-based cropping systems entomologists Eleventh cropping systems working group meeting 18-22 May 1981 PO Box 933 Manila Philippines p 132 133 144

105 International Rice Research Institute 1984 Cropping systems program Pages 368-373 in Annual report for 1983 PO Box 933 Manila Philippines

106 International Rice Research Institutc 1984 Upland rice in Asia Pages 45-68 in An overview of upland rice research Proceedings of the 1982 Bouakamp Ivory Coast upland rice workiiop loternational Rice Research Institute 11O Box 933 Manila Philippines

107 Irshad M 1977 Grasshoppers in rice in Pakistan lit Rice Res Newsl 2(3)8-9

108 Ito R and TOno 1964 Upland lice its culture and varieties in Japan lIternational Rice Cormmission loth Meeting of the Working Party on Rice Production and Protection 3-10 March 1964 Manila Philippines Food and Agriculture Organization Rome Italy 12 p

109 Jeffrey M I and V A Dyck 1983 Effect of moonlight on lightshytrap catches of brown planthopper lit Rice Res Newsl 8(6) 13

110 Jennings P R and T Alicia Pineda 1983 Management ofthe rice plnthoppers Sogatodsorivi(coIl in Latin America Presented at the meeting on stable plant resistance FAO and lIndocsian I)inectorate of Food Crops Protection Bali lndonesia 29 Novcniber-2 l)ccemshyber 9 p

IllJimenez Gomez J A 1979 Rice insect pests i lla ios ()riCntales lil Spanish] Pages 99-103 in Rice ciltivation Instituto (olotibiatto Agropecuario Compndio No 29

112 Jongeleen F JJ Soesanto and A Mockidjo 1979 Studies oti pathogens of two species of Scarabacidac froinIndonesia Z Angew Enlomnol 87412-417

113 Jouvenaz 1) PC S lofgren and W A Baiks 1981 Biological control of imported fire antsa review of cirrent knowledge Bull Entomol Soc Am 27203-208

114 Kainoh Y K Shiniiu S MarUani Y Taiaki 1980 lost finding behavior of tie rice bu lplotori+JchinvittjI)allas (liciniptera Corcidac) with special reference to diet pattern ol aggregation and feeding on the rice plant Appl Entomol Zool 15225-233

115 Kakkar K 0 P Sharma and P I Sharma 1977 Note on Ihteronyrchus liodresRedt (Coleopicra I)ynastidae) as asporadic pest of paddy and maie inHimachal Pradesh Corr Agric 156-57

116 Kalode M131 79 Major insect pest problems and their control in upland rice (Unpublishcd mineogriph CWI oJNaaiat lni Chandaratat (198 1)original not seen)

117 Kalshoven L i 98 I The pests of crops in I ldoniesia (translated by P A vatn der Laan) 1 T Ichtiar B3aru-vati Iloeve lakarta Indonesia 701 p

118 Kamal N Q 1981 Suppression of whitebacked planto ner Sogatella fircaifra (Horvath) aLId rice lealfolder (nhal -cis licdb lis (Guenee) populations by natural cnclmie hlI) tiesis Grcgorio Araneta University Foundation Victoncta Park Metto Manila Philippines 179 Ip

119 Karim A N M Reaul 1975 A species ofliealy bug Trioni i70s ceres new oii rice in the Philippines Rice Entoniol Newsl 247

120 Katanyukul Wand C Chandaratat 1981 Insect pestsof uplaid rice and their control Report firstFAOUNlFi Tliailand training course on improved cultural practices for tplad rice Chiang Mai Thailand 5-23 Oct 1981 Food and Agriculture Organiation Rome Italy 22 p

121 Kern F 1955 lhletwioa(I)uninemts) bhhit Biturm(Coleoptera) a new pest of sugarcane inVenezuela [inSpanish with Etnglish sunmary] Agron Trop (Maracay) 527-29 (Rev Appl Eitoniol A4572)

122 Khan K M and S Ghai 1974 White grubs ald their control in India Pesticides (1Bombay) 819-25

123 Kiritani K 1975 Integrated pest management of rice pests Pages 66-70 itt Proceedings platning workshop ott cooperative hIeld research in pest managementP SMotooka ed East-West Center Honololu Hawaii USA

124 Kisimoto R 1967 Ecology of insect vectors forecasting and chemical control Pages 243-255 in The virus diseases of tle rice plant Proceedings of a symposium atthe International Rice Research Institute April 1967 The Johns Hopkins Press altimore Maryshyland USA

125 Kisimoto R and V A Dyck 1976 Climate and rice insects Pages 367-391 in Climate and rice International Rice Research Institute PO Box 933 Manila Philippines

126 Kureha S S Kobayashi SAsari and T Ishihara 1977 Life history and control of the mole cricket Grrllotalpa africana Palisot de Beauvois [in Japanese English translation 7520 1RRI library] Proc Kanto-Tosan Plant Prot Soc 21192-197

127 Kushwaha K S 1981 The black beetle as a pest of rice in Haryana India Int Rice Res Newsl 6(4)14

128 Lawani S M 1982 A review of tile effects of various agronomic practices on cereal stem borers Trop Pest Manage 28266-276

129 Leuck D B 1966 Biology of the lesser cornstalk borer in South Georgia J Econ Entomol 59 797-801

130 lever R J A W 1955 Insect pests of the rice plant in Malaya Int Rice Comm Newsl 1612-21

131 Li Ching-Sing and Shin-Foon Chiu 1951 The study ofthe rice gall midge Iachydiplosis orvza Wood Mason [in Chinese with English summary] 1Taiwan Agric Res 21-13

132 Iitsinger J A 1984 Assesament of the need-based application for rice Proceedings TechnologyTransfer Workshop 15-16 March 1984 International Rice Research Institute 1PO Box 933 Manila Philippines 6 p

133 Litsinger J A 1984 Environmental diversity and insect pest abundance with reference to the Pacific Pages 285-307 in Proceedings sub-regional training course on methods of controlling diseases insects and other pests of plants in the South Pacific 4-20 Oct 1982 Vaini Kingdom of Tonga Nukualofa

134 Litsinger J A RF Apostol and M BObusan 1983 White grub Leucopholis irrorata (Coleoptera Scarabaeidae) pest status popula-tion dynamics and chemical conllol in arice-maize cropping pattern in the Philippines J Econ Entoinol 761133-1138

135 Litsinger J A C de la Cruz F Raymundo A T Barrion M D Lumaban J P Bandong M S Venugopal F Paragna and E Balete 1982 Insect pests and insecticide reponse of dry-seeded bunded rice Pages 179-193 in Report of workshop on cropping systems research in Asia International Rice Research Institute Los Bafos Philippine

136 Litsinger J A M 1) Luinaban J P Bandong A T Barrion W Katanyukul and N Panda 1978 Insect pest management in rainfcd lowland rice agroecosystms of tropical Asia International Rice Research Conference 17-21 April 1978 International Rice Research Institute Los Bafios Philippines

137 Litsinger J A and K Moody 1976 Integrated pest management in multiple cropping systems Pages 293-317 in R I Papendick P A Sanchez and G B Triplett eds Multiple cropping American Society of Agronomy Madison Wisconsin USA

138 Loevinsohn M E 1984 The ecology and control of rice pests in relation to the intensity and synchrony of cultivation Ph D thesis University of London 36) p

139 Loevinsohn M E J A Litsinger N Panda M A Austria R F Apostol C Andrion and J Bandong 1982 A basis for predicting rice insect pest abundance in cropping systems Paper presented at the 1982 International rice research conference on rice and cropping systems 19-23 April 1982 International Rice Research Institute Los Bafios Iaguna Philippines 25 p

140 Lopez A W 1930 Repot of the Entomology Department Pages 145-172 in Annual report of the Philippine Sugar Association 1929-30 Manila

141 Lopez AW 1931 Entomology Department Pages 227-266 in Annual report of the director or research 1930-31 Philippine Sugar Association Manila

142 Lopez A W 1932 Entomology Deparment Pages 252-279 in Annual report of the Philippine Sugar Association 1931-32 Manila

143 Lungibill P and G G Ainslee 1917 The lessercornstalk borer US Dep Agric Agric Res Serv Tech Bull 539 27 p

IRPS No 123 January 1987 39

144 McGuire J U Jr and BS Crandall 1967 Survey of insect pests and plant diseases ofselected food crops of Mexico Central America and Panama U SDepartment of Agriculture International Agriculshytural Development Service Agricultuial Researrh Service October 1967 11 p

145 McNeill Sand T R ESouthwood 1978 The role of nitrogen in the development of insectplant relations Pages 77-9 in lliochemical aspects of plant and animal coevolution J B Ilarbornu ed Academic Press London UK

146 Maebara H 1955 Spatial distribution of lastinstiarlanaeof purplish stem borer (Sesamia iqferens Walker) hibernating in an upland rice field Bull Kagoshima Univ Fac Agric 438-41

147 Mallick E H and N G Hajra 1977 Response of rice plant to defoliation treatments at different developmental stages Indian Agric 2151-56

148 Mansingh A 1971 Physiological classification of dormancies in insects Canl Entomol 103983-10)9

149 Martell C G 1974 Catalogue of plant feeding insects of Mexico [in Spanish] Fitofilo 27(69)1-176

150 Martinez 0 J 1982 Principal rice pests in Ilanos orientalis [in Spanish] P~ages 80-94 in Curso deact nalization de conocimientos en arroz Villaniencis Instituto (olonmbiano Agropecuario

151 Martins J F da Silva F Ferrcira A S Prabhu F 1 P Zimmermann 1980 Use of pesticides for the control of the main subterranean upland rice pests esqui Agropecu Bras 1553-u

152 Martins I F da Silva F Fe eira rod BdamSilveira Pinheiro I Q2 Damage simulation caused by defoliators of upland rice [in Portuguese with English summary] Pesqui Agropecu Bras 171113-1119

153 Martins 1F da Silva N(iyuen van Tan and 1Ida Silveira Pinheiro 1981 Stem borer resistance in upland rice associated with plant morphological characters [in Portuuce Iesqui Agropecu Bras 16187-192

154 Masaki S 1980 Sumnmer diapause Ann Rev Entomiol 251-25 155 Masuzawa T 11 Suwa and F Nakasoji 1983 l)ifkleences of

oviposition preferencc and survival rate of two skipper butterflies Parnara guttata and PIchlpidai iathiais (Iepidoptera HIesperiidac) on rice plant and cogon grass New Entomol 3247-56

156 Maurya D M and C P Vaish 1984 Upland rice in India Pages 69-92 in An overview oftpland rice research Proceedings ofthe 1982 Bouakamp Ivory Coast upland rice workshop International Rice Research Institute IO Box 933 Mnila Philippines

157 Menon M G R R N Katiyai S Kuar and S S Misra 1970 Termites attacking the paddy crop Sci Cult 36294-295

158 Miyahara Y and 1 Nishikubo 1967 Behavior of young nymphs of the red rice root aphid Iholahsipiun ri liabhdontinalisSasaki in relation to movement from the leaf to the roots of upland rice [in Japanese] Pioc Assoc Plant Prot Kyushu 1363-66

159 Mo ljoa Tjien 1952 White grub of upland rice (Holotrichiahelleri Brsk) [in Indonesian with English summary] Contrib Agric Res Stn Bogor No 13134 p

160 Mochida 0 and T Suryana 1979 The occurrence of the brown planthopper and ragged stunt on lowland and upland rice ill Central Java and logyakarta in tile late of January 1979 Entomnolgy Survey Report No 15 Central Research Institute of Agriculture (CRIA) Sukamandi Indonesia I February 1979 9 p

161Monet C 1979 Varietal resistance to upland rice pests [inI French] Pages 35-39 in Annual report 1977 Institut des Savanes Department des Cultures Vivrieres Bouak6 Ivory Coast

162 Monte 0 1942 A caterpillar ofrice-fields[in Portuguese] Biol6gico (Sao Paulo) 8161-163 (Rev Appl Entomol A31414)

163 Munroe I) D 1979 Pest management strategy for hill rice in four resettlement areas in Sarawak Malaysia Pages 89-102 i1 Guidelines for integrated control of rice insect pests Food and Agriculture Organization Plant Production and Protection Paper 14 Rome Italy 115 p

40 IRPS No 123 January 1987

164 Nakasuji F M Ishii and Jen-Zon Ito 1984 Rice skippers in Taiwan and their life histories ZZrC (Taipei) Tech Bull 82 10 p

165 Navas I) 1976 Fall armyworum in rice Pages 99-106 il Ecological animal control by habitat management Proc Tall Timbers Con-ference 28 Feb-I Mar 1974 Fall Timbers Res Stn Misc Publ 6 Tallahassee Horida USA

166 Nickel 1 1 1961 Biological control of rice stem borer A feasibility study Tcch Bull 2 International Rice Rcarch Institute PO Box 933 Manila Philippines IIl p

167 Nigan P M and R A Vernia 1985 Insect pests of uplaild rice in Uttar Pradesh Int Rice Res Newsl 10(6)22

168 Nilakhe S S 1985 -cological observations on spittle bugs with emphasis on their occirence in rice Pcsqtui Agropecu Bras 20407-414

169 Nixon G EJ 195 1 lie association of ats with aphids and coccids Commonwealth Iusti lcte of Entonology Iondon UK 36 p

170 Nurdin F an Ishak Manti 1985 Study oit seedling fly of upland rice Interlational Upland Rice Moitoring Tour Report 26 lebruary-2 MarLI Sukaranti West Sumatra

171 Oka 1 N 1979 Ciltural conttol of the brown planthopper Pages 357-369 in Brown planthopper threat to rice production in Asia International Rice Research Institutc 1p0 Box 933 Manila Philippines

172 Ohni M and A Vilnaii 1975 Rice insect pests of Italy biology of Sipha grcria (Kaltenbach) and its control [in Italian] Riso (Milan) 2459-71

173 Otanes F Q atnd 1 I Sison 194 1 Pests of rice Philipp J Agric 12211-259

174 Painter R I 1951 Insect icsistance in crop plants University Press of Kansits Lawrence Kansas USA 520 p

175 Pantua P C and 1A litsinger 1981 Arthropod abundance in continuous vs biannual rice cropping systeis Int Rice Res Ncwsl 6(2)20-21

176 Pathak M ) and V A l)yck 1975 Sttudics ott insect pests of upland rice Pages 186-191 in Major research in upland rice International Rice Research Institute PO Box 933 Ilanila Philippines

177 Patil B R B M IHasabe and 1) S Ajri 198 I Chemical control of white gritb lIditriihinhorraia Fabr intfesting groundntut paddy atnd sorghum Indian 1Entomol 43232-236

178 Pawar A 1) 1974 (ollection and identification of insect pests of rice rlerminal repirt lInternational Rice Research Institute PO Box 933 Manila Philippines 72 p (tnimco)

179 IerrajU A aid A S Reedy 1965 Observations ott diapausc itt tie steti borer ohrice Rice News Teller (lai-Mar)9-13

180 Pont A C 1972 A review of the Oriental species of theri ona Rondani (I)iptera Muscidac) of econtomic importance Pages 27-102 in Control of sorglium shoot fly M G lomtani and W R Young eds Oxford atd IH1 Ptthl Co New I)clhi India 324 p

181 Prasad S C 1PChand aid M 1PSingh 1983 Incidence of rice gall midge in the upland plateau region of Hihar Oryza 20235-236

182 Rai P S 198 0 Nisaga simpex damage to rice in the hill tracts of South India Int Rice kles News[ 5(I)12

183 Rathorc Y S 1969 Gonocplhahli dcprssunt Fabr as itpredator of (tilo zonwIlhis (Swinhoc) and other moths in field cage at Patnagar Uttar Pradesh Indian 1Entotnol 3127

184 Reed W 1983 Crop losses causcd by insect pists in the developing world Proceedings 10th Internatial Congress of Plant Protection lrighton UK 174-80

185 Reedy M V 1981 Ateactogasterinhens(Walker)a new root-grub pest of tipland paddy and inaize Curr Sci 501076

186 Reveno M A 1968 Catalogue of the insect pests of economic crops in Colombia [in Spaiish] Asociacion Latino Americana de Entomo-logia (AIAE) 129-33

187 Reyes G M 1957 Rice dwarf disease in the Philippines Food and Agriculture Organiiation Plant Prot Bull 617-19

188 Rice S E A A Grigarick and M 0 Way 1982 Effect of leaf and panicle feeding byarmyworm (Lepidoptera Noctuidae) larvae on rice grain yield Environ Entomol 7593-595

189 Risch S J 1) Andow and M A Altieri 1983 Agioccosystem diversity arnd pest control data tentative conclusions and nizw research dirctions Environ Entomol 12625-629

190 Rivera C T 1956 Three organic insecticides in thecontrol of insects affecting upland rice Philipp Agric 39465-472

191 Roffrey 1 1972 Locusta outbreaks in the Philippines Acrida 1177-188

192 Rojas R 1) 1941 The effect of pruning at varying ages upon the yield of Ramairio Phipp Agric 29851-860

193 Roonwal M i 1979 Ternite life and termite control in tropical South Asia Scientilic Pnbl odhpr India 139 p

194 Rosetto C1 F da Silva Martins N C Schmidt and L EAzzini 1978 Damage caused by pasturespittle bugs 4Dvoi)lavopictaandDJ schach) on rice lBragantia Nota 6 (Matcli)25-27

195 Rosztto C1- S S Neto I- Amantc I) Iink 1Garcia-Vieira D M de SoLza N V Bansa t and A M de Olivcira 1971 Rice pests of Brazil [in Portuguese] Pages 149-238 inrFA()-International Rice Commission 2nd meeting of the Rice Committee for the Americas 6-11 l)ecember 1971 Pelotas Rio Grande do Sul Brazil

196 Rothschild G H L 1969 Obseratiois on the armyworm Spodoptera matritia acronrctoiiles Gn (Lep Noctuidae) Bull Entomol Res 59143-160

197 Rothschild G 1- L 1974 Parasites of the armworm Spolopiera mauritiaacroncitoihes in Malaysia Fntornopliaga 19293-299

198 Sachan i N S K Gangwar and 1N Katujar 1980 Field crickets damaging cages in North Eastern IHill Region Indian J Entomol 42526-527

199 Sands 1) P A 1977 The biology and ecology of Lepocorisa (Hemiptera Alydidac) in Papua New Guinea Res Bull 18 Departshyment of Primary Industry Port Moresby Papua New Guinea 104 p

200 Sarawak 1968 titomology division Pages 97-106 in Report of the research branch of the Department of Altriculture 1966 Kuching (Rev AIppl Eintomol A56603)

201 Satpathy JM M S Das and K Naik 1977 Effect of multiple and mixed cropping on the incidence of scine important pestsI Entomol Res 178-85

202 Sauer IHF G 1939 Notes Ott Elasmpnlpualtsligniosellths Zeller (Lep Pyr) a serious pest ofcereals in Sao Paolo [in Portuguese] Arq Inst Biol 10199-206 (Rev Appl Entotnol A28378)

203 Satinders 1L A 1 SKing and C L Vargas S 1983 Crop pests of Central Anerica with t bibliography [in Spanish] Centro Agroshynotnico Tropical de Investigacion y Ensenanza Turrialba Costa Rica (CATIE) Tec Bol 9 90 p

204 Sauphanor B 1985 Some factors of upland rice tolerttce to stem borers in West Africa Inseu Sci Appl 6429434

205 Sen P atnd S Chakravorty 1976 Stentflics in Haritighata Nadia West Bengal India [it Rice Res Newsl 1(12) 15

206 Senapai B and 1M Satpathy 1983 The seedling fly Atherigona orrzae Malloch is a pest of rice in Orissa (India) J Entomol Res 777-79

207 Shhjahan M S Alatn M Ah nad and BN Islam 1983 A new scarabaeid beetle pest of dryland rice in Bangladesh Int Rice Res Ncwsl 8(I)16-17

208 Sharma G L S C Modgol and R C Gautam 1979 Effect of depth of submergence ott incidence of bacterial blight and white grub infestation in transplanted rice Int Rice Res Newsl 4(6)12-13

209 Sharma S 1985 Scarabaeid beetle outbreak itt upland rice Int Rice Res Newsl 1(1(l)24

210 Shiraki T 1917 The paddy borer Schoenobhits incertulas Walker [achaku Agric Exp Stn Formosa 256 p

211 Silva A de Brito and 11P Magalhaes 1981 Insect pests of rice [in Portuguese] Circular Tecnica No 22 EMBRAPA Centro de Pesquisa t ecuaria de Tropico Unido (CPAJU) Belem Brazil 14 p

212 Singh S A 1979 Emergence of rice leaf butterfly as a rice pest in Manipur India int Rice Res Newsl 4(4)16-17

213 SinghS A 1983 Changing trends of insect pestsof rice in Manipur Int Rice Res Newsl 8(f)12

214 Siwi S S and P H van Doesburg 1984 Lenovorisa Latreille in Indonesia (Heteroptera Coreidae Alydinae) Zool Meded (Leiden) 58117-129

215 Soejitno J 1977 Reaction of some upland rice varieties toseedling fly eltherikonaexigua Stein [in Indonesian] Siri Hana I Penyakit 630-35 Lembga I osat Fenclilian Pertanian Bogor Indonesia

216 Socnardi 1967 Insect pests of rice in Indonesia Pages675-683 in The major inse pests of thu rice plant Proceedings of a symposium at the Internatital Rice Rec Wi itt S pternber 1964 The Johns Hopkins Press Baltimore Maryland USA

217 Soto P E A T Petcz and I W Buddenhagen 1976 Survey of insect pests and diseases of rice in different ecological zones in Nigeria Rice Entomol Ncwsl 435-46

218 Soto 11E and Z Ziddiqi 1976 Screening for resistance to African rice insect pests Varietal improsetnent sCmi n r Bouak Ivory Coast 13-17 Sept 1976 West Africa Rice l)evelopment Association Monrovia Iiberia

219 Squire F A 1972 Entomological problems in Bolivia PANS (PestArtic News Sumr) 18249-268

220 Stapely 1 II 1978 The rice leaf roller in the Solomon Islands Int Rice Res Newsl 3(l)13

221 Suenaga H 1937 On aphids attacking the roots of rice Oyo-I)oubts Zasshi 9150-152 [in Japanese] (Rev Appl Entomol A25745)

222 Suenaga H 1952 On the seasonal abundance of three subterranean aphids on upland rice Bull Kyushu Agric Exp Sti 1249-262

223 Sutynso Yas and Dandi Soekaria 1964 Beberapa hana yang ttenperang tana man padi tanah kering [in Indonesian] Rapat kerja Padi Tanah Kering 6 sd 8 Juily 1964 Kaliurang 9 p

224 Tallamy I) W and R F Denno 1983 Alternative life history patterns in risky environments in example from lace bugs Pages 129-147 in Insect life history patterns R F l)cnio and If Dingle eds Spring-Verlag 225 p

225 Tamura 1 1939 Observations on some insect pests of field crops [in Japanese] 1 Plant Prot 26414-420 (Rev Appl EntomolA27-556)

226 Tamura and Suzuki 1964 Influences of the early-transplanting practice in rice culture upon development of the rice stem borer Chilo siql)ressalisWalker Bull Ilokuriku Natil Agric Exp Stn 761-94

227 Tan J P 1924 The rice root aphid Drowhi ihirsoia Philipp Agric 13277-288

228 Tanaka 1 1957 Taxonomy and distribution of some subterranean aphids injurious to upland rice in Japan with description of a new species [in Japanese English summary] Botyu-Kagaku 22168-176

229 Tanaka T 1961 The rice root aphids their ecology and control Spec [full College of Agriculture Utsunomiya University Japan F3 p

230 Tauber M IC A Tauber R Nechols and 1Obrycki 1983 Seasonal activity of parasitoids control by external internal and genetic factors Pages 87-108 itt I)iapause and life cycle strategies in insects V K Brown and I Ilodek eds )r W Junk The Hague Netherlands

231 Taylor W F 1972 Effects of artificial defoliation (simulating pest damage) on varieties of upland rice Exp Agric 879-83

232 feotia T P S and S Natud 1966 Biotoomics of the rice skipper Parnaraniathiis Fabricius (Lepidoptera Hesperiidac) Indian J Entomol 28181-186

233 Teran F 0 1971 Rice pests of Santa Cruz [in Spanish Ministerio de Agricultura Sita Cruz Bolivia Bull 45 8 p

234 Than Itun 1976 Population dynamics of the yellow rice borer Tryporza itcertulas (Walker) and its damage to the rice plant MS thesis University of the Philippines at Ios Bafios Laguna Philippines 92 p

235 Uichanco L B 1911 Pests Pages 55-74 it Rice Philipp Agric 11-200

IRPS No 123 January 1987 41

236 van der Laan P A 195 1 Life history of the rice seedling fly Contrib Gen Agric Res Stn Boor Indonesia 118 15 p

237 van der Laan 11A 1959 Correlation between rainfall in the dry season and the occurrence of the white rice borer (Scirpolhaga innotala WIk) in lava Entomol Fxp Appl 212-20

238 Van Driesche R G 1983 Meaning of percent parasitismin studies of insect parasitism Fnviron Entomol 121 I1- 122

239 Venugopal M S and A litsingcr 1984 Fffect ofcarbofuran on rice growth Prot Ecol 7313-317

240 Vietnam Directorate of Rural Affairs 1962 Annual work progress report on crop improvement program of rice sugarcane vegetable and field crops (lor the period from July 1961 to June 1962) (11) 299 p (Rev Appl Entomol A51526-527)

241 Wan M T K 1972 Observations on rice leaf- and planthoppers in Sarawak (Malaysian Borneo) Malay Agric 148308-335

242 West Africa Rice I)esclopnient Association 1978 Annual research report Vol 2 Statistics Monrovia Liberia

243 West Africa Rice Development Association 1979 Annual research report p47 Monrovia Liberia

244 Williams ) F and CS lofgren 1981 Fli I illy l-468 anew bait toxicant for control of the red imported lire at Fla Entomol 64472477

245 Williams I) 1 1970 The mealybugs OFlomoptera Coccoidea Iscudococcidac) of sugar-cane rice and sorghum Bull Entomol Res 0109-188

246 Williams D) 1 1982 Pulinaria icervi (Signoret) and its allies ott sugar cane and other grasses Btll Entoinol Res 72111-117

247 Wirjosuiardjo S 1976 Study ofchenmical cont rot of insect pests ott upland rice Imiu Pertanian (Jogjakarta) 1323-329

248 Wirjosuhardjo S and N Wirarta 1976 Chemical control of the white grub tIolorithiatlltriin upland rice in M uneng Irobolinggo Indonesia) [in Indonesian] Imut Pertanian (Jogjakarta) 1330-332

249 Yano K T Miyake and V F Eastop 1983 The biology and economic importance of rice aphids (Henuiptera Aphid idae) a review Bull Entomol Res 73539-566

250 Yelp D and P Rhodes 19(4 lhe success of drilled dryland rice iii the Fiji Islands International Rice Commission Working Party on Rice Production and Protection Manila Philippines 3-10 March1964 Food and Agriculture Organization Rome Italy 13 p

251 Young W R 1981 Fifty-five years of research on the sorghum shootfly Insect Sci Appi 23-9

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No 104 Faridpur acomputer-assisted instruction nmodel forrainfed lowland random coefficient model No 116 Consumer demand for rice grain quality in Thailand Indonesia andrice

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No 106 Rice grassy stunt virus 2 a new strain of rice grassy stunt in tire literature

Philippines No 118 IRRI-Korca collaborative project for the development of cold-No 107 Physical losses and quality deterioration in rice plstprodnCtion tolerant lines through anthei culture

systemis No 119 Problen soils as potenial areas for adverse soils-tolerant rice No 108 (olptblication ofIRRI materials a survey of translators and varieties in South and Ssuthcasi Asia

publishers No 120 Changes isinall-farm iLtireliing techii)hgy in Thailand and [he No 109 Classification of Philippine rainfall patterns Philippines No 110 Contributions of modern rice varieties to rutrition in Asia No 121 Landforms and modern rice varieties No Ill Changes in rice breeding in 10 Asian countries 1965-84 No 122 Yield stability and modern rice technology No 112 Design parameters affecting the performance of the IRRI-designed

axial-flow pump

Page 9: UPLANDRICE. INSECTl--1i M T -E!R.EOLOU t RTANCE ,AND …delphacid.s3.amazonaws.com › 6958.pdf · The significance of abiotic factors in upland rice insect ecology will be apparent

9 IRPS No 123 January 1987

Table I continued

Order Family Species Distribution Reference

Spodoptera exigua (Hubner) Spodoptera exem pta (Walker) Spodoptera ornithogalli (Guenee)

Africa Africa Latin America

32 217 32 33 36217 150

Mythimna loreyi (Duponchel) Asia 30 Mythimna separata (Walker) Mythimna (- Pseudaletia)latifascia [= adultera

Asia Latin America

178 195

(Schaus)] (Walker)Mythimna (=Pseudaletia) sequax (Fabriclus) Latin America 68 Mythimna rosellinea (Walker) Asia b Mythimna yu (Guenee) Asia b Mocisfrugalis (Fabricius) Asia 30 117 Mocis latipes (Guenee) Latin America 66 68 149 150 203

Platysenta (=Spodoptera) compta (Walker) Asia 219233A Barrion unpubl data

Agrotis ipsilon (Hufnagel) Latin America Africa

33 68 81 150 1)5 203

Agrotis spp Latin America 60 Achaeajanata (Linnaeus) EldanasaccharinaWalker

Asia Africa

117 36

Armyworm unspecified Asia 72 84 Pyralidae Cnaphalocrocis medinalis (Guenee)

Marasmia bilinealisHampson Asia Africa

30 4559 72 84233 36

Marasmia trapezalis(Guenee) Africa 32 217 Marasmia (=Sumia) exigua (Butler) Asia 220 Marasnianmralis (Walker) Asia b Marasiniapatnalis Bradley Marasiniaspp

Asia Asia

b b

Leaf roller unspecified M2liarphaseparatellaRagont

Asia Africa

72 12 57

Maliarphasp Asia b Chilo auricilius Dudgeon Asia b Chilo partellus (Swinhoe) Chilo polychrysus (Meyrick)

Africa Asia

12 92 59 72

Qhilo diffusilineus J de Joannis Africa 6 1229 Chilosuppressalis(Walker)Cibozacconius Blezzinski

Asia Africa

59 72 6 11 12

Cravubus spp Latin America 203 Acigona loftini (Dyar) Latin America 203 Acigona chrysographella(Kollar)Scirpophagaincertulas(Walker)

Asia Asia

82 59 72

Scirpophagainnotata (Walker) Asia 59 200 Scirpophaganiuella (Fabricius) Asia b Elasinopalpuslignosellus (Zeller) Latin America 28 60 61 66 81 150

Diatraeasaccharalis(Fabricius) Latin America 162195203219233 28 66 81 111 150 195

Rupela albincllaCramer Latin America 111

Gelechlidae

Stem borer unspecified Undetermined stem borer nr Maliarpha BrachniaarctraeaMayr

Asia Asia Asia

72 b 55

Brachmi spp Africa 32 Hesperiidae Parnaraguttata Bremer et Grey Asia b

Parnaranaso Fabricius Asia 55 Pelopidasagnaagna (Moore) Pelopidasmathias (Fabricius)

Asia Asia Africa

164 36 155217

Pelopidasconjuncta conjuncta (Herrich-Schaffer) Asia b Borbo fanta Evans Africa 6

Satyridac Melanitisleda ismene Cramer Asia 84213 Mycalesis asophisHew Asia 30

Diptera Cecldomyiidae Mycalesis spp Orseoliaoryzae (Wood-Masn)

Asia Asia

b 72 88 131 181

Orseoliaoryzivora Harris et Gagne Africa 61132218 Chloropidae Steleocerellus (=Mepachymens) ensifer (Thomson)

Oscinella spp Asia Asia

205 205

Gaurax sppChloropsoryzae Matsuntura (= kuwonae Aldrich)

Asia Asia

205 82

Mucldae Atherigonaoryzae Malloch Asia 30 82 117130 Atherigona exigua Stein Asia 82 117 178

10 IRPS No 123 January 1987

Table I continued Order Family

Diopsidae

Agronyzidae

lHymenoptera Formicidae

Species

Atherigona oritentalisSchiner Atherigona indica MallochDiopsis longicornis(= thoracica)Macqurt Diopsis spp Pseudonapnyza asiatica Spencer Pseudonapomyza spicata (Malloch) Agronoza orvzae MunakataAcromyren landoltibalzani(Emery) Acromyrmex landoltifracticornis(Forel)Acromyrnex heyeri Forel)Atta bisphaericaFore Atta capiguaraGoncalves Atta laeLigata (F Smi h1) Atta opacicepsBorgmeierSolenopsisgeminata (abricius)Pheidole sp Pheidolegeton diversus Jerdon Monomorium pharaonis (Linnaeus) Ants unspecified

Distribution Reference

Asia 82 Asia Africa 82205Africa 11 1248 Africa 217 Asia 213 Asia h Asia bLatin America 6168 195 Latin America 61 195 Latin America 61 68 195 Latin America 61 68 195 Latin America 14 61 68 195Latin America 61 68 195 Latin America 61 195 Asia b Asia b Asia 1) Asia b Asia 120

aG orlentalls formerly recorded in Asia as G africana thPhilippines reported in this study CSubterranean root feeders

subterranean nests Foraging ants will feed on germinatedseeds only when the supply of ungerminated seed is low However these ant specialists feed on a wide varietyplants and thus help control grassy weeds (39) This dual

of

pest benefit role extends from tending ants which protecthoneydew-excreting pest species including root aphids and mealybugs (workers repel potential predators and even selectively kill parasitized aphids) to predators of leaf- and stem-feeding rice pests such as leaffolders armywormswhite grubs and stem borers (1 17)

The upland rice environment is particularly well suited to ants because of the lack of flooding Frequent tillage perpetuates grasses and granivorous species that are highlyadapted to constructing new nests in recently tilled fieldsAnt nests however are relatively shallow and tilling with a moldboard plow can destroy them Also upland ricefields are uually small and isolated and a given area in a smallfield has more field borders than it has in a larger fieldallowing high rates of infiltration from more sedentaryspecies such as Plieidole that do not colonize tilled fields but nest in soil under shrubs and trees along field bordersPheidole workers forage into fields from borders and store seed in their nests In slash-and-burn or other no-tillage rice cultures nests would be preserved in the field and possiblylead to high rates of seed predation However granivorous species do not prefer forest habitats

Damage is characterized by reduced usually patchy plantstand (120) Seed-thieving ants greatly prefer rice but they can be selective regarding plants they collect and store Theyharvest more seed than they can consume so patches of germip ing surplus seed (such as Rottboellia) will sproutfrom their nests during the rainy season

In Latin America leaf-cutting ants (Aita spp and Acromyrmex spp) occur only in upland fields and defoliate

young rice plants (47 68) They take leaves to fungal gardensin underground nests where fungi predigest the plant material

Termites Subterranean termites of the family Termitidae which lack symbiotic protozoa to help digest plants are the most frequently mentioned termite pests of upland rice (1185 120) This family cultures fungi in special undergroundcells --- fungal combs made of half-digested plant material (117) The fungi inoculated onto the combs break down plant material into food that termits can digest The termites then consume the fungal combs These grasslandtermites build nests below the plow layer in uplandricefields The colonies attack living plants only when dead plant material is not available They will attack a droughtshystressed crop but prefer older plants having greater cellulose content They will damage newly planted crops where clean culture has removed vegetation

Infestations are worse in deep light-textured soils with low moisture content The first sign of damage is yellowingof older leaves (195) Termites feed on roots (the plantyellows then wilts and finally collapses) and germinatingseed (loss of stand) or move above ground at night to cutseedlings at ground level which they cover with soil for later consumption Soft-bodied termites are highly sensitive to desiccation They live within a self-contained system of airtight chambers and build surface tunnels linei with nudand body secretions (carton) to maintain more than 90 relative humidity Termites can be located by their tunnels

Termites apparently can withstand limited submergence (157) and can be a pest of dry lowland seedbeds (117)perhaps by having nests below perched water tables

Termites appear to do more damage in Latin America where large ricefields are plowed with tractors deprivingtermites of preferred vegetation and forcing them to live on

rice plants In Asia upland ricefields are typically small with Iprge field border areas (ecotones) where termites can forage for preferred food other than growing rice plants All grassland termites do not feed on rice In an upland area in Batangas Province Philippines ricefields are commonly infested by Hosiititernesluzonensis (Oshima) however this termite does not damage rice In the same province though Mrroternesgivus(Hagen)feeds on rice seedlings (178)

In Africa termites may damage rice even though fields are small and surrounded by perennial vegetation These termite species perhaps prefer rice over other hosts

Mole crickets Nymphs and adults are nocturnal and burrow through soil feeding on roots of a wide variety of plants or forage above ground as predators of insects Adults live 3-5 mo and are even cannibalistic Mole crickets prefer low-lyin moist upland soils with high organic matter (II 81) In the lowlands they inhabit rice bunds or nonflooded fields Adults are highly mobile and can leave a flooded field to locate a more suitable habitat Losses show as wilted plants (from root feeding) or reduced stands (120) Damage is more common near field borders where mole crickets relocate after tillage (126)

Fiehl crickets Nymphs and adults of field crickets have similar nocturnal habits and damage rice as much as mole crickets do Piles of weeds removed from fields attract them They make subterranean nests and tunnel through the soil to feed on roots Some species prefer seed to roots Others feed at the base of stems causing deadhearts

Tunnel entrances surrounded by excavated soil are easily seen Young plants are cut at ground level and stored in underground cell Field crickets tre highly polyphagous as are all soil pests (117) Sachan et al (198) reported that field crickets prefer maize to upland rice Like mole crickets field crickets cannot survive in standing water and are therefore more prevalent in upland fields

Whitegrubsandhackbeetles Scarab beetles that feed on living roots as larvae but not as adults are called white grubs or chafers (Melolonthinae Rutellinae) and those that feed as adults but not as larvae on plant crowns are called black beetles (Dynastinac) In the tropics chafers or white grubs have a I-yr life cycle starting with adults emerging from the soil about I mo after the first downpour of the rainy season (134 159) They lay eggs at the same time farmers sow upland rice The rice passes its most susceptible stage and damage is mostly avoided when the white grubs are small After several months the long-lived larvae are large enough for two or three to denude the root system of mature rice (223) This intensity of damage is rare but wilting occurs when root loss is combined with drought stress

The larvae need damp (not saturated or dry) soil to survive and pass the unfavorable dry season 05-2 m underground in aestivation in the moist soil The first heavy rains of the season (20-30 amd) stimulate the grubs to resume activity and after several weeks they develop into

IRPS No 123 January 1987 11

pupae and adults and eventually dig their way to the soil surface and fly to nearby trees to seek food and mates The adults of most white grub species feed only on trees but in some species notably Eutheolahumilis the adults defoliate and cut off stems at ground level (28) Putheola hidentata damages sugarcane fields in Venezuela (121) Floods kill white grubs (208) Grasslands can support large populations and therefore white grubs can be more abundant in newly planted upland ricefields that were previously fallow (81)

Black beetles live up to a year adults can attack a crop at any stage by burrowing into the soil (127) and feeding at the base of stems to cause whiteheads (207) Adults emerge with the early rains and are normally more abundant and do more damage on a young rice crop with small root systems (38 81)

The larvae feed only on dead organic matter in dryland fields and do not attack rice Adults arc highly mobile and although sensitive to flooding can invade a field soon after it drains (115)

Root aplhids Several genera of migratory aphids feed on the roots of upland rice (249) Populations build up more in light-textured soils with high percolation rates Damage has also been reported from dry seed heds of lowland rice (52) Tending ants -- Pheidhle Cren aster Tetramoriuw lsius Tapinoma -- are necessary for root aphids to multiply Aiits harbor aphids in their nests over winter or during unfavorable periods and relocate them on rice plants digging tunnels along the root systems to allow these softshybodied insects to penetrate the soil Most intensive studies on aphid life histories have been done in temperate environments (158 222 229) Root aphids fly to rice plants at the beginning of the rice season and pass through several generations continually developing winged forms to relocate on alternate grassy hosts until winter when they move to perennial hosts usually trees Yield loss occurs mainly through reduced tillering but infested plants become yellow and stunted and in extreme cases wilt (227) Aphid populations build up gradually so damage usually begins in the late vegetative and reproductive growth stages

Root-feeding ineal tugs More than half of the mealybug species or rice feed on roots (245) Being soft bodied mealybugs are not adapted to living underground but they survive because tending ants (the same species that tend root aphids) dig burrows for them and move them from plant to plant Well-drained soils ihelp mcalybugs survive Although six species are recorded as subterranean root pests of rice we found no report citing damage Root-feeding mealybugs are probably more prevalent on perennial grasses than on short-lived annual rice

The most common rice mealyb ig Brevennia rehi is a foliar feeder favored by lack ol rain apparently it is responsible for high yield loss in Northeast India and Bangladesh (N Panda pers comm as stated by IRRI [1061) This report needs confirmation Mealybug damage was only briefly mentioned in another report (156)

12 IRPS No 123 January 1987

Root weevils A few species of upland weevils (IHporneces Donacia Atactogaster) attack rice roots as larvae and feed on stems and leaves as adults Damage ispatchy The larvae are not adapted to submergence and are true upland soil weevils isopposed to the so-called water weevils They rarely become sufficiently abundant to cause damage Adults on rice foliage or the soil surface show that the roots are being attacked by the grubs Root weevils are highly polyphagous (185)

Root heeths Soil-inhabiting false wireworms Gono-cephalumt spp are reported in upland ricefields The adults and larvae normally feed on decaying organic matter but land preparation for upland rice removes these sources The larvae feed on seedling roots and nocturnal adults roam on the soil surface to cut off seedlings at ground level (82) This group of ground beetles is highly omnivorous adults even prey on stem borer larvae (183) True wireworms (Elate-ridae) feed on living roots but rice as a host has been reported Infrequently (117) In Latin America the highlypclyphagous larvae of Diabrotica rootworms are also recorded on rice

Root htus Upland rice seedlings can be attacked by the adults anI nlViphs of chinch bugs Blissusand Caenoblissus and bro n bug Scaptocoris which feed on the roots with theirsucking nouthparts (80 83) Three genera of burrower bugs A tms (eotonhus and Stiharopus -are common iil grasslands in the Philippines and may attack the first crop (rice or maize) planted after the dry season Chinch bugs are most abundant after a dry spell These insects are large and can kill seedlings or stunt and reduce tillering on older plants I hey prefer other hosts and rarely damage rice Adults can also feed oii developing rice grains (81)

(utworms Soil-dwelling noctuid larvae hide under-ground during the day to avoid predation by birds and bccone active at night above grour cutting off you ing plants at ground level (hence tile naiie cvtworn) arvaedrag severed plants into hurrows llie larvae have wide host ranges arid pupatc in the soillgrotis ipsilon occurs worldwide and appears to be the most common cutworm pest of upland environments but is more prevalent inlowland rice particularly in dry seedbeds

IFoliar pests Other groups of insects are adapted to upland environments because they pass one growth stage underground either as egg (in heinrietabolous species) or pupa (in holomnetabolous groups)

Rice seedlibig maggots AitJeriota flies occur in non-fhoodcd environments in Asia and Africa (180) Feeding of larvae one per iller causes deadhearts Flies as opposed to moths only secondarily evolved as pliytophagous pestsfroni a saprophagous Origin Athrnonalarvae secondarihv feed onidecaying tillers whereas larvae of stem borer moths actively Iced on living tillers to cause deadlicarts Seedling maggots will die if they cannot sever a tiller on which to feed

as it decays Plants die under heavy attack whereas lighter infestations cause stunting delayed maturity and ragged leaves (236)

Eggs are laid only on actively growing plants (49) aid damage occurs wiihin the first week after crop emergence(170) Larvae pupate more in stems than in soil Seedling maggot attack is highly seasonal (206) normally peaking 2-3 mo after the onset of the rainy season These pests possiblyaestivatc over the dry season Rice isonly one of their many graminaceous crop and weed hosts

The rice stem maggot (hlorops orzae is adapted to temperate upland and lowland rice growing regions of Asia In Japan and China upland rice grows in mountainous regions a habitat of this chloropid fly The larvae tunnel into tilestems The stem maggot is more damaging during the seedling stage larvac hibernate in grasses

File several dipterans reported as stem maggots in dryshyseeded lowland seedlings inIndia (205) could presumably attack upland rio

Leaf urinrs Icaf miners are common on wheat and maize but are rarely reported on upland rice (213) Three leaf miners are more prevalent on upland than lowland rice inthe Philippines Pvudonalomy a asicanh Vficataand lIAgromrza orzae attack at the seedling stage The larvae develop inside the tunnels they construct in the parenchyma tissue P Vpicata and Aorvzac prefer maizeEehtsine and other grasses to rice spicatacan become abundant on wheat

Leaf beetles Chrysomelid and coccinellid eaf bcetles scrape or otherwise remove leaf tissue as larvae or adults or both This damage accelerates desiccation of plants Tile larvae of sonic species are root feeders Flea beetles Epitrix and (lraetocnetna and rootworms l)iahroticaspp feed on rice foliage only as adults Their root-feeding larvae have hosts other than rice The adults and larvae of the hispa-like Oediolmlpa spp beetles of Brazil and (hmootriha (=EPfishyachra) beetle of Africa feed on upland rice leaves Oealioshypapa spp larvae are leaf miners (mootriha is also pshyvalent in lowland environments (11) Leaf beetles prefer vegetative rice (81)

The rice hispa Dicladipaarnigera is not reported on upland rice and prcers more aquatic habitats In Central Africa however I)cladispm viridicyaiea attacks uplandrice in the vegetative stage as well as lowland seedbeds 7richipasericea is reported on upland rice in West Africa and Madagascar (57)

The rice leaf beetle 1d1ea 01izae riiuech like Chioropsor|rzae is adapted to temperate regions of Ashi and is usually a pest on lowland rice but seldom oi inpland rice in the mountains (82) The adult hibernates on grasses

Armmworms Species of grassland-adapted SpoptloeraPlatysemta and A ithimna larvae create widespread epishydemics They can defoliate ricefields generally in patchesfrom early vegetative growth to harvest (41) Armywornms pupate inthe soil and highly favor upland rice (34) Adults

can disperse long distances to colonize even remote upland areas They become more abundant than their natural enemies particularly after favorable rains following a prolonged drought (196) The drought kills the armyworm and natural enemies (mainly parasites) but the armyworm can recolonize rapidly

The natural increase insoil fertility from mineralization of soil N over the drought period promotes luxuriant plant growth to foster a rapid buildup of these highly fecund species Weeding rice increases tilelikelihood of damage The parasites normally return to control the armyworms but not until after serious damage

Outbreaks on upland rice have been reported in Ghana (3) inzibar (38) Central Africa (36) lPanama (165) Brazil (28) Malaysia (196) and India (167) Defoliation may be severe often leaving only the base of stems Armyworm larvae hide during the day under litter or in soil cracks At night they ascend plants to feed Larvae pupate in the soil

Thrips Thrips frequent upland rice but damage is less than in the lowlands Thrips larvae and adults feed on leaf blades by rasping causing leaf rolling and stippling Thrips prefer vegetative stage rice and their survival increases during drought (heavy rain washes them from foliage) although drought stress usually outweighs thrips damage Drought and thrips cause leaf rolling in upland rice (235) but thrips have been overlooked perhaps becaLse of their small size Thrips are most numerous indry season irrigated rice The combination of favorable plant growth and lack of rainfall is ideal for thrips In contrast the lack of rainfall in upland rice hurts the crop and eventually the thrips whose numbers must decline

In [atin America Frankinilla rodeos attacks panicles before they emerge from the boot calusing sterile grains

Stem hug~s Black bugs are the most common group of rice stem-feeding -lemiptera in Asia The most frequently mentioned species ----Scotinophara coarctaia and Shrida - prefer aquatic habitats and therefore could not occur in upland rice as sometimes reported Upland species in the same gerurs - notably S tarsalis and S scotti -- are morphologically similar and remove sap from stems of grasses and upland rice The upland species are notably more abundant in upland rice planted near forests

In Latin America tile Tibraca ima-large pentatomid tiventris causes deadhearts and whitehcads on upland rice Both adults and nymphs remove sap from internodes of plants more than 3 wk old Tillers are killed by mechanical damage and by the entry of secondary microbial infection This damage is often confused with that of lepidopterous stem borers (233) Eggs are laid on the leaves and stems Nymphs are gregarious This species can also be found on irrigated rice Drought reduces population buildup

Stiem borers Lepidopterous and dipterous stem borers are widely recorded on upland rice but cultural practices and alternate hosts determine abtndance Except for Elasmolmll)uS Iinosellts numbers usually build up toward

IRPS No 123 January 1987 13

the end of the crop growth cycle Therefore late-maturing varieties staggered planting and lack offloodingfavorstem borer buildup Many stem borer species can lie dormant in the stUbbleafter harvest and theirsurvival is encouraged by panicle harvesting (leaving tall stems) and lack of tillage after harvest

Stem borers disperse and lay many eggs Many species are polyphagous however the monophagous MaliarpiasepashyratellaDiopsis Ivmgicornis Scirpophaga incertlas and S innotata can also be abundant on upland rice (hilo Suplressalisdevelops fasterand becomes la rger when reared on upland than on lowland rice (226)

The lesser corn stalk borer lasmopal tt Ittoselhs is semisubterranean and is perhaps the stern borer most adapted to upland rice(129) Highly polyphagous it infests maize peanut and cowpea It attacks seedling rice (one larva can kill tip to four plants before maximum tillering) (195) (hiv and Maliarphastem borers prefer older plants But Scirpophagaspp will attack young plants even in the nursery lasn opalls larvae tunnel into the stems at or below the soil level mausing dcadhcarts Larvae are not found in their turnnels inside sterns beca use vhei disturbed they retreat into cases made from soil particles bound by silk The cases areattached to the tunnel entrance (143) This behavior probably evolved as protection from natural enemies Larvae pupate in the soil The lesser corn stalk borer is more abundant during drought damage is often confused with that of drought (233) This species prefers sandy soils arid its distribution within a field may relate to soil texture (143)

Diatraeasaccharalis aside from the lesser corn stalk borer is frequently a stem borer pest in Latin America According to Teran (233) rice and maize are more preferred hosts than sugarcaine its namesake

Atcitonacrvsograpla is a less known upland rice stem borer in Asia with habits similar to Easinopall s (82) because it is not found in lowlands Acutra has grassy weed alternate hosts As a moth it is often confused with the dark-headed stern borer(hi polur(standstriped stern borer (hi sup)lpsrcais

Other common upland rice stem borers are also reported in the lowlands The dominant species in upland rice vary greatly with location In Uttar lPradesh India (78) and in Japan (123) the pink stem borer Sesamia htfirens is more prevalent hIKenya the principal rice stem borer is Chi partelus

(92) The lowland African sten borers Aialiarpha sepashyratellaDiopsis vngicornis and Sesamia calatnisnis are less abundant Oimupland than on lowland rice(217) but perhaps cause greater yield loss because plant injury is greater if combined with drought css (92) The stalk-eyed stern borer D)ivpsis is more prevalent in wet habitats (6 1I217)

In Central and West Africa Maliarpha and Sesamia calanistis are the dominant stern borers of upland rice (8 9 10 II) (lilo dihlfilineui and Chio partellus inhabit the

14 IRPS No 123 January 1987

upland savannas whereas Chilo zacconio prefers the lowlands (II) Aaliarpha is most abundant in the mangrove swamp habitats (91) coastal regions (16) or rain forest zones (II) of Africa and appears to be the homologue ofScirlpophaga incertulas The yellow stem borr can also be prevalent in upland rice (120 167) The sugarcane borer Eldanasaccharinais so far only a potential pest of uplandrice (91) despite Grist and evers (82) warning

Grasshotl)ers andI locitas Short-horned grasshoppers and locusts -- Locvta Patanrga Schistocerca - layeggs in the soil inhabit grasslands and develop into swarms to seriously damage upland rice from time to time (173) Largenumbers can destroy a field leaving only the stubble In low cmuoroer s giassnoppeirs an cut panicles (173) Farmers may not plant if locust swarms are imminent Prolonged dryweather followed by favorable rain favors development of large swarms (173) Usually breeding in di) grasslands theydisperse toattack crops including upland rice(191) Eggsaresusceptible to desiccation d uring drough t and toinundation during seasonal rains With favorable soil moisture over consecutive seasons natural enemies ca nnot prevent a largepopulation increase leading to migratory swarms (64)

Most grasshoppers on rice are adapted to uplands (107)but lowland rice grasshoppers iervgl(rlhusspp and Ora spp occur occasionally on upland rice (2 82) They lay eggson rice foliage Nymphs are scmiaquatic

The gryllid Lsctrtus coucinnus is a pest of lowland and deep water rice in Bangladesh and Thailand In the Philippines it damages upland rice grown in isolated pockets near lakes The gryllid nymphs and ad ults feed on the central portions of folrage leasing only the midrib and leaf nargins

Leal ler Pvralid moths whose larvae fold leaves to make a feeding shelter occur worldwide (217) but arc appareritly most common in Asia Leaffolde rsattacking rice in Asia are a complex of species (26) (Ctalhahcrcisinedinalis the best-known rice Ia flolde r in Asia may prefer grassy weeds over rice (86) It is a late colonizer of uplandrice U1arasmja eviua a lowland species (220) nreflers rice to grass specres (86) and colonizes upland rice during il early growth stages llarasnria patnalis is the most corn-rnonly encountered species in upland rice in the Philippines Perennial grasses are their alternate hosts and they can survive year-round in upland rice environments The leaf-folders 11arasmia ruralisand Alarasmia spp are least aburndant Dormancy is unknown in leaffolders Cnaplha-ocrocis medhnalis however is known as a migrant (90)Leaffolders remove photosynthetic tissue attack during the flag leaf stage is particularly injurious Popurlations are normally held in check by natural enemies but leaffolders become parlicularly abundant in conditions of high plantfertility and shade

Butterflies Skippers PelopidasInathiasand Borhofanta are more prevalent in African lowland than in uplandhabitats (6) but extensive studies in Japan and China

showed that Pelopidas mathias prefers an upland habitat (155) Parnaragutata seasonally migrates between lowland and upland areas and is equally abundant in both environshyments (164)

The green horned caterpillar Melanitis leda ismene is more abundant in lowlands (212) but our experience in the Philippines suggests that Pelopidas niathiasand 11elanifis leda ismene are often more abundant in uplands than in lowlands Bamboo iaunbuiva spp annually sustains P mathiaspopulations in the Philippine uplands Adults feed on nectar from flowers and migrate to upland rice during the late vegetative stage

iPi erlled h i-1 1d dill L tIC i 7-0 l(become numerous probably because of high egg predationEgg and larval parasitization are often low Butterflies can readily disperse long distances to seek renlote uplandricefields However they have low biotic potential and even it predation rates are low the rarely become abundant Upland environments with their diverse microhabitats can provide more favorable sites than lowland rice plains

tutterfly pests of rice have broad host ranges and become dormant Juring urnfavorable times of the year

IolhagrotslaidopwraThe larvae of many polyshyphagous moths are reported to defoliate upland rice tussock moths lsaispelnnatula Iawlia sUifita luproctisirgtwltaE mnorand P vathorrhoeaslug caterpillar

litotiahicolorhairy caterpillars Nisaga sinp vandA mata spp and wooly bears l)iacrisiaohliqtia and Crvatonotts gangis Many of these polyphgous larvac migrate and are transients in rice including HelhWhis arm iera and 1t zea highly pol phagous species (82)

Gall nidges The Asian rice gall midge Orseoliaorrzae (88 131) and its African counterpart 0 ortzivora(6 11 12 32217) prelerlowland to upland rice Gall midges in rainfed areas are prevalent only during tle rainy season and dispersepoorly Their alternate hosts (wild rices LeersiaIschaetnuan and Pasahmt)are mostlyaquatic grasses and are less likelyto be near upland rice areas In an area of mixed uplandfloating and rainfed lowland rice cultures in northern Thailand upland rice which is ilanted a month earlier was attacked firstbut subsequcent populations were much lower than in rainfed lowla id or floating rice Low numbers of gallmidges passed the dry season orr perennial weeds such as Paspahim (88)

Smot (hid(s Reports of aphids attacking upland rice plants are few (249) Without maize the maize leaf aphidRho alosihhunm maidis attacks upland rice in the Philipshypines The plum peach aphid IIrsteroneuraselariae was reported in Sierra Leerne toinfest leavesand tnriperied grain(7) Siphaglrctriaecaused transverse linrea r necrotic striae iir rice leaves in Italy (172) I-lea v rains held populations in check Rice is not a preferred host for shoot aphids

Leafhoper andI )anthol)pers Cicadellids and delshyphacidsdispersecasilyand readilycolonie upland ricefrom nearby lowland areas A study in Sarawak (241) showed

IRPS No 123 January 1987 15

similar species in upland and lowland rice Nephoiettix The principal vectors of rice virus diseases generally have species dominated The most prevalent species accounting narrow host ranges and alternate hosts in the grass family for 90-95 of the sweep net samples wre N virescens N are mostly annuals dying in the dr3 season Some virus umropictus Nilaparvia hgens Soatellafiircifera Cfana vectors such as BPH can disperse lontg distances but few do

sectraand Recilia dorsalisLowlands had higher popula- Th restriction of upland rice and weed hosts to the rainy tions Katanyukul and Chandaratat (120) monitored hopper season the small area planted to upland rice and the hopper numbers in upland and lowland fields of northern Thaiiand vectors having narrow host ranges all work against virus and found similar results A comparative study of plant- diseases in upland rice hoppers in Fiji showed more brown planthoppe BPH) and In Latin America Sogctdhs o(rzicola and S cuhanus whitebacked planthopper (WBPH) in wetland rice than in transmit hoja blanca virus disease but reports on upland lowland rice and hopperburn was prevalent in lowland rice are rare except in the favored upland areas of Colombia areas with standing water (89) Peru and Venezuela (35 42) S rricolaocurs in Brazil

Hopperburn in upland ricefields in Asia is rare (84) In on upland rice but hojaI blanca has not been reported (67) Batangas Philippines isolated fields of upland rice become S orzicola has a wide host ramgc (10) similar to WBPI- in hopperburned infrequently from WBPH A report from Fiji Asia WBPH however is not known to transmit any virus cites hopperburn from BPH and WBPH (250) In these two disease situations upland rice was grown on highly fertic soil and In Africa pale yellow mosaic virus is endemic to swampy N fertilizer was applied Hoppers respond to better nutrition lowland areas where rice ratoons year-round (19) and is Normally upland rice is grown on poor soil and receives no mechanically vectored mainly by hispid beetles A minor fertilizer vector a flea beetle Chaeiocnema spp is an upland rice

In Asia upland rice ras are normally not far from pest and upland ricetields near endemic areas occasionally lowland ricefields-- a source of hoppers Otherwise hopper contract the virus species with greater polyphagy than BPH or Nelhotettiv Spittle bugs Cercopid nymphs producc a protective virescens would be dominant frothy covering that looks like saliva In Latin America

In Africa leathoppers and planthoppers do not become spittle bugs damage upland rice In parts of Brazil where abundant on upland rice (6 II) probably because lowland upland ricefields are surrounded by pastures adults rice is not widely rlanted nearby The only reported migrating from grasses can damage rice severely and cause hopperburn as from Nilapanrata maeander in breeder complete hopperburn in a young rice crop within a week of plots receiving high N rates in Nigeria (98) Hoppers colonization (194 195) Dispersal to rice occurs even if however are a greter problem in Latin America where pastures are verdant and spittle bugs prefer pasture grasses large grassland areas breed polyphagous species (Gra- to rice (168) Nymphs are rarely abundant enough to phocephala spp Hortensia spp lEvitianus ohscurunervis damage rice Regular rainfall favors their development Balclutha spp and Draeculacephala spp) vhich then (233) disperse to cause hopperburn on seedling rice (68) Rosetto et al (195) reported Graphocep7hala spp and Hortensia spp Seed pests to be equally abundant on upland and lowland rice Alydid and pentatomid bugs are seed pests of upland rice

The leafliopper and planthopper epidemics in Asia that worldwide In Asia Leptocorisa sp prevail Their habits caused severe losses from hopperburn and vijus dicases make them particularly suitable for upland rice 1)thev feed over the past several decades occurred principally on on many grasses (and therefore can survive the earl y wet lowland rice Except in a few isolated cases (160) upland season before rice sets its grain) 2)adults are long-ied and rice has been spared from tungro grassy stunt and ragged mobile allowing them to find isolated plantings 0 rice stunt virus diseases vectorcd by green lcafhopper (GLH) (besides seed bugs only birds and rats appear to have shyand BPI] (123) Virus diseases have occurred in upland rice capability) and 3) adults aestivate in forested areas or where lowland rice was nearby Rice dwarf disease however sugarcane fields during the dry sca son when neither rice nor was first recorded on upland rice in the Philippines (187) grassy weeds are present (I199) Rice seed bugs concentrate Virus diseases can be perpetuated only on livir plants or on small-scale upland riceficlds because they can actively insect vectors and no living rice - plans tooned or search them out The prevalence of groves of trees characshyvolunteer - grows year-round in upland arcas excert in a teristic of upland environments ensures nearby aostivation few with 12 mo of rainfall sites keeping the seed bugs close by

The smaller brown planthopper Laodcelphax striatellu Leptocorisae are uncommon in lowland rice plains occurs in upland rice in Japan (18) It spreads black- because 1) their populations are diluted in a sea of rice streaked dwarf and stripe viruses between winter wheat and planted more or less at the same time (they ca n only feed on barley to lowland rice Early planting of lowland rice rice during the milk to hard dough stages) 2) they lack nurseries spreads these viruses but upland ice has remained aestivation sites and 3) weedy fallow areas are limited uninfected The lowands of Asia are dominated by L oratorius

16 IRPS No 123 January 1987

which lays its eggs hign on tile foliage (199) L acuta and L solonlonlesis oviposit at ground level the former on litter and the latter loose on the ground These egg laying habis explain theirtenvironmental prefcrencecs(214) In Japan the postdiapause adults of L chinensis migrate from moun-tainous areas to upland ricefields during floweiing (1 14)

Pentatornid bugs prevail in Latin America (8182 195) and in Africa pentatomid and coreid bugs arc equally cited (6 82) l)amage is often characterstic of the species (5)rcoris ventralis and Mnida spp are the common

pentatomid seed bugs of Philippine upland rice but they prefer lEchinochva spp grass seed to rice Vearaviritda occurs worldwide and hoften more abundant in the uplands(54) We saw few reports of seed bugs on upland rice but see(d bugs apparently opelrate cqually between by land and upland rice Good dispersal is necessary ior species that call feed for only 2-3 xon a crop and must first find those crops at ripening stages

Other pct species feed n flowers In Mato Grosso Brazil (194) and Santa Cruz Bolivia the panicle weevil Aeo-hariuia amllitarsis attacks rice during flowering (68 233)The larva e rice sten horers adutlts feed at the bases of spikelets caIusing empty grains Also inLatin America thecydnid -lkiituttatralusisa seed bug as anad tilt and nymph (194)

A number of beetles ecd on poilen - coccinellids lialrotia Aulac lhora and Monohpl-a (55) - but becatise riCe is self-pollinating and fcrtiliation occurs before pikelets open they pose no threat to yield

IST ECOLOGY

We now look at characteiistics of the life cycles of upland rice insect pests to lea rn hew each plays an adaptive role in upland rice ecology

Life history strategies Orzasativadoes not tolerate drodgh well so upland rice is highly seasonal normally grown in the wettest months of the year Usually rice is present for less than half of the year The rice-fiee fallow poses serious problems of survival to upland rice insects which have evolved att least four mechanisms to ovrcolne tle cvcl ic-l lack of a hostI) polyphagy 2) longevity 3) dormancy and 4) vagility (dispersal)

Poldt)agy lolyphagy is defined as having hosts of at least two botanical families oligophagy of more than one gents and ionophagy of only one ventus (40) - inour case Orvza Most wild rices however are illi tiC and theieiirc are normally far removed from upland rice hahitat othewise more would be alternate hosts for uplanJ iice insects Thte highly mornophagous rice pests shySciriol)haita hiceritlas Nilal)arvaiahiens Iand Nealo--tettiyvircsctws in Asia and Aalirpha secparatella and 1ioj)sis lngicornis inAfrica - occur on upland rice but

are more abundant in lowland culture They can specialize in rice because they are highly vagile and can attack all growth stages By specializing in rice monophagous rice pests can outcompete related but polyphagous speciesIntensive andI extensive lowland rice culture has favored monophagous species Riqwla alhinella (Cramer) the onlymonophagous species in Latin America is rarely reported on upland ice (I1)

Because of its limited temporal and spatial existence upland rice has not favored the evolution of specialized species Clhang(44)shows evidence that upland rice has only recently been cultured by man from the lowlands thus pests have had little time to fully adapt to the crop All but the lowland-adapted insect pest species have alternate planthosts to rice Root pests with limited mobility are highly polyphagous Pests that specialize in one growth stage eg secdling maggots seed bugs are oligophagous or polyphagous

Most alternat hosts of rice pests are annual grasseswhich also are seasonal Upla_rice is also a host but not the preferred one of the most highly polypliagous spcciessoil pests grass hoppers armvwarrs and Vearaviirielda

The d ry season poses igreat obstacle to ani upland riceinsects survival and only sonic perennial plant hosts are suitable food during the off-seasoin Plolyphagy allows upland rice insects to suLrx-ixe during thC rainy season when plants- both annual and perennial -areactivelygrowing and therefore more nutritious

Loiugevitr The ability of aiinsect to live 2-3 mo without undergoing dormancy helps species sich as root-feedingwhite grbs (as larvae) and secd-feeding bugs (as adtilt) to survive unfavorable periods White grubs arc not highly fecund and their life strategy is to improve survivorship of limited progeny Their subterranean habitat hides them from manv natural enemies lBtut root tissue is not highly nutritious so they ltilst eat great quantities and need a largeinsect biomass to digest it Other soil-inhabiting rootshyfeeding insects have short life cycles but are more fecund

Seed pests need greater longcvi vas adults to locate ahost at early grain dcclopment arnd give sedentary offspring time to develop Few groups of upland rice insects live longin active development Polyphagy and longevity only increase their ability to survive during the rainy seasonOther mechanisms are needed to survive the dry season

orantc In the tropics the nonrice season is the dry season while intemperate areas tile off-+season is winter Many upland rice insects undergo dormancy inlsummer (aestivation) or winter (hibernation) Dormancy is simplyinactivity during winters with prolonged temperatures below 10 C insects find shelter arnd cease movement Activity is resu med with warmer t empcera tu res A deep state of irnactivity termed diapauise occurs in sole species that prepare physiologically (havxing low metabolism and convcrting gonad s to fatty fuel reserves) for the unfavorable season (148) They enter diapaUse inanticipation of unshy

IRPS No 123 January 1987 17

favorable weather and resume activity after favorable Armyworms skippers and locusts actively travel hundreds weather begins usually in response to photoperiod or of kilometers in air fronts such as intertropical convergencethrough a delayed response to a stimulus such as heavy zones to descend onto upland rice fields far from their rainfall (237) Not all individuals of a population have the breeding grounds (31) Several generations can build ip on same threshold for environmental responses for entering or alternate hosts in tile grass family before rice is plantedterminating diapause -- this is seen as a mechanism to These large insects are active fliers Smaller insects such as prevent total mortality if a false cue appears such as a thrips aphids and planthoppers passively migrate long sudden warm spell in winter or rain in the middle of the dry distances by the wind season (154) Otherspecies ean readily travel tens of kilometers - stem

Probably more upland rice insects undL go dormancy borers leaffolders leafhoppes cutworms spittle bugs leaf than we realize Candidates are Atherigona spp root beetles seed bugs ants and termites aphids root weevils crickets leaf beetles thrips and seed The least dispersive species are white grabs mealybugSbugs Insects known to entervarious states of aestivation are seedling maggots crickets weevils and gall midgesleptocorisaspp adults (199) Maliarphascparatellalarvae Each species has evolved a unique set of attributes to (9) Scirpophaga innotata larvae (117) Chilo partelhtsv enable it to survive and adapt to the changing uplandlarvae (21) ScirpopThagainccetulas (179) white grub larvae environments Several evolutionary avenues lead to fitness (134) Patangasuccincia adults (6 82) and Schistocerca (Table 2) but there is no reason to believe one set of gregariaeggs (82) attributes is better than another (224)

Insects known to enter hibernation are LodeIphaDstriatellus nymphs (124) Chtio Tvlpressalis larivae (125) DroughtSesaia inferens larvae (146) but not S calavisis(2) Well-drained rainfed upland soils are subject to drought ifIDiaraea saccharalis larvae (75) Scirpophaga- tcerlttas rains do not fall within 2-3 wk Prolonged drought followedlarae(5 lacchrli s lae (75)saSlOhagasshoperseggs (82)and felo pidas athias pOpae (232) by favorable rains stimulates armyworm and locust outshybreaks but mealybtigs root aphids and thrips become

Lcpmotorisa oratorisand L acuta adults aggregate on numerous onlyafter dry spells for two reasons First heavy trees sugarcane or other shady moist sites L paluwanmsis rain normally kills soft-bodied foliage-feeding thrips and confined to Sulu and Palawan Philippines lives on the mealybugs and soil-inhabiting root aphids and root mealyshygrass Brachiariamutica (Forsk) Stapf in the absence of bugs Drought eliminates this source of mortality Second rice The adults can live for 5 mo When aestivating their the rice plant responds to drought by breaking down rate of metabolism drops and they do not feed They will proteins into soluble N compounds which enter the phloem however puncture plant tissues of their aestivation host in and are taken up by these sap-feeding insects (245) Greater search of water This behavior has led to errors of the host nutrition therefore leads to greater survivorship and range of rice bugs (I) Aestivation quickly terminates with fecundity rain probably meaning aestivation is not deep Small rice area

Some stem borers aestivate as mature larvae in rice Except in the highly mechanized large landholdings of stubble or in straw stacked as livestock feed However Latin America upland ricefieldstypically are patchy within moriality increases proportionally with the duration of the a highly diverse flora The small fields result in higher ratios dry season even if the stubble remains undisturbed If the of perimeter to area than with the typically larger lowland stubble is plowed for a crop following rice few stem borers ricefields Upland ricefields themselves may be intercroppedsurvive Rainfall terminates dormancy but it normally with cereals legumes or root crops Mixed intercroppingtakes several weeks for moths to develop and emerge with a wide variety of species also occ in tribal slash-andshy

Last-instar white grub larvae tunnel 1-2 m deep in the soil burn agriculture (50 74) to construct pupal chambers in which they aestivate The Upland ricefields tend to be small because of the labor larvae are very sensitive to dryness so pupal chambers are needed to clear land in slash-and-burn areas and to till land sealed to coierve body moisture Aestivation terminates to minimize weeds Rice requires more tillage than does with the first heavy rains but as in stem borer 3-4 wk pass maize because rice competes less well with weeds Uplandbefore adults develop rice is more a subsistence than a cash crop and small areas

Insects may respond differently in each region to cold (I ha) can feed a family temperature or drought Biotypes or loal populations may Small fields favor some insect pests such as some ant evolve (56) Local populations of Scirpophagaincertulasor species that can forage from more permanent field borders Patangasuccincta may have become adapted to prevailing Trees provide food and mating sites for white grubs (134) as conditions well as aestivation sites for LetTocorisa spp (199) Field

iagiliti The ability to disperse combined with high border grasses plus tIhose within a ricefield provide alternate fecundity orshort life cycles enable some upland rice pests to food for all but the highly monophagous upland rice better exploit the temporarily favorable upland habitat colonizers and a habitat for natural enemies

0 z

Table 2 Life history patterns of upland rice insects or insect groups

Host range Longevity Dormancy Vagility FertilityMonophagy Oligophagy Polyphagy Low Medium High Yes No Low Dispersive Migratory Low Medium High(Oryza spp) (grasses sedges) (angiosperms) (lt3 wk) (gt2 mo) (lt1 kin) (lt50 eggs) (gt500 eggs)

Ants xTermites x x x

Mole crickets x x x xx x xField crickets x xx

White grubs x x x xx xRoot aphids x x xxMealybugs x

x x x xxSeedling maggots x

x x x x xLeaf beetles

x xx xArmyworms x x xx x

Thrips x x xx xChilo spp x x xx xMaliarpha x x x x

xLocusts x x Xx x xGall midges x x xxiVephotettL iirescens x x xx x x xLeprocorisaspp x

x xLeaftolders x x x

Sogatellafurcifera x

xx x x x x x x

00

IRPS No 123 January 1Q87 19

All this appeals to favor insect pest buildup but the notwithstanding the release of stored nutrients in response contrary is true Loevinsohn (138) showed that insect pest to drought that temporarily benefits philoem feeders populations respond exponentially with the proportion of land devoted to rice up to about 75 of the area then the M POR ANCE I I P1A 1) RWI P1St S

rate of response declines In most upland rico areas tie rice crop occupies less than 50i of the area- therefore the Visits to upland ricefields normally reveal 1kw insect pests potential for population buildup is low This relationship Loevinsohn (138) and Locvinsohn ct al (139) compared may not hold for seed pests that can locate small ricefields insect abundance using annual light trap atchcs in upland

Another outcome of the highly diverse flora and phical rainfed lowland and irrigated lowland sites in the Philipshyenvironment of upland rice isthateach upland ricearca has pines Although only one upland site was studied data a unique composition of insect pests showed equal or fewer insects at the upland site than at the

For example in Tanauan Batangas Philippines the rainfcd lowland sites and fewer than at the irrigated sites main upland rice pests are rice leaffolder(RLF) armvworm (Table 3) M Lltiple regression analysis showed that the key Aythntnaseparata WBPH ants Soli psis gentitataand factors explaining insect abundance among a set of cropping white grub Leucopholis irrorata Sixty kilometers away in intensity variables were the number of rice crops grown per Real Quezon the main pests are seedling maggot Athe- year followed by area devoted to rice Table 3 shows that rigona orae flea beetle (Thacitcmnra hasalis thrips the upland rice site had only I rice crop perym rand that rice Stenchaetothrips hiurnmis and rice bug Leptocorisa was planted in only 20Vi of the area Fertiliter and oratorius In a third Philippine upland rice site in Claveria insecticide use had little bea ring on the trend The armters at Misamis Oriental Mindanao the main pests are ants the upland site applied 60 kg N ha higher than at most Solenopsis gemtinata seedling maggot Alieriona orvzae irrigated sites white grubs Holotrichia in)iUaiuoan t and Lucopholiv Insect occurrence at the upland site was omparable to irrorata stem borers Sesamia ifiArelr (hio auriciliusC that at the Cagayan rainfed lowland site parlicularly in 1982stpessallis Scirpophaga incertulas S innoiata and when drought prevented many farmers fron planting(only

4c ona chrrsurapella root aphids Tetraneura niriah- 40(of the rice area eventually was planted that year) M ore dunrinalisand Rhupaosiphunm riahbdminalis lcaffolders favorable weather occurred in 1980 and 1981 which Marasniapataltis and Chaphalocrocis Medinalis army- supported greater rice areas and consequently more insects worm MIthinna separata and rice bug Leltucorisa The most intensive site for insect pests was South Cotabato oratoriusThe great diversity of insect problems is typical Mindanao where farmers plant 23-25 rice crops per year fer upland rice and makes control efforts more difficult with irrigation The Zaragoa Nutva Ecija site also had particularly regarding breeding for insect resistance high pest abundance and at this site the fields were highly

asynchronous Asynchrony was the third most important Low yield potential Iactor in explaining rice insect abundance Most upland rice is grown on low-nutrient and mineral- Rice cropping intensity in upland areas is bound to toxic soils which when combined with eiratic rainfall remain low Only one rainid upland crop is possible per make upland rice a highly riskycrop for the farmerto invest year (Farmers in Claveria Misamis Oriental Mindanao in costly soil amendments and land management Often Philippines tried asecond upland ricecrop i 1984a ycarof upland rice isgrown far from markets making inputs even favorable rains but no harvest was possible b-cause of more expensive Farmers growing a subsistence crop hosi- drought blast and birds) Rice area isaso bound to remain tate to invest in costly inputs to raise yields low because of the high labor and power rcquirelmenlts to

Even having varieties that could double existing yield prepare the land (because of weeds)and because upland rice potential under present management levels would probably wvill not compete with lowland rice as a cash crop Some not prompt many farmers to use purchased insecticide upland rice sites with prolonged rainy seasons howeverare unless tile risk of crop failure were reduced such as it was planted asynchronously such as the Clacra site with irrigation systems in lowland rice The seasonal abundance of Upland rice insect pests

The Tanauan Batangas site is atypical for upland rice attracted toa light trap wasgrapied for crop year 1980-81 in The soils and rainfall pattern are favorable and because of Tanauan FBatangas (Fig I 2) W13P11 was he most its nearness to Manila farmers have cash resources ftm common rice hopper and its population peaked at the sales of vegetables to purchase tertilier They applied an midgrowth stage as it does in lowland rice D-Vac suction average of 60 kg Nha after particle initiation and obtained samples showed an earlier peak than that from a light trap 25-35 tha yields The high fertility isone reason why RLF indicating emigration W13PI usually emigrate after the armyworms and WBPH developed into large numbers vegetative stage Populations in 1980 reached levels that Farmers however used no insecticide on upland rice caused patches of slight yellowing in some fields WBPH although they did on vegetables probably cause hopperburn in Batangas They immigrate

The poor growing conditions fOr rice generally mean year-round as shown by low levels in the light trap BPH poor growth and low fecundity for insect pests fIeding on it however were scarce in the light trap and not collected with

Table 3 Abundance of rice insect pests and factors in rice cropping intensity of Philippine upland rainfed lowland and irrigated lowland locations where light traps were operated daily for atleast I yr -I

0Rainfed Rainfed lowland Irrigated lowlandupland

Pangasinan Cagayan lloilo Santa Maria Nueva Ecija South Cotabato Tanauan Caaringayan Caaringayan Bangag Bangag Bangag Buray

LagunaCale

Batangas Manaoag 1980-81 Santa Motiica 1982-83 Cabanatuan Zaragoza Avancena NaniamanSolana 1981-82 1982-83 Oton

1980-81 1979-80 Oton 1981-82 1981-82 Koronadal Koronadal1980-81 1979-80 1979-80 1983 1983

Insect (kerosene light trapNephotcttixcatch nolyr)virescens 650 1240 500 1800 330 210 3400 1600 160N nigropictus 210 - 360 18400 5500 2900- 140 90 20 - shy 1700 -Reciliadorsalis 710 - 3200 1700180 890 630 140 170 2400 800Nilaparratalugens 130 690 - - 4200 4600300 520 750 180 60 3300 1600 740 310Sogatellafiircifera 1700 31000 13000 8200310 210 3300 610 270 980 1600 6300 -Cyrrorhinus - 12000 6700600 540 3200 290 160 330 120 410Scirpophaga spp 50 410 620 - - 21000 12000150 180 220 270 30 1000Cnaphalocrocismedinalis 70 70 540 610 270 11000 6100200 140 100 6 60 250Chilo suppressalis 0 60 60 1900 300 200- - 30 20 0 6 3 160 -C auricilius - 0 00 - - 20 3 60 20 3 0Sesania inferens 0 - 80 60 3 80 10 51 -Cropping intensity _ -

Rice crops (noyr) 10 10 12 10 10 10 13Rice area (7c) 17 19 20 20 25 2320 85 85 60 70 40 85 85 90Fertilizer (kg Nha) 60 20 85 80 70 7020 0 0 0 30 30Insecticide (no of 40 60 40 300 04 04 0 03 300 13 13 30 50 40 33 33applicationscrop)

__________

IRPS No 123 January 1987 21

Light trapped Insects Drytnd __ 2(notrap-nght) Suctioned Insects (nom )

50 -)o 9 jO40- SOW ffurciWO Light trp

D-Vc suction

20 to 4

20

Egg porasitizahion (M) 75 60

300 SOOampReOftwciffr

45 30shy

15

-insects (notrop-night) 4 Nileporvata Iuen5

Egg parastization()

Light trapped insects (no trop-nght) Suchoned insects (no m 2 )

6 8 Cytorh~nuls hvidipennii 0

4 05 2

50 0 Insects (noltrop-night)40-ui Ipanh pe g

Moohase 1 Seasonal abundance measured by light trap

Rainfall(etod) and D-Vac suction sampler of whitebacked

to ~ hopper Nilaparvata lugens and lea ffolder60)-t Cnaphalocrocis mnedinalis andplthpe g20 L I II I I I parasites and mtrid predator Ortorhjntus livi-Apr May Jun Jul Aug Sep Oct Nov Dec JaIn Feb Mar dipennis set against cropping pattern daily1 9 80o 19 8 1rainfall and moon phase Tanauan Batangas

1980-81

the D- Vac Cyrtorhinus build up on the crop coterminous are collected in light traps during full moon (109)with WBPH buildup Cyrtorhinus disperses during the Because upland rice shares most of the lowland rice pestsrainy season RLF not readily collected in light traps was and those pest species are generally less impotant to uplandprevalent only during the rice crop rice than lowland rice there is a general belief that insect

Nephotettix virescens was more abundant than Nnigro- pests are less important onpicLus in the light trap collections and on the crop GLH upland than lowland rice Arcview of limited data on upland rice shows an averag of population peaked at the reproductive stage as did zigzag 10-21 yield loss from insect pests on a par with that ofleafhopper (ZLH) Crtorhinus tracked the populations of lowland uee determined by the same methodology (132) planthoppers and leafhoppers Both GLH species were (Table 4) McGuire and Crandall (144) made a similarcollected year-round in the light trap indicating immigra- estimate for Central America Cramer (51) howevertion from nearby irrigated rice areas Possibly because of the estimated 35 yieldlow populations loss in South America based on ainsect number did not correlate with similar pest complex - stem borers and seed bugs shymoon phase although reports indicate that more hoppers attacking irrigated rice in North America This estimate did

22 IRPS No 123 January 1987

Light trapped insects D~~nd (no trap-night )

St plht Suctined insects (nom 2

)

D-Voc suction 1520

1o

0 Lih trap 5 ~105 o5

00 Eggparositizotion()5040 NephOtgtha virneens

30 20

I010 A-Insects (notrop -night)

4 Nephatelh nigropccent i

2 o I vV^J - -- ^ VA____ f _

Light trapped insects (no trap-night) Suctioned insects (no rnE

10 -1208 ReCdO doRsol

6 15

4 10 052 iwxALAiamp 0

Light trapped insects (notrap-night) 2Suc oned nsects (nom )

8 CyrtorhmnusI6o nn]10 i0075

4 05

2 0 0

025

2 Seasonal abundance measured by light trap Roinfall (mmd) Moon phase and D-Vac suction sampler of green leafhoppersNephotettix virescens and N ngropicrus and I zigzag leafhopper Recilia dorsalis their egg 60

parasites and micid predator Cyrtorhinus livi-dipennis set against cropping pattern dailyrainfall and moon phase Tanauan Batangas1980-81

40

5 Lf Apr May Jun

1980 Jul Aug Sp Oct Nov I

-aMar

1981

not account for soil pests and was not based on field data in the absence of pests Carbofuran granules are phytotonicAkinsola (I1) cited mainly data from irrigated rice areas in to rice (239) The prevailing hypothesis is that carbofuranAfrica but estimated yield losses of upland rice between 15 prevents soil bacteria from consuming fertilizer The phytoshyand 30 from insects However losses in Thailand over a tonic effect is through greater availability of fertilizer to the4-yr period ranged from I to 13 averaging 5(106) plant Denitrifying bacteria are known to be more active inYield loss data however should be interpreted with aerobic than anaerobic soils therefore it may not be merecaution (184) The Philippine data were all derived from coincidence that the highest recorded yield losses (22-69)trials on farmers fields under farmers normal agronomic occurred in those trials with 12-30 kg basal Nha On themanagement practices except for Bukidnon and Capiz other hand contro with carbofuran granules may not be aswhere researchers used fertilizer Batangas farmers are good as a seed treatment against seed pests and perhapsatypical because ofhigh levels of fertilizer used The trials in yields would have been higher if a systemic seed treatmentBrazil were carried out at an experiment station but insecticide had been used conditions were similar to those of local farmers The 1979 and 1980 trials in Batangas had two full-

Carbofuran granules were used in every trial as a broad- protection treatments one using carbofuran granules andspectrum insecticide applied basally with the seed An the other using bendiocarb WP as a seed treatment Plotsobjection to using insecticides to measure yield losses isthat protected with bendiocarb yielded higher than those proshyinsecticidescandirectlyorindirectlystimulateplantgrowth tected with carbofuran granules The other trials in the

Table 4 Yield losses to insect pests and yield responses to insecticide in upland rice in the Philippines Brazil and Upper Volta

Yield (tha) Yield loss (5) Location Year Cultivar Protected

with Protected

without Check Total Vegetative Reproductive Ripening

Basal N (kgha) Reference

carbofuran carbofuran Protected No with carbofuran

carbofuran

Philippines Pili Camarines Sur Tanauan Batangas

Pangantucan Bukidnon

Dumarao t-apiz

BrazilGoias Minas Ge~ais

Upper Volta6

1975 1974 1974 1975 1975 1976 1977 1978

1979

1980

1979 1980 1979

1977 1978 1979

Bursiging puti Kinanda Dagge Kinanda Dagge Dagge Dagge Dagge UPL Ri5 Dagge UPL Ri5 Dagge UPL RiS UPL Ri5 UPL Ri5 UPL RiS

IAC47 IAC47 IAC47

14 33 11 36 35 31 30 27 37 26 40 30 40 42 46 35

19 21 09

--

-34 24 24 27 --

29 43 31 42 -

47 25

---

10 28 05 27 27 30 26 29 33 28 33 29 43 23 36 11

15 15 06

29 15 55 25 23

3 13 -7 11

-7 18

3 -7 45 22 69

24 28 35

--

21 -11 -20

4 -

-3

23 7

-2 -23 56

-

-

-__0

6 --

-

-23 14

5 0 8 2

22

-

--

0

-

--18 11

3 -10

4 10

7

--7 -

14 -

-1 6

10 -5

8 9 7

-

--

0 0 0 0 0 0 0 0 0 0 0

30 30 30

12 12 12

99 99

100 100 101 101 102 102 102 102 103 103 104 104 104

66 66 66

Farako-Ba

Mean

1977 1978

IRAT10 IRATIO

38 66

--

40 43

-5 35

21

-

10

--

--

--

- 242 243

Z

tJ

-w

24 IRPS No 123 January 1987

Philippines which had two treatments offering protection In the 9 Philippine trials where yield losses were measuredwith and without carbofuran granules relied on further by growth stage 10 of the yield loss (21) occurred at thecontrol with foliar spray and not on seed dressings The vegetative stage and 517 occurred at each of the reproshycarbofuran-protected plots had an 18yield los compared ductive and ripening stages Ibis result is similar to thewith only 10 with foliar sprayS This yield difference is pattern of yield loss in lowland rice measured by successiveprobably due to differences in protection against sown-seed treatments where the crop is unprotected at each growthpests (foliar spray do not control sown-seed pests) rather stage but proticted during the other growth stagesthan phytotonic effects became basal fertilizer wa used in However the insect complexes r sponsible for vegetativeonly 2 of the 10 comparisons stage yield losses in upland rice are entirely diftfrent than in

The recorded yield losses were not related to yield lowland rice The key vegetative stage pests in lowland ricepotential (Fig 3) One might expect higheryielding crops to arc aquatic whereas those in upland rice are sown--seedhave higher percentage of yield losses but this ws not tile (sown in soil) root and seedling pests This dift rence maycase One reason is that insect damage exacerbates plant explain why agricul torists have overlooked tile ilmportanccinjury from drought stress low yielding fields from d ought of upland rice insect pests Many reports of upland rice pestsstress would show disproportionate differences between in Asia ha Ve focused on major lowland species (46 108)insect-protected and -unprotected treatments Shoot and It isalso surprising that tei vegctative stage of upland rice root pests would be more responsible for losses from would record the highest yield losses becatuseseeral studiesdrought stress than sown-scd pests A second reason have shown that upland rice can readily recover from foliageconcerns seed pests which would have a greater impact on reinoval during that stage (117 152 165 188) Upland ricelow tillering low yielding varieties such as )aggc Low poscsses a hitgh obilitV to compensate for ca rly loss oftillering varieties nnot fill in the space created by removal foliage and will even be stinulated by foliage removal toof seeds oweer 1) R i 5 is high yielding because it tillersP1 produce a higher viel than plants ithout any foliageactively which can close the canopy to provide greater weed removed (63 192 231) [ie reason for this apparentcontrol discrepancy hovce is that yield loss at the vegetative

stage does not octUr frol leat arca loss but fron pestsYield loss () removing sown seed and feeding on devcloping tillers and roots -- damage that cannot be readily conpLtnsatCd for

70 - Measuring yield losses in upland rice cnvironrnents is further complicated by a report from Brazil that greater yield losses occurred in response to better insect control60 - =-09 as Insecticide seed treatments resuled in denser plant stands

(n=21) which in turn created a selectively more favorable environshyment for blast (69) This resuIt was parlticurla rlxexacerbated50 when cirbofuran granules were used in seed furrows Aside

ffrom a greater plant stand ca rbofuran iay have auigmented the N lecl and stimulated blast If so ca rbofuran should be40 - replaced with another insecticide or the seeding or fertilizerI rates decreased

Other methods to determine the impact of upland rice30 shy pests wvcrc to interview farners or to develop single speciesbull bull correlations of population levels and yield loss Feneira (66)2 NIM)st (75(1) Golas farmers reporteddid both in Brazil

20- insect losses in ufpland rice and 251 said those lossesS avcragcd 37(1 [his result is consistent with tie yield losses 0 0 measured by field trials in the same statc(Table4) For every

I0 4 of seedlings removC d by leaf-cutting ants a I 7yield loss is predicted Similarly Ferreira (66) found that ai

0 - average of 53 thrips per panicle before panicle initiation meant 3 times more unfillh d grains

-IO0 I 1 1 I I-101 0 2 3 4 5 6 7

Yield (tho) C)N [ROt IM FI IODS 3 Correlation between upland rice yield and yield lols caused by Upland rice insect pests may be controlled by culturalinsects measured in 21 trials in the Philippines Ivory Coast andBrazil genetic biological and chemical methods

IRPS No 123 January 1987 25

Cultural control dry season plowing will have little effect on them Plowing Crop husbandry to reduce upland insect pest populations when the soil is wet from the first monsoon rains will expose can be classified into practices effective in a single field and soil insects that tunnel close to the soil surface to predatory those effective only at the community level Cultural birds chickens dogs and even man (field crickets nole controls should come first in a pest management program crickets and white grubs a re delicacies in man tra ditional They have broad stable effects as pests have little pos- diets) sibility of overcoming them through selection of biotyps The greatest degree i1 inIscct control fron tillage comes (138) On the other hand some cultu ral controls work only from plowing soon after rice ha rvest when most soil insects with high labor or power inputs and connunity-wilt lie close to the soil surface anrd stern borers are in the rice methods require coordination of iany farniters stuble Whitegrubsare mature and tillage exposesthem to

Pantin time The luxury of being able to shift the time of predators Tirnelv control will prtec the crop following pla ntring upland rice is open onl hr la rniers in areas with a rice and lowr the populationIfor succeedinig years Plowingprolonged rainifall pa ttern lI copping intensity or underthe ice stulble helps it decompose killing sten borer rnechanied land preparatiot Gcnerally planlting as soon larvae inside as the rFiinv season begins will lower populatiois of 1110st [requent tillage destroys ait nests bit not termites insect pests Insect populationsarelo aftcra dlvseoll or Tillage and clean culture howes er rcliove food for winter fallow delayed plaiting lets them build tip on termites which then mtay attack a young rice crop Scarab alIternate hosts that gi vigorotsly with the first seasonial beetles also prefer to miposit in eceitly tilled fields Noshyratiris This first flush iof weds cros lItxuttriartlv with iceCult is dibbling faors the survival oftillace Clre Such rineralied N released during the dr season aid made stils i aels tnd insects thtt pttpate or lay eggs- in the soil

available to the plants with the rains It iliso igrtmitcally Zero tilla hio eerl conserves predators ofthe GIH (37) advanttgeous for the rice ciop to tap this trlLItlaI fertility nt it mcttoi lo lessen the impact o0 soil insects and grow vigorotsly to compete with wrus and to tolerate brotdcasting seed or planting in ftltrrows is prefc ired over pest infestation dihlirig in hills Crickets are large enotlgh to eat seedlings in

Larly plantngs terd to escape seedling maggolts Upland a hill atrd tillering normnally ca iriot fill in tlhe missing space rice in Bitangas is planted in Ma and ine itli the Iirst Oftcn dibbll rd is not well covered and iseasily found by rains Fairners are highly mtotiated to plant early tocscape foraging insets ioIdents and birds drlitght and typlotns ricar harct Arn Atrgtst planting Plan uit sii Incrcatsini sced density protects tile cropwould be evcely attacked by It riim orivzo At tti againrst seed and seelling pests especially ants particularly other Philippiie tipland sites Rea l Qteot antd (laelit if the crop will germinate and ermerge quickly M isainis Oriental seedlig mnaggots arnrige ric boe t se hr trftinll itI India rice intcrclopped witi cotton or farnes have a longer growing season and plnit 2 rin or pigeon pci had lower 6IIt and WBIlII poptrlations than mote after the ortset of the rainy season liaaiangas ftrmners rice alone (211) NfailC arid Urplald ir_ 2re coilltori in the plant ca rly becia use ofa shorter raiinfall period arnd the desi re Philippines he chatngC in pest status folii iniercropping is to lliarvet before the typhoor sea son peaks ltighly locatioll-specific (133) cmianarid the net elfc be either

Early plantings cornbind with early-iatiuring varictics nil bcnelficial ortdetriential (189) For example BIatangias provide higher crp tolerartce for white grubs Aestiatior farmers intercrip Itro in their ipland ricefields A liornshyterminates with the first rains but 5-6 moare ineeded for the nrii iotd-Ahis i (I iiiacius) which norlially feeds resting larvae to develop into beetles emerge and lay eggs on taro ill feed or miarbv rice plants Intercropping and for larvae to reach the danriigii g ird-instar stage probably would not afft0 seed pests A crluanion crop (134 In lataigas this normually iseiough tiite for harvest planted ith rice and haing gleater tolerarnce for root

It Brazil howecver delayed planting is a suggested dariage or being toxic nayact isi trapcrop to root-feeditig cultural control for the lesser cornstalk borer and insect pests Acial-fei i ig pests voultId be most affected by pests favored l drought (66 195) Cropping internsity in intrer ropping Shoot aphids leaf beetles sten borers and Brail is low and delayed plraing places the rice crop in the hopplers would be likely ca ntidates to note effcts On the niost stable rainfall period reducing likelihood old rought othlt halnd the corlipalior crop stch~ sitniaie Iiiv be more

In rilniy arca s rinmirsare roi atl to plant other crops benehited than rice from initecropping ( 137) before rice and althNgh it woultl be desirabh to avoid 11echi Weeuling luring the first month after clop infestations by planting rice cuatl lariners lack the labor cstablishent will force pcts such is anyvworris which and time to do so ol a low priority crp prefer grassy cetds ontor tle vorling rice Soil iampsecs will

77age Uplild rice Stils ae light textured and Call be mvnye to the rice crop Cleani citrltirc also lay lore territes plowed duiring the d rv seasor notl only to grin time for to attack I u cetincti inrice ceds i ay he le optinal planting with lie fitst Inilsoon riiains but also to desiccate solition 13) soil insects arid weeds such is ntsedge that foim rhizomes Irtlizr Itneatsing N Iciiliier oin hlwland rice has Htowever becta use most soil insects lie below die plow layer fAtwvored Irigher populains of pliithoppeiv Imlhoppers

26 IRPS No 123 January 1987

and leaffolde rs However because of the need to temper fertilizer use on upland rice o prevent blast fIrtilizer rates will be lower than is comonia r for lowlad rice causing relatixely less hopper and IaIfTfoide r population buildup However inupland rice areas without a history of blast such Is Batangas farmers apply high rates of N and conscqtrently the crop has high infcstation levels of WBPI]and RIF The fhrmrcrs leec insect buildup by splitting the fertili11r in0to two to three applicatiois bit do not applV fertilizer at the vegetati estage because this prodrces a tall crop which illeasily lodge during typhoons Greater crop fertility leads to greater insect survival (better nutrition) larger insects (to cat no re foliage) arid greater fec titd t (to lav nmre eggs) (128 145 171) Not all insect species respondequally hovr Increasing rates (if N l anrid K increased Dialraa s-acchralis and (iaeiurema flea beetle but reduced thrips (66) Zn rcdLuced deadheart density from the lesser cornIstalk borer

Fnldingi Although most Upland rice areas do nIot hawxcaccess to wa ter floodiig the fields is rRrrnriierdcd in areas of Brazil or root bug and white gib control ( 195)

Crop rotatioti Tillage before planting a crop after rc harvest will control stern boreis and other pests renain n the stubble arid will unearth soil pests Planting a non-gratriiaceous crop after rice is recommncrded for termite control (66) Crop rotation is less nttdcd in upland rice culture than in lowlands because atcrop-free period is assured even after harvest of crops following rice

t1ulchilt iryl plastic irli irig designed for weedcoitrol ii upland rice inJapan resulted in ntr highanri liy infestation of (ihilo sulytresvais (93 9-1 95) NIulched Upland rice grew more luxuriantlv and attracted more (Avipositing mothis than u lched rice Also the rice plant and stein borers imatoured 10rC quickly and a partialgeneratiori af vouing ista iNoverwintcred in the stubbleLotili--malitrt varieti Quickly maturing crops reduce the tmbCr of pCst generations that Calt build up Rapid crop establishment olan ca rl v-natu ring vaitvty if)pa 11icLa r will lessen white gib and stem borer damiage White grub larvae intle last iistar are highly destIruCtive so aii early-iiatri ng variety will escape serious root loss Stem borers build 1ip slowlyx in rice to ecormic abundant in a late-Maturing crop anrd caiuse whi tlieads Numibers of somespecies increase exponeitially with each succeeding genera-ti so a xarietyexhat atures I trowh earier thian another will hac less insect damage Early-naturing vaiicties however will not lessen seed pest daiage either ii the soil or oina standing crop

Strwhoi hAIJttam Inrscts tcd irig on thlie aerial portions of rice plats disperse from carly to late plantedfields AS iiiibCirs increase exponentially with eachinsect

gtscnrattm e eiglbrig f rmc~s1(uil upp wh rc generation pests build tIP where neighboring fihrTC e9r1sstagger their plantings II Batangas farmers plant uplandrice within I iro use 120-d s-arieties and plant at the start of the rainy season Ini Misarnis Oriental laricrs stagger their

plantings up to 3 mo apart and use a set of atieties that mature in 4-7 mo Insect pest danage particularly from stem borers is greater in Misarmis Oriental Also seedling maggots damage rice there because rainfall favors frequent planting of maic In Batangas rice escapes seedling maggots bCca usC of distinct drs season and synchron0us early planting with the onset of ains

Therefore combinations of cultural practices ea rlyplanting synchronous planing crop rotation and earlyshymaturing vaneties protect the rice crop against most insect pests

Ilmant resistance Beca tse upland rice has few specialized insect pests and many insect types attack the crop worldwide regional breeding and strong internatiornal cooperation are required

Sown-sced pests arc economically controlled by insectishycide seed treatment and should the relore receive lowxpriori ty inbreeding objectives

Root pests are normally cost ly to control with insecticides but finding resista t sonres mayvbe di fficult Tanaka (229) for exa -ple fi led to find varieties resistant to root aphids Selecting for large root bioniass perhaps should be the strategy for pests sLuch as Whlite grubs that renliove roots Fainter(174) howcvcr cites varietiesof crops tolerant ofor resistant t(i root pests Some sorghun varieties are high in cyanide so rice sorghun might incorporate broadshyspcctruri chemical resistance

Seedling vigor and drought tolerance are high prioritybreeding objectives so tupland rices should have large r root systernsand be able to tolerate ligherlevels of root loss Sonie root loss inolder plants is beneficial if it stimulates new root developnrut and therefore crihances intake of soil nutrients (53) The rice seedling inaggot rtiLotia spp may be

controlled thirough resistant aittics A hi rgc breeding effort on sorghniu has had fair success inideveloping arreties esrstait to Asoccata Rodani (251) A field trial comparing 10 upland ricc varieties showed differences aniong varieties va ring from II to 4(- damaged tilleis (215) Hovver resistarce to 1Iill-0r gtia species may rot cross ox-rti otier

Shiraki (210) reported that upland i-ice stens wcrc harder than lows-land rice stemsperhaps becaIusc (f a higher silicoricontent and thus were more resistant to yellow stern borer hc tortia Iow tillering (f uplaid iice xarneties prodtres a

higher percentage of infested tillers than would occor in higher tillering lowland iice under a siiilar egg density The corollary therefore is to select for high tilleriig upland rices

Stemii bor amcstheh main targes dobrecders inAfrica (571 204 )but nticl oifth e sc reen irig is dotie unde r lo wland co d t osrm s wih A h r~ o l pe i s n acorditions (11 96 218) As with lthrinrnwspecies mayhave to be dealt with independently however initial results show cross resistance between African and Asian species(92)

In Brazil screening Ior insect resistance focttses oil lou I pest problems Varieties tolerant of lasmolwalpus ligi-sellus and resistant to )iatraea saccharalis have been identified (70 153) Current work also emphasies 7iraca limativetntris and Thmialspis (=Iosis) flavouicta (66) These insect pests Ilt onlV citise IIigh yield losses tihey also ire difficult to control chemically ofr by other ilealls

ManV upland rice vaieties aic early maturing and are resistant to pests such as gall midge (I Xl ) Early mIturity is often a highly desirable trait to esuilpe pest buildup but it is not true resista nce bca usc ilthose va reties are plinted late they will be itsdamaged as ain early planted susc ible variety

P~erhaps tile cosiiiopolitall of all upland rice pestsimost are the sced pests (ripenling stagc) which pose a particular problem beta use scuds re ised IsIO for hitnit4ans Ayltoxic substatice introd tcld geticticllyilmay also be toxic to nal iniless it caln he placed old in thfe IdhuffSelectmi fo hairiness or lotng wns kill mIlct sthilrohleiiis dutilig threshing pa rtictfarl maitial threshinfl [hireshers ill complain of itcfimin atid itteascd (fust levels

Attempts itIRRI to fitid tcistatncc to the rice bug plorisa have ilotbeCnl prmitisilg lIIetIattonlids iorc

through the lenilna or palca hillhairs sedstVIor seeds wilh awns that deter bugs promote complaints from threshers Seed bugs liptocorisa and Stcnocoris enter the seed through the opening between tile lemna and palC but varieties with narrow openings are diifficult to mill

Breeding priorities should concentrate ott inotCI gruLtLIps that are most difficult to c nttrol by other meains Steil) horers appear to be the most widespread group that sigiiificantly dalag s rice aind is hard to manage by other Ilnca us

Biological control Very fe stutdic of mitatt~l eneties of rice pests in upland envirollnletits wee in tile wefouiid literature Therefore have not compiled a list of species Ve present a list of natural enemies iOtid in upland ricefields tuic latialln Batangas for some conitno iOldiar pest species ([itble 5)

Muich ofl thc lollowinle itfor ittiol cotlliCs rol Stutdies in the lowlatnds or roi crops otithcr thlin tice

Qu(ailtr o 11ll t11t( Nat trrtf lii hcoeits ight Ile used agaitist ile social insects tilt and termites Sollie predator specice live ill theirf nests n ilnic theirnllt ip-pearanIlCe aMRidbehavior IJtrouMdCing a mote aggressise ant species to displace pest ant codonies is a possibility (169) Patihogens as potential natural encnies have reccived little atteition (I13) Natural ncnuics asa group however hac not been toIl iporta it in Iegulating aintshown be and ternite numbers

The nocturnal behavior aid subteitmien habitat of lole crickets and field crickets protect them from many ntllral enemies There are only limited records of natural cnelics for these groups of pests some pathogens (87) but mainly

IRPS No 123 JattuIny I 271987

parasites - scelionids on eggsand sphecids on ilymphs and adults Sphecid wasps Larra carutmaria (SImitI hI L huzwuensis Rohwer and lsaltiuhta Willi ams speciali in mole crickets pa ralyzing them nd dragging the ninto thcir nests as food lot their voting ()tier sphCcid species Lins aurulenoa (Iabricius) lot maiac (Ashmead ) 1 lhshylssellaus (Smith) and Ml lal)oriosu (Snithf specialie in field clickets

White grUbsO the other hand hae Icti scriuns enuigh to have been extensively studied for cotttroi 1W nati al enemies( 112 140 141) Scoliid parasites C(ajsoMcispp have beetn intrtduced itto tilePhilippites for control of Ielxl ]udisiro la The rcsearchcts w Intttrdced themle during ill era claimed that the white gttionthreak as controlled (142) howevcr it isnormal for pest epidemics to subside and there was no diect evidence to indictte that tfe parasite was responsible A tmore promisitg titctllod might he to identify iso ltcttass prt~f tcartd apply hiactelaln fingal pathogens to tile soil Ihe lalattese beetle in North America has hcci coitrolled hv tite conmnleciall aailable hactcrial prcpa ration itacihlpu pav ofr milksdiscas (71) Spote prtyefott the soil itloted ilrtier reitiain actic fr Veas

Root aphid sardtnctly bugs readilslic tilib to predatots and parlsites once their tetndine ants are controlled The larvac and adtilts of a coccinellid beetle 1Scynms sp prey on llriadlouais in slash and burn upland rice Baits treated with insecticide may be used to control tending ants by killing their ourig (244)

Root-tkcditig blugs false wireoris wireorms and root wccils aic r ofsuch little ifilpoltatice that thenomttll role of liatural Cticniics hias tiot beei asscssed A possibility hoevrl tiglt he tit ItaIIitic niaCIMtodcs lintt1g11en Itese soil-itlhlfbitine mCtitodcs W5uhtfhave to be imiass Produiced

large miltibcr of niatltrll cnemics ircknown ft CltIoriS IC IpblCeii is thtt clttvortnlls ofonlie the crop soott alter land ptel)tritiOti a1tithe bcgit iniig of the raittv

Citsoln hfien tiatllral leniiiy populations aric lo Colltloil WOtuld iii ole mass prodtticing andirclcaslng key spccies

le egg stage is the fpirt (olthe Cedlitgt Ialot life cycle tI hich to f)ctis ftor tittlril elctit cottntrol hplraites o1 pleditors ttsit elleggs highly expoIsYCl I Sgg pailsites or predators stolld probahly e ittass andtbe produced released Littlh i toruialioll is a ailable ott what species to trV o ho to Iniss podLice thiCIi A etiloplhid wasp Icmihlaol sp effctiicly regushy

lates the leatf lcloalapolm2a spioanoiiitr heat andc rice antid should be collsctvcd

leaf beetles whCllose e am] eggs ate laid in the grutindhirO ould be motC difficutlt to control by niatulral enelV Mnll ipuhltiOn But species sUici IS c_gsthispid beetles ith

and larVle oilleascs ate vtilcratble to iparaites and pledators

Natural enemisc of tlirips otl rice are little known

28 IRPS No 123 January 1987

Table S Parasites recorded from upland rice insect pests in Tanauan Batangas Philippines 1977-84

Order Family Species Host stage

I lymttenoptera Braconidae Leaffolders Cnaphalocrocis Marasinia spy

Cardiochilesphilipplinensis Ashmead a Larva

Cotesia angustillasis(Gahan)Cotesia or cypris (Nixon)

Larva Larva

Cotesia or taeniaticornis(Nixon) Larva Cilonusinotakatae Munakata Chelonits SppAlacrocetrts philippinensis (Ashtmead)

Egg to larva Egg to larva Larva

Iclineumonidac Orgilts spp Isclutojppa hireator(Fabricius)

Larva Larva

Itoplectis naraigav(Ashmcad) Larva Tenelhtcha philippiensis (Ashmead) Larva Tenelchha stangli (Ashmead) a

irichomma cnaphalocrosis Uchida Larva Larva

Bethylidac Elasmidac

Goniozms or trianguli r Kieffer lasmtusalhopictus Crawford

Larva Larva

Diptera

Encyrtidac Chalcididac

Tachinidac

Elasmtus spp Copidosomopsisnacohiae (Fidy) Brach nteria evcarinata Gahan Biachyieria lasus (Walker) Ztgohothria ciliata (Wulp)

Larva Egg to larva Pupa Pupa Larva

Argyrophylx nigrotihialis(Blaranov) Larva

l-ymenoptera Braconidac Yellow stern borer Scirpopiaga incertulas

Braco chinensisSzeplegeti Larva CIthelotis spp Chelonus niunakatae Munakata Cotesi flaripes Cameron Stenoracon nicerilli (Bingham) a

Larva Egg to larva Larva Larva

Ichneumonidac Tropobracon schoenobii (Viereck) Amatroinorpiaaccepta metathoracica(Ashmiead)Frihorus sinicus (I lolmgren)

Larva Larva Larva

Ischnojoppaluteator (Fabricius) Larva Isotima nr dammeratiti Rohwer Larva Tenelucha philippinensis (Ashmead) a

Larva Tenehicha stangli (Ashmcad)Trichomma enaphalocrosis Uchida

Larva Lara

Chalcididac Eulophidac Trichogranmmatidae

Xanthopimnpla stemmator (Thunberg) Charops hrachYpterumn Gupta and Malteswary Brachtm eria sppTetrasticttus schoenohii Ferriere Trichorigranm chihunis Ishi

Larva to pupa Larva Pupa Egg Egg

Diptera

Scelionidac Pteromalidae Tachinidae

Tt ichograintnta aponiicuitt (Ashniead) Telenomits rowani (Gahan) a

Trichoinalopsis apattelctena (Crawford) Jeirhaea spp

Egg Fgg Egg pupa Larva

Z4gobothria ciliata (Wulp) Larva

Ilymenoptera lecneumonirlac Gol fringed borer (hilo auricilius

Trichomm enoptalocrosisUchida Larva

Ieioenoptera Braconidae

Aantholimlh steiniator (Thtinberg) a

Pink stet Ioret Sesamia inferens StoInacoil nicrilhi (Bingh am)

Loarva to pupa

larva

Diptera Ilyinenoptera

Armvtnyorms Mythimna separata Spodoptera mauritia acronyetoidesTachinidae Zvgohothriaatropiora( Robinea i -Desvoid y)Eulophidae It-phctrtiscitapadoe (Ashimead)

Larva Larva

(halcididae Braconidac

Brachpin ria spip (oesia spp

Larva larva

IHyntenoptera I)iptera

Rice skipper Ielop ida (halcididae Tachinidae

math ias aidgreen iortedci ciliarNIelan itis eda Brat lhincria sp iii matuqitatt Cameron Irgyrolhirlaxi trotlhiais(marumtv)a1

ismtne Larva Larva

Diptera Ilymenoptera

Tachnidae lelineutionidae

fuown se tilo per Mois frugalisIrgtrophrlax 1igrothiahiv 3 a rat ov) anthopimpa pmnctata (Iabricius)

Larva Larva to pupa

aMost dominant

IRPS No 123 January 1987 29

Normally predatory thrips prey on herbivorous thrips strepsiptera on both Spiders dominate tilenymphaladultNatural enemies of stem bugs --Scolinoplara and Tibraa predators Spittle bugs move to rice from pasture grasses - are in order of importance egg par~isites and predators and biocontrol would be successful only ifcarried out before fungal pathogens and nymphal adult parasites Egg tiley immigrate into ricefields parasites could either be introduced fron other areas or Biological control tactics for hemipterous seed bugs mass produced and released ould follow those outlined for stein bugs

Armyworms are normally lwld in hcck by the activities Coml)arison of environnewtns Natural enemies are of egg and larval parasites When these natural enemies fail perhaps faced with even greaterconstraints than insect pestsusually because of drought armyworns become epidemic to survive ind reproduce in upland environments TheyIt may be worthwhile therefore to release parasites during have adapted ani survived by having wide host ranges the the rainy months following a drought Virus diseases of ability to aestivate or hibernate over unfavorable seasonslarvae would beanotheravnue to exploreforarnyworms and the ability to disperse or by other mechanisms

Parasites have ben traditionallyconsidered for biolegical Comparing the most dominant and therefore the most control attempts against stein borers usually by rekasing adapted species of natural enemies collected in various exotic species( 166) This approach has not met with success environments in the Philippines we can understand how and new avenues should be explored The role of predators different rice environments are arid pathogens is little inderstood and deserves greater Anagqrms olptahiisa mymarid planthopper egg parasiteattention Egg parasitism rates are normally reasonably is cquallyadapted to all Philippine rice environments (Tablehigh but should be supplemented with effective egg 6) It is selective for planthoppers and even parasitizes eggspredators Orthopterans are potential egg predators of the n-iize planthopper Peregrinitsmaidis itt upland areas Metioche vittaticollis (Sta ) Anavipha nifiwnnis (Saus- Alavohis also parasitizes maize planthopper eggs but is sure) and Conocephahs hongipnni (de i-laan) feed on most prevalent on BPH in the irrigated wetlands (onashystem borer eggs etioche and Anaxil)ha specialize in eggs tocerus spp like A optalhilis on BPH and WBPI Iattacks with no hair such as Chilo spp The omnivorous (ono- the eggs of GLH in ell rie environments It is also found cephahsspecializes in eggs cov red with hairs it is a proven parasitizing the eggs ef the white rice licahopper (oimna egg predator of Scirlouphaga spp but has the discouraging spetra Oirosita naias a trichogrammatid egg parasitehabit of eating rice guin (65) attacks BIPH and WBPH in the wetlands and has no other

Grasshoppers have egg (scelionid) and nymphalad ult host A related species 0 aesopi specializes in G1-H in (nemestrinid tachinid and sarcophagid) v irasites Among wetland environments Dryinid strepsipt-an and pipunshypathogens protozoans have been recorded most frequently culid nymphal and adult parasites of planthoppers and Control of locust species by natural enemies could con- leafhoppers also specialize either in hopper species or centrate on their habitual breeding areas environments Among the dryinids Pseuhdgonatoipsflavi-

Leaffolders have rich complexes of natural enemies some feimur prefers the lowlands and BPH while P nuch has no adapted to upland rice (22 27) ranging from egg predators environmental prefirences but is adapted only to WBPH (gryllids coccinellids) to larval parasites (braconids ichncu- The main d ryinid species on GLH aptogonatop)usspp is monids) to larval prcdators (ants and carabids) to larval most dominant in rainfed lowlands and also attacks other pathogeiis (vira and fungal diseases) Efforts to augment leafhoppers within Amnrasca Cicadulinaand Bachaltuathese natural enemies should focus on the egg predators Strepsiptera are more dominant on plarithoppers than (rearing releasing)and viral or fungal pathogens (culturing lealhoppers The Elenchus species in the Philippines attack spraying) all species of planthoppers within the genera Sogawilla

Parasitization of the large conspicuous larvae of Sogatodels NilaparsataHarualia and Opiconsiva Elenshyskippers green horned caterpillars and other polyphagous cusrvasuntaisuiis adapted to the major rice environments Lepidoptera by tachinids and chalcids is normally low ELg but prefers BPH in rainfed arid WBPIH in wetland environshyparasites and predators are perhaps the key to their control ments Ilalictoph(utsmtnroeiand HVpectrus attack G LH

Gall midges are parasitized by pterormalids and platy- and COfimravcctra respectivelygasterids that aestivate within their larval host between Pipunculids are mainly parasitic oii lealhoppers Each seasons Shoot aphids are preyed upon by coccinellids species has a unique environmental preference Tomosshysyrphids and chrysopids varireIa oraetoraprefers Ne7 otetltix nigropictrs and

Planthoppers arid leafhoppers have egg parasites and Pilunwulhtsjavanensisattacks Dehoceplhahls spp which predators as well as nymphialad nlt parasitesand predators are more abundant in upland than irrigated lowland their rich complex of natural enemies includes fungal environments T suhvirescensattacks Nephotettix virescens pathogens The principal egg parasites are myrnarids and and prefers the lowlands Pilpuiulus nttillattls is an trichogramniatids Egg predators are norinally mirid plant upland rice parasite arid attacks only Nephotetix speciesbugs that suck out the yolk Nymphal ad nIt parasites are More than 17 recorded species of leaffolder parasites in dryinids on planthoppers pipunculids on lealhoppers and the Philippines attack species of C(naplwocrocis and

30 IRPS No 123 January 1987

Table 6 Dominant natural enemies of rice pests in upland rainfed lowland and irrigated lowland rice environments of the Philippines

Natural enemy

Host stage Species

Brown planthop-Parasite Egg

MymaridaeAnagrus optabilis(Perkins) Anagrusflaveolus Waterhouse

TrichogrammatidacOligosita naiasGirault

NymphAdult Dryinidac

Pseudogotatopuslaiifemttr Esaki et Hashimoto Elenchidae

Elenchus yasurnatsui Kifune and Hirashima

Environment

Upand Rainfed lowland Irrigated lowland

Ailaparata lugens

x x X x

x

x x

x X X Whitebacked planthoppers Sogatella furcifera and Sogatodes pusanusEgg

MymaridaeAnagrus optabilis(Perkins)

TrichogrammatidacOl~gositanaias Girault

NymphAdult Dryinidae

Pseudogonatopusnudus Perkins Elenchidae

Elenchus yasutatsui Kifune et Hirashima

Parasite Egg

MymaridaeGonatocerus spp

TrichogrammatidaeOligositaaesopi Girault

NymphAdult Pipunculidac

Tomosvaryella subviresccns Loew Tomosvaryella oryzaetora (Koizumi) Pipunclusmutillatus (de Meijere)

DryinidaeIlaplogonatopusspp

Egg Copidosomopsisnacoleiae (Eady)

Larva BraLonidae

Cardiochiles philipphnensisAshnead lchneumonidac

Trichiomnia cnaphalocrosisUchida Temelucha stangli (Ashmead)

Pupa Chalcididae

BrachymeriaexcarinataGahan Brachymerialasus (Walker) Brachymeria nr tachardiaegroup

Parasite Egg

Scelionidae Telenomus rowani (Gahan)

EulophidacTettasticusschoenobhi Ferriere

Larva Braconidae

Stenobraconnicevillei (Bingham) Iclhncurnonidac

Temelucha phiiippinensis(Ashinead)Temelucha stangli (Ashmead)

x x x

x x

X x X

x x Green leafhopper Nephotettix virescens

X x x

x x

X X X X

X

X Rice leaffolders CnaphalocrocisMarasmia

x X x

X

X X

x x x x x X x x x

Yellow stem borer Scirpophaga incertulas

X X x

x

x

x X XAinauromorphaaccepta metathoracica(Ashmead) x

continued on next page

IRPS No 123 January 1987 31

Table 6 cntinued

Natural enemy Environment

Host stage Species Upland Rainfed lowland Irrigated lowland

Larva-pupa Gold fringed borer Chilo attriciiuslchneumonidae

Xanthopimpla stemmator (Thunberg) x aPupa x

Pteromalidae Trichoinalopsis apanteloctena (Crawford) ax

Pink stem borerb Sesamnia inferensParasite

Larva Braconidae

Stenobracon nicevillei (Bingham) x b b Armyworms Mvthinna SpodopteraLarva

Tachinidac Zygobothriaarroptvora (Robineau-Desvoidy) x x

Braconidac Cotesia spp

X Skipper Pelopidas mathias and green horned caterpillar Melanitis leda ismeneF-gg

TrichogramrnatidacTrichogrannaspp x

Scelionidae Undetermined Scelionid

x Larva

Tachinidae Argyroph lax nitrotibialis (Baranov) X x

Pteromalidae Trichomnalopsis apanteloctena (Crawford)

X Predator

CoenagrionidaeAgriocnenis femina femina Brauer x xIschnura senegalensis tambur

Miridae CrtorhinsliidipennisReuter x x

Nabidac Stcnonabis tagalicus (Stal) x

VeliidaeAlicrovelia douglasi atrolineata (Bergroth) x

Coccinellidae Micraspis crocea (Mulsant) X x x

Vespidae Eumenes campaniformis (Fabricius) x

Formicidac Diaecamna spp xOdontoponera transversa (F Smith) x

Carabidae Chaenius spp OphioneaW- aJasciataSchmidt-Goebel

x

Ophionea ishii ishii Habu x x X

Gryllidav Anaxipta longipennis (Saussur-) x xMetioche ittaticollis (Sial) x x x

Tettigoniidae Conocephalus longipennis (de Haan) x x x

aNo rearing made on rainfed lowland bNo rearing made on rainfed lowland or Irrigated lowland

MrasmiaAmong the Laval parasites the braconid Cardio- lowlands and parasitizes oriental maize borer OsIriniachilesphilijpphensis is most adapted to an upland environ- furnacalis (Guenee) larvae The leaffolder pupal parasitesment It also parasitizes Hydelepta indicata a common not only occur equally in all environments but also haveleaffolder of legumes The ichneumonids Trichomma cna- wvide host ranges- parasitizing species of OsIrinia Marucaphalocrosis and Temehcha stangli occur more in the Hydelepta and Honiona - in maize and legumeslowlands T stangli is most adapted to irrigated lowlands The principal egg parasite of yeltow stem borer (YSB)and also parasitizes Chilo and Scirpophaga stem borer Telenomus rowani is widely adapted to all environmentslarvae T cnaphalocrosis is most prevalent in rainfed and has an unusual alternate host -Tabanus eggs (25)

32 IRPS No 123 January 1987

Tetrastictu schoenobii specializes only in YSB and occurs mainly in irrigated lowlands -- further proof that YSB is most adapted to the wetter environments

Larval parasites of YSB are highly environment-specific The braconid Stenohracon nicevillei occurs only in upland rice but also attacks the pink stem borer (PSB) Tieinchwa phiiiplPwnsis an ichneumonid occurs in upland rice and also parasitizes the larvae of (7hiu stqpresaisinatro-morpha accepta melaihoracica like 7 stangi is mnost adapted to the irrigated lowlands

The dark-headed stem borer(l)HSB) (ih pochit3sus was not collected in all rice environments and the gold fringed borer Chilo auricilius nd PSB were collected only in upland rice Xaantiholiplastenrnaloran ichneumonid larval parasite appears to be widely adapted whereas TrihomaslopisapanitI)cteiiaa pteromalid pupal parasite occurs mostly in wetlands and also parasitizes larvae of Chilo suppressais and Pelhuidas mathias

Armyworms Myit hinin and Spodujipera have two main larval parasites - a tachinid Zrgohothriaatropivora in rainfed environments and a braconid Cotesia spp in irrigated lowlands

The butterfly pests Pelopidas and 4elanitishave tricho-grammatid egg parasites found only in irrigated lowland environments

Two larval parasites show environmental preferences --Argrroph av nigrotihialis a tachinid in rainfed habitats and Trichonialopsisapantelkcnela in irrigated ricefields A limgrotibialisalsoattacks the sweet potato hornwormAgrius convolvtdi and T apanteloctena parasitizes DH S1B

Predators as a group show more distinct environmental preferences The aquatic damselflies Coenagrionidae are most abundant in the lowlands Cvriorhinus is also most adapted to the lowlands The nabid Stenonabis tagalicus occurs only in the uplands and isalso prevalent on legumes The coccinellid lady beetles wasps and ants are more prevalent in upland areas AlicrasIpis cit-ea larvae prey ona variety of aphids attacking legumes and maize Etunienes campanonia vespid mud wasp makes nests in trees and is therefore most adapted to the more botanically diverse uplands Soil-dwelling ants cannot tolerate flooding

Upland environments are habitats to arboreal carabid beetles Three species of Chlaeniusbeetles prey on leaffolder larvae and are perhaps more important than parasites (27)

The gryllids and tettigoniids are widely adapted egg predators not only on rice but also on maize and legumes They feed on eggs of most insects that are laid on leaves

The spidercommunity of upland rice environments isrich in species At one site (Batangas Philippines) 31 species have been recorded (23) Of the 176 spider species recorded in Philippine ricefields about one-half (82) occur in upland rice The spiderspecies of upland rice environments overlap more with those of rainfed lowland than with those of irrigated lowland rice (24) The same study showed that of the three environments irrigated rice has (ie greatest spider species diversity foiiowed by rainfed lowland and upland

However of the 10 most prevalent ricefield spiders 9 were abundant in upland rice showing wide adaptation All three spider guilds -- orb-web space-web and huting spiders shywere prevalent in upland ricefields There were however differences in environmental prelerences for some spider species

Of the orb-wcavers Tetragniathamanlihulatawas partishycila rlv abundant 7 japonica was low and Ltucatge dehcorata was absent in upland rice

As in other environments Aiypeiwnaifirtosana was the most dominant space-web spider however its relative numbers were lowerthan for lowland sites Among hunting spiders Ox|opes javanus and Lcosa leatcost fgla were more abundant and Lvcosa lpseudoanmnlata was less abundant in upland compared with lowland rice envishyroniments

Extensive rearing of rice insect pests in upland rainfed lowland and irrigated sites over an 8-yr period in the Philippines provided insight into the effectiveness of parasites as natural enemies (Table 7 8) Most of toe data are from large sample sizes taken over at least one crop which overcomes the pitfalls expressed by Van Driesche (238)

VSB egg parasitization was surprisingly similar across all environments Larval parasitization was highly dynamic among sites and even years within the same site but showed somewhat lowerlevels in uplandsthan in irrigated wetlands This would have been predicted for egg parasites as well since YSB is adapted to the lowlands and so must be its parasites This isevidence that Telenomus rowani is as good at dispersing as its main hosts Trowani clings onto the anal tufts of female YSB and parasitizes eggs as they are laid This phoresy may explain its wide adaptation The more specialized lowland YSB larval parasites therefore appear more effective than their upland counterparts with more alternate hosts

Low collection levels of PSB larvae especially in the wetlands make comparisons between environments difficult Low levels of larval parasifization were recorded in both rainfed environments Except in the Cagayan rainfed wetland site in 1980 low levels ef larval parasitization were evident at irrigated and rainfed sitesalike No differences by environment were evident Higher levels of parasitization occurred in Sarawak during a 1967 outbreak also from tachinids Rothschild (197) indicated that higher parasishytization occurred in upland rice areas

Skipper and green horned caterpillar larval parasitization rates were generally low and showed no environmental effect htighest activity occurred in Cagayan in 1980 (48 parasitization) Although larval parasitization levels of RLF were variable year to year activity appeared greater in the irrigated lowlands than in the uplands Again the highest levels were recorded in Cagayan in 1980 Cagayan has extensive grasslands and fallow around the rice areas and a relatively long rainfall period Rainfall in 1930 was good for crop growth and apparently for increase of larval parasites

IRPS No 123 January 1987 33

Table 7 Parasitization rates of rice planthoppers and green leafhopper in upland rainfed lowland and irrigated lowland environments Philipshypines 1976-840

Environment and site Year Egg parasitization () Nymphaladult parasitization ()

BPH W13P GLH BPII WBPH GLIH

UplandBatangas

Rainfed lowland

1976 1979 1980 1982

15 17 12

2(47)8(39)

4(51)

9(1800)14 (212)

12(304)

8 (101)6 (127)

9(116)

Iloilo

Cagayan

1976 1977 1978 1981 1980 1981 1982

0 (69)

11

44 48 40

-

-

46 55 44

0 (471)

11

52 56 49

0 (200)9 (214)

10 (1259)8(235)6 (900) 3(800)

-7 (462)

11 (270)3 (313)5 (900) 1 (800)

4 (600)10 (725)10 (362)12(870)5 (900) 3 (800)

Pangasinan

Irrigatedlowland

1984 1976 1978 1979 1982

13 (436) 13(379) 14 (457)

6(14)7 (197)

17(198) 11 (486) 18 (275)

3 (81)10 (274)3 (116)

16(871) 19(88)

28(211)9 (764)5 (647)7 (1204) 8(305)

Laguna 1977 11 7 21 1978 31 1979 14 4 4 28 1980 7 8 20 1981 1983

56 7(278)

26 16 (1650)

16 27(1921)

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R I Apostol unpublished 1lo1lo and Pangasinan 136 Cagayan BCanapi unpublished Liuna 175 I C Pantua unpublished 118 234 43 Carino and Shepard unpublished BPH = brown planthopper Nila parvara htgens W131Pi =whitebacked planthopper Sogatelia furcifera GLH = green leafhopper Nephotettrix virescens Figiires in parentheses are sample sizes

Tauher et al (230) reported that internal external and genetic factors influence the seasonal activity of parasitoids

Hopper egg parasites but not nymphaladult parasites were also parlicularly abundant at the Cagayan site possibly the egg parasites have more alternative hopper prey in the weedy areas than nymphaladult paisites

The seasonal dynamics of hopper egg parasites can be seen for upland rice in Batangas(Fig I 2) and for rainfed lowland lice in Pangasinan( 135)and Cagayan( 105) Levels of BPHWBPI-L and GLH egg parasitization were corn-parable in upland and irrigated lowland environments Except for some high rates of paraMsitization early in the season the parasitization rate tended to be steady in all envilonments However hopper nymphaladult parasi-tizution levels were more similar in the rainfed environments and lower than in irrigated rice

Overall parasites do not appear highly effective by themselves against upland or even lowland pests Among natural enemies parasites have attracted the most attention because they can be more readily assesscd

Predators and pathogens lend themselves better to management practices Predators the most important group of natural enemies in riccfields are difficult to quantify On the other hand pathogens appear less im-portant but can be readily cultured and disseminated

The natural enemy commulnity of upland rice is rich in species and differs significantly from that of lowland rice

These beneficial organisms must be conserved by applying insecticides judiciously particularly sprays Seed or soil placement of chemicals minimizes exposure to natural enemies The strategy to derive the greatest benefit from natural enemies isto allow the greatest number of beneficial species to thrive Then perhaps at least one species will be effective against each pest at any tine overcoming the variable abundance of each species within and between years

Programs to introduce exotic species or mass produce indigenous ones are ambitious and expensive because they require trained people on a sustained not ad hoc basis

Chemical control Insecticides offer rapid and efficient control of upland insect pests (190) but-should be used only after other control measures have been considered Insecticides are rarely used on upland rice because of cost (32) Upland rice yields are normally too low to justify the expenditure Also most upland rice is grown as a cash crop Spraying upland rice is more difficult than lowland rice because water is less accessible Government subsidy programs to provide pestishycides to tribal peoples for upland rice production have created problems of toxicity to humans because isolated people have had no experience to be able to handle pesticides safely

However ins-eticide use has beenjustified when a crop is

0Z

Table 8 Parasitization rates of lepidopterous pests in upland rainfed lowland and irrigated lowland environments Philippines 1976-84a

Parasitization (5)

Environment and site Year YSB GFB PSB Earcutting Swarming Skipper Greenborned RLF Brown larva larva caterpillar caterpillar caterpillar caterpillar larva semilooperEgg Larva larva larva larva larva larva

UplandBatangas 1977 - 0 (57) - 0 (63) - 0 (71) - 0 (106) 11(142) 2 (101)1978 43 (4697) 8(451) 5 (38) 3 (40) 6 (61) 2(58) 15 (57) 13 (129) 13 (1286) 4 (30)1979 59(4135) 12(320) 13 (11) 5(28) 9(45) 6(46) 7(23) 8(160) 9(1152) 8(26)

Rainfed lowland Iloilo 1976 - 0 (10) 0(10) 0 (7) 0 (19) 2 13) 15 (180) 5 (76)1978 0 (54) 0 (3) 0 (17) 9 (64) 4 (78) 3 (73) 0 (121) 0 (191)1979 64 (2350) 16 (84) 6 (17) 0(6) 18 (68) 25 (104)

1981 56 (1322) 18(304) 0(5) 12(51) 18(138)Pangasinan 1976 60 (7 7 )b 62(82) 0(111) 5 (39)

1978 58 (53)b 5(51) Cagayan 1979 59(713) 8 (101) 6 (34) 0(7)

1980 0(11) 4 (28) 15 (286)

62 ( 48(177) 61(140)1981 0(5) 12 (2) 13(45) 14 (92)

Irrigatedlowland Laguna 1974 62 8

1974-75 50 41 1975 43 72 1978 96 (623) 21(24) 33 (60)1979 83(2017) 18(110) 8(62) 11 (89) 22(400)1980 78 (1625) 20(75) 0 (16) 4 (72)1983 60

9 (80) 7 (270) 30

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R F Apostol unpublished Iloilo and Pangasnan 136 Cagayan B Canapi unpublished Laguna 175 P CPantua unpublished 118 234 143 Carino and Shepard unpublished YSB = yellow stem borer ScirpophagaincertulasGFB = gold fringed borer Chilo auricilius PSB = pink stem borerSesamia inferens earcutting caterpillar Alythimna separata swarming caterpillar Spcdoptera mauritia acronyctoides skipper Pelopidas mathias reen horned caterpillar Melanitls ledaismene RLF = rice leaffoldersCnaphalocrocismedinalis and Marasmiaspp brown semilooper Mocisfrugalis Figures in parentheses are sample sizes Egg masses

IRPS No 123 January 1987 35

Table 9 Effect of seeding rate and insecticide on plant density and Yield losses from insect pests in upland rice often areyield of Dagge upland rice Batangas Philippines 1976a comparable to those of lowland rainfed or irrigated rice

Seeding rate Insecticide Dosage Plant density Yield (0-Y30o) but usually from different pests The principal (kg aiha) seed treatment (kg aiha) (noIra-row) (tha) groups are soil pests feeding on sown seed and rootsat 14 DE followed by seedling pests These groups have largely gone

50 None - 35 a 20 a unnoticed leading researchers to conclude that stem borers50 Carbofuran F 10 38 ab 23 a and seed bugs are the most important pests (17) Yield loss 10 beCabofuranF100 None - 4551 c 23 a trials that pinpoint major pests need to includc seed

100 Dieldrin WP 10 52 cd 22 a treatments Granules applied in the furrow may not give150 None - 59 d 22 a adequate protection from many seed pests

aAv of 8 fields DE = days after crop emergence In a column means The floristic and pedological diversity of upland rice followed by a common letter are not significantly different at the environments is matched by the diversity of arthropods5level by D)MRT F = flowable Wi = wettable powder both pests and their natural enemies Irrigation has homoshy

genized lowland rice environments Flooded soils tend tothreatened by epidemics of locusts or armyworms New have the same properties and to eliminate soil pests The technology stressing efficiency has minimized insecticide spatial and temporal dominance of cultivated rice has cost benefited specialist pests

Seed treatment is normally inexpensive and can be Upland rice has no specialist arthropod species The economicallyjustified in -nanycases(66) Treating seed with monophagous pests on upland rice prefer the lowlands systemic insecticide (216) but not chlorinated hydrocarbons where rice evolved Rice in the uplands exists only because (247) is most effective in controlling seedling maggots of man and even though lowland rice predates upland riceFoliar sprays have to be repeated to obtain control and specialist species evolved Wheat maize and sorghum do often are washed off by frequent early season rains not have monophagous pests leading us to conclude that Granular insecticides are effective but required dosages are there is no evolutionary advantage for inssects to have too expensive to justify except under heavy infestation narrow host ranges in upland graminaceae(247) When directed against first-instar larvae banding Pests have adapted to the uplands not only by having granules in seed furrows (177) can be effective against white wider plant host ranges but also by having long life cyclesgrubs at low dosages (025 kg aii ha) (134) Seed treatments undergoing dormancy or being dispersive Each pest has a are not promising for white grub control (248) unique life history strategy suited to highly unpredictable

Seed treatment is also effective ainst ants and may be environments In agivenyear someinsect specics an better cheaper than increasing the seeding rate In Batangas exploit upland rice as a food source but no pest is a senious treating 50 kg seed ha was as effective as increasing the one every year on upland rice seeding rate to 100 kgha and using untreated seeds (Table The low yielding potential of upland rice means less 9) In Brazil seed treatments control termites and other soil quantity and lower quality of food for pests to exploit but insects including the lessercornstalk borer(66) Domiciano also less incentive for farmers to undertake control (58) however found seed treatments inconsistent in their The most effective control would be early and synchroshyeffect against high populations of lesser corn stalk borer nous planting of early-maturing upland rice varieties to

Baiting can be an inexpensive way of controlling ants prevent insect pest buildup Breeding for insect resistance seedling maggot flies mole crickets and field crickets Baits should initially focus on stem borers because they are verymade of locally available material and impregnated with difficult to control by other means Biocontrol efforts insecticide can be sparingly distributed taking adantage of should first stress conservation of natural enemies preshythe pests ability to disperse and encounter bait sites dators and pathogens rather than parasites Efficient

The new low-volume hand-held controlled-droplet chemical control includes seed treatment or baiting for seed sprayers may offer an advantage over high-volume knap- seedling and some root pests and using controlled dropletsack sprayes for controlling foliar pests Upland rice sprayers for spot treatments against foliar pests and seed canopies are more open than those in lowland rice and bugs when economic thresholds are reached better droplet penetration should result Controlled droplet applicators would be ideal for seedbugs and defoliators

CONCLUSION

Upland rice is attacked by a wider array of insect pests than is lowland rice mainly beca use of the addition of soil pests Generally population levels of lowland rice pests are lower in upland than lowland rice leading many people to conclude that insect pests are not important on upland rice

36 IRPS No 123 January 1987

REFERENCES CITED

1Abraham C C and K S R Mony 1978 Occurrence ofLeptocorisa acuta Fabr (Coreidae Hemiptera) as a pest of nutmeg trees J Bombay Nat ttistSoc 74553

2 Agyen-Sampong M 1975 HlierogrlphusdiaganensisKrauss (Orthop-tera Acrididae) a new pest of rice in Northern Ghana Ghana J Agric Sci 8249-253

3 Agyen-Samnpong M 1976 Varietal resistance and other inle-ratcd pest control components as factors of rice varietal improvement in West Africa Prcsented at the Second varietal improvement serinar 14-18 Sep 1976 Bouak Ivory Coast West Africa Rice Development Association Monrovia Liberia 18 p

4 Agyen-Sampong M 1977 Outbreak of Ihccrorrdnisorae in rice seedlings near Rokupr Sierra Leone lit Rice Res Newsl 2(6)16-17

5 Agyen-Sampong M 1980 Dirty panicles and rice yield reduction caused by hugs IntRice Res Newsl 5(l)11-12

6 Agyen-Sampong M 1982 The major pest problems of irrigated upland and mangrove swanip rice ecosystems ir the humid tropical Guinea savanna and Salel climate zones Pages 318-328 in Pro-ceedings training course cn concepts techniques and application of irtegrated pest management in rice inWest Africa 10-20 Jan 1982 Fendall Liberia West Africa Rice l)evelopioent Association Monrovia Liberia 505 p

7 Akibo-Betts V I and S A Raymundo 1978 Aphids as rice pests in Sierra Leone lit Rice Res Newsl 3(6)15-16

8 Akinsola F A 1975 Present status of different rice stemborers in parts of Nigeria iicL Entomo Newsl 328

9 Akinsola FA 1979 Ihe biology and ecology of rice stemborers in Nigeria Ph 1) thesis University ol lbadan Nigeria 217 p

10 Akinsola E A 1979 Guidelines of integrated management of insect pests of upland rice in West Africa Presented at the Seminar on integrated management of rice diseases and insect pests at Bobo-Dioulasso Upper Volta 17-22 Sep 1979 West Africa Rice Develop-ment Association Monrovia Liberia

II Akinsola IA 1984 Insect pests of upland rice in Africa Pages 301-305 in An overview of upland rice research Proceedings of the 1982 lIouak Iwsrv Coast upland rice workshop International Rice Research Institute 13O Box 933 Manila Philippines

12 Alam M S V TJohn and Kaung Zan 1985 Insect pests and diseases of rice in Africa Pages 67-82 in Rice improvement in Eastern Central and Southern Africa proceedings of the international rice workshop at Iusaka Zambia 9-19 April 1984

13 Alticri M A A van Schoonhovcn and J D Doll 1977 The ecological role of weeds in insect pest management systems a review illustiated with bean (Phaseohsvulgaris L)cropping systems PANS (Pest Artie News Summ) 23195-205

14 Anmante F 1967 The leaf-cutting ant Atia capiguaraa pest of pastures [in Portuguese] Biol6gico (Sao Paulo) 33113-120 (Rev Appl Entomol A56207)

15Andrade A C and D Puzzi 1951 Preliminary results of experi-merits on the control of Sculptcoris iastantus on sugarcane [in Portuguese] Bioulgico (Sao Paulo) 1744-49 (Rev Appl Entomol A41421422)

16 Appert J 1970 Maliarplia separaelha (rice stem borer) new observations of entomological problems of rice in Madagascar [in French] Agron Trop (Paris) 25329-334

17 Ariadeau Mand Z Harahap 1986 Relevant upland rice breeding objectives Pages 189-197 in Progress in upland rice research Proceedings of the 1985 Jakarta conference International Rice Research Institute 1O Box 933 Manila Philippines

18Asar S and Kisimoto 1975 Seasonal occurrence of the smaller brown planthopper in upland rice [in Japanese] Proc Kanto-Tosan Plant Prot Soc 2228

19 Bakker W 1974 Characterization and ecological aspects of rice yellow mottle virus in Kenya Meded Landbouwhogesh Wage-ningen 152 p

20 Balut F F 1970 White grubs attacking rice Oryza ativa L [in Portuguese] Biol6gico (So Paulo) 36321-322

21 Banerjee S N and L M Pramanik 1967 The lepidopterous stalk borers of rice and their life cycles in the tropics pages 103-124 in The major insect pests ofthe rice plant Proceedings ofa symposium at the International Rice Research Institute September 1964 The Johns Hopkins Press Baltimore Maryland USA

22 Barrion A T and J A Litsinger 1980 Ants a natural enemy of cliaiarac in dryland rice IntRice Res Newsl

5(4)22-23 23 Barrion A T and J A litsinger 1981 The spider fauna of

Philippine rice agroecosystem 1Dryland Philipp Entomnol 5139-166

24 Barrion A T and 1 A Iitsinger 1984 The spider fauna of Philippine rice agroccosystcms II Welland Philipp Eiitomol 611-37

25 Barrion A T1 and J A Litsinger 1984 lihanus (Diptera Tabanidae) eggs an alternative host of rice stei borer (SI3) egg parasite Telenoonis digns (Iliymenoptcra Scelionidae) lit Rice Res Newvsl 9(6)19

26 Barrion A T and J A Litsinger 1985 Identilication of rice leaffolders by wing markings Int Rice Res Newsl 10( 1)24

27 Barrion A T and J A Litsinger 1985 Chctiosspp (Coleoptera Carabidae) a leaffolder ILF)predator lit Rice Res Newsl 10(1)21

28 Bertels A 1970 Insect pests of rice and theircontrol [in Portuguese] Pelotas Brazil Instituto de Expcrimentacao Agropccuararias do Sul Circular 43 24 p

29 Bonzi S M 1982 Ctilo (flisilineus J de Joannis (Lepidoptera Pyralidae) stem borer of irrigated cereals in Upper Volta LioFrench] Agron Trop Riu Rizic Cult Vivrieres Trop 37207-209

30 Bourke T V T L Fenner J N LStihick G Baker EHassan D F OSullivan and C S li 1977 Insect pest survey for the year ending 30th June 1969 Papta New Guinea )epartment of Agrishyculture Stock and Fisheries Port Moresby 57 p

31 Bowden 1 1973 Migration of pests in the tropics Meded Rijksfac Landbouwwet Gent 38785-796

32 Breniere J 1973 The main rice parasites in West Africa and their control Pages 27-42 in Proceedings seminar on plant protection for the rice crop West Africa Rice Development Association Monrovia Liberia

33 Breniere J 1976 The principal insect pests of rice in West Africa and their control West Africa Rice Development Association Monrovia Liberia 52 p

34 Brown ES 1962 The African arinyworn Spoiolptera eXI)Ina (Walker) (Lepidoptera Noctuidae) a review of the literature Cornshymonwealth Institute of Entomology London UK57 p

35 Brown F B 1968 Upland rice in Latin America FAO-International Rice Commission 12th Session of the Working Party on Rice Production and Protection 2-14 Sept Peradeniya Sri lanka Food and Agriculture Organization Rome Italy 6 p

36 Buyckx F J 1 1962 Disease and insect pests of crops inCongo Rwanda and Burundi [iil French] Publications dc LInstilut National pour lestude Agrononiquedu Congo M Weissenbruck S A Brussels Belgium 708 p

37 Cariiio F 0 1981 Role of natural enentics inpopulation suppression and pest management of green rice lealhoppers Ph D thesis University ofthe Philippines at Los Bafitos PhilippinesLaguna 136 -

38 Carpente A J 1983 The development of rice cultivation and extension of vegetable productin in Zanzibar Final Report Vol II1 October 1983 Food and Agricultme Organization Ronte Italy

39 Carroll C P and S1 Risch 1983 Tropical annual cropping systems ant ecology Environ Manage 751-57

4(0 Cates R G 198 1 Host plant predictability and the feeding patterns of monophagous oligophagous and polyphagous insect herbivores Oecologia (Berl) 48319-326

IRPS No 123 January 1987 37

41 Cendcafia S M and F B Caora 1967 Insect pests of rice in the 62 Ene 1 C 1981 I)iiorldhol)ierusihilis Slater (licteroplera Philippines Pages 591-616 in The najor insect pests ofthe rice plant lygacidue Illissinac) a new rice pest front Nigeria lit Rice Res Proceedings of a symposium at the International Rice Research Newsl N3)15 Institute September 1964 The Johns Hopkins Press Baltimore 63 Fugenio T S 1965 rhe effect oi e-vCgetation on rice production Maryland USA 729 p and the possibility of overcoming it by leaf pruning MS thesis

42 Centro Ihternacional de Agricklittira Tropical 1984 Upland rice in University ol the Philippines at Los Baijos ILaguna Ihilippins 69 p Iatin Anerica Pages 93-119 hi An os ervies of uplaind rice research 04 Farrow R A 1975 The Alrican nmigratory locust ii is main Proceedings of the 1982 Boinakamp Ivory Coast upland rice workshop outbreak area of the middle Niger qlinlitatise studies of solitary International Rice Research Institute P0 Blox 933 Manila populations in relation to environmental factors Locusla No II Philippines 198 p

43 Chandra G 1979 Taxonomy ad hn riononlics ofthe imect parasites of 65 Fennal R G 1943 Food crop investigation Windward and rice Icalloppers and planthoppers in the Pihilippines and their Ieeward Islands First report Report onl work dunring the periodiiiportairce iii natural biol)gical cointrol Philipp Fntoltol 4119-139 April 1942-April 1943 Imperial College of Tropical Agriculture

44 Chang T T 1976 The originii colut ion enlIivation dissemination Trinidad 16 p (Rev of Appl Fintornol A3222-25) and diersificition of Asian and African rices -uphytica 25425-44 1 66 Ferreira F 1983 Integrated pest ritaageiret of upland rice [ill

45 Chatierjee P B 1979 Rice leaffolder attacks in India lit Rice Res Portuguese English summary] Pages 323-341 in Upland riceculttre Newsl 4(3)21 factors affecting production E Ferreira T Yamada and F-

46 Cheng Cbien-pan 19(4 Upland rice in Taiwan International Rice Malmolta eds Institutoda Potassae Foslato Institnto Internacional Cnmmission Wiorking Party oit Rice Production ard Protection I()th da Potassa Piracicaha Sao Paulo lraid Meeting 3-10 March 1964 Manila Philippines Food and Agricul- 67 Ferreira E arid F dl Silva Martins 1979 Occurrence of ture Organiaion Rome Italy 5 p Sogatohs orvziola (Muir) inupland rice in Goias Ilrail lit Rice

47 Chcrrett 1 M and 1) 1Peregrine 1976 A review of the status of Rcs Ncwssl 4(4)18 leaf-cutting ants and their control Ann Appl Biol 84124-128 68 Ferreira E and 1F da Silva Martiris 1984 Insect pestof rice ill

48 Chipeta F M 1974 Pests of rice in Malawi Pages 90-95 in Brazil and their control Jill Portugucsej FMIlRAtPA-CNPAF Proceedings Sith eastern Africa cereals research conference 10-15 Goiania Brmiil 67 p March 1974 DRB Mialda ed Ministry of Agriculture lilongwe 69 Ferreira FI F ca Silha Miartins S S Nero and 1 It P de Malawi Carvallio 1982 M tltiplc regression analysis todeterimine the effect of

49 Chu Yau-i 1979 The d atiaging period of the rice seedling fly insect populatiois ott upland rice yield [inIPortuguese Pesqui (Atlterigotiavxigu Stcin) (NItiscide Diptera) oil cori) at Fast Java Agropecu Bras 17671-675 Indonesia Plant Prot Bill 21403413 70 Ferreira F 1F da Silva Martins and F 1PZimmermann 1979

50 Conklin H C 1957 IHaninoo agriculture a report on aln integral Rice varietal resistance to tle lesser corn stalk borer [in Port uguese] systent of shi fti rig cult ivationi in the Pltilippincs Food and Agriculture Pesqui Agropccu Biras 14317-321 Organization Rome Italy 209 p 71 Fleming W F 1968 Biological control of lie hapanese beetle U S

51 Cramer H II 1967 Plant proection and world crop production Dep Agric RLs Sers [ech Bull 1383 78 p Pilaniensclhutz-Nachr 201-524 72 Food and Agriculture Organiation of the United Nations 1982

52 l)ani R C and Majunidar 1978 Prelininary observations on tie Country report surmimary Pages 27-37 in Report on the first incidence of root aphids on differeit rice cultnrcs Sci Cult 4488-89 FAOIUN)IPhail and regional training coiuirse in iniproved ctltural

53 Davidson R I 1979 lffects of root feeding on foliage yield Pages practices for upland rice 5-23 Oct 1981 Northern Region Agrculturd117-120 in T K Ci osb and R B Pottinger eds Proceedings of the Developririt Center Chiang Mai Food and Agricuilture Organiiashy2nd Australian conlerence on grassland invertebrate ecology Govt tion Rome Italy 37 n Printer Wellington New Zealand 73 Franco do Ariaral S 195 1 Residual action of insecticides in turfy

54 l)ean G 1W 1976 Rice insect pests inlaos Int Rice Res Newsl soil--the results of a field experiment [in Portuguese] Biol6gico (Sao 1(2)15 Paulo) 17209-214 (Rev Appl Entomol A41423)

55 I)ean G J W 1978 Insect pests of rice in Laos PANS 24(3)280-289 74 Freeman ) 1970 Report onithe Iban london School of -coroiics 56 )ingle H 1984 Behavior genes and lifc histories conplex adapta- Monographs on Social Anthropology No 41 Athlane Press

tions to uncertain environments Pages 169-194 in A new ecology London UK 317 p novel approaches to interactive systems P W Price C N Slobod- 75 Fuchs T W1 A Harding and 1W Smithlr 1979 Induction and chikoff and W S Gaud cdsl ohn Wiley New York USA termination oldiapause in the sugarcane borer Ann Flitonlol Soc

57 Dobelmann-JP and M Falais 1968 Upland rice cultivation ill tie Am 72271-274 North-western part of Madagascar in 1966-1967 [in French English 76 (hbriel B1) 1975 A review tf the major insect pests of some tpland summary] Trop Agron 2313-21 crops in tlc Philippines Pages 86-111 in Ad Htoc panel of experts on

58 Doniiciano N L 1979 Chemical control of tie lesser corn stalk pest disease arnd weed problems in some rainfed crops Bangkok (lolsonno~paloIgnowslhts) (Zeller 1848) (I-epid(ptera Ihycitid ae) Thailand Food aid Agriculture Organization Roire Italy

on rice in Ponta Gressa Parara 1977-1978 [in Portuguese] Pages 77 Garg D K and N K Shal 1983 White grub outbreak on rainfed 291-303 in Anais 1 Reuniao de tecnicos cmiirizicultura do estado de dryland rice in Uttar Pradeslh Int Rice Res Ncwsl 81(4)16-17 Sao Paulo-Campinas Cati Brazil 78 Garg I) K and IP Tandon 1982 Major insect pests of rice on hilly

59 Dunsmore 1 R 1970 Investigations on tlie varieties pests and tracts of Uttar Pradesh India lilt Rice Res Newsl 7(11)11-12diseases of upland rice (hill padi) in Sarawak Malaysia Int Rice 79 Garrity 1) P 1983 Upland rice research an international bibliograpy Comm Newsl 1929-35 1975-1982 Iternational Rice Research hnstitnte PO Box 933

60 Elias 1R1965 The new decline of rice seedlings is not caused by Manila Philippines 121 p parasites [in Portuguese] Biol6gico (Sao Paulo) 313-6 80 Ghauri M S K 1982 A new subspecies of Iinorh)rone bull

61 Empresa Brasiliera de Pesquisa Agropecuaria 1984 Upland rice itt cornlus Slater (Hemiptera Iygaeidae) front Papua New Guinea on Brazil Pages 121-134 it Av overview of upland rice research rice and carpet grass Ihll lEntoniol Res 72133-137 Proceedings ofthe 1982 Blouak Ivory Coast Upland rice workshop 81 Gonzales F 0 PArregoces R I ihernandef and OT Parada International Rice Research Institute PO Box 933 Manila 1983 Insect and imite pests of rice arid their control in Latin America Philippines [in Spanish] la Federacion Nacional de Arrocerts (Fedearroz)

Bogota Colombia 60 n

38 IRPS No 123 January 1987

82 Grist D H and R J A W Lever 1969 Pests of rice Longmans London UK520 p

83 Hale P R 1979 Two subterranean pests of upland rice in Papua New Guirnca IntRice Res Newsl 4(2)16-17

84 Hale P R and J T Hale 1975 Insect pests of rice in the New Guinea Islands Rice Entomol Newsl 34

85 Harris W V 1969 Termites as pests of crops and trees Common-wealth Institute of Entomology London UK 41 p

86 Hattori 1 1968 A method to distinguish between grass leaf roller (Ctaplha rocis medinalisGuenee) and the rice leaf roller (Susurtia exigua Butler) [in Japanese] Plant Prot (Japan) 23167-170

87 Henry I E and 1A Onsager 1984 Experimental control of the mormon cricket Iahrosshnph by NJsena locustae(Aficrospora Microsporidae) a protozoan parasite of grasshoppers (Ort Acri-didae) Entomophaga 27197-201

88 Hidaka 1 P Vungsilabulr and SKadkao 1974 Studies on ecology and control of the rice gall midge in Thailand Tech Bull 6 TropicalAgricultural Research Center Tokyo Japan 113 p

89 Hincklcy A 1) 1963 Ecology and control of rice planthoppers in Fiji Bull Entomrol Res 53467 481

90 Hirao J 1982 Ecology and chtInical control of the rice leaffolder Jpn Pestic Inf 4114-17

91 Ho DT 1984 Sten borer incifience in rice ecosystems in KenyaEast Africa Pages 307-320 in A overview of upland rice research Proceedings of the 1982 Bouac Ivory Coast upland rice workshop International Rice Research Institute PO Box 933 Manila Philippines

92 Ho )T J 1Njokah and J GKibuka 1983 Studies on rice sten borers in Kenya with emphasis on ifaliarpiaipiraellRag Insect Sci Appl 465-73

93 Ichihara I T Ito A NIatsuo and M Sawada 1971 Chemical control of the rice striped stem borer in upland rice grown in plastic mulch [iii Japanese] Proc Kanto-Tosan Plant Prot Soc 1882

94 IclhiharaIand A Matsuo 1969 Damage caused by rice striped stett borer to upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1686-87

95 Ichhara I atid A MIatsun11970 Conside ration oft lie extraordinary prevalence of rice striped stem borer in upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1778-79

96 Institut de Recherches Agrolnliqes Iropicale et des Cultures Vivrieres 1978 Insect resistance Pages 12-13 in Upland rice 1978 atnual report Bouak Ivory Coast 27 p

97 International Institute for Tropical Agriculture 1983 Aphids itt rice roots I ITA Res Briefs 4(1-3)4-5

98 International Institite fi Tropical Agriculture 1983 Hopperburn in rice caused by brown plathopper IITA Res Briefs 4(l-3)6-7

99 International Rice Research Institute 1975 Annual report for 1974 PO Box 933 Manila Philippines p 216

100 International Rice Restarch Institute 1976 Annual repiort for 1975 PO Box 933 Manila Philippines p 228

101 International Rice Research Institute 1979 Cropping systems program Component technology development and evaluation Page 404 i Annual report for 1978 11O Box 933 Manila Philippines

102 International Rice Research Institute Department of Entomology 1981) Insecticid evaluation 1979 Tables 73 74 PO Box 933Manila Philipes

103 International Rice Research Institute Department of Entonmology 1981 Insecticide evalomtion 1980 Table 81 POBox 933 Manila Philippines

104 International Rice Research Institute 1981 Report of a meeting of Asian rice-based cropping systems entomologists Eleventh cropping systems working group meeting 18-22 May 1981 PO Box 933 Manila Philippines p 132 133 144

105 International Rice Research Institute 1984 Cropping systems program Pages 368-373 in Annual report for 1983 PO Box 933 Manila Philippines

106 International Rice Research Institutc 1984 Upland rice in Asia Pages 45-68 in An overview of upland rice research Proceedings of the 1982 Bouakamp Ivory Coast upland rice workiiop loternational Rice Research Institute 11O Box 933 Manila Philippines

107 Irshad M 1977 Grasshoppers in rice in Pakistan lit Rice Res Newsl 2(3)8-9

108 Ito R and TOno 1964 Upland lice its culture and varieties in Japan lIternational Rice Cormmission loth Meeting of the Working Party on Rice Production and Protection 3-10 March 1964 Manila Philippines Food and Agriculture Organization Rome Italy 12 p

109 Jeffrey M I and V A Dyck 1983 Effect of moonlight on lightshytrap catches of brown planthopper lit Rice Res Newsl 8(6) 13

110 Jennings P R and T Alicia Pineda 1983 Management ofthe rice plnthoppers Sogatodsorivi(coIl in Latin America Presented at the meeting on stable plant resistance FAO and lIndocsian I)inectorate of Food Crops Protection Bali lndonesia 29 Novcniber-2 l)ccemshyber 9 p

IllJimenez Gomez J A 1979 Rice insect pests i lla ios ()riCntales lil Spanish] Pages 99-103 in Rice ciltivation Instituto (olotibiatto Agropecuario Compndio No 29

112 Jongeleen F JJ Soesanto and A Mockidjo 1979 Studies oti pathogens of two species of Scarabacidac froinIndonesia Z Angew Enlomnol 87412-417

113 Jouvenaz 1) PC S lofgren and W A Baiks 1981 Biological control of imported fire antsa review of cirrent knowledge Bull Entomol Soc Am 27203-208

114 Kainoh Y K Shiniiu S MarUani Y Taiaki 1980 lost finding behavior of tie rice bu lplotori+JchinvittjI)allas (liciniptera Corcidac) with special reference to diet pattern ol aggregation and feeding on the rice plant Appl Entomol Zool 15225-233

115 Kakkar K 0 P Sharma and P I Sharma 1977 Note on Ihteronyrchus liodresRedt (Coleopicra I)ynastidae) as asporadic pest of paddy and maie inHimachal Pradesh Corr Agric 156-57

116 Kalode M131 79 Major insect pest problems and their control in upland rice (Unpublishcd mineogriph CWI oJNaaiat lni Chandaratat (198 1)original not seen)

117 Kalshoven L i 98 I The pests of crops in I ldoniesia (translated by P A vatn der Laan) 1 T Ichtiar B3aru-vati Iloeve lakarta Indonesia 701 p

118 Kamal N Q 1981 Suppression of whitebacked planto ner Sogatella fircaifra (Horvath) aLId rice lealfolder (nhal -cis licdb lis (Guenee) populations by natural cnclmie hlI) tiesis Grcgorio Araneta University Foundation Victoncta Park Metto Manila Philippines 179 Ip

119 Karim A N M Reaul 1975 A species ofliealy bug Trioni i70s ceres new oii rice in the Philippines Rice Entoniol Newsl 247

120 Katanyukul Wand C Chandaratat 1981 Insect pestsof uplaid rice and their control Report firstFAOUNlFi Tliailand training course on improved cultural practices for tplad rice Chiang Mai Thailand 5-23 Oct 1981 Food and Agriculture Organiation Rome Italy 22 p

121 Kern F 1955 lhletwioa(I)uninemts) bhhit Biturm(Coleoptera) a new pest of sugarcane inVenezuela [inSpanish with Etnglish sunmary] Agron Trop (Maracay) 527-29 (Rev Appl Eitoniol A4572)

122 Khan K M and S Ghai 1974 White grubs ald their control in India Pesticides (1Bombay) 819-25

123 Kiritani K 1975 Integrated pest management of rice pests Pages 66-70 itt Proceedings platning workshop ott cooperative hIeld research in pest managementP SMotooka ed East-West Center Honololu Hawaii USA

124 Kisimoto R 1967 Ecology of insect vectors forecasting and chemical control Pages 243-255 in The virus diseases of tle rice plant Proceedings of a symposium atthe International Rice Research Institute April 1967 The Johns Hopkins Press altimore Maryshyland USA

125 Kisimoto R and V A Dyck 1976 Climate and rice insects Pages 367-391 in Climate and rice International Rice Research Institute PO Box 933 Manila Philippines

126 Kureha S S Kobayashi SAsari and T Ishihara 1977 Life history and control of the mole cricket Grrllotalpa africana Palisot de Beauvois [in Japanese English translation 7520 1RRI library] Proc Kanto-Tosan Plant Prot Soc 21192-197

127 Kushwaha K S 1981 The black beetle as a pest of rice in Haryana India Int Rice Res Newsl 6(4)14

128 Lawani S M 1982 A review of tile effects of various agronomic practices on cereal stem borers Trop Pest Manage 28266-276

129 Leuck D B 1966 Biology of the lesser cornstalk borer in South Georgia J Econ Entomol 59 797-801

130 lever R J A W 1955 Insect pests of the rice plant in Malaya Int Rice Comm Newsl 1612-21

131 Li Ching-Sing and Shin-Foon Chiu 1951 The study ofthe rice gall midge Iachydiplosis orvza Wood Mason [in Chinese with English summary] 1Taiwan Agric Res 21-13

132 Iitsinger J A 1984 Assesament of the need-based application for rice Proceedings TechnologyTransfer Workshop 15-16 March 1984 International Rice Research Institute 1PO Box 933 Manila Philippines 6 p

133 Litsinger J A 1984 Environmental diversity and insect pest abundance with reference to the Pacific Pages 285-307 in Proceedings sub-regional training course on methods of controlling diseases insects and other pests of plants in the South Pacific 4-20 Oct 1982 Vaini Kingdom of Tonga Nukualofa

134 Litsinger J A RF Apostol and M BObusan 1983 White grub Leucopholis irrorata (Coleoptera Scarabaeidae) pest status popula-tion dynamics and chemical conllol in arice-maize cropping pattern in the Philippines J Econ Entoinol 761133-1138

135 Litsinger J A C de la Cruz F Raymundo A T Barrion M D Lumaban J P Bandong M S Venugopal F Paragna and E Balete 1982 Insect pests and insecticide reponse of dry-seeded bunded rice Pages 179-193 in Report of workshop on cropping systems research in Asia International Rice Research Institute Los Bafos Philippine

136 Litsinger J A M 1) Luinaban J P Bandong A T Barrion W Katanyukul and N Panda 1978 Insect pest management in rainfcd lowland rice agroecosystms of tropical Asia International Rice Research Conference 17-21 April 1978 International Rice Research Institute Los Bafios Philippines

137 Litsinger J A and K Moody 1976 Integrated pest management in multiple cropping systems Pages 293-317 in R I Papendick P A Sanchez and G B Triplett eds Multiple cropping American Society of Agronomy Madison Wisconsin USA

138 Loevinsohn M E 1984 The ecology and control of rice pests in relation to the intensity and synchrony of cultivation Ph D thesis University of London 36) p

139 Loevinsohn M E J A Litsinger N Panda M A Austria R F Apostol C Andrion and J Bandong 1982 A basis for predicting rice insect pest abundance in cropping systems Paper presented at the 1982 International rice research conference on rice and cropping systems 19-23 April 1982 International Rice Research Institute Los Bafios Iaguna Philippines 25 p

140 Lopez A W 1930 Repot of the Entomology Department Pages 145-172 in Annual report of the Philippine Sugar Association 1929-30 Manila

141 Lopez AW 1931 Entomology Department Pages 227-266 in Annual report of the director or research 1930-31 Philippine Sugar Association Manila

142 Lopez A W 1932 Entomology Deparment Pages 252-279 in Annual report of the Philippine Sugar Association 1931-32 Manila

143 Lungibill P and G G Ainslee 1917 The lessercornstalk borer US Dep Agric Agric Res Serv Tech Bull 539 27 p

IRPS No 123 January 1987 39

144 McGuire J U Jr and BS Crandall 1967 Survey of insect pests and plant diseases ofselected food crops of Mexico Central America and Panama U SDepartment of Agriculture International Agriculshytural Development Service Agricultuial Researrh Service October 1967 11 p

145 McNeill Sand T R ESouthwood 1978 The role of nitrogen in the development of insectplant relations Pages 77-9 in lliochemical aspects of plant and animal coevolution J B Ilarbornu ed Academic Press London UK

146 Maebara H 1955 Spatial distribution of lastinstiarlanaeof purplish stem borer (Sesamia iqferens Walker) hibernating in an upland rice field Bull Kagoshima Univ Fac Agric 438-41

147 Mallick E H and N G Hajra 1977 Response of rice plant to defoliation treatments at different developmental stages Indian Agric 2151-56

148 Mansingh A 1971 Physiological classification of dormancies in insects Canl Entomol 103983-10)9

149 Martell C G 1974 Catalogue of plant feeding insects of Mexico [in Spanish] Fitofilo 27(69)1-176

150 Martinez 0 J 1982 Principal rice pests in Ilanos orientalis [in Spanish] P~ages 80-94 in Curso deact nalization de conocimientos en arroz Villaniencis Instituto (olonmbiano Agropecuario

151 Martins J F da Silva F Ferrcira A S Prabhu F 1 P Zimmermann 1980 Use of pesticides for the control of the main subterranean upland rice pests esqui Agropecu Bras 1553-u

152 Martins I F da Silva F Fe eira rod BdamSilveira Pinheiro I Q2 Damage simulation caused by defoliators of upland rice [in Portuguese with English summary] Pesqui Agropecu Bras 171113-1119

153 Martins 1F da Silva N(iyuen van Tan and 1Ida Silveira Pinheiro 1981 Stem borer resistance in upland rice associated with plant morphological characters [in Portuuce Iesqui Agropecu Bras 16187-192

154 Masaki S 1980 Sumnmer diapause Ann Rev Entomiol 251-25 155 Masuzawa T 11 Suwa and F Nakasoji 1983 l)ifkleences of

oviposition preferencc and survival rate of two skipper butterflies Parnara guttata and PIchlpidai iathiais (Iepidoptera HIesperiidac) on rice plant and cogon grass New Entomol 3247-56

156 Maurya D M and C P Vaish 1984 Upland rice in India Pages 69-92 in An overview oftpland rice research Proceedings ofthe 1982 Bouakamp Ivory Coast upland rice workshop International Rice Research Institute IO Box 933 Mnila Philippines

157 Menon M G R R N Katiyai S Kuar and S S Misra 1970 Termites attacking the paddy crop Sci Cult 36294-295

158 Miyahara Y and 1 Nishikubo 1967 Behavior of young nymphs of the red rice root aphid Iholahsipiun ri liabhdontinalisSasaki in relation to movement from the leaf to the roots of upland rice [in Japanese] Pioc Assoc Plant Prot Kyushu 1363-66

159 Mo ljoa Tjien 1952 White grub of upland rice (Holotrichiahelleri Brsk) [in Indonesian with English summary] Contrib Agric Res Stn Bogor No 13134 p

160 Mochida 0 and T Suryana 1979 The occurrence of the brown planthopper and ragged stunt on lowland and upland rice ill Central Java and logyakarta in tile late of January 1979 Entomnolgy Survey Report No 15 Central Research Institute of Agriculture (CRIA) Sukamandi Indonesia I February 1979 9 p

161Monet C 1979 Varietal resistance to upland rice pests [inI French] Pages 35-39 in Annual report 1977 Institut des Savanes Department des Cultures Vivrieres Bouak6 Ivory Coast

162 Monte 0 1942 A caterpillar ofrice-fields[in Portuguese] Biol6gico (Sao Paulo) 8161-163 (Rev Appl Entomol A31414)

163 Munroe I) D 1979 Pest management strategy for hill rice in four resettlement areas in Sarawak Malaysia Pages 89-102 i1 Guidelines for integrated control of rice insect pests Food and Agriculture Organization Plant Production and Protection Paper 14 Rome Italy 115 p

40 IRPS No 123 January 1987

164 Nakasuji F M Ishii and Jen-Zon Ito 1984 Rice skippers in Taiwan and their life histories ZZrC (Taipei) Tech Bull 82 10 p

165 Navas I) 1976 Fall armyworum in rice Pages 99-106 il Ecological animal control by habitat management Proc Tall Timbers Con-ference 28 Feb-I Mar 1974 Fall Timbers Res Stn Misc Publ 6 Tallahassee Horida USA

166 Nickel 1 1 1961 Biological control of rice stem borer A feasibility study Tcch Bull 2 International Rice Rcarch Institute PO Box 933 Manila Philippines IIl p

167 Nigan P M and R A Vernia 1985 Insect pests of uplaild rice in Uttar Pradesh Int Rice Res Newsl 10(6)22

168 Nilakhe S S 1985 -cological observations on spittle bugs with emphasis on their occirence in rice Pcsqtui Agropecu Bras 20407-414

169 Nixon G EJ 195 1 lie association of ats with aphids and coccids Commonwealth Iusti lcte of Entonology Iondon UK 36 p

170 Nurdin F an Ishak Manti 1985 Study oit seedling fly of upland rice Interlational Upland Rice Moitoring Tour Report 26 lebruary-2 MarLI Sukaranti West Sumatra

171 Oka 1 N 1979 Ciltural conttol of the brown planthopper Pages 357-369 in Brown planthopper threat to rice production in Asia International Rice Research Institutc 1p0 Box 933 Manila Philippines

172 Ohni M and A Vilnaii 1975 Rice insect pests of Italy biology of Sipha grcria (Kaltenbach) and its control [in Italian] Riso (Milan) 2459-71

173 Otanes F Q atnd 1 I Sison 194 1 Pests of rice Philipp J Agric 12211-259

174 Painter R I 1951 Insect icsistance in crop plants University Press of Kansits Lawrence Kansas USA 520 p

175 Pantua P C and 1A litsinger 1981 Arthropod abundance in continuous vs biannual rice cropping systeis Int Rice Res Ncwsl 6(2)20-21

176 Pathak M ) and V A l)yck 1975 Sttudics ott insect pests of upland rice Pages 186-191 in Major research in upland rice International Rice Research Institute PO Box 933 Ilanila Philippines

177 Patil B R B M IHasabe and 1) S Ajri 198 I Chemical control of white gritb lIditriihinhorraia Fabr intfesting groundntut paddy atnd sorghum Indian 1Entomol 43232-236

178 Pawar A 1) 1974 (ollection and identification of insect pests of rice rlerminal repirt lInternational Rice Research Institute PO Box 933 Manila Philippines 72 p (tnimco)

179 IerrajU A aid A S Reedy 1965 Observations ott diapausc itt tie steti borer ohrice Rice News Teller (lai-Mar)9-13

180 Pont A C 1972 A review of the Oriental species of theri ona Rondani (I)iptera Muscidac) of econtomic importance Pages 27-102 in Control of sorglium shoot fly M G lomtani and W R Young eds Oxford atd IH1 Ptthl Co New I)clhi India 324 p

181 Prasad S C 1PChand aid M 1PSingh 1983 Incidence of rice gall midge in the upland plateau region of Hihar Oryza 20235-236

182 Rai P S 198 0 Nisaga simpex damage to rice in the hill tracts of South India Int Rice kles News[ 5(I)12

183 Rathorc Y S 1969 Gonocplhahli dcprssunt Fabr as itpredator of (tilo zonwIlhis (Swinhoc) and other moths in field cage at Patnagar Uttar Pradesh Indian 1Entotnol 3127

184 Reed W 1983 Crop losses causcd by insect pists in the developing world Proceedings 10th Internatial Congress of Plant Protection lrighton UK 174-80

185 Reedy M V 1981 Ateactogasterinhens(Walker)a new root-grub pest of tipland paddy and inaize Curr Sci 501076

186 Reveno M A 1968 Catalogue of the insect pests of economic crops in Colombia [in Spaiish] Asociacion Latino Americana de Entomo-logia (AIAE) 129-33

187 Reyes G M 1957 Rice dwarf disease in the Philippines Food and Agriculture Organiiation Plant Prot Bull 617-19

188 Rice S E A A Grigarick and M 0 Way 1982 Effect of leaf and panicle feeding byarmyworm (Lepidoptera Noctuidae) larvae on rice grain yield Environ Entomol 7593-595

189 Risch S J 1) Andow and M A Altieri 1983 Agioccosystem diversity arnd pest control data tentative conclusions and nizw research dirctions Environ Entomol 12625-629

190 Rivera C T 1956 Three organic insecticides in thecontrol of insects affecting upland rice Philipp Agric 39465-472

191 Roffrey 1 1972 Locusta outbreaks in the Philippines Acrida 1177-188

192 Rojas R 1) 1941 The effect of pruning at varying ages upon the yield of Ramairio Phipp Agric 29851-860

193 Roonwal M i 1979 Ternite life and termite control in tropical South Asia Scientilic Pnbl odhpr India 139 p

194 Rosetto C1 F da Silva Martins N C Schmidt and L EAzzini 1978 Damage caused by pasturespittle bugs 4Dvoi)lavopictaandDJ schach) on rice lBragantia Nota 6 (Matcli)25-27

195 Rosztto C1- S S Neto I- Amantc I) Iink 1Garcia-Vieira D M de SoLza N V Bansa t and A M de Olivcira 1971 Rice pests of Brazil [in Portuguese] Pages 149-238 inrFA()-International Rice Commission 2nd meeting of the Rice Committee for the Americas 6-11 l)ecember 1971 Pelotas Rio Grande do Sul Brazil

196 Rothschild G H L 1969 Obseratiois on the armyworm Spodoptera matritia acronrctoiiles Gn (Lep Noctuidae) Bull Entomol Res 59143-160

197 Rothschild G 1- L 1974 Parasites of the armworm Spolopiera mauritiaacroncitoihes in Malaysia Fntornopliaga 19293-299

198 Sachan i N S K Gangwar and 1N Katujar 1980 Field crickets damaging cages in North Eastern IHill Region Indian J Entomol 42526-527

199 Sands 1) P A 1977 The biology and ecology of Lepocorisa (Hemiptera Alydidac) in Papua New Guinea Res Bull 18 Departshyment of Primary Industry Port Moresby Papua New Guinea 104 p

200 Sarawak 1968 titomology division Pages 97-106 in Report of the research branch of the Department of Altriculture 1966 Kuching (Rev AIppl Eintomol A56603)

201 Satpathy JM M S Das and K Naik 1977 Effect of multiple and mixed cropping on the incidence of scine important pestsI Entomol Res 178-85

202 Sauer IHF G 1939 Notes Ott Elasmpnlpualtsligniosellths Zeller (Lep Pyr) a serious pest ofcereals in Sao Paolo [in Portuguese] Arq Inst Biol 10199-206 (Rev Appl Entotnol A28378)

203 Satinders 1L A 1 SKing and C L Vargas S 1983 Crop pests of Central Anerica with t bibliography [in Spanish] Centro Agroshynotnico Tropical de Investigacion y Ensenanza Turrialba Costa Rica (CATIE) Tec Bol 9 90 p

204 Sauphanor B 1985 Some factors of upland rice tolerttce to stem borers in West Africa Inseu Sci Appl 6429434

205 Sen P atnd S Chakravorty 1976 Stentflics in Haritighata Nadia West Bengal India [it Rice Res Newsl 1(12) 15

206 Senapai B and 1M Satpathy 1983 The seedling fly Atherigona orrzae Malloch is a pest of rice in Orissa (India) J Entomol Res 777-79

207 Shhjahan M S Alatn M Ah nad and BN Islam 1983 A new scarabaeid beetle pest of dryland rice in Bangladesh Int Rice Res Ncwsl 8(I)16-17

208 Sharma G L S C Modgol and R C Gautam 1979 Effect of depth of submergence ott incidence of bacterial blight and white grub infestation in transplanted rice Int Rice Res Newsl 4(6)12-13

209 Sharma S 1985 Scarabaeid beetle outbreak itt upland rice Int Rice Res Newsl 1(1(l)24

210 Shiraki T 1917 The paddy borer Schoenobhits incertulas Walker [achaku Agric Exp Stn Formosa 256 p

211 Silva A de Brito and 11P Magalhaes 1981 Insect pests of rice [in Portuguese] Circular Tecnica No 22 EMBRAPA Centro de Pesquisa t ecuaria de Tropico Unido (CPAJU) Belem Brazil 14 p

212 Singh S A 1979 Emergence of rice leaf butterfly as a rice pest in Manipur India int Rice Res Newsl 4(4)16-17

213 SinghS A 1983 Changing trends of insect pestsof rice in Manipur Int Rice Res Newsl 8(f)12

214 Siwi S S and P H van Doesburg 1984 Lenovorisa Latreille in Indonesia (Heteroptera Coreidae Alydinae) Zool Meded (Leiden) 58117-129

215 Soejitno J 1977 Reaction of some upland rice varieties toseedling fly eltherikonaexigua Stein [in Indonesian] Siri Hana I Penyakit 630-35 Lembga I osat Fenclilian Pertanian Bogor Indonesia

216 Socnardi 1967 Insect pests of rice in Indonesia Pages675-683 in The major inse pests of thu rice plant Proceedings of a symposium at the Internatital Rice Rec Wi itt S pternber 1964 The Johns Hopkins Press Baltimore Maryland USA

217 Soto P E A T Petcz and I W Buddenhagen 1976 Survey of insect pests and diseases of rice in different ecological zones in Nigeria Rice Entomol Ncwsl 435-46

218 Soto 11E and Z Ziddiqi 1976 Screening for resistance to African rice insect pests Varietal improsetnent sCmi n r Bouak Ivory Coast 13-17 Sept 1976 West Africa Rice l)evelopment Association Monrovia Iiberia

219 Squire F A 1972 Entomological problems in Bolivia PANS (PestArtic News Sumr) 18249-268

220 Stapely 1 II 1978 The rice leaf roller in the Solomon Islands Int Rice Res Newsl 3(l)13

221 Suenaga H 1937 On aphids attacking the roots of rice Oyo-I)oubts Zasshi 9150-152 [in Japanese] (Rev Appl Entomol A25745)

222 Suenaga H 1952 On the seasonal abundance of three subterranean aphids on upland rice Bull Kyushu Agric Exp Sti 1249-262

223 Sutynso Yas and Dandi Soekaria 1964 Beberapa hana yang ttenperang tana man padi tanah kering [in Indonesian] Rapat kerja Padi Tanah Kering 6 sd 8 Juily 1964 Kaliurang 9 p

224 Tallamy I) W and R F Denno 1983 Alternative life history patterns in risky environments in example from lace bugs Pages 129-147 in Insect life history patterns R F l)cnio and If Dingle eds Spring-Verlag 225 p

225 Tamura 1 1939 Observations on some insect pests of field crops [in Japanese] 1 Plant Prot 26414-420 (Rev Appl EntomolA27-556)

226 Tamura and Suzuki 1964 Influences of the early-transplanting practice in rice culture upon development of the rice stem borer Chilo siql)ressalisWalker Bull Ilokuriku Natil Agric Exp Stn 761-94

227 Tan J P 1924 The rice root aphid Drowhi ihirsoia Philipp Agric 13277-288

228 Tanaka 1 1957 Taxonomy and distribution of some subterranean aphids injurious to upland rice in Japan with description of a new species [in Japanese English summary] Botyu-Kagaku 22168-176

229 Tanaka T 1961 The rice root aphids their ecology and control Spec [full College of Agriculture Utsunomiya University Japan F3 p

230 Tauber M IC A Tauber R Nechols and 1Obrycki 1983 Seasonal activity of parasitoids control by external internal and genetic factors Pages 87-108 itt I)iapause and life cycle strategies in insects V K Brown and I Ilodek eds )r W Junk The Hague Netherlands

231 Taylor W F 1972 Effects of artificial defoliation (simulating pest damage) on varieties of upland rice Exp Agric 879-83

232 feotia T P S and S Natud 1966 Biotoomics of the rice skipper Parnaraniathiis Fabricius (Lepidoptera Hesperiidac) Indian J Entomol 28181-186

233 Teran F 0 1971 Rice pests of Santa Cruz [in Spanish Ministerio de Agricultura Sita Cruz Bolivia Bull 45 8 p

234 Than Itun 1976 Population dynamics of the yellow rice borer Tryporza itcertulas (Walker) and its damage to the rice plant MS thesis University of the Philippines at Ios Bafios Laguna Philippines 92 p

235 Uichanco L B 1911 Pests Pages 55-74 it Rice Philipp Agric 11-200

IRPS No 123 January 1987 41

236 van der Laan P A 195 1 Life history of the rice seedling fly Contrib Gen Agric Res Stn Boor Indonesia 118 15 p

237 van der Laan 11A 1959 Correlation between rainfall in the dry season and the occurrence of the white rice borer (Scirpolhaga innotala WIk) in lava Entomol Fxp Appl 212-20

238 Van Driesche R G 1983 Meaning of percent parasitismin studies of insect parasitism Fnviron Entomol 121 I1- 122

239 Venugopal M S and A litsingcr 1984 Fffect ofcarbofuran on rice growth Prot Ecol 7313-317

240 Vietnam Directorate of Rural Affairs 1962 Annual work progress report on crop improvement program of rice sugarcane vegetable and field crops (lor the period from July 1961 to June 1962) (11) 299 p (Rev Appl Entomol A51526-527)

241 Wan M T K 1972 Observations on rice leaf- and planthoppers in Sarawak (Malaysian Borneo) Malay Agric 148308-335

242 West Africa Rice I)esclopnient Association 1978 Annual research report Vol 2 Statistics Monrovia Liberia

243 West Africa Rice Development Association 1979 Annual research report p47 Monrovia Liberia

244 Williams ) F and CS lofgren 1981 Fli I illy l-468 anew bait toxicant for control of the red imported lire at Fla Entomol 64472477

245 Williams I) 1 1970 The mealybugs OFlomoptera Coccoidea Iscudococcidac) of sugar-cane rice and sorghum Bull Entomol Res 0109-188

246 Williams D) 1 1982 Pulinaria icervi (Signoret) and its allies ott sugar cane and other grasses Btll Entoinol Res 72111-117

247 Wirjosuiardjo S 1976 Study ofchenmical cont rot of insect pests ott upland rice Imiu Pertanian (Jogjakarta) 1323-329

248 Wirjosuhardjo S and N Wirarta 1976 Chemical control of the white grub tIolorithiatlltriin upland rice in M uneng Irobolinggo Indonesia) [in Indonesian] Imut Pertanian (Jogjakarta) 1330-332

249 Yano K T Miyake and V F Eastop 1983 The biology and economic importance of rice aphids (Henuiptera Aphid idae) a review Bull Entomol Res 73539-566

250 Yelp D and P Rhodes 19(4 lhe success of drilled dryland rice iii the Fiji Islands International Rice Commission Working Party on Rice Production and Protection Manila Philippines 3-10 March1964 Food and Agriculture Organization Rome Italy 13 p

251 Young W R 1981 Fifty-five years of research on the sorghum shootfly Insect Sci Appi 23-9

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ISSN 05-3862Other papers in this series TITLES OF NUMBERS 1-45 ARE LISTED ON THE LAST PAGE OF NO46 THOSE OF NUMBERS 46-70 ARE ON THE LAST PAGE OF NO 71-80AND THOSE OF NUMBERS 71-1O0ARE ON

THE LAST PAGE OF NO 101-122

No 101The economics of hybrid rice production in (hina No 113 Boron toxicity in rice No 102 Rice ratooning No 114 Energy analysis rice production systems and rice research No 103 Growth and development of the (Jeep water rice plant No 115 Production risk and optimal fertilizer rates an application of the

No 104 Faridpur acomputer-assisted instruction nmodel forrainfed lowland random coefficient model No 116 Consumer demand for rice grain quality in Thailand Indonesia andrice

No 105 A readingand listening comprehcnsion test inEnglish for non native the Philippines speakers applying for tiraining at IRIRI No 117 Morphological changes in rice panicle development a review of

No 106 Rice grassy stunt virus 2 a new strain of rice grassy stunt in tire literature

Philippines No 118 IRRI-Korca collaborative project for the development of cold-No 107 Physical losses and quality deterioration in rice plstprodnCtion tolerant lines through anthei culture

systemis No 119 Problen soils as potenial areas for adverse soils-tolerant rice No 108 (olptblication ofIRRI materials a survey of translators and varieties in South and Ssuthcasi Asia

publishers No 120 Changes isinall-farm iLtireliing techii)hgy in Thailand and [he No 109 Classification of Philippine rainfall patterns Philippines No 110 Contributions of modern rice varieties to rutrition in Asia No 121 Landforms and modern rice varieties No Ill Changes in rice breeding in 10 Asian countries 1965-84 No 122 Yield stability and modern rice technology No 112 Design parameters affecting the performance of the IRRI-designed

axial-flow pump

Page 10: UPLANDRICE. INSECTl--1i M T -E!R.EOLOU t RTANCE ,AND …delphacid.s3.amazonaws.com › 6958.pdf · The significance of abiotic factors in upland rice insect ecology will be apparent

10 IRPS No 123 January 1987

Table I continued Order Family

Diopsidae

Agronyzidae

lHymenoptera Formicidae

Species

Atherigona oritentalisSchiner Atherigona indica MallochDiopsis longicornis(= thoracica)Macqurt Diopsis spp Pseudonapnyza asiatica Spencer Pseudonapomyza spicata (Malloch) Agronoza orvzae MunakataAcromyren landoltibalzani(Emery) Acromyrmex landoltifracticornis(Forel)Acromyrnex heyeri Forel)Atta bisphaericaFore Atta capiguaraGoncalves Atta laeLigata (F Smi h1) Atta opacicepsBorgmeierSolenopsisgeminata (abricius)Pheidole sp Pheidolegeton diversus Jerdon Monomorium pharaonis (Linnaeus) Ants unspecified

Distribution Reference

Asia 82 Asia Africa 82205Africa 11 1248 Africa 217 Asia 213 Asia h Asia bLatin America 6168 195 Latin America 61 195 Latin America 61 68 195 Latin America 61 68 195 Latin America 14 61 68 195Latin America 61 68 195 Latin America 61 195 Asia b Asia b Asia 1) Asia b Asia 120

aG orlentalls formerly recorded in Asia as G africana thPhilippines reported in this study CSubterranean root feeders

subterranean nests Foraging ants will feed on germinatedseeds only when the supply of ungerminated seed is low However these ant specialists feed on a wide varietyplants and thus help control grassy weeds (39) This dual

of

pest benefit role extends from tending ants which protecthoneydew-excreting pest species including root aphids and mealybugs (workers repel potential predators and even selectively kill parasitized aphids) to predators of leaf- and stem-feeding rice pests such as leaffolders armywormswhite grubs and stem borers (1 17)

The upland rice environment is particularly well suited to ants because of the lack of flooding Frequent tillage perpetuates grasses and granivorous species that are highlyadapted to constructing new nests in recently tilled fieldsAnt nests however are relatively shallow and tilling with a moldboard plow can destroy them Also upland ricefields are uually small and isolated and a given area in a smallfield has more field borders than it has in a larger fieldallowing high rates of infiltration from more sedentaryspecies such as Plieidole that do not colonize tilled fields but nest in soil under shrubs and trees along field bordersPheidole workers forage into fields from borders and store seed in their nests In slash-and-burn or other no-tillage rice cultures nests would be preserved in the field and possiblylead to high rates of seed predation However granivorous species do not prefer forest habitats

Damage is characterized by reduced usually patchy plantstand (120) Seed-thieving ants greatly prefer rice but they can be selective regarding plants they collect and store Theyharvest more seed than they can consume so patches of germip ing surplus seed (such as Rottboellia) will sproutfrom their nests during the rainy season

In Latin America leaf-cutting ants (Aita spp and Acromyrmex spp) occur only in upland fields and defoliate

young rice plants (47 68) They take leaves to fungal gardensin underground nests where fungi predigest the plant material

Termites Subterranean termites of the family Termitidae which lack symbiotic protozoa to help digest plants are the most frequently mentioned termite pests of upland rice (1185 120) This family cultures fungi in special undergroundcells --- fungal combs made of half-digested plant material (117) The fungi inoculated onto the combs break down plant material into food that termits can digest The termites then consume the fungal combs These grasslandtermites build nests below the plow layer in uplandricefields The colonies attack living plants only when dead plant material is not available They will attack a droughtshystressed crop but prefer older plants having greater cellulose content They will damage newly planted crops where clean culture has removed vegetation

Infestations are worse in deep light-textured soils with low moisture content The first sign of damage is yellowingof older leaves (195) Termites feed on roots (the plantyellows then wilts and finally collapses) and germinatingseed (loss of stand) or move above ground at night to cutseedlings at ground level which they cover with soil for later consumption Soft-bodied termites are highly sensitive to desiccation They live within a self-contained system of airtight chambers and build surface tunnels linei with nudand body secretions (carton) to maintain more than 90 relative humidity Termites can be located by their tunnels

Termites apparently can withstand limited submergence (157) and can be a pest of dry lowland seedbeds (117)perhaps by having nests below perched water tables

Termites appear to do more damage in Latin America where large ricefields are plowed with tractors deprivingtermites of preferred vegetation and forcing them to live on

rice plants In Asia upland ricefields are typically small with Iprge field border areas (ecotones) where termites can forage for preferred food other than growing rice plants All grassland termites do not feed on rice In an upland area in Batangas Province Philippines ricefields are commonly infested by Hosiititernesluzonensis (Oshima) however this termite does not damage rice In the same province though Mrroternesgivus(Hagen)feeds on rice seedlings (178)

In Africa termites may damage rice even though fields are small and surrounded by perennial vegetation These termite species perhaps prefer rice over other hosts

Mole crickets Nymphs and adults are nocturnal and burrow through soil feeding on roots of a wide variety of plants or forage above ground as predators of insects Adults live 3-5 mo and are even cannibalistic Mole crickets prefer low-lyin moist upland soils with high organic matter (II 81) In the lowlands they inhabit rice bunds or nonflooded fields Adults are highly mobile and can leave a flooded field to locate a more suitable habitat Losses show as wilted plants (from root feeding) or reduced stands (120) Damage is more common near field borders where mole crickets relocate after tillage (126)

Fiehl crickets Nymphs and adults of field crickets have similar nocturnal habits and damage rice as much as mole crickets do Piles of weeds removed from fields attract them They make subterranean nests and tunnel through the soil to feed on roots Some species prefer seed to roots Others feed at the base of stems causing deadhearts

Tunnel entrances surrounded by excavated soil are easily seen Young plants are cut at ground level and stored in underground cell Field crickets tre highly polyphagous as are all soil pests (117) Sachan et al (198) reported that field crickets prefer maize to upland rice Like mole crickets field crickets cannot survive in standing water and are therefore more prevalent in upland fields

Whitegrubsandhackbeetles Scarab beetles that feed on living roots as larvae but not as adults are called white grubs or chafers (Melolonthinae Rutellinae) and those that feed as adults but not as larvae on plant crowns are called black beetles (Dynastinac) In the tropics chafers or white grubs have a I-yr life cycle starting with adults emerging from the soil about I mo after the first downpour of the rainy season (134 159) They lay eggs at the same time farmers sow upland rice The rice passes its most susceptible stage and damage is mostly avoided when the white grubs are small After several months the long-lived larvae are large enough for two or three to denude the root system of mature rice (223) This intensity of damage is rare but wilting occurs when root loss is combined with drought stress

The larvae need damp (not saturated or dry) soil to survive and pass the unfavorable dry season 05-2 m underground in aestivation in the moist soil The first heavy rains of the season (20-30 amd) stimulate the grubs to resume activity and after several weeks they develop into

IRPS No 123 January 1987 11

pupae and adults and eventually dig their way to the soil surface and fly to nearby trees to seek food and mates The adults of most white grub species feed only on trees but in some species notably Eutheolahumilis the adults defoliate and cut off stems at ground level (28) Putheola hidentata damages sugarcane fields in Venezuela (121) Floods kill white grubs (208) Grasslands can support large populations and therefore white grubs can be more abundant in newly planted upland ricefields that were previously fallow (81)

Black beetles live up to a year adults can attack a crop at any stage by burrowing into the soil (127) and feeding at the base of stems to cause whiteheads (207) Adults emerge with the early rains and are normally more abundant and do more damage on a young rice crop with small root systems (38 81)

The larvae feed only on dead organic matter in dryland fields and do not attack rice Adults arc highly mobile and although sensitive to flooding can invade a field soon after it drains (115)

Root aplhids Several genera of migratory aphids feed on the roots of upland rice (249) Populations build up more in light-textured soils with high percolation rates Damage has also been reported from dry seed heds of lowland rice (52) Tending ants -- Pheidhle Cren aster Tetramoriuw lsius Tapinoma -- are necessary for root aphids to multiply Aiits harbor aphids in their nests over winter or during unfavorable periods and relocate them on rice plants digging tunnels along the root systems to allow these softshybodied insects to penetrate the soil Most intensive studies on aphid life histories have been done in temperate environments (158 222 229) Root aphids fly to rice plants at the beginning of the rice season and pass through several generations continually developing winged forms to relocate on alternate grassy hosts until winter when they move to perennial hosts usually trees Yield loss occurs mainly through reduced tillering but infested plants become yellow and stunted and in extreme cases wilt (227) Aphid populations build up gradually so damage usually begins in the late vegetative and reproductive growth stages

Root-feeding ineal tugs More than half of the mealybug species or rice feed on roots (245) Being soft bodied mealybugs are not adapted to living underground but they survive because tending ants (the same species that tend root aphids) dig burrows for them and move them from plant to plant Well-drained soils ihelp mcalybugs survive Although six species are recorded as subterranean root pests of rice we found no report citing damage Root-feeding mealybugs are probably more prevalent on perennial grasses than on short-lived annual rice

The most common rice mealyb ig Brevennia rehi is a foliar feeder favored by lack ol rain apparently it is responsible for high yield loss in Northeast India and Bangladesh (N Panda pers comm as stated by IRRI [1061) This report needs confirmation Mealybug damage was only briefly mentioned in another report (156)

12 IRPS No 123 January 1987

Root weevils A few species of upland weevils (IHporneces Donacia Atactogaster) attack rice roots as larvae and feed on stems and leaves as adults Damage ispatchy The larvae are not adapted to submergence and are true upland soil weevils isopposed to the so-called water weevils They rarely become sufficiently abundant to cause damage Adults on rice foliage or the soil surface show that the roots are being attacked by the grubs Root weevils are highly polyphagous (185)

Root heeths Soil-inhabiting false wireworms Gono-cephalumt spp are reported in upland ricefields The adults and larvae normally feed on decaying organic matter but land preparation for upland rice removes these sources The larvae feed on seedling roots and nocturnal adults roam on the soil surface to cut off seedlings at ground level (82) This group of ground beetles is highly omnivorous adults even prey on stem borer larvae (183) True wireworms (Elate-ridae) feed on living roots but rice as a host has been reported Infrequently (117) In Latin America the highlypclyphagous larvae of Diabrotica rootworms are also recorded on rice

Root htus Upland rice seedlings can be attacked by the adults anI nlViphs of chinch bugs Blissusand Caenoblissus and bro n bug Scaptocoris which feed on the roots with theirsucking nouthparts (80 83) Three genera of burrower bugs A tms (eotonhus and Stiharopus -are common iil grasslands in the Philippines and may attack the first crop (rice or maize) planted after the dry season Chinch bugs are most abundant after a dry spell These insects are large and can kill seedlings or stunt and reduce tillering on older plants I hey prefer other hosts and rarely damage rice Adults can also feed oii developing rice grains (81)

(utworms Soil-dwelling noctuid larvae hide under-ground during the day to avoid predation by birds and bccone active at night above grour cutting off you ing plants at ground level (hence tile naiie cvtworn) arvaedrag severed plants into hurrows llie larvae have wide host ranges arid pupatc in the soillgrotis ipsilon occurs worldwide and appears to be the most common cutworm pest of upland environments but is more prevalent inlowland rice particularly in dry seedbeds

IFoliar pests Other groups of insects are adapted to upland environments because they pass one growth stage underground either as egg (in heinrietabolous species) or pupa (in holomnetabolous groups)

Rice seedlibig maggots AitJeriota flies occur in non-fhoodcd environments in Asia and Africa (180) Feeding of larvae one per iller causes deadhearts Flies as opposed to moths only secondarily evolved as pliytophagous pestsfroni a saprophagous Origin Athrnonalarvae secondarihv feed onidecaying tillers whereas larvae of stem borer moths actively Iced on living tillers to cause deadlicarts Seedling maggots will die if they cannot sever a tiller on which to feed

as it decays Plants die under heavy attack whereas lighter infestations cause stunting delayed maturity and ragged leaves (236)

Eggs are laid only on actively growing plants (49) aid damage occurs wiihin the first week after crop emergence(170) Larvae pupate more in stems than in soil Seedling maggot attack is highly seasonal (206) normally peaking 2-3 mo after the onset of the rainy season These pests possiblyaestivatc over the dry season Rice isonly one of their many graminaceous crop and weed hosts

The rice stem maggot (hlorops orzae is adapted to temperate upland and lowland rice growing regions of Asia In Japan and China upland rice grows in mountainous regions a habitat of this chloropid fly The larvae tunnel into tilestems The stem maggot is more damaging during the seedling stage larvac hibernate in grasses

File several dipterans reported as stem maggots in dryshyseeded lowland seedlings inIndia (205) could presumably attack upland rio

Leaf urinrs Icaf miners are common on wheat and maize but are rarely reported on upland rice (213) Three leaf miners are more prevalent on upland than lowland rice inthe Philippines Pvudonalomy a asicanh Vficataand lIAgromrza orzae attack at the seedling stage The larvae develop inside the tunnels they construct in the parenchyma tissue P Vpicata and Aorvzac prefer maizeEehtsine and other grasses to rice spicatacan become abundant on wheat

Leaf beetles Chrysomelid and coccinellid eaf bcetles scrape or otherwise remove leaf tissue as larvae or adults or both This damage accelerates desiccation of plants Tile larvae of sonic species are root feeders Flea beetles Epitrix and (lraetocnetna and rootworms l)iahroticaspp feed on rice foliage only as adults Their root-feeding larvae have hosts other than rice The adults and larvae of the hispa-like Oediolmlpa spp beetles of Brazil and (hmootriha (=EPfishyachra) beetle of Africa feed on upland rice leaves Oealioshypapa spp larvae are leaf miners (mootriha is also pshyvalent in lowland environments (11) Leaf beetles prefer vegetative rice (81)

The rice hispa Dicladipaarnigera is not reported on upland rice and prcers more aquatic habitats In Central Africa however I)cladispm viridicyaiea attacks uplandrice in the vegetative stage as well as lowland seedbeds 7richipasericea is reported on upland rice in West Africa and Madagascar (57)

The rice leaf beetle 1d1ea 01izae riiuech like Chioropsor|rzae is adapted to temperate regions of Ashi and is usually a pest on lowland rice but seldom oi inpland rice in the mountains (82) The adult hibernates on grasses

Armmworms Species of grassland-adapted SpoptloeraPlatysemta and A ithimna larvae create widespread epishydemics They can defoliate ricefields generally in patchesfrom early vegetative growth to harvest (41) Armywornms pupate inthe soil and highly favor upland rice (34) Adults

can disperse long distances to colonize even remote upland areas They become more abundant than their natural enemies particularly after favorable rains following a prolonged drought (196) The drought kills the armyworm and natural enemies (mainly parasites) but the armyworm can recolonize rapidly

The natural increase insoil fertility from mineralization of soil N over the drought period promotes luxuriant plant growth to foster a rapid buildup of these highly fecund species Weeding rice increases tilelikelihood of damage The parasites normally return to control the armyworms but not until after serious damage

Outbreaks on upland rice have been reported in Ghana (3) inzibar (38) Central Africa (36) lPanama (165) Brazil (28) Malaysia (196) and India (167) Defoliation may be severe often leaving only the base of stems Armyworm larvae hide during the day under litter or in soil cracks At night they ascend plants to feed Larvae pupate in the soil

Thrips Thrips frequent upland rice but damage is less than in the lowlands Thrips larvae and adults feed on leaf blades by rasping causing leaf rolling and stippling Thrips prefer vegetative stage rice and their survival increases during drought (heavy rain washes them from foliage) although drought stress usually outweighs thrips damage Drought and thrips cause leaf rolling in upland rice (235) but thrips have been overlooked perhaps becaLse of their small size Thrips are most numerous indry season irrigated rice The combination of favorable plant growth and lack of rainfall is ideal for thrips In contrast the lack of rainfall in upland rice hurts the crop and eventually the thrips whose numbers must decline

In [atin America Frankinilla rodeos attacks panicles before they emerge from the boot calusing sterile grains

Stem hug~s Black bugs are the most common group of rice stem-feeding -lemiptera in Asia The most frequently mentioned species ----Scotinophara coarctaia and Shrida - prefer aquatic habitats and therefore could not occur in upland rice as sometimes reported Upland species in the same gerurs - notably S tarsalis and S scotti -- are morphologically similar and remove sap from stems of grasses and upland rice The upland species are notably more abundant in upland rice planted near forests

In Latin America tile Tibraca ima-large pentatomid tiventris causes deadhearts and whitehcads on upland rice Both adults and nymphs remove sap from internodes of plants more than 3 wk old Tillers are killed by mechanical damage and by the entry of secondary microbial infection This damage is often confused with that of lepidopterous stem borers (233) Eggs are laid on the leaves and stems Nymphs are gregarious This species can also be found on irrigated rice Drought reduces population buildup

Stiem borers Lepidopterous and dipterous stem borers are widely recorded on upland rice but cultural practices and alternate hosts determine abtndance Except for Elasmolmll)uS Iinosellts numbers usually build up toward

IRPS No 123 January 1987 13

the end of the crop growth cycle Therefore late-maturing varieties staggered planting and lack offloodingfavorstem borer buildup Many stem borer species can lie dormant in the stUbbleafter harvest and theirsurvival is encouraged by panicle harvesting (leaving tall stems) and lack of tillage after harvest

Stem borers disperse and lay many eggs Many species are polyphagous however the monophagous MaliarpiasepashyratellaDiopsis Ivmgicornis Scirpophaga incertlas and S innotata can also be abundant on upland rice (hilo Suplressalisdevelops fasterand becomes la rger when reared on upland than on lowland rice (226)

The lesser corn stalk borer lasmopal tt Ittoselhs is semisubterranean and is perhaps the stern borer most adapted to upland rice(129) Highly polyphagous it infests maize peanut and cowpea It attacks seedling rice (one larva can kill tip to four plants before maximum tillering) (195) (hiv and Maliarphastem borers prefer older plants But Scirpophagaspp will attack young plants even in the nursery lasn opalls larvae tunnel into the stems at or below the soil level mausing dcadhcarts Larvae are not found in their turnnels inside sterns beca use vhei disturbed they retreat into cases made from soil particles bound by silk The cases areattached to the tunnel entrance (143) This behavior probably evolved as protection from natural enemies Larvae pupate in the soil The lesser corn stalk borer is more abundant during drought damage is often confused with that of drought (233) This species prefers sandy soils arid its distribution within a field may relate to soil texture (143)

Diatraeasaccharalis aside from the lesser corn stalk borer is frequently a stem borer pest in Latin America According to Teran (233) rice and maize are more preferred hosts than sugarcaine its namesake

Atcitonacrvsograpla is a less known upland rice stem borer in Asia with habits similar to Easinopall s (82) because it is not found in lowlands Acutra has grassy weed alternate hosts As a moth it is often confused with the dark-headed stern borer(hi polur(standstriped stern borer (hi sup)lpsrcais

Other common upland rice stem borers are also reported in the lowlands The dominant species in upland rice vary greatly with location In Uttar lPradesh India (78) and in Japan (123) the pink stem borer Sesamia htfirens is more prevalent hIKenya the principal rice stem borer is Chi partelus

(92) The lowland African sten borers Aialiarpha sepashyratellaDiopsis vngicornis and Sesamia calatnisnis are less abundant Oimupland than on lowland rice(217) but perhaps cause greater yield loss because plant injury is greater if combined with drought css (92) The stalk-eyed stern borer D)ivpsis is more prevalent in wet habitats (6 1I217)

In Central and West Africa Maliarpha and Sesamia calanistis are the dominant stern borers of upland rice (8 9 10 II) (lilo dihlfilineui and Chio partellus inhabit the

14 IRPS No 123 January 1987

upland savannas whereas Chilo zacconio prefers the lowlands (II) Aaliarpha is most abundant in the mangrove swamp habitats (91) coastal regions (16) or rain forest zones (II) of Africa and appears to be the homologue ofScirlpophaga incertulas The yellow stem borr can also be prevalent in upland rice (120 167) The sugarcane borer Eldanasaccharinais so far only a potential pest of uplandrice (91) despite Grist and evers (82) warning

Grasshotl)ers andI locitas Short-horned grasshoppers and locusts -- Locvta Patanrga Schistocerca - layeggs in the soil inhabit grasslands and develop into swarms to seriously damage upland rice from time to time (173) Largenumbers can destroy a field leaving only the stubble In low cmuoroer s giassnoppeirs an cut panicles (173) Farmers may not plant if locust swarms are imminent Prolonged dryweather followed by favorable rain favors development of large swarms (173) Usually breeding in di) grasslands theydisperse toattack crops including upland rice(191) Eggsaresusceptible to desiccation d uring drough t and toinundation during seasonal rains With favorable soil moisture over consecutive seasons natural enemies ca nnot prevent a largepopulation increase leading to migratory swarms (64)

Most grasshoppers on rice are adapted to uplands (107)but lowland rice grasshoppers iervgl(rlhusspp and Ora spp occur occasionally on upland rice (2 82) They lay eggson rice foliage Nymphs are scmiaquatic

The gryllid Lsctrtus coucinnus is a pest of lowland and deep water rice in Bangladesh and Thailand In the Philippines it damages upland rice grown in isolated pockets near lakes The gryllid nymphs and ad ults feed on the central portions of folrage leasing only the midrib and leaf nargins

Leal ler Pvralid moths whose larvae fold leaves to make a feeding shelter occur worldwide (217) but arc appareritly most common in Asia Leaffolde rsattacking rice in Asia are a complex of species (26) (Ctalhahcrcisinedinalis the best-known rice Ia flolde r in Asia may prefer grassy weeds over rice (86) It is a late colonizer of uplandrice U1arasmja eviua a lowland species (220) nreflers rice to grass specres (86) and colonizes upland rice during il early growth stages llarasnria patnalis is the most corn-rnonly encountered species in upland rice in the Philippines Perennial grasses are their alternate hosts and they can survive year-round in upland rice environments The leaf-folders 11arasmia ruralisand Alarasmia spp are least aburndant Dormancy is unknown in leaffolders Cnaplha-ocrocis medhnalis however is known as a migrant (90)Leaffolders remove photosynthetic tissue attack during the flag leaf stage is particularly injurious Popurlations are normally held in check by natural enemies but leaffolders become parlicularly abundant in conditions of high plantfertility and shade

Butterflies Skippers PelopidasInathiasand Borhofanta are more prevalent in African lowland than in uplandhabitats (6) but extensive studies in Japan and China

showed that Pelopidas mathias prefers an upland habitat (155) Parnaragutata seasonally migrates between lowland and upland areas and is equally abundant in both environshyments (164)

The green horned caterpillar Melanitis leda ismene is more abundant in lowlands (212) but our experience in the Philippines suggests that Pelopidas niathiasand 11elanifis leda ismene are often more abundant in uplands than in lowlands Bamboo iaunbuiva spp annually sustains P mathiaspopulations in the Philippine uplands Adults feed on nectar from flowers and migrate to upland rice during the late vegetative stage

iPi erlled h i-1 1d dill L tIC i 7-0 l(become numerous probably because of high egg predationEgg and larval parasitization are often low Butterflies can readily disperse long distances to seek renlote uplandricefields However they have low biotic potential and even it predation rates are low the rarely become abundant Upland environments with their diverse microhabitats can provide more favorable sites than lowland rice plains

tutterfly pests of rice have broad host ranges and become dormant Juring urnfavorable times of the year

IolhagrotslaidopwraThe larvae of many polyshyphagous moths are reported to defoliate upland rice tussock moths lsaispelnnatula Iawlia sUifita luproctisirgtwltaE mnorand P vathorrhoeaslug caterpillar

litotiahicolorhairy caterpillars Nisaga sinp vandA mata spp and wooly bears l)iacrisiaohliqtia and Crvatonotts gangis Many of these polyphgous larvac migrate and are transients in rice including HelhWhis arm iera and 1t zea highly pol phagous species (82)

Gall nidges The Asian rice gall midge Orseoliaorrzae (88 131) and its African counterpart 0 ortzivora(6 11 12 32217) prelerlowland to upland rice Gall midges in rainfed areas are prevalent only during tle rainy season and dispersepoorly Their alternate hosts (wild rices LeersiaIschaetnuan and Pasahmt)are mostlyaquatic grasses and are less likelyto be near upland rice areas In an area of mixed uplandfloating and rainfed lowland rice cultures in northern Thailand upland rice which is ilanted a month earlier was attacked firstbut subsequcent populations were much lower than in rainfed lowla id or floating rice Low numbers of gallmidges passed the dry season orr perennial weeds such as Paspahim (88)

Smot (hid(s Reports of aphids attacking upland rice plants are few (249) Without maize the maize leaf aphidRho alosihhunm maidis attacks upland rice in the Philipshypines The plum peach aphid IIrsteroneuraselariae was reported in Sierra Leerne toinfest leavesand tnriperied grain(7) Siphaglrctriaecaused transverse linrea r necrotic striae iir rice leaves in Italy (172) I-lea v rains held populations in check Rice is not a preferred host for shoot aphids

Leafhoper andI )anthol)pers Cicadellids and delshyphacidsdispersecasilyand readilycolonie upland ricefrom nearby lowland areas A study in Sarawak (241) showed

IRPS No 123 January 1987 15

similar species in upland and lowland rice Nephoiettix The principal vectors of rice virus diseases generally have species dominated The most prevalent species accounting narrow host ranges and alternate hosts in the grass family for 90-95 of the sweep net samples wre N virescens N are mostly annuals dying in the dr3 season Some virus umropictus Nilaparvia hgens Soatellafiircifera Cfana vectors such as BPH can disperse lontg distances but few do

sectraand Recilia dorsalisLowlands had higher popula- Th restriction of upland rice and weed hosts to the rainy tions Katanyukul and Chandaratat (120) monitored hopper season the small area planted to upland rice and the hopper numbers in upland and lowland fields of northern Thaiiand vectors having narrow host ranges all work against virus and found similar results A comparative study of plant- diseases in upland rice hoppers in Fiji showed more brown planthoppe BPH) and In Latin America Sogctdhs o(rzicola and S cuhanus whitebacked planthopper (WBPH) in wetland rice than in transmit hoja blanca virus disease but reports on upland lowland rice and hopperburn was prevalent in lowland rice are rare except in the favored upland areas of Colombia areas with standing water (89) Peru and Venezuela (35 42) S rricolaocurs in Brazil

Hopperburn in upland ricefields in Asia is rare (84) In on upland rice but hojaI blanca has not been reported (67) Batangas Philippines isolated fields of upland rice become S orzicola has a wide host ramgc (10) similar to WBPI- in hopperburned infrequently from WBPH A report from Fiji Asia WBPH however is not known to transmit any virus cites hopperburn from BPH and WBPH (250) In these two disease situations upland rice was grown on highly fertic soil and In Africa pale yellow mosaic virus is endemic to swampy N fertilizer was applied Hoppers respond to better nutrition lowland areas where rice ratoons year-round (19) and is Normally upland rice is grown on poor soil and receives no mechanically vectored mainly by hispid beetles A minor fertilizer vector a flea beetle Chaeiocnema spp is an upland rice

In Asia upland rice ras are normally not far from pest and upland ricetields near endemic areas occasionally lowland ricefields-- a source of hoppers Otherwise hopper contract the virus species with greater polyphagy than BPH or Nelhotettiv Spittle bugs Cercopid nymphs producc a protective virescens would be dominant frothy covering that looks like saliva In Latin America

In Africa leathoppers and planthoppers do not become spittle bugs damage upland rice In parts of Brazil where abundant on upland rice (6 II) probably because lowland upland ricefields are surrounded by pastures adults rice is not widely rlanted nearby The only reported migrating from grasses can damage rice severely and cause hopperburn as from Nilapanrata maeander in breeder complete hopperburn in a young rice crop within a week of plots receiving high N rates in Nigeria (98) Hoppers colonization (194 195) Dispersal to rice occurs even if however are a greter problem in Latin America where pastures are verdant and spittle bugs prefer pasture grasses large grassland areas breed polyphagous species (Gra- to rice (168) Nymphs are rarely abundant enough to phocephala spp Hortensia spp lEvitianus ohscurunervis damage rice Regular rainfall favors their development Balclutha spp and Draeculacephala spp) vhich then (233) disperse to cause hopperburn on seedling rice (68) Rosetto et al (195) reported Graphocep7hala spp and Hortensia spp Seed pests to be equally abundant on upland and lowland rice Alydid and pentatomid bugs are seed pests of upland rice

The leafliopper and planthopper epidemics in Asia that worldwide In Asia Leptocorisa sp prevail Their habits caused severe losses from hopperburn and vijus dicases make them particularly suitable for upland rice 1)thev feed over the past several decades occurred principally on on many grasses (and therefore can survive the earl y wet lowland rice Except in a few isolated cases (160) upland season before rice sets its grain) 2)adults are long-ied and rice has been spared from tungro grassy stunt and ragged mobile allowing them to find isolated plantings 0 rice stunt virus diseases vectorcd by green lcafhopper (GLH) (besides seed bugs only birds and rats appear to have shyand BPI] (123) Virus diseases have occurred in upland rice capability) and 3) adults aestivate in forested areas or where lowland rice was nearby Rice dwarf disease however sugarcane fields during the dry sca son when neither rice nor was first recorded on upland rice in the Philippines (187) grassy weeds are present (I199) Rice seed bugs concentrate Virus diseases can be perpetuated only on livir plants or on small-scale upland riceficlds because they can actively insect vectors and no living rice - plans tooned or search them out The prevalence of groves of trees characshyvolunteer - grows year-round in upland arcas excert in a teristic of upland environments ensures nearby aostivation few with 12 mo of rainfall sites keeping the seed bugs close by

The smaller brown planthopper Laodcelphax striatellu Leptocorisae are uncommon in lowland rice plains occurs in upland rice in Japan (18) It spreads black- because 1) their populations are diluted in a sea of rice streaked dwarf and stripe viruses between winter wheat and planted more or less at the same time (they ca n only feed on barley to lowland rice Early planting of lowland rice rice during the milk to hard dough stages) 2) they lack nurseries spreads these viruses but upland ice has remained aestivation sites and 3) weedy fallow areas are limited uninfected The lowands of Asia are dominated by L oratorius

16 IRPS No 123 January 1987

which lays its eggs hign on tile foliage (199) L acuta and L solonlonlesis oviposit at ground level the former on litter and the latter loose on the ground These egg laying habis explain theirtenvironmental prefcrencecs(214) In Japan the postdiapause adults of L chinensis migrate from moun-tainous areas to upland ricefields during floweiing (1 14)

Pentatornid bugs prevail in Latin America (8182 195) and in Africa pentatomid and coreid bugs arc equally cited (6 82) l)amage is often characterstic of the species (5)rcoris ventralis and Mnida spp are the common

pentatomid seed bugs of Philippine upland rice but they prefer lEchinochva spp grass seed to rice Vearaviritda occurs worldwide and hoften more abundant in the uplands(54) We saw few reports of seed bugs on upland rice but see(d bugs apparently opelrate cqually between by land and upland rice Good dispersal is necessary ior species that call feed for only 2-3 xon a crop and must first find those crops at ripening stages

Other pct species feed n flowers In Mato Grosso Brazil (194) and Santa Cruz Bolivia the panicle weevil Aeo-hariuia amllitarsis attacks rice during flowering (68 233)The larva e rice sten horers adutlts feed at the bases of spikelets caIusing empty grains Also inLatin America thecydnid -lkiituttatralusisa seed bug as anad tilt and nymph (194)

A number of beetles ecd on poilen - coccinellids lialrotia Aulac lhora and Monohpl-a (55) - but becatise riCe is self-pollinating and fcrtiliation occurs before pikelets open they pose no threat to yield

IST ECOLOGY

We now look at characteiistics of the life cycles of upland rice insect pests to lea rn hew each plays an adaptive role in upland rice ecology

Life history strategies Orzasativadoes not tolerate drodgh well so upland rice is highly seasonal normally grown in the wettest months of the year Usually rice is present for less than half of the year The rice-fiee fallow poses serious problems of survival to upland rice insects which have evolved att least four mechanisms to ovrcolne tle cvcl ic-l lack of a hostI) polyphagy 2) longevity 3) dormancy and 4) vagility (dispersal)

Poldt)agy lolyphagy is defined as having hosts of at least two botanical families oligophagy of more than one gents and ionophagy of only one ventus (40) - inour case Orvza Most wild rices however are illi tiC and theieiirc are normally far removed from upland rice hahitat othewise more would be alternate hosts for uplanJ iice insects Thte highly mornophagous rice pests shySciriol)haita hiceritlas Nilal)arvaiahiens Iand Nealo--tettiyvircsctws in Asia and Aalirpha secparatella and 1ioj)sis lngicornis inAfrica - occur on upland rice but

are more abundant in lowland culture They can specialize in rice because they are highly vagile and can attack all growth stages By specializing in rice monophagous rice pests can outcompete related but polyphagous speciesIntensive andI extensive lowland rice culture has favored monophagous species Riqwla alhinella (Cramer) the onlymonophagous species in Latin America is rarely reported on upland ice (I1)

Because of its limited temporal and spatial existence upland rice has not favored the evolution of specialized species Clhang(44)shows evidence that upland rice has only recently been cultured by man from the lowlands thus pests have had little time to fully adapt to the crop All but the lowland-adapted insect pest species have alternate planthosts to rice Root pests with limited mobility are highly polyphagous Pests that specialize in one growth stage eg secdling maggots seed bugs are oligophagous or polyphagous

Most alternat hosts of rice pests are annual grasseswhich also are seasonal Upla_rice is also a host but not the preferred one of the most highly polypliagous spcciessoil pests grass hoppers armvwarrs and Vearaviirielda

The d ry season poses igreat obstacle to ani upland riceinsects survival and only sonic perennial plant hosts are suitable food during the off-seasoin Plolyphagy allows upland rice insects to suLrx-ixe during thC rainy season when plants- both annual and perennial -areactivelygrowing and therefore more nutritious

Loiugevitr The ability of aiinsect to live 2-3 mo without undergoing dormancy helps species sich as root-feedingwhite grbs (as larvae) and secd-feeding bugs (as adtilt) to survive unfavorable periods White grubs arc not highly fecund and their life strategy is to improve survivorship of limited progeny Their subterranean habitat hides them from manv natural enemies lBtut root tissue is not highly nutritious so they ltilst eat great quantities and need a largeinsect biomass to digest it Other soil-inhabiting rootshyfeeding insects have short life cycles but are more fecund

Seed pests need greater longcvi vas adults to locate ahost at early grain dcclopment arnd give sedentary offspring time to develop Few groups of upland rice insects live longin active development Polyphagy and longevity only increase their ability to survive during the rainy seasonOther mechanisms are needed to survive the dry season

orantc In the tropics the nonrice season is the dry season while intemperate areas tile off-+season is winter Many upland rice insects undergo dormancy inlsummer (aestivation) or winter (hibernation) Dormancy is simplyinactivity during winters with prolonged temperatures below 10 C insects find shelter arnd cease movement Activity is resu med with warmer t empcera tu res A deep state of irnactivity termed diapauise occurs in sole species that prepare physiologically (havxing low metabolism and convcrting gonad s to fatty fuel reserves) for the unfavorable season (148) They enter diapaUse inanticipation of unshy

IRPS No 123 January 1987 17

favorable weather and resume activity after favorable Armyworms skippers and locusts actively travel hundreds weather begins usually in response to photoperiod or of kilometers in air fronts such as intertropical convergencethrough a delayed response to a stimulus such as heavy zones to descend onto upland rice fields far from their rainfall (237) Not all individuals of a population have the breeding grounds (31) Several generations can build ip on same threshold for environmental responses for entering or alternate hosts in tile grass family before rice is plantedterminating diapause -- this is seen as a mechanism to These large insects are active fliers Smaller insects such as prevent total mortality if a false cue appears such as a thrips aphids and planthoppers passively migrate long sudden warm spell in winter or rain in the middle of the dry distances by the wind season (154) Otherspecies ean readily travel tens of kilometers - stem

Probably more upland rice insects undL go dormancy borers leaffolders leafhoppes cutworms spittle bugs leaf than we realize Candidates are Atherigona spp root beetles seed bugs ants and termites aphids root weevils crickets leaf beetles thrips and seed The least dispersive species are white grabs mealybugSbugs Insects known to entervarious states of aestivation are seedling maggots crickets weevils and gall midgesleptocorisaspp adults (199) Maliarphascparatellalarvae Each species has evolved a unique set of attributes to (9) Scirpophaga innotata larvae (117) Chilo partelhtsv enable it to survive and adapt to the changing uplandlarvae (21) ScirpopThagainccetulas (179) white grub larvae environments Several evolutionary avenues lead to fitness (134) Patangasuccincia adults (6 82) and Schistocerca (Table 2) but there is no reason to believe one set of gregariaeggs (82) attributes is better than another (224)

Insects known to enter hibernation are LodeIphaDstriatellus nymphs (124) Chtio Tvlpressalis larivae (125) DroughtSesaia inferens larvae (146) but not S calavisis(2) Well-drained rainfed upland soils are subject to drought ifIDiaraea saccharalis larvae (75) Scirpophaga- tcerlttas rains do not fall within 2-3 wk Prolonged drought followedlarae(5 lacchrli s lae (75)saSlOhagasshoperseggs (82)and felo pidas athias pOpae (232) by favorable rains stimulates armyworm and locust outshybreaks but mealybtigs root aphids and thrips become

Lcpmotorisa oratorisand L acuta adults aggregate on numerous onlyafter dry spells for two reasons First heavy trees sugarcane or other shady moist sites L paluwanmsis rain normally kills soft-bodied foliage-feeding thrips and confined to Sulu and Palawan Philippines lives on the mealybugs and soil-inhabiting root aphids and root mealyshygrass Brachiariamutica (Forsk) Stapf in the absence of bugs Drought eliminates this source of mortality Second rice The adults can live for 5 mo When aestivating their the rice plant responds to drought by breaking down rate of metabolism drops and they do not feed They will proteins into soluble N compounds which enter the phloem however puncture plant tissues of their aestivation host in and are taken up by these sap-feeding insects (245) Greater search of water This behavior has led to errors of the host nutrition therefore leads to greater survivorship and range of rice bugs (I) Aestivation quickly terminates with fecundity rain probably meaning aestivation is not deep Small rice area

Some stem borers aestivate as mature larvae in rice Except in the highly mechanized large landholdings of stubble or in straw stacked as livestock feed However Latin America upland ricefieldstypically are patchy within moriality increases proportionally with the duration of the a highly diverse flora The small fields result in higher ratios dry season even if the stubble remains undisturbed If the of perimeter to area than with the typically larger lowland stubble is plowed for a crop following rice few stem borers ricefields Upland ricefields themselves may be intercroppedsurvive Rainfall terminates dormancy but it normally with cereals legumes or root crops Mixed intercroppingtakes several weeks for moths to develop and emerge with a wide variety of species also occ in tribal slash-andshy

Last-instar white grub larvae tunnel 1-2 m deep in the soil burn agriculture (50 74) to construct pupal chambers in which they aestivate The Upland ricefields tend to be small because of the labor larvae are very sensitive to dryness so pupal chambers are needed to clear land in slash-and-burn areas and to till land sealed to coierve body moisture Aestivation terminates to minimize weeds Rice requires more tillage than does with the first heavy rains but as in stem borer 3-4 wk pass maize because rice competes less well with weeds Uplandbefore adults develop rice is more a subsistence than a cash crop and small areas

Insects may respond differently in each region to cold (I ha) can feed a family temperature or drought Biotypes or loal populations may Small fields favor some insect pests such as some ant evolve (56) Local populations of Scirpophagaincertulasor species that can forage from more permanent field borders Patangasuccincta may have become adapted to prevailing Trees provide food and mating sites for white grubs (134) as conditions well as aestivation sites for LetTocorisa spp (199) Field

iagiliti The ability to disperse combined with high border grasses plus tIhose within a ricefield provide alternate fecundity orshort life cycles enable some upland rice pests to food for all but the highly monophagous upland rice better exploit the temporarily favorable upland habitat colonizers and a habitat for natural enemies

0 z

Table 2 Life history patterns of upland rice insects or insect groups

Host range Longevity Dormancy Vagility FertilityMonophagy Oligophagy Polyphagy Low Medium High Yes No Low Dispersive Migratory Low Medium High(Oryza spp) (grasses sedges) (angiosperms) (lt3 wk) (gt2 mo) (lt1 kin) (lt50 eggs) (gt500 eggs)

Ants xTermites x x x

Mole crickets x x x xx x xField crickets x xx

White grubs x x x xx xRoot aphids x x xxMealybugs x

x x x xxSeedling maggots x

x x x x xLeaf beetles

x xx xArmyworms x x xx x

Thrips x x xx xChilo spp x x xx xMaliarpha x x x x

xLocusts x x Xx x xGall midges x x xxiVephotettL iirescens x x xx x x xLeprocorisaspp x

x xLeaftolders x x x

Sogatellafurcifera x

xx x x x x x x

00

IRPS No 123 January 1Q87 19

All this appeals to favor insect pest buildup but the notwithstanding the release of stored nutrients in response contrary is true Loevinsohn (138) showed that insect pest to drought that temporarily benefits philoem feeders populations respond exponentially with the proportion of land devoted to rice up to about 75 of the area then the M POR ANCE I I P1A 1) RWI P1St S

rate of response declines In most upland rico areas tie rice crop occupies less than 50i of the area- therefore the Visits to upland ricefields normally reveal 1kw insect pests potential for population buildup is low This relationship Loevinsohn (138) and Locvinsohn ct al (139) compared may not hold for seed pests that can locate small ricefields insect abundance using annual light trap atchcs in upland

Another outcome of the highly diverse flora and phical rainfed lowland and irrigated lowland sites in the Philipshyenvironment of upland rice isthateach upland ricearca has pines Although only one upland site was studied data a unique composition of insect pests showed equal or fewer insects at the upland site than at the

For example in Tanauan Batangas Philippines the rainfcd lowland sites and fewer than at the irrigated sites main upland rice pests are rice leaffolder(RLF) armvworm (Table 3) M Lltiple regression analysis showed that the key Aythntnaseparata WBPH ants Soli psis gentitataand factors explaining insect abundance among a set of cropping white grub Leucopholis irrorata Sixty kilometers away in intensity variables were the number of rice crops grown per Real Quezon the main pests are seedling maggot Athe- year followed by area devoted to rice Table 3 shows that rigona orae flea beetle (Thacitcmnra hasalis thrips the upland rice site had only I rice crop perym rand that rice Stenchaetothrips hiurnmis and rice bug Leptocorisa was planted in only 20Vi of the area Fertiliter and oratorius In a third Philippine upland rice site in Claveria insecticide use had little bea ring on the trend The armters at Misamis Oriental Mindanao the main pests are ants the upland site applied 60 kg N ha higher than at most Solenopsis gemtinata seedling maggot Alieriona orvzae irrigated sites white grubs Holotrichia in)iUaiuoan t and Lucopholiv Insect occurrence at the upland site was omparable to irrorata stem borers Sesamia ifiArelr (hio auriciliusC that at the Cagayan rainfed lowland site parlicularly in 1982stpessallis Scirpophaga incertulas S innoiata and when drought prevented many farmers fron planting(only

4c ona chrrsurapella root aphids Tetraneura niriah- 40(of the rice area eventually was planted that year) M ore dunrinalisand Rhupaosiphunm riahbdminalis lcaffolders favorable weather occurred in 1980 and 1981 which Marasniapataltis and Chaphalocrocis Medinalis army- supported greater rice areas and consequently more insects worm MIthinna separata and rice bug Leltucorisa The most intensive site for insect pests was South Cotabato oratoriusThe great diversity of insect problems is typical Mindanao where farmers plant 23-25 rice crops per year fer upland rice and makes control efforts more difficult with irrigation The Zaragoa Nutva Ecija site also had particularly regarding breeding for insect resistance high pest abundance and at this site the fields were highly

asynchronous Asynchrony was the third most important Low yield potential Iactor in explaining rice insect abundance Most upland rice is grown on low-nutrient and mineral- Rice cropping intensity in upland areas is bound to toxic soils which when combined with eiratic rainfall remain low Only one rainid upland crop is possible per make upland rice a highly riskycrop for the farmerto invest year (Farmers in Claveria Misamis Oriental Mindanao in costly soil amendments and land management Often Philippines tried asecond upland ricecrop i 1984a ycarof upland rice isgrown far from markets making inputs even favorable rains but no harvest was possible b-cause of more expensive Farmers growing a subsistence crop hosi- drought blast and birds) Rice area isaso bound to remain tate to invest in costly inputs to raise yields low because of the high labor and power rcquirelmenlts to

Even having varieties that could double existing yield prepare the land (because of weeds)and because upland rice potential under present management levels would probably wvill not compete with lowland rice as a cash crop Some not prompt many farmers to use purchased insecticide upland rice sites with prolonged rainy seasons howeverare unless tile risk of crop failure were reduced such as it was planted asynchronously such as the Clacra site with irrigation systems in lowland rice The seasonal abundance of Upland rice insect pests

The Tanauan Batangas site is atypical for upland rice attracted toa light trap wasgrapied for crop year 1980-81 in The soils and rainfall pattern are favorable and because of Tanauan FBatangas (Fig I 2) W13P11 was he most its nearness to Manila farmers have cash resources ftm common rice hopper and its population peaked at the sales of vegetables to purchase tertilier They applied an midgrowth stage as it does in lowland rice D-Vac suction average of 60 kg Nha after particle initiation and obtained samples showed an earlier peak than that from a light trap 25-35 tha yields The high fertility isone reason why RLF indicating emigration W13PI usually emigrate after the armyworms and WBPH developed into large numbers vegetative stage Populations in 1980 reached levels that Farmers however used no insecticide on upland rice caused patches of slight yellowing in some fields WBPH although they did on vegetables probably cause hopperburn in Batangas They immigrate

The poor growing conditions fOr rice generally mean year-round as shown by low levels in the light trap BPH poor growth and low fecundity for insect pests fIeding on it however were scarce in the light trap and not collected with

Table 3 Abundance of rice insect pests and factors in rice cropping intensity of Philippine upland rainfed lowland and irrigated lowland locations where light traps were operated daily for atleast I yr -I

0Rainfed Rainfed lowland Irrigated lowlandupland

Pangasinan Cagayan lloilo Santa Maria Nueva Ecija South Cotabato Tanauan Caaringayan Caaringayan Bangag Bangag Bangag Buray

LagunaCale

Batangas Manaoag 1980-81 Santa Motiica 1982-83 Cabanatuan Zaragoza Avancena NaniamanSolana 1981-82 1982-83 Oton

1980-81 1979-80 Oton 1981-82 1981-82 Koronadal Koronadal1980-81 1979-80 1979-80 1983 1983

Insect (kerosene light trapNephotcttixcatch nolyr)virescens 650 1240 500 1800 330 210 3400 1600 160N nigropictus 210 - 360 18400 5500 2900- 140 90 20 - shy 1700 -Reciliadorsalis 710 - 3200 1700180 890 630 140 170 2400 800Nilaparratalugens 130 690 - - 4200 4600300 520 750 180 60 3300 1600 740 310Sogatellafiircifera 1700 31000 13000 8200310 210 3300 610 270 980 1600 6300 -Cyrrorhinus - 12000 6700600 540 3200 290 160 330 120 410Scirpophaga spp 50 410 620 - - 21000 12000150 180 220 270 30 1000Cnaphalocrocismedinalis 70 70 540 610 270 11000 6100200 140 100 6 60 250Chilo suppressalis 0 60 60 1900 300 200- - 30 20 0 6 3 160 -C auricilius - 0 00 - - 20 3 60 20 3 0Sesania inferens 0 - 80 60 3 80 10 51 -Cropping intensity _ -

Rice crops (noyr) 10 10 12 10 10 10 13Rice area (7c) 17 19 20 20 25 2320 85 85 60 70 40 85 85 90Fertilizer (kg Nha) 60 20 85 80 70 7020 0 0 0 30 30Insecticide (no of 40 60 40 300 04 04 0 03 300 13 13 30 50 40 33 33applicationscrop)

__________

IRPS No 123 January 1987 21

Light trapped Insects Drytnd __ 2(notrap-nght) Suctioned Insects (nom )

50 -)o 9 jO40- SOW ffurciWO Light trp

D-Vc suction

20 to 4

20

Egg porasitizahion (M) 75 60

300 SOOampReOftwciffr

45 30shy

15

-insects (notrop-night) 4 Nileporvata Iuen5

Egg parastization()

Light trapped insects (no trop-nght) Suchoned insects (no m 2 )

6 8 Cytorh~nuls hvidipennii 0

4 05 2

50 0 Insects (noltrop-night)40-ui Ipanh pe g

Moohase 1 Seasonal abundance measured by light trap

Rainfall(etod) and D-Vac suction sampler of whitebacked

to ~ hopper Nilaparvata lugens and lea ffolder60)-t Cnaphalocrocis mnedinalis andplthpe g20 L I II I I I parasites and mtrid predator Ortorhjntus livi-Apr May Jun Jul Aug Sep Oct Nov Dec JaIn Feb Mar dipennis set against cropping pattern daily1 9 80o 19 8 1rainfall and moon phase Tanauan Batangas

1980-81

the D- Vac Cyrtorhinus build up on the crop coterminous are collected in light traps during full moon (109)with WBPH buildup Cyrtorhinus disperses during the Because upland rice shares most of the lowland rice pestsrainy season RLF not readily collected in light traps was and those pest species are generally less impotant to uplandprevalent only during the rice crop rice than lowland rice there is a general belief that insect

Nephotettix virescens was more abundant than Nnigro- pests are less important onpicLus in the light trap collections and on the crop GLH upland than lowland rice Arcview of limited data on upland rice shows an averag of population peaked at the reproductive stage as did zigzag 10-21 yield loss from insect pests on a par with that ofleafhopper (ZLH) Crtorhinus tracked the populations of lowland uee determined by the same methodology (132) planthoppers and leafhoppers Both GLH species were (Table 4) McGuire and Crandall (144) made a similarcollected year-round in the light trap indicating immigra- estimate for Central America Cramer (51) howevertion from nearby irrigated rice areas Possibly because of the estimated 35 yieldlow populations loss in South America based on ainsect number did not correlate with similar pest complex - stem borers and seed bugs shymoon phase although reports indicate that more hoppers attacking irrigated rice in North America This estimate did

22 IRPS No 123 January 1987

Light trapped insects D~~nd (no trap-night )

St plht Suctined insects (nom 2

)

D-Voc suction 1520

1o

0 Lih trap 5 ~105 o5

00 Eggparositizotion()5040 NephOtgtha virneens

30 20

I010 A-Insects (notrop -night)

4 Nephatelh nigropccent i

2 o I vV^J - -- ^ VA____ f _

Light trapped insects (no trap-night) Suctioned insects (no rnE

10 -1208 ReCdO doRsol

6 15

4 10 052 iwxALAiamp 0

Light trapped insects (notrap-night) 2Suc oned nsects (nom )

8 CyrtorhmnusI6o nn]10 i0075

4 05

2 0 0

025

2 Seasonal abundance measured by light trap Roinfall (mmd) Moon phase and D-Vac suction sampler of green leafhoppersNephotettix virescens and N ngropicrus and I zigzag leafhopper Recilia dorsalis their egg 60

parasites and micid predator Cyrtorhinus livi-dipennis set against cropping pattern dailyrainfall and moon phase Tanauan Batangas1980-81

40

5 Lf Apr May Jun

1980 Jul Aug Sp Oct Nov I

-aMar

1981

not account for soil pests and was not based on field data in the absence of pests Carbofuran granules are phytotonicAkinsola (I1) cited mainly data from irrigated rice areas in to rice (239) The prevailing hypothesis is that carbofuranAfrica but estimated yield losses of upland rice between 15 prevents soil bacteria from consuming fertilizer The phytoshyand 30 from insects However losses in Thailand over a tonic effect is through greater availability of fertilizer to the4-yr period ranged from I to 13 averaging 5(106) plant Denitrifying bacteria are known to be more active inYield loss data however should be interpreted with aerobic than anaerobic soils therefore it may not be merecaution (184) The Philippine data were all derived from coincidence that the highest recorded yield losses (22-69)trials on farmers fields under farmers normal agronomic occurred in those trials with 12-30 kg basal Nha On themanagement practices except for Bukidnon and Capiz other hand contro with carbofuran granules may not be aswhere researchers used fertilizer Batangas farmers are good as a seed treatment against seed pests and perhapsatypical because ofhigh levels of fertilizer used The trials in yields would have been higher if a systemic seed treatmentBrazil were carried out at an experiment station but insecticide had been used conditions were similar to those of local farmers The 1979 and 1980 trials in Batangas had two full-

Carbofuran granules were used in every trial as a broad- protection treatments one using carbofuran granules andspectrum insecticide applied basally with the seed An the other using bendiocarb WP as a seed treatment Plotsobjection to using insecticides to measure yield losses isthat protected with bendiocarb yielded higher than those proshyinsecticidescandirectlyorindirectlystimulateplantgrowth tected with carbofuran granules The other trials in the

Table 4 Yield losses to insect pests and yield responses to insecticide in upland rice in the Philippines Brazil and Upper Volta

Yield (tha) Yield loss (5) Location Year Cultivar Protected

with Protected

without Check Total Vegetative Reproductive Ripening

Basal N (kgha) Reference

carbofuran carbofuran Protected No with carbofuran

carbofuran

Philippines Pili Camarines Sur Tanauan Batangas

Pangantucan Bukidnon

Dumarao t-apiz

BrazilGoias Minas Ge~ais

Upper Volta6

1975 1974 1974 1975 1975 1976 1977 1978

1979

1980

1979 1980 1979

1977 1978 1979

Bursiging puti Kinanda Dagge Kinanda Dagge Dagge Dagge Dagge UPL Ri5 Dagge UPL Ri5 Dagge UPL RiS UPL Ri5 UPL Ri5 UPL RiS

IAC47 IAC47 IAC47

14 33 11 36 35 31 30 27 37 26 40 30 40 42 46 35

19 21 09

--

-34 24 24 27 --

29 43 31 42 -

47 25

---

10 28 05 27 27 30 26 29 33 28 33 29 43 23 36 11

15 15 06

29 15 55 25 23

3 13 -7 11

-7 18

3 -7 45 22 69

24 28 35

--

21 -11 -20

4 -

-3

23 7

-2 -23 56

-

-

-__0

6 --

-

-23 14

5 0 8 2

22

-

--

0

-

--18 11

3 -10

4 10

7

--7 -

14 -

-1 6

10 -5

8 9 7

-

--

0 0 0 0 0 0 0 0 0 0 0

30 30 30

12 12 12

99 99

100 100 101 101 102 102 102 102 103 103 104 104 104

66 66 66

Farako-Ba

Mean

1977 1978

IRAT10 IRATIO

38 66

--

40 43

-5 35

21

-

10

--

--

--

- 242 243

Z

tJ

-w

24 IRPS No 123 January 1987

Philippines which had two treatments offering protection In the 9 Philippine trials where yield losses were measuredwith and without carbofuran granules relied on further by growth stage 10 of the yield loss (21) occurred at thecontrol with foliar spray and not on seed dressings The vegetative stage and 517 occurred at each of the reproshycarbofuran-protected plots had an 18yield los compared ductive and ripening stages Ibis result is similar to thewith only 10 with foliar sprayS This yield difference is pattern of yield loss in lowland rice measured by successiveprobably due to differences in protection against sown-seed treatments where the crop is unprotected at each growthpests (foliar spray do not control sown-seed pests) rather stage but proticted during the other growth stagesthan phytotonic effects became basal fertilizer wa used in However the insect complexes r sponsible for vegetativeonly 2 of the 10 comparisons stage yield losses in upland rice are entirely diftfrent than in

The recorded yield losses were not related to yield lowland rice The key vegetative stage pests in lowland ricepotential (Fig 3) One might expect higheryielding crops to arc aquatic whereas those in upland rice are sown--seedhave higher percentage of yield losses but this ws not tile (sown in soil) root and seedling pests This dift rence maycase One reason is that insect damage exacerbates plant explain why agricul torists have overlooked tile ilmportanccinjury from drought stress low yielding fields from d ought of upland rice insect pests Many reports of upland rice pestsstress would show disproportionate differences between in Asia ha Ve focused on major lowland species (46 108)insect-protected and -unprotected treatments Shoot and It isalso surprising that tei vegctative stage of upland rice root pests would be more responsible for losses from would record the highest yield losses becatuseseeral studiesdrought stress than sown-scd pests A second reason have shown that upland rice can readily recover from foliageconcerns seed pests which would have a greater impact on reinoval during that stage (117 152 165 188) Upland ricelow tillering low yielding varieties such as )aggc Low poscsses a hitgh obilitV to compensate for ca rly loss oftillering varieties nnot fill in the space created by removal foliage and will even be stinulated by foliage removal toof seeds oweer 1) R i 5 is high yielding because it tillersP1 produce a higher viel than plants ithout any foliageactively which can close the canopy to provide greater weed removed (63 192 231) [ie reason for this apparentcontrol discrepancy hovce is that yield loss at the vegetative

stage does not octUr frol leat arca loss but fron pestsYield loss () removing sown seed and feeding on devcloping tillers and roots -- damage that cannot be readily conpLtnsatCd for

70 - Measuring yield losses in upland rice cnvironrnents is further complicated by a report from Brazil that greater yield losses occurred in response to better insect control60 - =-09 as Insecticide seed treatments resuled in denser plant stands

(n=21) which in turn created a selectively more favorable environshyment for blast (69) This resuIt was parlticurla rlxexacerbated50 when cirbofuran granules were used in seed furrows Aside

ffrom a greater plant stand ca rbofuran iay have auigmented the N lecl and stimulated blast If so ca rbofuran should be40 - replaced with another insecticide or the seeding or fertilizerI rates decreased

Other methods to determine the impact of upland rice30 shy pests wvcrc to interview farners or to develop single speciesbull bull correlations of population levels and yield loss Feneira (66)2 NIM)st (75(1) Golas farmers reporteddid both in Brazil

20- insect losses in ufpland rice and 251 said those lossesS avcragcd 37(1 [his result is consistent with tie yield losses 0 0 measured by field trials in the same statc(Table4) For every

I0 4 of seedlings removC d by leaf-cutting ants a I 7yield loss is predicted Similarly Ferreira (66) found that ai

0 - average of 53 thrips per panicle before panicle initiation meant 3 times more unfillh d grains

-IO0 I 1 1 I I-101 0 2 3 4 5 6 7

Yield (tho) C)N [ROt IM FI IODS 3 Correlation between upland rice yield and yield lols caused by Upland rice insect pests may be controlled by culturalinsects measured in 21 trials in the Philippines Ivory Coast andBrazil genetic biological and chemical methods

IRPS No 123 January 1987 25

Cultural control dry season plowing will have little effect on them Plowing Crop husbandry to reduce upland insect pest populations when the soil is wet from the first monsoon rains will expose can be classified into practices effective in a single field and soil insects that tunnel close to the soil surface to predatory those effective only at the community level Cultural birds chickens dogs and even man (field crickets nole controls should come first in a pest management program crickets and white grubs a re delicacies in man tra ditional They have broad stable effects as pests have little pos- diets) sibility of overcoming them through selection of biotyps The greatest degree i1 inIscct control fron tillage comes (138) On the other hand some cultu ral controls work only from plowing soon after rice ha rvest when most soil insects with high labor or power inputs and connunity-wilt lie close to the soil surface anrd stern borers are in the rice methods require coordination of iany farniters stuble Whitegrubsare mature and tillage exposesthem to

Pantin time The luxury of being able to shift the time of predators Tirnelv control will prtec the crop following pla ntring upland rice is open onl hr la rniers in areas with a rice and lowr the populationIfor succeedinig years Plowingprolonged rainifall pa ttern lI copping intensity or underthe ice stulble helps it decompose killing sten borer rnechanied land preparatiot Gcnerally planlting as soon larvae inside as the rFiinv season begins will lower populatiois of 1110st [requent tillage destroys ait nests bit not termites insect pests Insect populationsarelo aftcra dlvseoll or Tillage and clean culture howes er rcliove food for winter fallow delayed plaiting lets them build tip on termites which then mtay attack a young rice crop Scarab alIternate hosts that gi vigorotsly with the first seasonial beetles also prefer to miposit in eceitly tilled fields Noshyratiris This first flush iof weds cros lItxuttriartlv with iceCult is dibbling faors the survival oftillace Clre Such rineralied N released during the dr season aid made stils i aels tnd insects thtt pttpate or lay eggs- in the soil

available to the plants with the rains It iliso igrtmitcally Zero tilla hio eerl conserves predators ofthe GIH (37) advanttgeous for the rice ciop to tap this trlLItlaI fertility nt it mcttoi lo lessen the impact o0 soil insects and grow vigorotsly to compete with wrus and to tolerate brotdcasting seed or planting in ftltrrows is prefc ired over pest infestation dihlirig in hills Crickets are large enotlgh to eat seedlings in

Larly plantngs terd to escape seedling maggolts Upland a hill atrd tillering normnally ca iriot fill in tlhe missing space rice in Bitangas is planted in Ma and ine itli the Iirst Oftcn dibbll rd is not well covered and iseasily found by rains Fairners are highly mtotiated to plant early tocscape foraging insets ioIdents and birds drlitght and typlotns ricar harct Arn Atrgtst planting Plan uit sii Incrcatsini sced density protects tile cropwould be evcely attacked by It riim orivzo At tti againrst seed and seelling pests especially ants particularly other Philippiie tipland sites Rea l Qteot antd (laelit if the crop will germinate and ermerge quickly M isainis Oriental seedlig mnaggots arnrige ric boe t se hr trftinll itI India rice intcrclopped witi cotton or farnes have a longer growing season and plnit 2 rin or pigeon pci had lower 6IIt and WBIlII poptrlations than mote after the ortset of the rainy season liaaiangas ftrmners rice alone (211) NfailC arid Urplald ir_ 2re coilltori in the plant ca rly becia use ofa shorter raiinfall period arnd the desi re Philippines he chatngC in pest status folii iniercropping is to lliarvet before the typhoor sea son peaks ltighly locatioll-specific (133) cmianarid the net elfc be either

Early plantings cornbind with early-iatiuring varictics nil bcnelficial ortdetriential (189) For example BIatangias provide higher crp tolerartce for white grubs Aestiatior farmers intercrip Itro in their ipland ricefields A liornshyterminates with the first rains but 5-6 moare ineeded for the nrii iotd-Ahis i (I iiiacius) which norlially feeds resting larvae to develop into beetles emerge and lay eggs on taro ill feed or miarbv rice plants Intercropping and for larvae to reach the danriigii g ird-instar stage probably would not afft0 seed pests A crluanion crop (134 In lataigas this normually iseiough tiite for harvest planted ith rice and haing gleater tolerarnce for root

It Brazil howecver delayed planting is a suggested dariage or being toxic nayact isi trapcrop to root-feeditig cultural control for the lesser cornstalk borer and insect pests Acial-fei i ig pests voultId be most affected by pests favored l drought (66 195) Cropping internsity in intrer ropping Shoot aphids leaf beetles sten borers and Brail is low and delayed plraing places the rice crop in the hopplers would be likely ca ntidates to note effcts On the niost stable rainfall period reducing likelihood old rought othlt halnd the corlipalior crop stch~ sitniaie Iiiv be more

In rilniy arca s rinmirsare roi atl to plant other crops benehited than rice from initecropping ( 137) before rice and althNgh it woultl be desirabh to avoid 11echi Weeuling luring the first month after clop infestations by planting rice cuatl lariners lack the labor cstablishent will force pcts such is anyvworris which and time to do so ol a low priority crp prefer grassy cetds ontor tle vorling rice Soil iampsecs will

77age Uplild rice Stils ae light textured and Call be mvnye to the rice crop Cleani citrltirc also lay lore territes plowed duiring the d rv seasor notl only to grin time for to attack I u cetincti inrice ceds i ay he le optinal planting with lie fitst Inilsoon riiains but also to desiccate solition 13) soil insects arid weeds such is ntsedge that foim rhizomes Irtlizr Itneatsing N Iciiliier oin hlwland rice has Htowever becta use most soil insects lie below die plow layer fAtwvored Irigher populains of pliithoppeiv Imlhoppers

26 IRPS No 123 January 1987

and leaffolde rs However because of the need to temper fertilizer use on upland rice o prevent blast fIrtilizer rates will be lower than is comonia r for lowlad rice causing relatixely less hopper and IaIfTfoide r population buildup However inupland rice areas without a history of blast such Is Batangas farmers apply high rates of N and conscqtrently the crop has high infcstation levels of WBPI]and RIF The fhrmrcrs leec insect buildup by splitting the fertili11r in0to two to three applicatiois bit do not applV fertilizer at the vegetati estage because this prodrces a tall crop which illeasily lodge during typhoons Greater crop fertility leads to greater insect survival (better nutrition) larger insects (to cat no re foliage) arid greater fec titd t (to lav nmre eggs) (128 145 171) Not all insect species respondequally hovr Increasing rates (if N l anrid K increased Dialraa s-acchralis and (iaeiurema flea beetle but reduced thrips (66) Zn rcdLuced deadheart density from the lesser cornIstalk borer

Fnldingi Although most Upland rice areas do nIot hawxcaccess to wa ter floodiig the fields is rRrrnriierdcd in areas of Brazil or root bug and white gib control ( 195)

Crop rotatioti Tillage before planting a crop after rc harvest will control stern boreis and other pests renain n the stubble arid will unearth soil pests Planting a non-gratriiaceous crop after rice is recommncrded for termite control (66) Crop rotation is less nttdcd in upland rice culture than in lowlands because atcrop-free period is assured even after harvest of crops following rice

t1ulchilt iryl plastic irli irig designed for weedcoitrol ii upland rice inJapan resulted in ntr highanri liy infestation of (ihilo sulytresvais (93 9-1 95) NIulched Upland rice grew more luxuriantlv and attracted more (Avipositing mothis than u lched rice Also the rice plant and stein borers imatoured 10rC quickly and a partialgeneratiori af vouing ista iNoverwintcred in the stubbleLotili--malitrt varieti Quickly maturing crops reduce the tmbCr of pCst generations that Calt build up Rapid crop establishment olan ca rl v-natu ring vaitvty if)pa 11icLa r will lessen white gib and stem borer damiage White grub larvae intle last iistar are highly destIruCtive so aii early-iiatri ng variety will escape serious root loss Stem borers build 1ip slowlyx in rice to ecormic abundant in a late-Maturing crop anrd caiuse whi tlieads Numibers of somespecies increase exponeitially with each succeeding genera-ti so a xarietyexhat atures I trowh earier thian another will hac less insect damage Early-naturing vaiicties however will not lessen seed pest daiage either ii the soil or oina standing crop

Strwhoi hAIJttam Inrscts tcd irig on thlie aerial portions of rice plats disperse from carly to late plantedfields AS iiiibCirs increase exponentially with eachinsect

gtscnrattm e eiglbrig f rmc~s1(uil upp wh rc generation pests build tIP where neighboring fihrTC e9r1sstagger their plantings II Batangas farmers plant uplandrice within I iro use 120-d s-arieties and plant at the start of the rainy season Ini Misarnis Oriental laricrs stagger their

plantings up to 3 mo apart and use a set of atieties that mature in 4-7 mo Insect pest danage particularly from stem borers is greater in Misarmis Oriental Also seedling maggots damage rice there because rainfall favors frequent planting of maic In Batangas rice escapes seedling maggots bCca usC of distinct drs season and synchron0us early planting with the onset of ains

Therefore combinations of cultural practices ea rlyplanting synchronous planing crop rotation and earlyshymaturing vaneties protect the rice crop against most insect pests

Ilmant resistance Beca tse upland rice has few specialized insect pests and many insect types attack the crop worldwide regional breeding and strong internatiornal cooperation are required

Sown-sced pests arc economically controlled by insectishycide seed treatment and should the relore receive lowxpriori ty inbreeding objectives

Root pests are normally cost ly to control with insecticides but finding resista t sonres mayvbe di fficult Tanaka (229) for exa -ple fi led to find varieties resistant to root aphids Selecting for large root bioniass perhaps should be the strategy for pests sLuch as Whlite grubs that renliove roots Fainter(174) howcvcr cites varietiesof crops tolerant ofor resistant t(i root pests Some sorghun varieties are high in cyanide so rice sorghun might incorporate broadshyspcctruri chemical resistance

Seedling vigor and drought tolerance are high prioritybreeding objectives so tupland rices should have large r root systernsand be able to tolerate ligherlevels of root loss Sonie root loss inolder plants is beneficial if it stimulates new root developnrut and therefore crihances intake of soil nutrients (53) The rice seedling inaggot rtiLotia spp may be

controlled thirough resistant aittics A hi rgc breeding effort on sorghniu has had fair success inideveloping arreties esrstait to Asoccata Rodani (251) A field trial comparing 10 upland ricc varieties showed differences aniong varieties va ring from II to 4(- damaged tilleis (215) Hovver resistarce to 1Iill-0r gtia species may rot cross ox-rti otier

Shiraki (210) reported that upland i-ice stens wcrc harder than lows-land rice stemsperhaps becaIusc (f a higher silicoricontent and thus were more resistant to yellow stern borer hc tortia Iow tillering (f uplaid iice xarneties prodtres a

higher percentage of infested tillers than would occor in higher tillering lowland iice under a siiilar egg density The corollary therefore is to select for high tilleriig upland rices

Stemii bor amcstheh main targes dobrecders inAfrica (571 204 )but nticl oifth e sc reen irig is dotie unde r lo wland co d t osrm s wih A h r~ o l pe i s n acorditions (11 96 218) As with lthrinrnwspecies mayhave to be dealt with independently however initial results show cross resistance between African and Asian species(92)

In Brazil screening Ior insect resistance focttses oil lou I pest problems Varieties tolerant of lasmolwalpus ligi-sellus and resistant to )iatraea saccharalis have been identified (70 153) Current work also emphasies 7iraca limativetntris and Thmialspis (=Iosis) flavouicta (66) These insect pests Ilt onlV citise IIigh yield losses tihey also ire difficult to control chemically ofr by other ilealls

ManV upland rice vaieties aic early maturing and are resistant to pests such as gall midge (I Xl ) Early mIturity is often a highly desirable trait to esuilpe pest buildup but it is not true resista nce bca usc ilthose va reties are plinted late they will be itsdamaged as ain early planted susc ible variety

P~erhaps tile cosiiiopolitall of all upland rice pestsimost are the sced pests (ripenling stagc) which pose a particular problem beta use scuds re ised IsIO for hitnit4ans Ayltoxic substatice introd tcld geticticllyilmay also be toxic to nal iniless it caln he placed old in thfe IdhuffSelectmi fo hairiness or lotng wns kill mIlct sthilrohleiiis dutilig threshing pa rtictfarl maitial threshinfl [hireshers ill complain of itcfimin atid itteascd (fust levels

Attempts itIRRI to fitid tcistatncc to the rice bug plorisa have ilotbeCnl prmitisilg lIIetIattonlids iorc

through the lenilna or palca hillhairs sedstVIor seeds wilh awns that deter bugs promote complaints from threshers Seed bugs liptocorisa and Stcnocoris enter the seed through the opening between tile lemna and palC but varieties with narrow openings are diifficult to mill

Breeding priorities should concentrate ott inotCI gruLtLIps that are most difficult to c nttrol by other meains Steil) horers appear to be the most widespread group that sigiiificantly dalag s rice aind is hard to manage by other Ilnca us

Biological control Very fe stutdic of mitatt~l eneties of rice pests in upland envirollnletits wee in tile wefouiid literature Therefore have not compiled a list of species Ve present a list of natural enemies iOtid in upland ricefields tuic latialln Batangas for some conitno iOldiar pest species ([itble 5)

Muich ofl thc lollowinle itfor ittiol cotlliCs rol Stutdies in the lowlatnds or roi crops otithcr thlin tice

Qu(ailtr o 11ll t11t( Nat trrtf lii hcoeits ight Ile used agaitist ile social insects tilt and termites Sollie predator specice live ill theirf nests n ilnic theirnllt ip-pearanIlCe aMRidbehavior IJtrouMdCing a mote aggressise ant species to displace pest ant codonies is a possibility (169) Patihogens as potential natural encnies have reccived little atteition (I13) Natural ncnuics asa group however hac not been toIl iporta it in Iegulating aintshown be and ternite numbers

The nocturnal behavior aid subteitmien habitat of lole crickets and field crickets protect them from many ntllral enemies There are only limited records of natural cnelics for these groups of pests some pathogens (87) but mainly

IRPS No 123 JattuIny I 271987

parasites - scelionids on eggsand sphecids on ilymphs and adults Sphecid wasps Larra carutmaria (SImitI hI L huzwuensis Rohwer and lsaltiuhta Willi ams speciali in mole crickets pa ralyzing them nd dragging the ninto thcir nests as food lot their voting ()tier sphCcid species Lins aurulenoa (Iabricius) lot maiac (Ashmead ) 1 lhshylssellaus (Smith) and Ml lal)oriosu (Snithf specialie in field clickets

White grUbsO the other hand hae Icti scriuns enuigh to have been extensively studied for cotttroi 1W nati al enemies( 112 140 141) Scoliid parasites C(ajsoMcispp have beetn intrtduced itto tilePhilippites for control of Ielxl ]udisiro la The rcsearchcts w Intttrdced themle during ill era claimed that the white gttionthreak as controlled (142) howevcr it isnormal for pest epidemics to subside and there was no diect evidence to indictte that tfe parasite was responsible A tmore promisitg titctllod might he to identify iso ltcttass prt~f tcartd apply hiactelaln fingal pathogens to tile soil Ihe lalattese beetle in North America has hcci coitrolled hv tite conmnleciall aailable hactcrial prcpa ration itacihlpu pav ofr milksdiscas (71) Spote prtyefott the soil itloted ilrtier reitiain actic fr Veas

Root aphid sardtnctly bugs readilslic tilib to predatots and parlsites once their tetndine ants are controlled The larvac and adtilts of a coccinellid beetle 1Scynms sp prey on llriadlouais in slash and burn upland rice Baits treated with insecticide may be used to control tending ants by killing their ourig (244)

Root-tkcditig blugs false wireoris wireorms and root wccils aic r ofsuch little ifilpoltatice that thenomttll role of liatural Cticniics hias tiot beei asscssed A possibility hoevrl tiglt he tit ItaIIitic niaCIMtodcs lintt1g11en Itese soil-itlhlfbitine mCtitodcs W5uhtfhave to be imiass Produiced

large miltibcr of niatltrll cnemics ircknown ft CltIoriS IC IpblCeii is thtt clttvortnlls ofonlie the crop soott alter land ptel)tritiOti a1tithe bcgit iniig of the raittv

Citsoln hfien tiatllral leniiiy populations aric lo Colltloil WOtuld iii ole mass prodtticing andirclcaslng key spccies

le egg stage is the fpirt (olthe Cedlitgt Ialot life cycle tI hich to f)ctis ftor tittlril elctit cottntrol hplraites o1 pleditors ttsit elleggs highly expoIsYCl I Sgg pailsites or predators stolld probahly e ittass andtbe produced released Littlh i toruialioll is a ailable ott what species to trV o ho to Iniss podLice thiCIi A etiloplhid wasp Icmihlaol sp effctiicly regushy

lates the leatf lcloalapolm2a spioanoiiitr heat andc rice antid should be collsctvcd

leaf beetles whCllose e am] eggs ate laid in the grutindhirO ould be motC difficutlt to control by niatulral enelV Mnll ipuhltiOn But species sUici IS c_gsthispid beetles ith

and larVle oilleascs ate vtilcratble to iparaites and pledators

Natural enemisc of tlirips otl rice are little known

28 IRPS No 123 January 1987

Table S Parasites recorded from upland rice insect pests in Tanauan Batangas Philippines 1977-84

Order Family Species Host stage

I lymttenoptera Braconidae Leaffolders Cnaphalocrocis Marasinia spy

Cardiochilesphilipplinensis Ashmead a Larva

Cotesia angustillasis(Gahan)Cotesia or cypris (Nixon)

Larva Larva

Cotesia or taeniaticornis(Nixon) Larva Cilonusinotakatae Munakata Chelonits SppAlacrocetrts philippinensis (Ashtmead)

Egg to larva Egg to larva Larva

Iclineumonidac Orgilts spp Isclutojppa hireator(Fabricius)

Larva Larva

Itoplectis naraigav(Ashmcad) Larva Tenelhtcha philippiensis (Ashmead) Larva Tenelchha stangli (Ashmead) a

irichomma cnaphalocrosis Uchida Larva Larva

Bethylidac Elasmidac

Goniozms or trianguli r Kieffer lasmtusalhopictus Crawford

Larva Larva

Diptera

Encyrtidac Chalcididac

Tachinidac

Elasmtus spp Copidosomopsisnacohiae (Fidy) Brach nteria evcarinata Gahan Biachyieria lasus (Walker) Ztgohothria ciliata (Wulp)

Larva Egg to larva Pupa Pupa Larva

Argyrophylx nigrotihialis(Blaranov) Larva

l-ymenoptera Braconidac Yellow stern borer Scirpopiaga incertulas

Braco chinensisSzeplegeti Larva CIthelotis spp Chelonus niunakatae Munakata Cotesi flaripes Cameron Stenoracon nicerilli (Bingham) a

Larva Egg to larva Larva Larva

Ichneumonidac Tropobracon schoenobii (Viereck) Amatroinorpiaaccepta metathoracica(Ashmiead)Frihorus sinicus (I lolmgren)

Larva Larva Larva

Ischnojoppaluteator (Fabricius) Larva Isotima nr dammeratiti Rohwer Larva Tenelucha philippinensis (Ashmead) a

Larva Tenehicha stangli (Ashmcad)Trichomma enaphalocrosis Uchida

Larva Lara

Chalcididac Eulophidac Trichogranmmatidae

Xanthopimnpla stemmator (Thunberg) Charops hrachYpterumn Gupta and Malteswary Brachtm eria sppTetrasticttus schoenohii Ferriere Trichorigranm chihunis Ishi

Larva to pupa Larva Pupa Egg Egg

Diptera

Scelionidac Pteromalidae Tachinidae

Tt ichograintnta aponiicuitt (Ashniead) Telenomits rowani (Gahan) a

Trichoinalopsis apattelctena (Crawford) Jeirhaea spp

Egg Fgg Egg pupa Larva

Z4gobothria ciliata (Wulp) Larva

Ilymenoptera lecneumonirlac Gol fringed borer (hilo auricilius

Trichomm enoptalocrosisUchida Larva

Ieioenoptera Braconidae

Aantholimlh steiniator (Thtinberg) a

Pink stet Ioret Sesamia inferens StoInacoil nicrilhi (Bingh am)

Loarva to pupa

larva

Diptera Ilyinenoptera

Armvtnyorms Mythimna separata Spodoptera mauritia acronyetoidesTachinidae Zvgohothriaatropiora( Robinea i -Desvoid y)Eulophidae It-phctrtiscitapadoe (Ashimead)

Larva Larva

(halcididae Braconidac

Brachpin ria spip (oesia spp

Larva larva

IHyntenoptera I)iptera

Rice skipper Ielop ida (halcididae Tachinidae

math ias aidgreen iortedci ciliarNIelan itis eda Brat lhincria sp iii matuqitatt Cameron Irgyrolhirlaxi trotlhiais(marumtv)a1

ismtne Larva Larva

Diptera Ilymenoptera

Tachnidae lelineutionidae

fuown se tilo per Mois frugalisIrgtrophrlax 1igrothiahiv 3 a rat ov) anthopimpa pmnctata (Iabricius)

Larva Larva to pupa

aMost dominant

IRPS No 123 January 1987 29

Normally predatory thrips prey on herbivorous thrips strepsiptera on both Spiders dominate tilenymphaladultNatural enemies of stem bugs --Scolinoplara and Tibraa predators Spittle bugs move to rice from pasture grasses - are in order of importance egg par~isites and predators and biocontrol would be successful only ifcarried out before fungal pathogens and nymphal adult parasites Egg tiley immigrate into ricefields parasites could either be introduced fron other areas or Biological control tactics for hemipterous seed bugs mass produced and released ould follow those outlined for stein bugs

Armyworms are normally lwld in hcck by the activities Coml)arison of environnewtns Natural enemies are of egg and larval parasites When these natural enemies fail perhaps faced with even greaterconstraints than insect pestsusually because of drought armyworns become epidemic to survive ind reproduce in upland environments TheyIt may be worthwhile therefore to release parasites during have adapted ani survived by having wide host ranges the the rainy months following a drought Virus diseases of ability to aestivate or hibernate over unfavorable seasonslarvae would beanotheravnue to exploreforarnyworms and the ability to disperse or by other mechanisms

Parasites have ben traditionallyconsidered for biolegical Comparing the most dominant and therefore the most control attempts against stein borers usually by rekasing adapted species of natural enemies collected in various exotic species( 166) This approach has not met with success environments in the Philippines we can understand how and new avenues should be explored The role of predators different rice environments are arid pathogens is little inderstood and deserves greater Anagqrms olptahiisa mymarid planthopper egg parasiteattention Egg parasitism rates are normally reasonably is cquallyadapted to all Philippine rice environments (Tablehigh but should be supplemented with effective egg 6) It is selective for planthoppers and even parasitizes eggspredators Orthopterans are potential egg predators of the n-iize planthopper Peregrinitsmaidis itt upland areas Metioche vittaticollis (Sta ) Anavipha nifiwnnis (Saus- Alavohis also parasitizes maize planthopper eggs but is sure) and Conocephahs hongipnni (de i-laan) feed on most prevalent on BPH in the irrigated wetlands (onashystem borer eggs etioche and Anaxil)ha specialize in eggs tocerus spp like A optalhilis on BPH and WBPI Iattacks with no hair such as Chilo spp The omnivorous (ono- the eggs of GLH in ell rie environments It is also found cephahsspecializes in eggs cov red with hairs it is a proven parasitizing the eggs ef the white rice licahopper (oimna egg predator of Scirlouphaga spp but has the discouraging spetra Oirosita naias a trichogrammatid egg parasitehabit of eating rice guin (65) attacks BIPH and WBPH in the wetlands and has no other

Grasshoppers have egg (scelionid) and nymphalad ult host A related species 0 aesopi specializes in G1-H in (nemestrinid tachinid and sarcophagid) v irasites Among wetland environments Dryinid strepsipt-an and pipunshypathogens protozoans have been recorded most frequently culid nymphal and adult parasites of planthoppers and Control of locust species by natural enemies could con- leafhoppers also specialize either in hopper species or centrate on their habitual breeding areas environments Among the dryinids Pseuhdgonatoipsflavi-

Leaffolders have rich complexes of natural enemies some feimur prefers the lowlands and BPH while P nuch has no adapted to upland rice (22 27) ranging from egg predators environmental prefirences but is adapted only to WBPH (gryllids coccinellids) to larval parasites (braconids ichncu- The main d ryinid species on GLH aptogonatop)usspp is monids) to larval prcdators (ants and carabids) to larval most dominant in rainfed lowlands and also attacks other pathogeiis (vira and fungal diseases) Efforts to augment leafhoppers within Amnrasca Cicadulinaand Bachaltuathese natural enemies should focus on the egg predators Strepsiptera are more dominant on plarithoppers than (rearing releasing)and viral or fungal pathogens (culturing lealhoppers The Elenchus species in the Philippines attack spraying) all species of planthoppers within the genera Sogawilla

Parasitization of the large conspicuous larvae of Sogatodels NilaparsataHarualia and Opiconsiva Elenshyskippers green horned caterpillars and other polyphagous cusrvasuntaisuiis adapted to the major rice environments Lepidoptera by tachinids and chalcids is normally low ELg but prefers BPH in rainfed arid WBPIH in wetland environshyparasites and predators are perhaps the key to their control ments Ilalictoph(utsmtnroeiand HVpectrus attack G LH

Gall midges are parasitized by pterormalids and platy- and COfimravcctra respectivelygasterids that aestivate within their larval host between Pipunculids are mainly parasitic oii lealhoppers Each seasons Shoot aphids are preyed upon by coccinellids species has a unique environmental preference Tomosshysyrphids and chrysopids varireIa oraetoraprefers Ne7 otetltix nigropictrs and

Planthoppers arid leafhoppers have egg parasites and Pilunwulhtsjavanensisattacks Dehoceplhahls spp which predators as well as nymphialad nlt parasitesand predators are more abundant in upland than irrigated lowland their rich complex of natural enemies includes fungal environments T suhvirescensattacks Nephotettix virescens pathogens The principal egg parasites are myrnarids and and prefers the lowlands Pilpuiulus nttillattls is an trichogramniatids Egg predators are norinally mirid plant upland rice parasite arid attacks only Nephotetix speciesbugs that suck out the yolk Nymphal ad nIt parasites are More than 17 recorded species of leaffolder parasites in dryinids on planthoppers pipunculids on lealhoppers and the Philippines attack species of C(naplwocrocis and

30 IRPS No 123 January 1987

Table 6 Dominant natural enemies of rice pests in upland rainfed lowland and irrigated lowland rice environments of the Philippines

Natural enemy

Host stage Species

Brown planthop-Parasite Egg

MymaridaeAnagrus optabilis(Perkins) Anagrusflaveolus Waterhouse

TrichogrammatidacOligosita naiasGirault

NymphAdult Dryinidac

Pseudogotatopuslaiifemttr Esaki et Hashimoto Elenchidae

Elenchus yasurnatsui Kifune and Hirashima

Environment

Upand Rainfed lowland Irrigated lowland

Ailaparata lugens

x x X x

x

x x

x X X Whitebacked planthoppers Sogatella furcifera and Sogatodes pusanusEgg

MymaridaeAnagrus optabilis(Perkins)

TrichogrammatidacOl~gositanaias Girault

NymphAdult Dryinidae

Pseudogonatopusnudus Perkins Elenchidae

Elenchus yasutatsui Kifune et Hirashima

Parasite Egg

MymaridaeGonatocerus spp

TrichogrammatidaeOligositaaesopi Girault

NymphAdult Pipunculidac

Tomosvaryella subviresccns Loew Tomosvaryella oryzaetora (Koizumi) Pipunclusmutillatus (de Meijere)

DryinidaeIlaplogonatopusspp

Egg Copidosomopsisnacoleiae (Eady)

Larva BraLonidae

Cardiochiles philipphnensisAshnead lchneumonidac

Trichiomnia cnaphalocrosisUchida Temelucha stangli (Ashmead)

Pupa Chalcididae

BrachymeriaexcarinataGahan Brachymerialasus (Walker) Brachymeria nr tachardiaegroup

Parasite Egg

Scelionidae Telenomus rowani (Gahan)

EulophidacTettasticusschoenobhi Ferriere

Larva Braconidae

Stenobraconnicevillei (Bingham) Iclhncurnonidac

Temelucha phiiippinensis(Ashinead)Temelucha stangli (Ashmead)

x x x

x x

X x X

x x Green leafhopper Nephotettix virescens

X x x

x x

X X X X

X

X Rice leaffolders CnaphalocrocisMarasmia

x X x

X

X X

x x x x x X x x x

Yellow stem borer Scirpophaga incertulas

X X x

x

x

x X XAinauromorphaaccepta metathoracica(Ashmead) x

continued on next page

IRPS No 123 January 1987 31

Table 6 cntinued

Natural enemy Environment

Host stage Species Upland Rainfed lowland Irrigated lowland

Larva-pupa Gold fringed borer Chilo attriciiuslchneumonidae

Xanthopimpla stemmator (Thunberg) x aPupa x

Pteromalidae Trichoinalopsis apanteloctena (Crawford) ax

Pink stem borerb Sesamnia inferensParasite

Larva Braconidae

Stenobracon nicevillei (Bingham) x b b Armyworms Mvthinna SpodopteraLarva

Tachinidac Zygobothriaarroptvora (Robineau-Desvoidy) x x

Braconidac Cotesia spp

X Skipper Pelopidas mathias and green horned caterpillar Melanitis leda ismeneF-gg

TrichogramrnatidacTrichogrannaspp x

Scelionidae Undetermined Scelionid

x Larva

Tachinidae Argyroph lax nitrotibialis (Baranov) X x

Pteromalidae Trichomnalopsis apanteloctena (Crawford)

X Predator

CoenagrionidaeAgriocnenis femina femina Brauer x xIschnura senegalensis tambur

Miridae CrtorhinsliidipennisReuter x x

Nabidac Stcnonabis tagalicus (Stal) x

VeliidaeAlicrovelia douglasi atrolineata (Bergroth) x

Coccinellidae Micraspis crocea (Mulsant) X x x

Vespidae Eumenes campaniformis (Fabricius) x

Formicidac Diaecamna spp xOdontoponera transversa (F Smith) x

Carabidae Chaenius spp OphioneaW- aJasciataSchmidt-Goebel

x

Ophionea ishii ishii Habu x x X

Gryllidav Anaxipta longipennis (Saussur-) x xMetioche ittaticollis (Sial) x x x

Tettigoniidae Conocephalus longipennis (de Haan) x x x

aNo rearing made on rainfed lowland bNo rearing made on rainfed lowland or Irrigated lowland

MrasmiaAmong the Laval parasites the braconid Cardio- lowlands and parasitizes oriental maize borer OsIriniachilesphilijpphensis is most adapted to an upland environ- furnacalis (Guenee) larvae The leaffolder pupal parasitesment It also parasitizes Hydelepta indicata a common not only occur equally in all environments but also haveleaffolder of legumes The ichneumonids Trichomma cna- wvide host ranges- parasitizing species of OsIrinia Marucaphalocrosis and Temehcha stangli occur more in the Hydelepta and Honiona - in maize and legumeslowlands T stangli is most adapted to irrigated lowlands The principal egg parasite of yeltow stem borer (YSB)and also parasitizes Chilo and Scirpophaga stem borer Telenomus rowani is widely adapted to all environmentslarvae T cnaphalocrosis is most prevalent in rainfed and has an unusual alternate host -Tabanus eggs (25)

32 IRPS No 123 January 1987

Tetrastictu schoenobii specializes only in YSB and occurs mainly in irrigated lowlands -- further proof that YSB is most adapted to the wetter environments

Larval parasites of YSB are highly environment-specific The braconid Stenohracon nicevillei occurs only in upland rice but also attacks the pink stem borer (PSB) Tieinchwa phiiiplPwnsis an ichneumonid occurs in upland rice and also parasitizes the larvae of (7hiu stqpresaisinatro-morpha accepta melaihoracica like 7 stangi is mnost adapted to the irrigated lowlands

The dark-headed stem borer(l)HSB) (ih pochit3sus was not collected in all rice environments and the gold fringed borer Chilo auricilius nd PSB were collected only in upland rice Xaantiholiplastenrnaloran ichneumonid larval parasite appears to be widely adapted whereas TrihomaslopisapanitI)cteiiaa pteromalid pupal parasite occurs mostly in wetlands and also parasitizes larvae of Chilo suppressais and Pelhuidas mathias

Armyworms Myit hinin and Spodujipera have two main larval parasites - a tachinid Zrgohothriaatropivora in rainfed environments and a braconid Cotesia spp in irrigated lowlands

The butterfly pests Pelopidas and 4elanitishave tricho-grammatid egg parasites found only in irrigated lowland environments

Two larval parasites show environmental preferences --Argrroph av nigrotihialis a tachinid in rainfed habitats and Trichonialopsisapantelkcnela in irrigated ricefields A limgrotibialisalsoattacks the sweet potato hornwormAgrius convolvtdi and T apanteloctena parasitizes DH S1B

Predators as a group show more distinct environmental preferences The aquatic damselflies Coenagrionidae are most abundant in the lowlands Cvriorhinus is also most adapted to the lowlands The nabid Stenonabis tagalicus occurs only in the uplands and isalso prevalent on legumes The coccinellid lady beetles wasps and ants are more prevalent in upland areas AlicrasIpis cit-ea larvae prey ona variety of aphids attacking legumes and maize Etunienes campanonia vespid mud wasp makes nests in trees and is therefore most adapted to the more botanically diverse uplands Soil-dwelling ants cannot tolerate flooding

Upland environments are habitats to arboreal carabid beetles Three species of Chlaeniusbeetles prey on leaffolder larvae and are perhaps more important than parasites (27)

The gryllids and tettigoniids are widely adapted egg predators not only on rice but also on maize and legumes They feed on eggs of most insects that are laid on leaves

The spidercommunity of upland rice environments isrich in species At one site (Batangas Philippines) 31 species have been recorded (23) Of the 176 spider species recorded in Philippine ricefields about one-half (82) occur in upland rice The spiderspecies of upland rice environments overlap more with those of rainfed lowland than with those of irrigated lowland rice (24) The same study showed that of the three environments irrigated rice has (ie greatest spider species diversity foiiowed by rainfed lowland and upland

However of the 10 most prevalent ricefield spiders 9 were abundant in upland rice showing wide adaptation All three spider guilds -- orb-web space-web and huting spiders shywere prevalent in upland ricefields There were however differences in environmental prelerences for some spider species

Of the orb-wcavers Tetragniathamanlihulatawas partishycila rlv abundant 7 japonica was low and Ltucatge dehcorata was absent in upland rice

As in other environments Aiypeiwnaifirtosana was the most dominant space-web spider however its relative numbers were lowerthan for lowland sites Among hunting spiders Ox|opes javanus and Lcosa leatcost fgla were more abundant and Lvcosa lpseudoanmnlata was less abundant in upland compared with lowland rice envishyroniments

Extensive rearing of rice insect pests in upland rainfed lowland and irrigated sites over an 8-yr period in the Philippines provided insight into the effectiveness of parasites as natural enemies (Table 7 8) Most of toe data are from large sample sizes taken over at least one crop which overcomes the pitfalls expressed by Van Driesche (238)

VSB egg parasitization was surprisingly similar across all environments Larval parasitization was highly dynamic among sites and even years within the same site but showed somewhat lowerlevels in uplandsthan in irrigated wetlands This would have been predicted for egg parasites as well since YSB is adapted to the lowlands and so must be its parasites This isevidence that Telenomus rowani is as good at dispersing as its main hosts Trowani clings onto the anal tufts of female YSB and parasitizes eggs as they are laid This phoresy may explain its wide adaptation The more specialized lowland YSB larval parasites therefore appear more effective than their upland counterparts with more alternate hosts

Low collection levels of PSB larvae especially in the wetlands make comparisons between environments difficult Low levels of larval parasifization were recorded in both rainfed environments Except in the Cagayan rainfed wetland site in 1980 low levels ef larval parasitization were evident at irrigated and rainfed sitesalike No differences by environment were evident Higher levels of parasitization occurred in Sarawak during a 1967 outbreak also from tachinids Rothschild (197) indicated that higher parasishytization occurred in upland rice areas

Skipper and green horned caterpillar larval parasitization rates were generally low and showed no environmental effect htighest activity occurred in Cagayan in 1980 (48 parasitization) Although larval parasitization levels of RLF were variable year to year activity appeared greater in the irrigated lowlands than in the uplands Again the highest levels were recorded in Cagayan in 1980 Cagayan has extensive grasslands and fallow around the rice areas and a relatively long rainfall period Rainfall in 1930 was good for crop growth and apparently for increase of larval parasites

IRPS No 123 January 1987 33

Table 7 Parasitization rates of rice planthoppers and green leafhopper in upland rainfed lowland and irrigated lowland environments Philipshypines 1976-840

Environment and site Year Egg parasitization () Nymphaladult parasitization ()

BPH W13P GLH BPII WBPH GLIH

UplandBatangas

Rainfed lowland

1976 1979 1980 1982

15 17 12

2(47)8(39)

4(51)

9(1800)14 (212)

12(304)

8 (101)6 (127)

9(116)

Iloilo

Cagayan

1976 1977 1978 1981 1980 1981 1982

0 (69)

11

44 48 40

-

-

46 55 44

0 (471)

11

52 56 49

0 (200)9 (214)

10 (1259)8(235)6 (900) 3(800)

-7 (462)

11 (270)3 (313)5 (900) 1 (800)

4 (600)10 (725)10 (362)12(870)5 (900) 3 (800)

Pangasinan

Irrigatedlowland

1984 1976 1978 1979 1982

13 (436) 13(379) 14 (457)

6(14)7 (197)

17(198) 11 (486) 18 (275)

3 (81)10 (274)3 (116)

16(871) 19(88)

28(211)9 (764)5 (647)7 (1204) 8(305)

Laguna 1977 11 7 21 1978 31 1979 14 4 4 28 1980 7 8 20 1981 1983

56 7(278)

26 16 (1650)

16 27(1921)

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R I Apostol unpublished 1lo1lo and Pangasinan 136 Cagayan BCanapi unpublished Liuna 175 I C Pantua unpublished 118 234 43 Carino and Shepard unpublished BPH = brown planthopper Nila parvara htgens W131Pi =whitebacked planthopper Sogatelia furcifera GLH = green leafhopper Nephotettrix virescens Figiires in parentheses are sample sizes

Tauher et al (230) reported that internal external and genetic factors influence the seasonal activity of parasitoids

Hopper egg parasites but not nymphaladult parasites were also parlicularly abundant at the Cagayan site possibly the egg parasites have more alternative hopper prey in the weedy areas than nymphaladult paisites

The seasonal dynamics of hopper egg parasites can be seen for upland rice in Batangas(Fig I 2) and for rainfed lowland lice in Pangasinan( 135)and Cagayan( 105) Levels of BPHWBPI-L and GLH egg parasitization were corn-parable in upland and irrigated lowland environments Except for some high rates of paraMsitization early in the season the parasitization rate tended to be steady in all envilonments However hopper nymphaladult parasi-tizution levels were more similar in the rainfed environments and lower than in irrigated rice

Overall parasites do not appear highly effective by themselves against upland or even lowland pests Among natural enemies parasites have attracted the most attention because they can be more readily assesscd

Predators and pathogens lend themselves better to management practices Predators the most important group of natural enemies in riccfields are difficult to quantify On the other hand pathogens appear less im-portant but can be readily cultured and disseminated

The natural enemy commulnity of upland rice is rich in species and differs significantly from that of lowland rice

These beneficial organisms must be conserved by applying insecticides judiciously particularly sprays Seed or soil placement of chemicals minimizes exposure to natural enemies The strategy to derive the greatest benefit from natural enemies isto allow the greatest number of beneficial species to thrive Then perhaps at least one species will be effective against each pest at any tine overcoming the variable abundance of each species within and between years

Programs to introduce exotic species or mass produce indigenous ones are ambitious and expensive because they require trained people on a sustained not ad hoc basis

Chemical control Insecticides offer rapid and efficient control of upland insect pests (190) but-should be used only after other control measures have been considered Insecticides are rarely used on upland rice because of cost (32) Upland rice yields are normally too low to justify the expenditure Also most upland rice is grown as a cash crop Spraying upland rice is more difficult than lowland rice because water is less accessible Government subsidy programs to provide pestishycides to tribal peoples for upland rice production have created problems of toxicity to humans because isolated people have had no experience to be able to handle pesticides safely

However ins-eticide use has beenjustified when a crop is

0Z

Table 8 Parasitization rates of lepidopterous pests in upland rainfed lowland and irrigated lowland environments Philippines 1976-84a

Parasitization (5)

Environment and site Year YSB GFB PSB Earcutting Swarming Skipper Greenborned RLF Brown larva larva caterpillar caterpillar caterpillar caterpillar larva semilooperEgg Larva larva larva larva larva larva

UplandBatangas 1977 - 0 (57) - 0 (63) - 0 (71) - 0 (106) 11(142) 2 (101)1978 43 (4697) 8(451) 5 (38) 3 (40) 6 (61) 2(58) 15 (57) 13 (129) 13 (1286) 4 (30)1979 59(4135) 12(320) 13 (11) 5(28) 9(45) 6(46) 7(23) 8(160) 9(1152) 8(26)

Rainfed lowland Iloilo 1976 - 0 (10) 0(10) 0 (7) 0 (19) 2 13) 15 (180) 5 (76)1978 0 (54) 0 (3) 0 (17) 9 (64) 4 (78) 3 (73) 0 (121) 0 (191)1979 64 (2350) 16 (84) 6 (17) 0(6) 18 (68) 25 (104)

1981 56 (1322) 18(304) 0(5) 12(51) 18(138)Pangasinan 1976 60 (7 7 )b 62(82) 0(111) 5 (39)

1978 58 (53)b 5(51) Cagayan 1979 59(713) 8 (101) 6 (34) 0(7)

1980 0(11) 4 (28) 15 (286)

62 ( 48(177) 61(140)1981 0(5) 12 (2) 13(45) 14 (92)

Irrigatedlowland Laguna 1974 62 8

1974-75 50 41 1975 43 72 1978 96 (623) 21(24) 33 (60)1979 83(2017) 18(110) 8(62) 11 (89) 22(400)1980 78 (1625) 20(75) 0 (16) 4 (72)1983 60

9 (80) 7 (270) 30

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R F Apostol unpublished Iloilo and Pangasnan 136 Cagayan B Canapi unpublished Laguna 175 P CPantua unpublished 118 234 143 Carino and Shepard unpublished YSB = yellow stem borer ScirpophagaincertulasGFB = gold fringed borer Chilo auricilius PSB = pink stem borerSesamia inferens earcutting caterpillar Alythimna separata swarming caterpillar Spcdoptera mauritia acronyctoides skipper Pelopidas mathias reen horned caterpillar Melanitls ledaismene RLF = rice leaffoldersCnaphalocrocismedinalis and Marasmiaspp brown semilooper Mocisfrugalis Figures in parentheses are sample sizes Egg masses

IRPS No 123 January 1987 35

Table 9 Effect of seeding rate and insecticide on plant density and Yield losses from insect pests in upland rice often areyield of Dagge upland rice Batangas Philippines 1976a comparable to those of lowland rainfed or irrigated rice

Seeding rate Insecticide Dosage Plant density Yield (0-Y30o) but usually from different pests The principal (kg aiha) seed treatment (kg aiha) (noIra-row) (tha) groups are soil pests feeding on sown seed and rootsat 14 DE followed by seedling pests These groups have largely gone

50 None - 35 a 20 a unnoticed leading researchers to conclude that stem borers50 Carbofuran F 10 38 ab 23 a and seed bugs are the most important pests (17) Yield loss 10 beCabofuranF100 None - 4551 c 23 a trials that pinpoint major pests need to includc seed

100 Dieldrin WP 10 52 cd 22 a treatments Granules applied in the furrow may not give150 None - 59 d 22 a adequate protection from many seed pests

aAv of 8 fields DE = days after crop emergence In a column means The floristic and pedological diversity of upland rice followed by a common letter are not significantly different at the environments is matched by the diversity of arthropods5level by D)MRT F = flowable Wi = wettable powder both pests and their natural enemies Irrigation has homoshy

genized lowland rice environments Flooded soils tend tothreatened by epidemics of locusts or armyworms New have the same properties and to eliminate soil pests The technology stressing efficiency has minimized insecticide spatial and temporal dominance of cultivated rice has cost benefited specialist pests

Seed treatment is normally inexpensive and can be Upland rice has no specialist arthropod species The economicallyjustified in -nanycases(66) Treating seed with monophagous pests on upland rice prefer the lowlands systemic insecticide (216) but not chlorinated hydrocarbons where rice evolved Rice in the uplands exists only because (247) is most effective in controlling seedling maggots of man and even though lowland rice predates upland riceFoliar sprays have to be repeated to obtain control and specialist species evolved Wheat maize and sorghum do often are washed off by frequent early season rains not have monophagous pests leading us to conclude that Granular insecticides are effective but required dosages are there is no evolutionary advantage for inssects to have too expensive to justify except under heavy infestation narrow host ranges in upland graminaceae(247) When directed against first-instar larvae banding Pests have adapted to the uplands not only by having granules in seed furrows (177) can be effective against white wider plant host ranges but also by having long life cyclesgrubs at low dosages (025 kg aii ha) (134) Seed treatments undergoing dormancy or being dispersive Each pest has a are not promising for white grub control (248) unique life history strategy suited to highly unpredictable

Seed treatment is also effective ainst ants and may be environments In agivenyear someinsect specics an better cheaper than increasing the seeding rate In Batangas exploit upland rice as a food source but no pest is a senious treating 50 kg seed ha was as effective as increasing the one every year on upland rice seeding rate to 100 kgha and using untreated seeds (Table The low yielding potential of upland rice means less 9) In Brazil seed treatments control termites and other soil quantity and lower quality of food for pests to exploit but insects including the lessercornstalk borer(66) Domiciano also less incentive for farmers to undertake control (58) however found seed treatments inconsistent in their The most effective control would be early and synchroshyeffect against high populations of lesser corn stalk borer nous planting of early-maturing upland rice varieties to

Baiting can be an inexpensive way of controlling ants prevent insect pest buildup Breeding for insect resistance seedling maggot flies mole crickets and field crickets Baits should initially focus on stem borers because they are verymade of locally available material and impregnated with difficult to control by other means Biocontrol efforts insecticide can be sparingly distributed taking adantage of should first stress conservation of natural enemies preshythe pests ability to disperse and encounter bait sites dators and pathogens rather than parasites Efficient

The new low-volume hand-held controlled-droplet chemical control includes seed treatment or baiting for seed sprayers may offer an advantage over high-volume knap- seedling and some root pests and using controlled dropletsack sprayes for controlling foliar pests Upland rice sprayers for spot treatments against foliar pests and seed canopies are more open than those in lowland rice and bugs when economic thresholds are reached better droplet penetration should result Controlled droplet applicators would be ideal for seedbugs and defoliators

CONCLUSION

Upland rice is attacked by a wider array of insect pests than is lowland rice mainly beca use of the addition of soil pests Generally population levels of lowland rice pests are lower in upland than lowland rice leading many people to conclude that insect pests are not important on upland rice

36 IRPS No 123 January 1987

REFERENCES CITED

1Abraham C C and K S R Mony 1978 Occurrence ofLeptocorisa acuta Fabr (Coreidae Hemiptera) as a pest of nutmeg trees J Bombay Nat ttistSoc 74553

2 Agyen-Sampong M 1975 HlierogrlphusdiaganensisKrauss (Orthop-tera Acrididae) a new pest of rice in Northern Ghana Ghana J Agric Sci 8249-253

3 Agyen-Samnpong M 1976 Varietal resistance and other inle-ratcd pest control components as factors of rice varietal improvement in West Africa Prcsented at the Second varietal improvement serinar 14-18 Sep 1976 Bouak Ivory Coast West Africa Rice Development Association Monrovia Liberia 18 p

4 Agyen-Sampong M 1977 Outbreak of Ihccrorrdnisorae in rice seedlings near Rokupr Sierra Leone lit Rice Res Newsl 2(6)16-17

5 Agyen-Sampong M 1980 Dirty panicles and rice yield reduction caused by hugs IntRice Res Newsl 5(l)11-12

6 Agyen-Sampong M 1982 The major pest problems of irrigated upland and mangrove swanip rice ecosystems ir the humid tropical Guinea savanna and Salel climate zones Pages 318-328 in Pro-ceedings training course cn concepts techniques and application of irtegrated pest management in rice inWest Africa 10-20 Jan 1982 Fendall Liberia West Africa Rice l)evelopioent Association Monrovia Liberia 505 p

7 Akibo-Betts V I and S A Raymundo 1978 Aphids as rice pests in Sierra Leone lit Rice Res Newsl 3(6)15-16

8 Akinsola F A 1975 Present status of different rice stemborers in parts of Nigeria iicL Entomo Newsl 328

9 Akinsola FA 1979 Ihe biology and ecology of rice stemborers in Nigeria Ph 1) thesis University ol lbadan Nigeria 217 p

10 Akinsola E A 1979 Guidelines of integrated management of insect pests of upland rice in West Africa Presented at the Seminar on integrated management of rice diseases and insect pests at Bobo-Dioulasso Upper Volta 17-22 Sep 1979 West Africa Rice Develop-ment Association Monrovia Liberia

II Akinsola IA 1984 Insect pests of upland rice in Africa Pages 301-305 in An overview of upland rice research Proceedings of the 1982 lIouak Iwsrv Coast upland rice workshop International Rice Research Institute 13O Box 933 Manila Philippines

12 Alam M S V TJohn and Kaung Zan 1985 Insect pests and diseases of rice in Africa Pages 67-82 in Rice improvement in Eastern Central and Southern Africa proceedings of the international rice workshop at Iusaka Zambia 9-19 April 1984

13 Alticri M A A van Schoonhovcn and J D Doll 1977 The ecological role of weeds in insect pest management systems a review illustiated with bean (Phaseohsvulgaris L)cropping systems PANS (Pest Artie News Summ) 23195-205

14 Anmante F 1967 The leaf-cutting ant Atia capiguaraa pest of pastures [in Portuguese] Biol6gico (Sao Paulo) 33113-120 (Rev Appl Entomol A56207)

15Andrade A C and D Puzzi 1951 Preliminary results of experi-merits on the control of Sculptcoris iastantus on sugarcane [in Portuguese] Bioulgico (Sao Paulo) 1744-49 (Rev Appl Entomol A41421422)

16 Appert J 1970 Maliarplia separaelha (rice stem borer) new observations of entomological problems of rice in Madagascar [in French] Agron Trop (Paris) 25329-334

17 Ariadeau Mand Z Harahap 1986 Relevant upland rice breeding objectives Pages 189-197 in Progress in upland rice research Proceedings of the 1985 Jakarta conference International Rice Research Institute 1O Box 933 Manila Philippines

18Asar S and Kisimoto 1975 Seasonal occurrence of the smaller brown planthopper in upland rice [in Japanese] Proc Kanto-Tosan Plant Prot Soc 2228

19 Bakker W 1974 Characterization and ecological aspects of rice yellow mottle virus in Kenya Meded Landbouwhogesh Wage-ningen 152 p

20 Balut F F 1970 White grubs attacking rice Oryza ativa L [in Portuguese] Biol6gico (So Paulo) 36321-322

21 Banerjee S N and L M Pramanik 1967 The lepidopterous stalk borers of rice and their life cycles in the tropics pages 103-124 in The major insect pests ofthe rice plant Proceedings ofa symposium at the International Rice Research Institute September 1964 The Johns Hopkins Press Baltimore Maryland USA

22 Barrion A T and J A Litsinger 1980 Ants a natural enemy of cliaiarac in dryland rice IntRice Res Newsl

5(4)22-23 23 Barrion A T and J A litsinger 1981 The spider fauna of

Philippine rice agroecosystem 1Dryland Philipp Entomnol 5139-166

24 Barrion A T and 1 A Iitsinger 1984 The spider fauna of Philippine rice agroccosystcms II Welland Philipp Eiitomol 611-37

25 Barrion A T1 and J A Litsinger 1984 lihanus (Diptera Tabanidae) eggs an alternative host of rice stei borer (SI3) egg parasite Telenoonis digns (Iliymenoptcra Scelionidae) lit Rice Res Newvsl 9(6)19

26 Barrion A T and J A Litsinger 1985 Identilication of rice leaffolders by wing markings Int Rice Res Newsl 10( 1)24

27 Barrion A T and J A Litsinger 1985 Chctiosspp (Coleoptera Carabidae) a leaffolder ILF)predator lit Rice Res Newsl 10(1)21

28 Bertels A 1970 Insect pests of rice and theircontrol [in Portuguese] Pelotas Brazil Instituto de Expcrimentacao Agropccuararias do Sul Circular 43 24 p

29 Bonzi S M 1982 Ctilo (flisilineus J de Joannis (Lepidoptera Pyralidae) stem borer of irrigated cereals in Upper Volta LioFrench] Agron Trop Riu Rizic Cult Vivrieres Trop 37207-209

30 Bourke T V T L Fenner J N LStihick G Baker EHassan D F OSullivan and C S li 1977 Insect pest survey for the year ending 30th June 1969 Papta New Guinea )epartment of Agrishyculture Stock and Fisheries Port Moresby 57 p

31 Bowden 1 1973 Migration of pests in the tropics Meded Rijksfac Landbouwwet Gent 38785-796

32 Breniere J 1973 The main rice parasites in West Africa and their control Pages 27-42 in Proceedings seminar on plant protection for the rice crop West Africa Rice Development Association Monrovia Liberia

33 Breniere J 1976 The principal insect pests of rice in West Africa and their control West Africa Rice Development Association Monrovia Liberia 52 p

34 Brown ES 1962 The African arinyworn Spoiolptera eXI)Ina (Walker) (Lepidoptera Noctuidae) a review of the literature Cornshymonwealth Institute of Entomology London UK57 p

35 Brown F B 1968 Upland rice in Latin America FAO-International Rice Commission 12th Session of the Working Party on Rice Production and Protection 2-14 Sept Peradeniya Sri lanka Food and Agriculture Organization Rome Italy 6 p

36 Buyckx F J 1 1962 Disease and insect pests of crops inCongo Rwanda and Burundi [iil French] Publications dc LInstilut National pour lestude Agrononiquedu Congo M Weissenbruck S A Brussels Belgium 708 p

37 Cariiio F 0 1981 Role of natural enentics inpopulation suppression and pest management of green rice lealhoppers Ph D thesis University ofthe Philippines at Los Bafitos PhilippinesLaguna 136 -

38 Carpente A J 1983 The development of rice cultivation and extension of vegetable productin in Zanzibar Final Report Vol II1 October 1983 Food and Agricultme Organization Ronte Italy

39 Carroll C P and S1 Risch 1983 Tropical annual cropping systems ant ecology Environ Manage 751-57

4(0 Cates R G 198 1 Host plant predictability and the feeding patterns of monophagous oligophagous and polyphagous insect herbivores Oecologia (Berl) 48319-326

IRPS No 123 January 1987 37

41 Cendcafia S M and F B Caora 1967 Insect pests of rice in the 62 Ene 1 C 1981 I)iiorldhol)ierusihilis Slater (licteroplera Philippines Pages 591-616 in The najor insect pests ofthe rice plant lygacidue Illissinac) a new rice pest front Nigeria lit Rice Res Proceedings of a symposium at the International Rice Research Newsl N3)15 Institute September 1964 The Johns Hopkins Press Baltimore 63 Fugenio T S 1965 rhe effect oi e-vCgetation on rice production Maryland USA 729 p and the possibility of overcoming it by leaf pruning MS thesis

42 Centro Ihternacional de Agricklittira Tropical 1984 Upland rice in University ol the Philippines at Los Baijos ILaguna Ihilippins 69 p Iatin Anerica Pages 93-119 hi An os ervies of uplaind rice research 04 Farrow R A 1975 The Alrican nmigratory locust ii is main Proceedings of the 1982 Boinakamp Ivory Coast upland rice workshop outbreak area of the middle Niger qlinlitatise studies of solitary International Rice Research Institute P0 Blox 933 Manila populations in relation to environmental factors Locusla No II Philippines 198 p

43 Chandra G 1979 Taxonomy ad hn riononlics ofthe imect parasites of 65 Fennal R G 1943 Food crop investigation Windward and rice Icalloppers and planthoppers in the Pihilippines and their Ieeward Islands First report Report onl work dunring the periodiiiportairce iii natural biol)gical cointrol Philipp Fntoltol 4119-139 April 1942-April 1943 Imperial College of Tropical Agriculture

44 Chang T T 1976 The originii colut ion enlIivation dissemination Trinidad 16 p (Rev of Appl Fintornol A3222-25) and diersificition of Asian and African rices -uphytica 25425-44 1 66 Ferreira F 1983 Integrated pest ritaageiret of upland rice [ill

45 Chatierjee P B 1979 Rice leaffolder attacks in India lit Rice Res Portuguese English summary] Pages 323-341 in Upland riceculttre Newsl 4(3)21 factors affecting production E Ferreira T Yamada and F-

46 Cheng Cbien-pan 19(4 Upland rice in Taiwan International Rice Malmolta eds Institutoda Potassae Foslato Institnto Internacional Cnmmission Wiorking Party oit Rice Production ard Protection I()th da Potassa Piracicaha Sao Paulo lraid Meeting 3-10 March 1964 Manila Philippines Food and Agricul- 67 Ferreira E arid F dl Silva Martins 1979 Occurrence of ture Organiaion Rome Italy 5 p Sogatohs orvziola (Muir) inupland rice in Goias Ilrail lit Rice

47 Chcrrett 1 M and 1) 1Peregrine 1976 A review of the status of Rcs Ncwssl 4(4)18 leaf-cutting ants and their control Ann Appl Biol 84124-128 68 Ferreira E and 1F da Silva Martiris 1984 Insect pestof rice ill

48 Chipeta F M 1974 Pests of rice in Malawi Pages 90-95 in Brazil and their control Jill Portugucsej FMIlRAtPA-CNPAF Proceedings Sith eastern Africa cereals research conference 10-15 Goiania Brmiil 67 p March 1974 DRB Mialda ed Ministry of Agriculture lilongwe 69 Ferreira FI F ca Silha Miartins S S Nero and 1 It P de Malawi Carvallio 1982 M tltiplc regression analysis todeterimine the effect of

49 Chu Yau-i 1979 The d atiaging period of the rice seedling fly insect populatiois ott upland rice yield [inIPortuguese Pesqui (Atlterigotiavxigu Stcin) (NItiscide Diptera) oil cori) at Fast Java Agropecu Bras 17671-675 Indonesia Plant Prot Bill 21403413 70 Ferreira F 1F da Silva Martins and F 1PZimmermann 1979

50 Conklin H C 1957 IHaninoo agriculture a report on aln integral Rice varietal resistance to tle lesser corn stalk borer [in Port uguese] systent of shi fti rig cult ivationi in the Pltilippincs Food and Agriculture Pesqui Agropccu Biras 14317-321 Organization Rome Italy 209 p 71 Fleming W F 1968 Biological control of lie hapanese beetle U S

51 Cramer H II 1967 Plant proection and world crop production Dep Agric RLs Sers [ech Bull 1383 78 p Pilaniensclhutz-Nachr 201-524 72 Food and Agriculture Organiation of the United Nations 1982

52 l)ani R C and Majunidar 1978 Prelininary observations on tie Country report surmimary Pages 27-37 in Report on the first incidence of root aphids on differeit rice cultnrcs Sci Cult 4488-89 FAOIUN)IPhail and regional training coiuirse in iniproved ctltural

53 Davidson R I 1979 lffects of root feeding on foliage yield Pages practices for upland rice 5-23 Oct 1981 Northern Region Agrculturd117-120 in T K Ci osb and R B Pottinger eds Proceedings of the Developririt Center Chiang Mai Food and Agricuilture Organiiashy2nd Australian conlerence on grassland invertebrate ecology Govt tion Rome Italy 37 n Printer Wellington New Zealand 73 Franco do Ariaral S 195 1 Residual action of insecticides in turfy

54 l)ean G 1W 1976 Rice insect pests inlaos Int Rice Res Newsl soil--the results of a field experiment [in Portuguese] Biol6gico (Sao 1(2)15 Paulo) 17209-214 (Rev Appl Entomol A41423)

55 I)ean G J W 1978 Insect pests of rice in Laos PANS 24(3)280-289 74 Freeman ) 1970 Report onithe Iban london School of -coroiics 56 )ingle H 1984 Behavior genes and lifc histories conplex adapta- Monographs on Social Anthropology No 41 Athlane Press

tions to uncertain environments Pages 169-194 in A new ecology London UK 317 p novel approaches to interactive systems P W Price C N Slobod- 75 Fuchs T W1 A Harding and 1W Smithlr 1979 Induction and chikoff and W S Gaud cdsl ohn Wiley New York USA termination oldiapause in the sugarcane borer Ann Flitonlol Soc

57 Dobelmann-JP and M Falais 1968 Upland rice cultivation ill tie Am 72271-274 North-western part of Madagascar in 1966-1967 [in French English 76 (hbriel B1) 1975 A review tf the major insect pests of some tpland summary] Trop Agron 2313-21 crops in tlc Philippines Pages 86-111 in Ad Htoc panel of experts on

58 Doniiciano N L 1979 Chemical control of tie lesser corn stalk pest disease arnd weed problems in some rainfed crops Bangkok (lolsonno~paloIgnowslhts) (Zeller 1848) (I-epid(ptera Ihycitid ae) Thailand Food aid Agriculture Organization Roire Italy

on rice in Ponta Gressa Parara 1977-1978 [in Portuguese] Pages 77 Garg D K and N K Shal 1983 White grub outbreak on rainfed 291-303 in Anais 1 Reuniao de tecnicos cmiirizicultura do estado de dryland rice in Uttar Pradeslh Int Rice Res Ncwsl 81(4)16-17 Sao Paulo-Campinas Cati Brazil 78 Garg I) K and IP Tandon 1982 Major insect pests of rice on hilly

59 Dunsmore 1 R 1970 Investigations on tlie varieties pests and tracts of Uttar Pradesh India lilt Rice Res Newsl 7(11)11-12diseases of upland rice (hill padi) in Sarawak Malaysia Int Rice 79 Garrity 1) P 1983 Upland rice research an international bibliograpy Comm Newsl 1929-35 1975-1982 Iternational Rice Research hnstitnte PO Box 933

60 Elias 1R1965 The new decline of rice seedlings is not caused by Manila Philippines 121 p parasites [in Portuguese] Biol6gico (Sao Paulo) 313-6 80 Ghauri M S K 1982 A new subspecies of Iinorh)rone bull

61 Empresa Brasiliera de Pesquisa Agropecuaria 1984 Upland rice itt cornlus Slater (Hemiptera Iygaeidae) front Papua New Guinea on Brazil Pages 121-134 it Av overview of upland rice research rice and carpet grass Ihll lEntoniol Res 72133-137 Proceedings ofthe 1982 Blouak Ivory Coast Upland rice workshop 81 Gonzales F 0 PArregoces R I ihernandef and OT Parada International Rice Research Institute PO Box 933 Manila 1983 Insect and imite pests of rice arid their control in Latin America Philippines [in Spanish] la Federacion Nacional de Arrocerts (Fedearroz)

Bogota Colombia 60 n

38 IRPS No 123 January 1987

82 Grist D H and R J A W Lever 1969 Pests of rice Longmans London UK520 p

83 Hale P R 1979 Two subterranean pests of upland rice in Papua New Guirnca IntRice Res Newsl 4(2)16-17

84 Hale P R and J T Hale 1975 Insect pests of rice in the New Guinea Islands Rice Entomol Newsl 34

85 Harris W V 1969 Termites as pests of crops and trees Common-wealth Institute of Entomology London UK 41 p

86 Hattori 1 1968 A method to distinguish between grass leaf roller (Ctaplha rocis medinalisGuenee) and the rice leaf roller (Susurtia exigua Butler) [in Japanese] Plant Prot (Japan) 23167-170

87 Henry I E and 1A Onsager 1984 Experimental control of the mormon cricket Iahrosshnph by NJsena locustae(Aficrospora Microsporidae) a protozoan parasite of grasshoppers (Ort Acri-didae) Entomophaga 27197-201

88 Hidaka 1 P Vungsilabulr and SKadkao 1974 Studies on ecology and control of the rice gall midge in Thailand Tech Bull 6 TropicalAgricultural Research Center Tokyo Japan 113 p

89 Hincklcy A 1) 1963 Ecology and control of rice planthoppers in Fiji Bull Entomrol Res 53467 481

90 Hirao J 1982 Ecology and chtInical control of the rice leaffolder Jpn Pestic Inf 4114-17

91 Ho DT 1984 Sten borer incifience in rice ecosystems in KenyaEast Africa Pages 307-320 in A overview of upland rice research Proceedings of the 1982 Bouac Ivory Coast upland rice workshop International Rice Research Institute PO Box 933 Manila Philippines

92 Ho )T J 1Njokah and J GKibuka 1983 Studies on rice sten borers in Kenya with emphasis on ifaliarpiaipiraellRag Insect Sci Appl 465-73

93 Ichihara I T Ito A NIatsuo and M Sawada 1971 Chemical control of the rice striped stem borer in upland rice grown in plastic mulch [iii Japanese] Proc Kanto-Tosan Plant Prot Soc 1882

94 IclhiharaIand A Matsuo 1969 Damage caused by rice striped stett borer to upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1686-87

95 Ichhara I atid A MIatsun11970 Conside ration oft lie extraordinary prevalence of rice striped stem borer in upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1778-79

96 Institut de Recherches Agrolnliqes Iropicale et des Cultures Vivrieres 1978 Insect resistance Pages 12-13 in Upland rice 1978 atnual report Bouak Ivory Coast 27 p

97 International Institute for Tropical Agriculture 1983 Aphids itt rice roots I ITA Res Briefs 4(1-3)4-5

98 International Institite fi Tropical Agriculture 1983 Hopperburn in rice caused by brown plathopper IITA Res Briefs 4(l-3)6-7

99 International Rice Research Institute 1975 Annual report for 1974 PO Box 933 Manila Philippines p 216

100 International Rice Restarch Institute 1976 Annual repiort for 1975 PO Box 933 Manila Philippines p 228

101 International Rice Research Institute 1979 Cropping systems program Component technology development and evaluation Page 404 i Annual report for 1978 11O Box 933 Manila Philippines

102 International Rice Research Institute Department of Entomology 1981) Insecticid evaluation 1979 Tables 73 74 PO Box 933Manila Philipes

103 International Rice Research Institute Department of Entonmology 1981 Insecticide evalomtion 1980 Table 81 POBox 933 Manila Philippines

104 International Rice Research Institute 1981 Report of a meeting of Asian rice-based cropping systems entomologists Eleventh cropping systems working group meeting 18-22 May 1981 PO Box 933 Manila Philippines p 132 133 144

105 International Rice Research Institute 1984 Cropping systems program Pages 368-373 in Annual report for 1983 PO Box 933 Manila Philippines

106 International Rice Research Institutc 1984 Upland rice in Asia Pages 45-68 in An overview of upland rice research Proceedings of the 1982 Bouakamp Ivory Coast upland rice workiiop loternational Rice Research Institute 11O Box 933 Manila Philippines

107 Irshad M 1977 Grasshoppers in rice in Pakistan lit Rice Res Newsl 2(3)8-9

108 Ito R and TOno 1964 Upland lice its culture and varieties in Japan lIternational Rice Cormmission loth Meeting of the Working Party on Rice Production and Protection 3-10 March 1964 Manila Philippines Food and Agriculture Organization Rome Italy 12 p

109 Jeffrey M I and V A Dyck 1983 Effect of moonlight on lightshytrap catches of brown planthopper lit Rice Res Newsl 8(6) 13

110 Jennings P R and T Alicia Pineda 1983 Management ofthe rice plnthoppers Sogatodsorivi(coIl in Latin America Presented at the meeting on stable plant resistance FAO and lIndocsian I)inectorate of Food Crops Protection Bali lndonesia 29 Novcniber-2 l)ccemshyber 9 p

IllJimenez Gomez J A 1979 Rice insect pests i lla ios ()riCntales lil Spanish] Pages 99-103 in Rice ciltivation Instituto (olotibiatto Agropecuario Compndio No 29

112 Jongeleen F JJ Soesanto and A Mockidjo 1979 Studies oti pathogens of two species of Scarabacidac froinIndonesia Z Angew Enlomnol 87412-417

113 Jouvenaz 1) PC S lofgren and W A Baiks 1981 Biological control of imported fire antsa review of cirrent knowledge Bull Entomol Soc Am 27203-208

114 Kainoh Y K Shiniiu S MarUani Y Taiaki 1980 lost finding behavior of tie rice bu lplotori+JchinvittjI)allas (liciniptera Corcidac) with special reference to diet pattern ol aggregation and feeding on the rice plant Appl Entomol Zool 15225-233

115 Kakkar K 0 P Sharma and P I Sharma 1977 Note on Ihteronyrchus liodresRedt (Coleopicra I)ynastidae) as asporadic pest of paddy and maie inHimachal Pradesh Corr Agric 156-57

116 Kalode M131 79 Major insect pest problems and their control in upland rice (Unpublishcd mineogriph CWI oJNaaiat lni Chandaratat (198 1)original not seen)

117 Kalshoven L i 98 I The pests of crops in I ldoniesia (translated by P A vatn der Laan) 1 T Ichtiar B3aru-vati Iloeve lakarta Indonesia 701 p

118 Kamal N Q 1981 Suppression of whitebacked planto ner Sogatella fircaifra (Horvath) aLId rice lealfolder (nhal -cis licdb lis (Guenee) populations by natural cnclmie hlI) tiesis Grcgorio Araneta University Foundation Victoncta Park Metto Manila Philippines 179 Ip

119 Karim A N M Reaul 1975 A species ofliealy bug Trioni i70s ceres new oii rice in the Philippines Rice Entoniol Newsl 247

120 Katanyukul Wand C Chandaratat 1981 Insect pestsof uplaid rice and their control Report firstFAOUNlFi Tliailand training course on improved cultural practices for tplad rice Chiang Mai Thailand 5-23 Oct 1981 Food and Agriculture Organiation Rome Italy 22 p

121 Kern F 1955 lhletwioa(I)uninemts) bhhit Biturm(Coleoptera) a new pest of sugarcane inVenezuela [inSpanish with Etnglish sunmary] Agron Trop (Maracay) 527-29 (Rev Appl Eitoniol A4572)

122 Khan K M and S Ghai 1974 White grubs ald their control in India Pesticides (1Bombay) 819-25

123 Kiritani K 1975 Integrated pest management of rice pests Pages 66-70 itt Proceedings platning workshop ott cooperative hIeld research in pest managementP SMotooka ed East-West Center Honololu Hawaii USA

124 Kisimoto R 1967 Ecology of insect vectors forecasting and chemical control Pages 243-255 in The virus diseases of tle rice plant Proceedings of a symposium atthe International Rice Research Institute April 1967 The Johns Hopkins Press altimore Maryshyland USA

125 Kisimoto R and V A Dyck 1976 Climate and rice insects Pages 367-391 in Climate and rice International Rice Research Institute PO Box 933 Manila Philippines

126 Kureha S S Kobayashi SAsari and T Ishihara 1977 Life history and control of the mole cricket Grrllotalpa africana Palisot de Beauvois [in Japanese English translation 7520 1RRI library] Proc Kanto-Tosan Plant Prot Soc 21192-197

127 Kushwaha K S 1981 The black beetle as a pest of rice in Haryana India Int Rice Res Newsl 6(4)14

128 Lawani S M 1982 A review of tile effects of various agronomic practices on cereal stem borers Trop Pest Manage 28266-276

129 Leuck D B 1966 Biology of the lesser cornstalk borer in South Georgia J Econ Entomol 59 797-801

130 lever R J A W 1955 Insect pests of the rice plant in Malaya Int Rice Comm Newsl 1612-21

131 Li Ching-Sing and Shin-Foon Chiu 1951 The study ofthe rice gall midge Iachydiplosis orvza Wood Mason [in Chinese with English summary] 1Taiwan Agric Res 21-13

132 Iitsinger J A 1984 Assesament of the need-based application for rice Proceedings TechnologyTransfer Workshop 15-16 March 1984 International Rice Research Institute 1PO Box 933 Manila Philippines 6 p

133 Litsinger J A 1984 Environmental diversity and insect pest abundance with reference to the Pacific Pages 285-307 in Proceedings sub-regional training course on methods of controlling diseases insects and other pests of plants in the South Pacific 4-20 Oct 1982 Vaini Kingdom of Tonga Nukualofa

134 Litsinger J A RF Apostol and M BObusan 1983 White grub Leucopholis irrorata (Coleoptera Scarabaeidae) pest status popula-tion dynamics and chemical conllol in arice-maize cropping pattern in the Philippines J Econ Entoinol 761133-1138

135 Litsinger J A C de la Cruz F Raymundo A T Barrion M D Lumaban J P Bandong M S Venugopal F Paragna and E Balete 1982 Insect pests and insecticide reponse of dry-seeded bunded rice Pages 179-193 in Report of workshop on cropping systems research in Asia International Rice Research Institute Los Bafos Philippine

136 Litsinger J A M 1) Luinaban J P Bandong A T Barrion W Katanyukul and N Panda 1978 Insect pest management in rainfcd lowland rice agroecosystms of tropical Asia International Rice Research Conference 17-21 April 1978 International Rice Research Institute Los Bafios Philippines

137 Litsinger J A and K Moody 1976 Integrated pest management in multiple cropping systems Pages 293-317 in R I Papendick P A Sanchez and G B Triplett eds Multiple cropping American Society of Agronomy Madison Wisconsin USA

138 Loevinsohn M E 1984 The ecology and control of rice pests in relation to the intensity and synchrony of cultivation Ph D thesis University of London 36) p

139 Loevinsohn M E J A Litsinger N Panda M A Austria R F Apostol C Andrion and J Bandong 1982 A basis for predicting rice insect pest abundance in cropping systems Paper presented at the 1982 International rice research conference on rice and cropping systems 19-23 April 1982 International Rice Research Institute Los Bafios Iaguna Philippines 25 p

140 Lopez A W 1930 Repot of the Entomology Department Pages 145-172 in Annual report of the Philippine Sugar Association 1929-30 Manila

141 Lopez AW 1931 Entomology Department Pages 227-266 in Annual report of the director or research 1930-31 Philippine Sugar Association Manila

142 Lopez A W 1932 Entomology Deparment Pages 252-279 in Annual report of the Philippine Sugar Association 1931-32 Manila

143 Lungibill P and G G Ainslee 1917 The lessercornstalk borer US Dep Agric Agric Res Serv Tech Bull 539 27 p

IRPS No 123 January 1987 39

144 McGuire J U Jr and BS Crandall 1967 Survey of insect pests and plant diseases ofselected food crops of Mexico Central America and Panama U SDepartment of Agriculture International Agriculshytural Development Service Agricultuial Researrh Service October 1967 11 p

145 McNeill Sand T R ESouthwood 1978 The role of nitrogen in the development of insectplant relations Pages 77-9 in lliochemical aspects of plant and animal coevolution J B Ilarbornu ed Academic Press London UK

146 Maebara H 1955 Spatial distribution of lastinstiarlanaeof purplish stem borer (Sesamia iqferens Walker) hibernating in an upland rice field Bull Kagoshima Univ Fac Agric 438-41

147 Mallick E H and N G Hajra 1977 Response of rice plant to defoliation treatments at different developmental stages Indian Agric 2151-56

148 Mansingh A 1971 Physiological classification of dormancies in insects Canl Entomol 103983-10)9

149 Martell C G 1974 Catalogue of plant feeding insects of Mexico [in Spanish] Fitofilo 27(69)1-176

150 Martinez 0 J 1982 Principal rice pests in Ilanos orientalis [in Spanish] P~ages 80-94 in Curso deact nalization de conocimientos en arroz Villaniencis Instituto (olonmbiano Agropecuario

151 Martins J F da Silva F Ferrcira A S Prabhu F 1 P Zimmermann 1980 Use of pesticides for the control of the main subterranean upland rice pests esqui Agropecu Bras 1553-u

152 Martins I F da Silva F Fe eira rod BdamSilveira Pinheiro I Q2 Damage simulation caused by defoliators of upland rice [in Portuguese with English summary] Pesqui Agropecu Bras 171113-1119

153 Martins 1F da Silva N(iyuen van Tan and 1Ida Silveira Pinheiro 1981 Stem borer resistance in upland rice associated with plant morphological characters [in Portuuce Iesqui Agropecu Bras 16187-192

154 Masaki S 1980 Sumnmer diapause Ann Rev Entomiol 251-25 155 Masuzawa T 11 Suwa and F Nakasoji 1983 l)ifkleences of

oviposition preferencc and survival rate of two skipper butterflies Parnara guttata and PIchlpidai iathiais (Iepidoptera HIesperiidac) on rice plant and cogon grass New Entomol 3247-56

156 Maurya D M and C P Vaish 1984 Upland rice in India Pages 69-92 in An overview oftpland rice research Proceedings ofthe 1982 Bouakamp Ivory Coast upland rice workshop International Rice Research Institute IO Box 933 Mnila Philippines

157 Menon M G R R N Katiyai S Kuar and S S Misra 1970 Termites attacking the paddy crop Sci Cult 36294-295

158 Miyahara Y and 1 Nishikubo 1967 Behavior of young nymphs of the red rice root aphid Iholahsipiun ri liabhdontinalisSasaki in relation to movement from the leaf to the roots of upland rice [in Japanese] Pioc Assoc Plant Prot Kyushu 1363-66

159 Mo ljoa Tjien 1952 White grub of upland rice (Holotrichiahelleri Brsk) [in Indonesian with English summary] Contrib Agric Res Stn Bogor No 13134 p

160 Mochida 0 and T Suryana 1979 The occurrence of the brown planthopper and ragged stunt on lowland and upland rice ill Central Java and logyakarta in tile late of January 1979 Entomnolgy Survey Report No 15 Central Research Institute of Agriculture (CRIA) Sukamandi Indonesia I February 1979 9 p

161Monet C 1979 Varietal resistance to upland rice pests [inI French] Pages 35-39 in Annual report 1977 Institut des Savanes Department des Cultures Vivrieres Bouak6 Ivory Coast

162 Monte 0 1942 A caterpillar ofrice-fields[in Portuguese] Biol6gico (Sao Paulo) 8161-163 (Rev Appl Entomol A31414)

163 Munroe I) D 1979 Pest management strategy for hill rice in four resettlement areas in Sarawak Malaysia Pages 89-102 i1 Guidelines for integrated control of rice insect pests Food and Agriculture Organization Plant Production and Protection Paper 14 Rome Italy 115 p

40 IRPS No 123 January 1987

164 Nakasuji F M Ishii and Jen-Zon Ito 1984 Rice skippers in Taiwan and their life histories ZZrC (Taipei) Tech Bull 82 10 p

165 Navas I) 1976 Fall armyworum in rice Pages 99-106 il Ecological animal control by habitat management Proc Tall Timbers Con-ference 28 Feb-I Mar 1974 Fall Timbers Res Stn Misc Publ 6 Tallahassee Horida USA

166 Nickel 1 1 1961 Biological control of rice stem borer A feasibility study Tcch Bull 2 International Rice Rcarch Institute PO Box 933 Manila Philippines IIl p

167 Nigan P M and R A Vernia 1985 Insect pests of uplaild rice in Uttar Pradesh Int Rice Res Newsl 10(6)22

168 Nilakhe S S 1985 -cological observations on spittle bugs with emphasis on their occirence in rice Pcsqtui Agropecu Bras 20407-414

169 Nixon G EJ 195 1 lie association of ats with aphids and coccids Commonwealth Iusti lcte of Entonology Iondon UK 36 p

170 Nurdin F an Ishak Manti 1985 Study oit seedling fly of upland rice Interlational Upland Rice Moitoring Tour Report 26 lebruary-2 MarLI Sukaranti West Sumatra

171 Oka 1 N 1979 Ciltural conttol of the brown planthopper Pages 357-369 in Brown planthopper threat to rice production in Asia International Rice Research Institutc 1p0 Box 933 Manila Philippines

172 Ohni M and A Vilnaii 1975 Rice insect pests of Italy biology of Sipha grcria (Kaltenbach) and its control [in Italian] Riso (Milan) 2459-71

173 Otanes F Q atnd 1 I Sison 194 1 Pests of rice Philipp J Agric 12211-259

174 Painter R I 1951 Insect icsistance in crop plants University Press of Kansits Lawrence Kansas USA 520 p

175 Pantua P C and 1A litsinger 1981 Arthropod abundance in continuous vs biannual rice cropping systeis Int Rice Res Ncwsl 6(2)20-21

176 Pathak M ) and V A l)yck 1975 Sttudics ott insect pests of upland rice Pages 186-191 in Major research in upland rice International Rice Research Institute PO Box 933 Ilanila Philippines

177 Patil B R B M IHasabe and 1) S Ajri 198 I Chemical control of white gritb lIditriihinhorraia Fabr intfesting groundntut paddy atnd sorghum Indian 1Entomol 43232-236

178 Pawar A 1) 1974 (ollection and identification of insect pests of rice rlerminal repirt lInternational Rice Research Institute PO Box 933 Manila Philippines 72 p (tnimco)

179 IerrajU A aid A S Reedy 1965 Observations ott diapausc itt tie steti borer ohrice Rice News Teller (lai-Mar)9-13

180 Pont A C 1972 A review of the Oriental species of theri ona Rondani (I)iptera Muscidac) of econtomic importance Pages 27-102 in Control of sorglium shoot fly M G lomtani and W R Young eds Oxford atd IH1 Ptthl Co New I)clhi India 324 p

181 Prasad S C 1PChand aid M 1PSingh 1983 Incidence of rice gall midge in the upland plateau region of Hihar Oryza 20235-236

182 Rai P S 198 0 Nisaga simpex damage to rice in the hill tracts of South India Int Rice kles News[ 5(I)12

183 Rathorc Y S 1969 Gonocplhahli dcprssunt Fabr as itpredator of (tilo zonwIlhis (Swinhoc) and other moths in field cage at Patnagar Uttar Pradesh Indian 1Entotnol 3127

184 Reed W 1983 Crop losses causcd by insect pists in the developing world Proceedings 10th Internatial Congress of Plant Protection lrighton UK 174-80

185 Reedy M V 1981 Ateactogasterinhens(Walker)a new root-grub pest of tipland paddy and inaize Curr Sci 501076

186 Reveno M A 1968 Catalogue of the insect pests of economic crops in Colombia [in Spaiish] Asociacion Latino Americana de Entomo-logia (AIAE) 129-33

187 Reyes G M 1957 Rice dwarf disease in the Philippines Food and Agriculture Organiiation Plant Prot Bull 617-19

188 Rice S E A A Grigarick and M 0 Way 1982 Effect of leaf and panicle feeding byarmyworm (Lepidoptera Noctuidae) larvae on rice grain yield Environ Entomol 7593-595

189 Risch S J 1) Andow and M A Altieri 1983 Agioccosystem diversity arnd pest control data tentative conclusions and nizw research dirctions Environ Entomol 12625-629

190 Rivera C T 1956 Three organic insecticides in thecontrol of insects affecting upland rice Philipp Agric 39465-472

191 Roffrey 1 1972 Locusta outbreaks in the Philippines Acrida 1177-188

192 Rojas R 1) 1941 The effect of pruning at varying ages upon the yield of Ramairio Phipp Agric 29851-860

193 Roonwal M i 1979 Ternite life and termite control in tropical South Asia Scientilic Pnbl odhpr India 139 p

194 Rosetto C1 F da Silva Martins N C Schmidt and L EAzzini 1978 Damage caused by pasturespittle bugs 4Dvoi)lavopictaandDJ schach) on rice lBragantia Nota 6 (Matcli)25-27

195 Rosztto C1- S S Neto I- Amantc I) Iink 1Garcia-Vieira D M de SoLza N V Bansa t and A M de Olivcira 1971 Rice pests of Brazil [in Portuguese] Pages 149-238 inrFA()-International Rice Commission 2nd meeting of the Rice Committee for the Americas 6-11 l)ecember 1971 Pelotas Rio Grande do Sul Brazil

196 Rothschild G H L 1969 Obseratiois on the armyworm Spodoptera matritia acronrctoiiles Gn (Lep Noctuidae) Bull Entomol Res 59143-160

197 Rothschild G 1- L 1974 Parasites of the armworm Spolopiera mauritiaacroncitoihes in Malaysia Fntornopliaga 19293-299

198 Sachan i N S K Gangwar and 1N Katujar 1980 Field crickets damaging cages in North Eastern IHill Region Indian J Entomol 42526-527

199 Sands 1) P A 1977 The biology and ecology of Lepocorisa (Hemiptera Alydidac) in Papua New Guinea Res Bull 18 Departshyment of Primary Industry Port Moresby Papua New Guinea 104 p

200 Sarawak 1968 titomology division Pages 97-106 in Report of the research branch of the Department of Altriculture 1966 Kuching (Rev AIppl Eintomol A56603)

201 Satpathy JM M S Das and K Naik 1977 Effect of multiple and mixed cropping on the incidence of scine important pestsI Entomol Res 178-85

202 Sauer IHF G 1939 Notes Ott Elasmpnlpualtsligniosellths Zeller (Lep Pyr) a serious pest ofcereals in Sao Paolo [in Portuguese] Arq Inst Biol 10199-206 (Rev Appl Entotnol A28378)

203 Satinders 1L A 1 SKing and C L Vargas S 1983 Crop pests of Central Anerica with t bibliography [in Spanish] Centro Agroshynotnico Tropical de Investigacion y Ensenanza Turrialba Costa Rica (CATIE) Tec Bol 9 90 p

204 Sauphanor B 1985 Some factors of upland rice tolerttce to stem borers in West Africa Inseu Sci Appl 6429434

205 Sen P atnd S Chakravorty 1976 Stentflics in Haritighata Nadia West Bengal India [it Rice Res Newsl 1(12) 15

206 Senapai B and 1M Satpathy 1983 The seedling fly Atherigona orrzae Malloch is a pest of rice in Orissa (India) J Entomol Res 777-79

207 Shhjahan M S Alatn M Ah nad and BN Islam 1983 A new scarabaeid beetle pest of dryland rice in Bangladesh Int Rice Res Ncwsl 8(I)16-17

208 Sharma G L S C Modgol and R C Gautam 1979 Effect of depth of submergence ott incidence of bacterial blight and white grub infestation in transplanted rice Int Rice Res Newsl 4(6)12-13

209 Sharma S 1985 Scarabaeid beetle outbreak itt upland rice Int Rice Res Newsl 1(1(l)24

210 Shiraki T 1917 The paddy borer Schoenobhits incertulas Walker [achaku Agric Exp Stn Formosa 256 p

211 Silva A de Brito and 11P Magalhaes 1981 Insect pests of rice [in Portuguese] Circular Tecnica No 22 EMBRAPA Centro de Pesquisa t ecuaria de Tropico Unido (CPAJU) Belem Brazil 14 p

212 Singh S A 1979 Emergence of rice leaf butterfly as a rice pest in Manipur India int Rice Res Newsl 4(4)16-17

213 SinghS A 1983 Changing trends of insect pestsof rice in Manipur Int Rice Res Newsl 8(f)12

214 Siwi S S and P H van Doesburg 1984 Lenovorisa Latreille in Indonesia (Heteroptera Coreidae Alydinae) Zool Meded (Leiden) 58117-129

215 Soejitno J 1977 Reaction of some upland rice varieties toseedling fly eltherikonaexigua Stein [in Indonesian] Siri Hana I Penyakit 630-35 Lembga I osat Fenclilian Pertanian Bogor Indonesia

216 Socnardi 1967 Insect pests of rice in Indonesia Pages675-683 in The major inse pests of thu rice plant Proceedings of a symposium at the Internatital Rice Rec Wi itt S pternber 1964 The Johns Hopkins Press Baltimore Maryland USA

217 Soto P E A T Petcz and I W Buddenhagen 1976 Survey of insect pests and diseases of rice in different ecological zones in Nigeria Rice Entomol Ncwsl 435-46

218 Soto 11E and Z Ziddiqi 1976 Screening for resistance to African rice insect pests Varietal improsetnent sCmi n r Bouak Ivory Coast 13-17 Sept 1976 West Africa Rice l)evelopment Association Monrovia Iiberia

219 Squire F A 1972 Entomological problems in Bolivia PANS (PestArtic News Sumr) 18249-268

220 Stapely 1 II 1978 The rice leaf roller in the Solomon Islands Int Rice Res Newsl 3(l)13

221 Suenaga H 1937 On aphids attacking the roots of rice Oyo-I)oubts Zasshi 9150-152 [in Japanese] (Rev Appl Entomol A25745)

222 Suenaga H 1952 On the seasonal abundance of three subterranean aphids on upland rice Bull Kyushu Agric Exp Sti 1249-262

223 Sutynso Yas and Dandi Soekaria 1964 Beberapa hana yang ttenperang tana man padi tanah kering [in Indonesian] Rapat kerja Padi Tanah Kering 6 sd 8 Juily 1964 Kaliurang 9 p

224 Tallamy I) W and R F Denno 1983 Alternative life history patterns in risky environments in example from lace bugs Pages 129-147 in Insect life history patterns R F l)cnio and If Dingle eds Spring-Verlag 225 p

225 Tamura 1 1939 Observations on some insect pests of field crops [in Japanese] 1 Plant Prot 26414-420 (Rev Appl EntomolA27-556)

226 Tamura and Suzuki 1964 Influences of the early-transplanting practice in rice culture upon development of the rice stem borer Chilo siql)ressalisWalker Bull Ilokuriku Natil Agric Exp Stn 761-94

227 Tan J P 1924 The rice root aphid Drowhi ihirsoia Philipp Agric 13277-288

228 Tanaka 1 1957 Taxonomy and distribution of some subterranean aphids injurious to upland rice in Japan with description of a new species [in Japanese English summary] Botyu-Kagaku 22168-176

229 Tanaka T 1961 The rice root aphids their ecology and control Spec [full College of Agriculture Utsunomiya University Japan F3 p

230 Tauber M IC A Tauber R Nechols and 1Obrycki 1983 Seasonal activity of parasitoids control by external internal and genetic factors Pages 87-108 itt I)iapause and life cycle strategies in insects V K Brown and I Ilodek eds )r W Junk The Hague Netherlands

231 Taylor W F 1972 Effects of artificial defoliation (simulating pest damage) on varieties of upland rice Exp Agric 879-83

232 feotia T P S and S Natud 1966 Biotoomics of the rice skipper Parnaraniathiis Fabricius (Lepidoptera Hesperiidac) Indian J Entomol 28181-186

233 Teran F 0 1971 Rice pests of Santa Cruz [in Spanish Ministerio de Agricultura Sita Cruz Bolivia Bull 45 8 p

234 Than Itun 1976 Population dynamics of the yellow rice borer Tryporza itcertulas (Walker) and its damage to the rice plant MS thesis University of the Philippines at Ios Bafios Laguna Philippines 92 p

235 Uichanco L B 1911 Pests Pages 55-74 it Rice Philipp Agric 11-200

IRPS No 123 January 1987 41

236 van der Laan P A 195 1 Life history of the rice seedling fly Contrib Gen Agric Res Stn Boor Indonesia 118 15 p

237 van der Laan 11A 1959 Correlation between rainfall in the dry season and the occurrence of the white rice borer (Scirpolhaga innotala WIk) in lava Entomol Fxp Appl 212-20

238 Van Driesche R G 1983 Meaning of percent parasitismin studies of insect parasitism Fnviron Entomol 121 I1- 122

239 Venugopal M S and A litsingcr 1984 Fffect ofcarbofuran on rice growth Prot Ecol 7313-317

240 Vietnam Directorate of Rural Affairs 1962 Annual work progress report on crop improvement program of rice sugarcane vegetable and field crops (lor the period from July 1961 to June 1962) (11) 299 p (Rev Appl Entomol A51526-527)

241 Wan M T K 1972 Observations on rice leaf- and planthoppers in Sarawak (Malaysian Borneo) Malay Agric 148308-335

242 West Africa Rice I)esclopnient Association 1978 Annual research report Vol 2 Statistics Monrovia Liberia

243 West Africa Rice Development Association 1979 Annual research report p47 Monrovia Liberia

244 Williams ) F and CS lofgren 1981 Fli I illy l-468 anew bait toxicant for control of the red imported lire at Fla Entomol 64472477

245 Williams I) 1 1970 The mealybugs OFlomoptera Coccoidea Iscudococcidac) of sugar-cane rice and sorghum Bull Entomol Res 0109-188

246 Williams D) 1 1982 Pulinaria icervi (Signoret) and its allies ott sugar cane and other grasses Btll Entoinol Res 72111-117

247 Wirjosuiardjo S 1976 Study ofchenmical cont rot of insect pests ott upland rice Imiu Pertanian (Jogjakarta) 1323-329

248 Wirjosuhardjo S and N Wirarta 1976 Chemical control of the white grub tIolorithiatlltriin upland rice in M uneng Irobolinggo Indonesia) [in Indonesian] Imut Pertanian (Jogjakarta) 1330-332

249 Yano K T Miyake and V F Eastop 1983 The biology and economic importance of rice aphids (Henuiptera Aphid idae) a review Bull Entomol Res 73539-566

250 Yelp D and P Rhodes 19(4 lhe success of drilled dryland rice iii the Fiji Islands International Rice Commission Working Party on Rice Production and Protection Manila Philippines 3-10 March1964 Food and Agriculture Organization Rome Italy 13 p

251 Young W R 1981 Fifty-five years of research on the sorghum shootfly Insect Sci Appi 23-9

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No 101The economics of hybrid rice production in (hina No 113 Boron toxicity in rice No 102 Rice ratooning No 114 Energy analysis rice production systems and rice research No 103 Growth and development of the (Jeep water rice plant No 115 Production risk and optimal fertilizer rates an application of the

No 104 Faridpur acomputer-assisted instruction nmodel forrainfed lowland random coefficient model No 116 Consumer demand for rice grain quality in Thailand Indonesia andrice

No 105 A readingand listening comprehcnsion test inEnglish for non native the Philippines speakers applying for tiraining at IRIRI No 117 Morphological changes in rice panicle development a review of

No 106 Rice grassy stunt virus 2 a new strain of rice grassy stunt in tire literature

Philippines No 118 IRRI-Korca collaborative project for the development of cold-No 107 Physical losses and quality deterioration in rice plstprodnCtion tolerant lines through anthei culture

systemis No 119 Problen soils as potenial areas for adverse soils-tolerant rice No 108 (olptblication ofIRRI materials a survey of translators and varieties in South and Ssuthcasi Asia

publishers No 120 Changes isinall-farm iLtireliing techii)hgy in Thailand and [he No 109 Classification of Philippine rainfall patterns Philippines No 110 Contributions of modern rice varieties to rutrition in Asia No 121 Landforms and modern rice varieties No Ill Changes in rice breeding in 10 Asian countries 1965-84 No 122 Yield stability and modern rice technology No 112 Design parameters affecting the performance of the IRRI-designed

axial-flow pump

Page 11: UPLANDRICE. INSECTl--1i M T -E!R.EOLOU t RTANCE ,AND …delphacid.s3.amazonaws.com › 6958.pdf · The significance of abiotic factors in upland rice insect ecology will be apparent

rice plants In Asia upland ricefields are typically small with Iprge field border areas (ecotones) where termites can forage for preferred food other than growing rice plants All grassland termites do not feed on rice In an upland area in Batangas Province Philippines ricefields are commonly infested by Hosiititernesluzonensis (Oshima) however this termite does not damage rice In the same province though Mrroternesgivus(Hagen)feeds on rice seedlings (178)

In Africa termites may damage rice even though fields are small and surrounded by perennial vegetation These termite species perhaps prefer rice over other hosts

Mole crickets Nymphs and adults are nocturnal and burrow through soil feeding on roots of a wide variety of plants or forage above ground as predators of insects Adults live 3-5 mo and are even cannibalistic Mole crickets prefer low-lyin moist upland soils with high organic matter (II 81) In the lowlands they inhabit rice bunds or nonflooded fields Adults are highly mobile and can leave a flooded field to locate a more suitable habitat Losses show as wilted plants (from root feeding) or reduced stands (120) Damage is more common near field borders where mole crickets relocate after tillage (126)

Fiehl crickets Nymphs and adults of field crickets have similar nocturnal habits and damage rice as much as mole crickets do Piles of weeds removed from fields attract them They make subterranean nests and tunnel through the soil to feed on roots Some species prefer seed to roots Others feed at the base of stems causing deadhearts

Tunnel entrances surrounded by excavated soil are easily seen Young plants are cut at ground level and stored in underground cell Field crickets tre highly polyphagous as are all soil pests (117) Sachan et al (198) reported that field crickets prefer maize to upland rice Like mole crickets field crickets cannot survive in standing water and are therefore more prevalent in upland fields

Whitegrubsandhackbeetles Scarab beetles that feed on living roots as larvae but not as adults are called white grubs or chafers (Melolonthinae Rutellinae) and those that feed as adults but not as larvae on plant crowns are called black beetles (Dynastinac) In the tropics chafers or white grubs have a I-yr life cycle starting with adults emerging from the soil about I mo after the first downpour of the rainy season (134 159) They lay eggs at the same time farmers sow upland rice The rice passes its most susceptible stage and damage is mostly avoided when the white grubs are small After several months the long-lived larvae are large enough for two or three to denude the root system of mature rice (223) This intensity of damage is rare but wilting occurs when root loss is combined with drought stress

The larvae need damp (not saturated or dry) soil to survive and pass the unfavorable dry season 05-2 m underground in aestivation in the moist soil The first heavy rains of the season (20-30 amd) stimulate the grubs to resume activity and after several weeks they develop into

IRPS No 123 January 1987 11

pupae and adults and eventually dig their way to the soil surface and fly to nearby trees to seek food and mates The adults of most white grub species feed only on trees but in some species notably Eutheolahumilis the adults defoliate and cut off stems at ground level (28) Putheola hidentata damages sugarcane fields in Venezuela (121) Floods kill white grubs (208) Grasslands can support large populations and therefore white grubs can be more abundant in newly planted upland ricefields that were previously fallow (81)

Black beetles live up to a year adults can attack a crop at any stage by burrowing into the soil (127) and feeding at the base of stems to cause whiteheads (207) Adults emerge with the early rains and are normally more abundant and do more damage on a young rice crop with small root systems (38 81)

The larvae feed only on dead organic matter in dryland fields and do not attack rice Adults arc highly mobile and although sensitive to flooding can invade a field soon after it drains (115)

Root aplhids Several genera of migratory aphids feed on the roots of upland rice (249) Populations build up more in light-textured soils with high percolation rates Damage has also been reported from dry seed heds of lowland rice (52) Tending ants -- Pheidhle Cren aster Tetramoriuw lsius Tapinoma -- are necessary for root aphids to multiply Aiits harbor aphids in their nests over winter or during unfavorable periods and relocate them on rice plants digging tunnels along the root systems to allow these softshybodied insects to penetrate the soil Most intensive studies on aphid life histories have been done in temperate environments (158 222 229) Root aphids fly to rice plants at the beginning of the rice season and pass through several generations continually developing winged forms to relocate on alternate grassy hosts until winter when they move to perennial hosts usually trees Yield loss occurs mainly through reduced tillering but infested plants become yellow and stunted and in extreme cases wilt (227) Aphid populations build up gradually so damage usually begins in the late vegetative and reproductive growth stages

Root-feeding ineal tugs More than half of the mealybug species or rice feed on roots (245) Being soft bodied mealybugs are not adapted to living underground but they survive because tending ants (the same species that tend root aphids) dig burrows for them and move them from plant to plant Well-drained soils ihelp mcalybugs survive Although six species are recorded as subterranean root pests of rice we found no report citing damage Root-feeding mealybugs are probably more prevalent on perennial grasses than on short-lived annual rice

The most common rice mealyb ig Brevennia rehi is a foliar feeder favored by lack ol rain apparently it is responsible for high yield loss in Northeast India and Bangladesh (N Panda pers comm as stated by IRRI [1061) This report needs confirmation Mealybug damage was only briefly mentioned in another report (156)

12 IRPS No 123 January 1987

Root weevils A few species of upland weevils (IHporneces Donacia Atactogaster) attack rice roots as larvae and feed on stems and leaves as adults Damage ispatchy The larvae are not adapted to submergence and are true upland soil weevils isopposed to the so-called water weevils They rarely become sufficiently abundant to cause damage Adults on rice foliage or the soil surface show that the roots are being attacked by the grubs Root weevils are highly polyphagous (185)

Root heeths Soil-inhabiting false wireworms Gono-cephalumt spp are reported in upland ricefields The adults and larvae normally feed on decaying organic matter but land preparation for upland rice removes these sources The larvae feed on seedling roots and nocturnal adults roam on the soil surface to cut off seedlings at ground level (82) This group of ground beetles is highly omnivorous adults even prey on stem borer larvae (183) True wireworms (Elate-ridae) feed on living roots but rice as a host has been reported Infrequently (117) In Latin America the highlypclyphagous larvae of Diabrotica rootworms are also recorded on rice

Root htus Upland rice seedlings can be attacked by the adults anI nlViphs of chinch bugs Blissusand Caenoblissus and bro n bug Scaptocoris which feed on the roots with theirsucking nouthparts (80 83) Three genera of burrower bugs A tms (eotonhus and Stiharopus -are common iil grasslands in the Philippines and may attack the first crop (rice or maize) planted after the dry season Chinch bugs are most abundant after a dry spell These insects are large and can kill seedlings or stunt and reduce tillering on older plants I hey prefer other hosts and rarely damage rice Adults can also feed oii developing rice grains (81)

(utworms Soil-dwelling noctuid larvae hide under-ground during the day to avoid predation by birds and bccone active at night above grour cutting off you ing plants at ground level (hence tile naiie cvtworn) arvaedrag severed plants into hurrows llie larvae have wide host ranges arid pupatc in the soillgrotis ipsilon occurs worldwide and appears to be the most common cutworm pest of upland environments but is more prevalent inlowland rice particularly in dry seedbeds

IFoliar pests Other groups of insects are adapted to upland environments because they pass one growth stage underground either as egg (in heinrietabolous species) or pupa (in holomnetabolous groups)

Rice seedlibig maggots AitJeriota flies occur in non-fhoodcd environments in Asia and Africa (180) Feeding of larvae one per iller causes deadhearts Flies as opposed to moths only secondarily evolved as pliytophagous pestsfroni a saprophagous Origin Athrnonalarvae secondarihv feed onidecaying tillers whereas larvae of stem borer moths actively Iced on living tillers to cause deadlicarts Seedling maggots will die if they cannot sever a tiller on which to feed

as it decays Plants die under heavy attack whereas lighter infestations cause stunting delayed maturity and ragged leaves (236)

Eggs are laid only on actively growing plants (49) aid damage occurs wiihin the first week after crop emergence(170) Larvae pupate more in stems than in soil Seedling maggot attack is highly seasonal (206) normally peaking 2-3 mo after the onset of the rainy season These pests possiblyaestivatc over the dry season Rice isonly one of their many graminaceous crop and weed hosts

The rice stem maggot (hlorops orzae is adapted to temperate upland and lowland rice growing regions of Asia In Japan and China upland rice grows in mountainous regions a habitat of this chloropid fly The larvae tunnel into tilestems The stem maggot is more damaging during the seedling stage larvac hibernate in grasses

File several dipterans reported as stem maggots in dryshyseeded lowland seedlings inIndia (205) could presumably attack upland rio

Leaf urinrs Icaf miners are common on wheat and maize but are rarely reported on upland rice (213) Three leaf miners are more prevalent on upland than lowland rice inthe Philippines Pvudonalomy a asicanh Vficataand lIAgromrza orzae attack at the seedling stage The larvae develop inside the tunnels they construct in the parenchyma tissue P Vpicata and Aorvzac prefer maizeEehtsine and other grasses to rice spicatacan become abundant on wheat

Leaf beetles Chrysomelid and coccinellid eaf bcetles scrape or otherwise remove leaf tissue as larvae or adults or both This damage accelerates desiccation of plants Tile larvae of sonic species are root feeders Flea beetles Epitrix and (lraetocnetna and rootworms l)iahroticaspp feed on rice foliage only as adults Their root-feeding larvae have hosts other than rice The adults and larvae of the hispa-like Oediolmlpa spp beetles of Brazil and (hmootriha (=EPfishyachra) beetle of Africa feed on upland rice leaves Oealioshypapa spp larvae are leaf miners (mootriha is also pshyvalent in lowland environments (11) Leaf beetles prefer vegetative rice (81)

The rice hispa Dicladipaarnigera is not reported on upland rice and prcers more aquatic habitats In Central Africa however I)cladispm viridicyaiea attacks uplandrice in the vegetative stage as well as lowland seedbeds 7richipasericea is reported on upland rice in West Africa and Madagascar (57)

The rice leaf beetle 1d1ea 01izae riiuech like Chioropsor|rzae is adapted to temperate regions of Ashi and is usually a pest on lowland rice but seldom oi inpland rice in the mountains (82) The adult hibernates on grasses

Armmworms Species of grassland-adapted SpoptloeraPlatysemta and A ithimna larvae create widespread epishydemics They can defoliate ricefields generally in patchesfrom early vegetative growth to harvest (41) Armywornms pupate inthe soil and highly favor upland rice (34) Adults

can disperse long distances to colonize even remote upland areas They become more abundant than their natural enemies particularly after favorable rains following a prolonged drought (196) The drought kills the armyworm and natural enemies (mainly parasites) but the armyworm can recolonize rapidly

The natural increase insoil fertility from mineralization of soil N over the drought period promotes luxuriant plant growth to foster a rapid buildup of these highly fecund species Weeding rice increases tilelikelihood of damage The parasites normally return to control the armyworms but not until after serious damage

Outbreaks on upland rice have been reported in Ghana (3) inzibar (38) Central Africa (36) lPanama (165) Brazil (28) Malaysia (196) and India (167) Defoliation may be severe often leaving only the base of stems Armyworm larvae hide during the day under litter or in soil cracks At night they ascend plants to feed Larvae pupate in the soil

Thrips Thrips frequent upland rice but damage is less than in the lowlands Thrips larvae and adults feed on leaf blades by rasping causing leaf rolling and stippling Thrips prefer vegetative stage rice and their survival increases during drought (heavy rain washes them from foliage) although drought stress usually outweighs thrips damage Drought and thrips cause leaf rolling in upland rice (235) but thrips have been overlooked perhaps becaLse of their small size Thrips are most numerous indry season irrigated rice The combination of favorable plant growth and lack of rainfall is ideal for thrips In contrast the lack of rainfall in upland rice hurts the crop and eventually the thrips whose numbers must decline

In [atin America Frankinilla rodeos attacks panicles before they emerge from the boot calusing sterile grains

Stem hug~s Black bugs are the most common group of rice stem-feeding -lemiptera in Asia The most frequently mentioned species ----Scotinophara coarctaia and Shrida - prefer aquatic habitats and therefore could not occur in upland rice as sometimes reported Upland species in the same gerurs - notably S tarsalis and S scotti -- are morphologically similar and remove sap from stems of grasses and upland rice The upland species are notably more abundant in upland rice planted near forests

In Latin America tile Tibraca ima-large pentatomid tiventris causes deadhearts and whitehcads on upland rice Both adults and nymphs remove sap from internodes of plants more than 3 wk old Tillers are killed by mechanical damage and by the entry of secondary microbial infection This damage is often confused with that of lepidopterous stem borers (233) Eggs are laid on the leaves and stems Nymphs are gregarious This species can also be found on irrigated rice Drought reduces population buildup

Stiem borers Lepidopterous and dipterous stem borers are widely recorded on upland rice but cultural practices and alternate hosts determine abtndance Except for Elasmolmll)uS Iinosellts numbers usually build up toward

IRPS No 123 January 1987 13

the end of the crop growth cycle Therefore late-maturing varieties staggered planting and lack offloodingfavorstem borer buildup Many stem borer species can lie dormant in the stUbbleafter harvest and theirsurvival is encouraged by panicle harvesting (leaving tall stems) and lack of tillage after harvest

Stem borers disperse and lay many eggs Many species are polyphagous however the monophagous MaliarpiasepashyratellaDiopsis Ivmgicornis Scirpophaga incertlas and S innotata can also be abundant on upland rice (hilo Suplressalisdevelops fasterand becomes la rger when reared on upland than on lowland rice (226)

The lesser corn stalk borer lasmopal tt Ittoselhs is semisubterranean and is perhaps the stern borer most adapted to upland rice(129) Highly polyphagous it infests maize peanut and cowpea It attacks seedling rice (one larva can kill tip to four plants before maximum tillering) (195) (hiv and Maliarphastem borers prefer older plants But Scirpophagaspp will attack young plants even in the nursery lasn opalls larvae tunnel into the stems at or below the soil level mausing dcadhcarts Larvae are not found in their turnnels inside sterns beca use vhei disturbed they retreat into cases made from soil particles bound by silk The cases areattached to the tunnel entrance (143) This behavior probably evolved as protection from natural enemies Larvae pupate in the soil The lesser corn stalk borer is more abundant during drought damage is often confused with that of drought (233) This species prefers sandy soils arid its distribution within a field may relate to soil texture (143)

Diatraeasaccharalis aside from the lesser corn stalk borer is frequently a stem borer pest in Latin America According to Teran (233) rice and maize are more preferred hosts than sugarcaine its namesake

Atcitonacrvsograpla is a less known upland rice stem borer in Asia with habits similar to Easinopall s (82) because it is not found in lowlands Acutra has grassy weed alternate hosts As a moth it is often confused with the dark-headed stern borer(hi polur(standstriped stern borer (hi sup)lpsrcais

Other common upland rice stem borers are also reported in the lowlands The dominant species in upland rice vary greatly with location In Uttar lPradesh India (78) and in Japan (123) the pink stem borer Sesamia htfirens is more prevalent hIKenya the principal rice stem borer is Chi partelus

(92) The lowland African sten borers Aialiarpha sepashyratellaDiopsis vngicornis and Sesamia calatnisnis are less abundant Oimupland than on lowland rice(217) but perhaps cause greater yield loss because plant injury is greater if combined with drought css (92) The stalk-eyed stern borer D)ivpsis is more prevalent in wet habitats (6 1I217)

In Central and West Africa Maliarpha and Sesamia calanistis are the dominant stern borers of upland rice (8 9 10 II) (lilo dihlfilineui and Chio partellus inhabit the

14 IRPS No 123 January 1987

upland savannas whereas Chilo zacconio prefers the lowlands (II) Aaliarpha is most abundant in the mangrove swamp habitats (91) coastal regions (16) or rain forest zones (II) of Africa and appears to be the homologue ofScirlpophaga incertulas The yellow stem borr can also be prevalent in upland rice (120 167) The sugarcane borer Eldanasaccharinais so far only a potential pest of uplandrice (91) despite Grist and evers (82) warning

Grasshotl)ers andI locitas Short-horned grasshoppers and locusts -- Locvta Patanrga Schistocerca - layeggs in the soil inhabit grasslands and develop into swarms to seriously damage upland rice from time to time (173) Largenumbers can destroy a field leaving only the stubble In low cmuoroer s giassnoppeirs an cut panicles (173) Farmers may not plant if locust swarms are imminent Prolonged dryweather followed by favorable rain favors development of large swarms (173) Usually breeding in di) grasslands theydisperse toattack crops including upland rice(191) Eggsaresusceptible to desiccation d uring drough t and toinundation during seasonal rains With favorable soil moisture over consecutive seasons natural enemies ca nnot prevent a largepopulation increase leading to migratory swarms (64)

Most grasshoppers on rice are adapted to uplands (107)but lowland rice grasshoppers iervgl(rlhusspp and Ora spp occur occasionally on upland rice (2 82) They lay eggson rice foliage Nymphs are scmiaquatic

The gryllid Lsctrtus coucinnus is a pest of lowland and deep water rice in Bangladesh and Thailand In the Philippines it damages upland rice grown in isolated pockets near lakes The gryllid nymphs and ad ults feed on the central portions of folrage leasing only the midrib and leaf nargins

Leal ler Pvralid moths whose larvae fold leaves to make a feeding shelter occur worldwide (217) but arc appareritly most common in Asia Leaffolde rsattacking rice in Asia are a complex of species (26) (Ctalhahcrcisinedinalis the best-known rice Ia flolde r in Asia may prefer grassy weeds over rice (86) It is a late colonizer of uplandrice U1arasmja eviua a lowland species (220) nreflers rice to grass specres (86) and colonizes upland rice during il early growth stages llarasnria patnalis is the most corn-rnonly encountered species in upland rice in the Philippines Perennial grasses are their alternate hosts and they can survive year-round in upland rice environments The leaf-folders 11arasmia ruralisand Alarasmia spp are least aburndant Dormancy is unknown in leaffolders Cnaplha-ocrocis medhnalis however is known as a migrant (90)Leaffolders remove photosynthetic tissue attack during the flag leaf stage is particularly injurious Popurlations are normally held in check by natural enemies but leaffolders become parlicularly abundant in conditions of high plantfertility and shade

Butterflies Skippers PelopidasInathiasand Borhofanta are more prevalent in African lowland than in uplandhabitats (6) but extensive studies in Japan and China

showed that Pelopidas mathias prefers an upland habitat (155) Parnaragutata seasonally migrates between lowland and upland areas and is equally abundant in both environshyments (164)

The green horned caterpillar Melanitis leda ismene is more abundant in lowlands (212) but our experience in the Philippines suggests that Pelopidas niathiasand 11elanifis leda ismene are often more abundant in uplands than in lowlands Bamboo iaunbuiva spp annually sustains P mathiaspopulations in the Philippine uplands Adults feed on nectar from flowers and migrate to upland rice during the late vegetative stage

iPi erlled h i-1 1d dill L tIC i 7-0 l(become numerous probably because of high egg predationEgg and larval parasitization are often low Butterflies can readily disperse long distances to seek renlote uplandricefields However they have low biotic potential and even it predation rates are low the rarely become abundant Upland environments with their diverse microhabitats can provide more favorable sites than lowland rice plains

tutterfly pests of rice have broad host ranges and become dormant Juring urnfavorable times of the year

IolhagrotslaidopwraThe larvae of many polyshyphagous moths are reported to defoliate upland rice tussock moths lsaispelnnatula Iawlia sUifita luproctisirgtwltaE mnorand P vathorrhoeaslug caterpillar

litotiahicolorhairy caterpillars Nisaga sinp vandA mata spp and wooly bears l)iacrisiaohliqtia and Crvatonotts gangis Many of these polyphgous larvac migrate and are transients in rice including HelhWhis arm iera and 1t zea highly pol phagous species (82)

Gall nidges The Asian rice gall midge Orseoliaorrzae (88 131) and its African counterpart 0 ortzivora(6 11 12 32217) prelerlowland to upland rice Gall midges in rainfed areas are prevalent only during tle rainy season and dispersepoorly Their alternate hosts (wild rices LeersiaIschaetnuan and Pasahmt)are mostlyaquatic grasses and are less likelyto be near upland rice areas In an area of mixed uplandfloating and rainfed lowland rice cultures in northern Thailand upland rice which is ilanted a month earlier was attacked firstbut subsequcent populations were much lower than in rainfed lowla id or floating rice Low numbers of gallmidges passed the dry season orr perennial weeds such as Paspahim (88)

Smot (hid(s Reports of aphids attacking upland rice plants are few (249) Without maize the maize leaf aphidRho alosihhunm maidis attacks upland rice in the Philipshypines The plum peach aphid IIrsteroneuraselariae was reported in Sierra Leerne toinfest leavesand tnriperied grain(7) Siphaglrctriaecaused transverse linrea r necrotic striae iir rice leaves in Italy (172) I-lea v rains held populations in check Rice is not a preferred host for shoot aphids

Leafhoper andI )anthol)pers Cicadellids and delshyphacidsdispersecasilyand readilycolonie upland ricefrom nearby lowland areas A study in Sarawak (241) showed

IRPS No 123 January 1987 15

similar species in upland and lowland rice Nephoiettix The principal vectors of rice virus diseases generally have species dominated The most prevalent species accounting narrow host ranges and alternate hosts in the grass family for 90-95 of the sweep net samples wre N virescens N are mostly annuals dying in the dr3 season Some virus umropictus Nilaparvia hgens Soatellafiircifera Cfana vectors such as BPH can disperse lontg distances but few do

sectraand Recilia dorsalisLowlands had higher popula- Th restriction of upland rice and weed hosts to the rainy tions Katanyukul and Chandaratat (120) monitored hopper season the small area planted to upland rice and the hopper numbers in upland and lowland fields of northern Thaiiand vectors having narrow host ranges all work against virus and found similar results A comparative study of plant- diseases in upland rice hoppers in Fiji showed more brown planthoppe BPH) and In Latin America Sogctdhs o(rzicola and S cuhanus whitebacked planthopper (WBPH) in wetland rice than in transmit hoja blanca virus disease but reports on upland lowland rice and hopperburn was prevalent in lowland rice are rare except in the favored upland areas of Colombia areas with standing water (89) Peru and Venezuela (35 42) S rricolaocurs in Brazil

Hopperburn in upland ricefields in Asia is rare (84) In on upland rice but hojaI blanca has not been reported (67) Batangas Philippines isolated fields of upland rice become S orzicola has a wide host ramgc (10) similar to WBPI- in hopperburned infrequently from WBPH A report from Fiji Asia WBPH however is not known to transmit any virus cites hopperburn from BPH and WBPH (250) In these two disease situations upland rice was grown on highly fertic soil and In Africa pale yellow mosaic virus is endemic to swampy N fertilizer was applied Hoppers respond to better nutrition lowland areas where rice ratoons year-round (19) and is Normally upland rice is grown on poor soil and receives no mechanically vectored mainly by hispid beetles A minor fertilizer vector a flea beetle Chaeiocnema spp is an upland rice

In Asia upland rice ras are normally not far from pest and upland ricetields near endemic areas occasionally lowland ricefields-- a source of hoppers Otherwise hopper contract the virus species with greater polyphagy than BPH or Nelhotettiv Spittle bugs Cercopid nymphs producc a protective virescens would be dominant frothy covering that looks like saliva In Latin America

In Africa leathoppers and planthoppers do not become spittle bugs damage upland rice In parts of Brazil where abundant on upland rice (6 II) probably because lowland upland ricefields are surrounded by pastures adults rice is not widely rlanted nearby The only reported migrating from grasses can damage rice severely and cause hopperburn as from Nilapanrata maeander in breeder complete hopperburn in a young rice crop within a week of plots receiving high N rates in Nigeria (98) Hoppers colonization (194 195) Dispersal to rice occurs even if however are a greter problem in Latin America where pastures are verdant and spittle bugs prefer pasture grasses large grassland areas breed polyphagous species (Gra- to rice (168) Nymphs are rarely abundant enough to phocephala spp Hortensia spp lEvitianus ohscurunervis damage rice Regular rainfall favors their development Balclutha spp and Draeculacephala spp) vhich then (233) disperse to cause hopperburn on seedling rice (68) Rosetto et al (195) reported Graphocep7hala spp and Hortensia spp Seed pests to be equally abundant on upland and lowland rice Alydid and pentatomid bugs are seed pests of upland rice

The leafliopper and planthopper epidemics in Asia that worldwide In Asia Leptocorisa sp prevail Their habits caused severe losses from hopperburn and vijus dicases make them particularly suitable for upland rice 1)thev feed over the past several decades occurred principally on on many grasses (and therefore can survive the earl y wet lowland rice Except in a few isolated cases (160) upland season before rice sets its grain) 2)adults are long-ied and rice has been spared from tungro grassy stunt and ragged mobile allowing them to find isolated plantings 0 rice stunt virus diseases vectorcd by green lcafhopper (GLH) (besides seed bugs only birds and rats appear to have shyand BPI] (123) Virus diseases have occurred in upland rice capability) and 3) adults aestivate in forested areas or where lowland rice was nearby Rice dwarf disease however sugarcane fields during the dry sca son when neither rice nor was first recorded on upland rice in the Philippines (187) grassy weeds are present (I199) Rice seed bugs concentrate Virus diseases can be perpetuated only on livir plants or on small-scale upland riceficlds because they can actively insect vectors and no living rice - plans tooned or search them out The prevalence of groves of trees characshyvolunteer - grows year-round in upland arcas excert in a teristic of upland environments ensures nearby aostivation few with 12 mo of rainfall sites keeping the seed bugs close by

The smaller brown planthopper Laodcelphax striatellu Leptocorisae are uncommon in lowland rice plains occurs in upland rice in Japan (18) It spreads black- because 1) their populations are diluted in a sea of rice streaked dwarf and stripe viruses between winter wheat and planted more or less at the same time (they ca n only feed on barley to lowland rice Early planting of lowland rice rice during the milk to hard dough stages) 2) they lack nurseries spreads these viruses but upland ice has remained aestivation sites and 3) weedy fallow areas are limited uninfected The lowands of Asia are dominated by L oratorius

16 IRPS No 123 January 1987

which lays its eggs hign on tile foliage (199) L acuta and L solonlonlesis oviposit at ground level the former on litter and the latter loose on the ground These egg laying habis explain theirtenvironmental prefcrencecs(214) In Japan the postdiapause adults of L chinensis migrate from moun-tainous areas to upland ricefields during floweiing (1 14)

Pentatornid bugs prevail in Latin America (8182 195) and in Africa pentatomid and coreid bugs arc equally cited (6 82) l)amage is often characterstic of the species (5)rcoris ventralis and Mnida spp are the common

pentatomid seed bugs of Philippine upland rice but they prefer lEchinochva spp grass seed to rice Vearaviritda occurs worldwide and hoften more abundant in the uplands(54) We saw few reports of seed bugs on upland rice but see(d bugs apparently opelrate cqually between by land and upland rice Good dispersal is necessary ior species that call feed for only 2-3 xon a crop and must first find those crops at ripening stages

Other pct species feed n flowers In Mato Grosso Brazil (194) and Santa Cruz Bolivia the panicle weevil Aeo-hariuia amllitarsis attacks rice during flowering (68 233)The larva e rice sten horers adutlts feed at the bases of spikelets caIusing empty grains Also inLatin America thecydnid -lkiituttatralusisa seed bug as anad tilt and nymph (194)

A number of beetles ecd on poilen - coccinellids lialrotia Aulac lhora and Monohpl-a (55) - but becatise riCe is self-pollinating and fcrtiliation occurs before pikelets open they pose no threat to yield

IST ECOLOGY

We now look at characteiistics of the life cycles of upland rice insect pests to lea rn hew each plays an adaptive role in upland rice ecology

Life history strategies Orzasativadoes not tolerate drodgh well so upland rice is highly seasonal normally grown in the wettest months of the year Usually rice is present for less than half of the year The rice-fiee fallow poses serious problems of survival to upland rice insects which have evolved att least four mechanisms to ovrcolne tle cvcl ic-l lack of a hostI) polyphagy 2) longevity 3) dormancy and 4) vagility (dispersal)

Poldt)agy lolyphagy is defined as having hosts of at least two botanical families oligophagy of more than one gents and ionophagy of only one ventus (40) - inour case Orvza Most wild rices however are illi tiC and theieiirc are normally far removed from upland rice hahitat othewise more would be alternate hosts for uplanJ iice insects Thte highly mornophagous rice pests shySciriol)haita hiceritlas Nilal)arvaiahiens Iand Nealo--tettiyvircsctws in Asia and Aalirpha secparatella and 1ioj)sis lngicornis inAfrica - occur on upland rice but

are more abundant in lowland culture They can specialize in rice because they are highly vagile and can attack all growth stages By specializing in rice monophagous rice pests can outcompete related but polyphagous speciesIntensive andI extensive lowland rice culture has favored monophagous species Riqwla alhinella (Cramer) the onlymonophagous species in Latin America is rarely reported on upland ice (I1)

Because of its limited temporal and spatial existence upland rice has not favored the evolution of specialized species Clhang(44)shows evidence that upland rice has only recently been cultured by man from the lowlands thus pests have had little time to fully adapt to the crop All but the lowland-adapted insect pest species have alternate planthosts to rice Root pests with limited mobility are highly polyphagous Pests that specialize in one growth stage eg secdling maggots seed bugs are oligophagous or polyphagous

Most alternat hosts of rice pests are annual grasseswhich also are seasonal Upla_rice is also a host but not the preferred one of the most highly polypliagous spcciessoil pests grass hoppers armvwarrs and Vearaviirielda

The d ry season poses igreat obstacle to ani upland riceinsects survival and only sonic perennial plant hosts are suitable food during the off-seasoin Plolyphagy allows upland rice insects to suLrx-ixe during thC rainy season when plants- both annual and perennial -areactivelygrowing and therefore more nutritious

Loiugevitr The ability of aiinsect to live 2-3 mo without undergoing dormancy helps species sich as root-feedingwhite grbs (as larvae) and secd-feeding bugs (as adtilt) to survive unfavorable periods White grubs arc not highly fecund and their life strategy is to improve survivorship of limited progeny Their subterranean habitat hides them from manv natural enemies lBtut root tissue is not highly nutritious so they ltilst eat great quantities and need a largeinsect biomass to digest it Other soil-inhabiting rootshyfeeding insects have short life cycles but are more fecund

Seed pests need greater longcvi vas adults to locate ahost at early grain dcclopment arnd give sedentary offspring time to develop Few groups of upland rice insects live longin active development Polyphagy and longevity only increase their ability to survive during the rainy seasonOther mechanisms are needed to survive the dry season

orantc In the tropics the nonrice season is the dry season while intemperate areas tile off-+season is winter Many upland rice insects undergo dormancy inlsummer (aestivation) or winter (hibernation) Dormancy is simplyinactivity during winters with prolonged temperatures below 10 C insects find shelter arnd cease movement Activity is resu med with warmer t empcera tu res A deep state of irnactivity termed diapauise occurs in sole species that prepare physiologically (havxing low metabolism and convcrting gonad s to fatty fuel reserves) for the unfavorable season (148) They enter diapaUse inanticipation of unshy

IRPS No 123 January 1987 17

favorable weather and resume activity after favorable Armyworms skippers and locusts actively travel hundreds weather begins usually in response to photoperiod or of kilometers in air fronts such as intertropical convergencethrough a delayed response to a stimulus such as heavy zones to descend onto upland rice fields far from their rainfall (237) Not all individuals of a population have the breeding grounds (31) Several generations can build ip on same threshold for environmental responses for entering or alternate hosts in tile grass family before rice is plantedterminating diapause -- this is seen as a mechanism to These large insects are active fliers Smaller insects such as prevent total mortality if a false cue appears such as a thrips aphids and planthoppers passively migrate long sudden warm spell in winter or rain in the middle of the dry distances by the wind season (154) Otherspecies ean readily travel tens of kilometers - stem

Probably more upland rice insects undL go dormancy borers leaffolders leafhoppes cutworms spittle bugs leaf than we realize Candidates are Atherigona spp root beetles seed bugs ants and termites aphids root weevils crickets leaf beetles thrips and seed The least dispersive species are white grabs mealybugSbugs Insects known to entervarious states of aestivation are seedling maggots crickets weevils and gall midgesleptocorisaspp adults (199) Maliarphascparatellalarvae Each species has evolved a unique set of attributes to (9) Scirpophaga innotata larvae (117) Chilo partelhtsv enable it to survive and adapt to the changing uplandlarvae (21) ScirpopThagainccetulas (179) white grub larvae environments Several evolutionary avenues lead to fitness (134) Patangasuccincia adults (6 82) and Schistocerca (Table 2) but there is no reason to believe one set of gregariaeggs (82) attributes is better than another (224)

Insects known to enter hibernation are LodeIphaDstriatellus nymphs (124) Chtio Tvlpressalis larivae (125) DroughtSesaia inferens larvae (146) but not S calavisis(2) Well-drained rainfed upland soils are subject to drought ifIDiaraea saccharalis larvae (75) Scirpophaga- tcerlttas rains do not fall within 2-3 wk Prolonged drought followedlarae(5 lacchrli s lae (75)saSlOhagasshoperseggs (82)and felo pidas athias pOpae (232) by favorable rains stimulates armyworm and locust outshybreaks but mealybtigs root aphids and thrips become

Lcpmotorisa oratorisand L acuta adults aggregate on numerous onlyafter dry spells for two reasons First heavy trees sugarcane or other shady moist sites L paluwanmsis rain normally kills soft-bodied foliage-feeding thrips and confined to Sulu and Palawan Philippines lives on the mealybugs and soil-inhabiting root aphids and root mealyshygrass Brachiariamutica (Forsk) Stapf in the absence of bugs Drought eliminates this source of mortality Second rice The adults can live for 5 mo When aestivating their the rice plant responds to drought by breaking down rate of metabolism drops and they do not feed They will proteins into soluble N compounds which enter the phloem however puncture plant tissues of their aestivation host in and are taken up by these sap-feeding insects (245) Greater search of water This behavior has led to errors of the host nutrition therefore leads to greater survivorship and range of rice bugs (I) Aestivation quickly terminates with fecundity rain probably meaning aestivation is not deep Small rice area

Some stem borers aestivate as mature larvae in rice Except in the highly mechanized large landholdings of stubble or in straw stacked as livestock feed However Latin America upland ricefieldstypically are patchy within moriality increases proportionally with the duration of the a highly diverse flora The small fields result in higher ratios dry season even if the stubble remains undisturbed If the of perimeter to area than with the typically larger lowland stubble is plowed for a crop following rice few stem borers ricefields Upland ricefields themselves may be intercroppedsurvive Rainfall terminates dormancy but it normally with cereals legumes or root crops Mixed intercroppingtakes several weeks for moths to develop and emerge with a wide variety of species also occ in tribal slash-andshy

Last-instar white grub larvae tunnel 1-2 m deep in the soil burn agriculture (50 74) to construct pupal chambers in which they aestivate The Upland ricefields tend to be small because of the labor larvae are very sensitive to dryness so pupal chambers are needed to clear land in slash-and-burn areas and to till land sealed to coierve body moisture Aestivation terminates to minimize weeds Rice requires more tillage than does with the first heavy rains but as in stem borer 3-4 wk pass maize because rice competes less well with weeds Uplandbefore adults develop rice is more a subsistence than a cash crop and small areas

Insects may respond differently in each region to cold (I ha) can feed a family temperature or drought Biotypes or loal populations may Small fields favor some insect pests such as some ant evolve (56) Local populations of Scirpophagaincertulasor species that can forage from more permanent field borders Patangasuccincta may have become adapted to prevailing Trees provide food and mating sites for white grubs (134) as conditions well as aestivation sites for LetTocorisa spp (199) Field

iagiliti The ability to disperse combined with high border grasses plus tIhose within a ricefield provide alternate fecundity orshort life cycles enable some upland rice pests to food for all but the highly monophagous upland rice better exploit the temporarily favorable upland habitat colonizers and a habitat for natural enemies

0 z

Table 2 Life history patterns of upland rice insects or insect groups

Host range Longevity Dormancy Vagility FertilityMonophagy Oligophagy Polyphagy Low Medium High Yes No Low Dispersive Migratory Low Medium High(Oryza spp) (grasses sedges) (angiosperms) (lt3 wk) (gt2 mo) (lt1 kin) (lt50 eggs) (gt500 eggs)

Ants xTermites x x x

Mole crickets x x x xx x xField crickets x xx

White grubs x x x xx xRoot aphids x x xxMealybugs x

x x x xxSeedling maggots x

x x x x xLeaf beetles

x xx xArmyworms x x xx x

Thrips x x xx xChilo spp x x xx xMaliarpha x x x x

xLocusts x x Xx x xGall midges x x xxiVephotettL iirescens x x xx x x xLeprocorisaspp x

x xLeaftolders x x x

Sogatellafurcifera x

xx x x x x x x

00

IRPS No 123 January 1Q87 19

All this appeals to favor insect pest buildup but the notwithstanding the release of stored nutrients in response contrary is true Loevinsohn (138) showed that insect pest to drought that temporarily benefits philoem feeders populations respond exponentially with the proportion of land devoted to rice up to about 75 of the area then the M POR ANCE I I P1A 1) RWI P1St S

rate of response declines In most upland rico areas tie rice crop occupies less than 50i of the area- therefore the Visits to upland ricefields normally reveal 1kw insect pests potential for population buildup is low This relationship Loevinsohn (138) and Locvinsohn ct al (139) compared may not hold for seed pests that can locate small ricefields insect abundance using annual light trap atchcs in upland

Another outcome of the highly diverse flora and phical rainfed lowland and irrigated lowland sites in the Philipshyenvironment of upland rice isthateach upland ricearca has pines Although only one upland site was studied data a unique composition of insect pests showed equal or fewer insects at the upland site than at the

For example in Tanauan Batangas Philippines the rainfcd lowland sites and fewer than at the irrigated sites main upland rice pests are rice leaffolder(RLF) armvworm (Table 3) M Lltiple regression analysis showed that the key Aythntnaseparata WBPH ants Soli psis gentitataand factors explaining insect abundance among a set of cropping white grub Leucopholis irrorata Sixty kilometers away in intensity variables were the number of rice crops grown per Real Quezon the main pests are seedling maggot Athe- year followed by area devoted to rice Table 3 shows that rigona orae flea beetle (Thacitcmnra hasalis thrips the upland rice site had only I rice crop perym rand that rice Stenchaetothrips hiurnmis and rice bug Leptocorisa was planted in only 20Vi of the area Fertiliter and oratorius In a third Philippine upland rice site in Claveria insecticide use had little bea ring on the trend The armters at Misamis Oriental Mindanao the main pests are ants the upland site applied 60 kg N ha higher than at most Solenopsis gemtinata seedling maggot Alieriona orvzae irrigated sites white grubs Holotrichia in)iUaiuoan t and Lucopholiv Insect occurrence at the upland site was omparable to irrorata stem borers Sesamia ifiArelr (hio auriciliusC that at the Cagayan rainfed lowland site parlicularly in 1982stpessallis Scirpophaga incertulas S innoiata and when drought prevented many farmers fron planting(only

4c ona chrrsurapella root aphids Tetraneura niriah- 40(of the rice area eventually was planted that year) M ore dunrinalisand Rhupaosiphunm riahbdminalis lcaffolders favorable weather occurred in 1980 and 1981 which Marasniapataltis and Chaphalocrocis Medinalis army- supported greater rice areas and consequently more insects worm MIthinna separata and rice bug Leltucorisa The most intensive site for insect pests was South Cotabato oratoriusThe great diversity of insect problems is typical Mindanao where farmers plant 23-25 rice crops per year fer upland rice and makes control efforts more difficult with irrigation The Zaragoa Nutva Ecija site also had particularly regarding breeding for insect resistance high pest abundance and at this site the fields were highly

asynchronous Asynchrony was the third most important Low yield potential Iactor in explaining rice insect abundance Most upland rice is grown on low-nutrient and mineral- Rice cropping intensity in upland areas is bound to toxic soils which when combined with eiratic rainfall remain low Only one rainid upland crop is possible per make upland rice a highly riskycrop for the farmerto invest year (Farmers in Claveria Misamis Oriental Mindanao in costly soil amendments and land management Often Philippines tried asecond upland ricecrop i 1984a ycarof upland rice isgrown far from markets making inputs even favorable rains but no harvest was possible b-cause of more expensive Farmers growing a subsistence crop hosi- drought blast and birds) Rice area isaso bound to remain tate to invest in costly inputs to raise yields low because of the high labor and power rcquirelmenlts to

Even having varieties that could double existing yield prepare the land (because of weeds)and because upland rice potential under present management levels would probably wvill not compete with lowland rice as a cash crop Some not prompt many farmers to use purchased insecticide upland rice sites with prolonged rainy seasons howeverare unless tile risk of crop failure were reduced such as it was planted asynchronously such as the Clacra site with irrigation systems in lowland rice The seasonal abundance of Upland rice insect pests

The Tanauan Batangas site is atypical for upland rice attracted toa light trap wasgrapied for crop year 1980-81 in The soils and rainfall pattern are favorable and because of Tanauan FBatangas (Fig I 2) W13P11 was he most its nearness to Manila farmers have cash resources ftm common rice hopper and its population peaked at the sales of vegetables to purchase tertilier They applied an midgrowth stage as it does in lowland rice D-Vac suction average of 60 kg Nha after particle initiation and obtained samples showed an earlier peak than that from a light trap 25-35 tha yields The high fertility isone reason why RLF indicating emigration W13PI usually emigrate after the armyworms and WBPH developed into large numbers vegetative stage Populations in 1980 reached levels that Farmers however used no insecticide on upland rice caused patches of slight yellowing in some fields WBPH although they did on vegetables probably cause hopperburn in Batangas They immigrate

The poor growing conditions fOr rice generally mean year-round as shown by low levels in the light trap BPH poor growth and low fecundity for insect pests fIeding on it however were scarce in the light trap and not collected with

Table 3 Abundance of rice insect pests and factors in rice cropping intensity of Philippine upland rainfed lowland and irrigated lowland locations where light traps were operated daily for atleast I yr -I

0Rainfed Rainfed lowland Irrigated lowlandupland

Pangasinan Cagayan lloilo Santa Maria Nueva Ecija South Cotabato Tanauan Caaringayan Caaringayan Bangag Bangag Bangag Buray

LagunaCale

Batangas Manaoag 1980-81 Santa Motiica 1982-83 Cabanatuan Zaragoza Avancena NaniamanSolana 1981-82 1982-83 Oton

1980-81 1979-80 Oton 1981-82 1981-82 Koronadal Koronadal1980-81 1979-80 1979-80 1983 1983

Insect (kerosene light trapNephotcttixcatch nolyr)virescens 650 1240 500 1800 330 210 3400 1600 160N nigropictus 210 - 360 18400 5500 2900- 140 90 20 - shy 1700 -Reciliadorsalis 710 - 3200 1700180 890 630 140 170 2400 800Nilaparratalugens 130 690 - - 4200 4600300 520 750 180 60 3300 1600 740 310Sogatellafiircifera 1700 31000 13000 8200310 210 3300 610 270 980 1600 6300 -Cyrrorhinus - 12000 6700600 540 3200 290 160 330 120 410Scirpophaga spp 50 410 620 - - 21000 12000150 180 220 270 30 1000Cnaphalocrocismedinalis 70 70 540 610 270 11000 6100200 140 100 6 60 250Chilo suppressalis 0 60 60 1900 300 200- - 30 20 0 6 3 160 -C auricilius - 0 00 - - 20 3 60 20 3 0Sesania inferens 0 - 80 60 3 80 10 51 -Cropping intensity _ -

Rice crops (noyr) 10 10 12 10 10 10 13Rice area (7c) 17 19 20 20 25 2320 85 85 60 70 40 85 85 90Fertilizer (kg Nha) 60 20 85 80 70 7020 0 0 0 30 30Insecticide (no of 40 60 40 300 04 04 0 03 300 13 13 30 50 40 33 33applicationscrop)

__________

IRPS No 123 January 1987 21

Light trapped Insects Drytnd __ 2(notrap-nght) Suctioned Insects (nom )

50 -)o 9 jO40- SOW ffurciWO Light trp

D-Vc suction

20 to 4

20

Egg porasitizahion (M) 75 60

300 SOOampReOftwciffr

45 30shy

15

-insects (notrop-night) 4 Nileporvata Iuen5

Egg parastization()

Light trapped insects (no trop-nght) Suchoned insects (no m 2 )

6 8 Cytorh~nuls hvidipennii 0

4 05 2

50 0 Insects (noltrop-night)40-ui Ipanh pe g

Moohase 1 Seasonal abundance measured by light trap

Rainfall(etod) and D-Vac suction sampler of whitebacked

to ~ hopper Nilaparvata lugens and lea ffolder60)-t Cnaphalocrocis mnedinalis andplthpe g20 L I II I I I parasites and mtrid predator Ortorhjntus livi-Apr May Jun Jul Aug Sep Oct Nov Dec JaIn Feb Mar dipennis set against cropping pattern daily1 9 80o 19 8 1rainfall and moon phase Tanauan Batangas

1980-81

the D- Vac Cyrtorhinus build up on the crop coterminous are collected in light traps during full moon (109)with WBPH buildup Cyrtorhinus disperses during the Because upland rice shares most of the lowland rice pestsrainy season RLF not readily collected in light traps was and those pest species are generally less impotant to uplandprevalent only during the rice crop rice than lowland rice there is a general belief that insect

Nephotettix virescens was more abundant than Nnigro- pests are less important onpicLus in the light trap collections and on the crop GLH upland than lowland rice Arcview of limited data on upland rice shows an averag of population peaked at the reproductive stage as did zigzag 10-21 yield loss from insect pests on a par with that ofleafhopper (ZLH) Crtorhinus tracked the populations of lowland uee determined by the same methodology (132) planthoppers and leafhoppers Both GLH species were (Table 4) McGuire and Crandall (144) made a similarcollected year-round in the light trap indicating immigra- estimate for Central America Cramer (51) howevertion from nearby irrigated rice areas Possibly because of the estimated 35 yieldlow populations loss in South America based on ainsect number did not correlate with similar pest complex - stem borers and seed bugs shymoon phase although reports indicate that more hoppers attacking irrigated rice in North America This estimate did

22 IRPS No 123 January 1987

Light trapped insects D~~nd (no trap-night )

St plht Suctined insects (nom 2

)

D-Voc suction 1520

1o

0 Lih trap 5 ~105 o5

00 Eggparositizotion()5040 NephOtgtha virneens

30 20

I010 A-Insects (notrop -night)

4 Nephatelh nigropccent i

2 o I vV^J - -- ^ VA____ f _

Light trapped insects (no trap-night) Suctioned insects (no rnE

10 -1208 ReCdO doRsol

6 15

4 10 052 iwxALAiamp 0

Light trapped insects (notrap-night) 2Suc oned nsects (nom )

8 CyrtorhmnusI6o nn]10 i0075

4 05

2 0 0

025

2 Seasonal abundance measured by light trap Roinfall (mmd) Moon phase and D-Vac suction sampler of green leafhoppersNephotettix virescens and N ngropicrus and I zigzag leafhopper Recilia dorsalis their egg 60

parasites and micid predator Cyrtorhinus livi-dipennis set against cropping pattern dailyrainfall and moon phase Tanauan Batangas1980-81

40

5 Lf Apr May Jun

1980 Jul Aug Sp Oct Nov I

-aMar

1981

not account for soil pests and was not based on field data in the absence of pests Carbofuran granules are phytotonicAkinsola (I1) cited mainly data from irrigated rice areas in to rice (239) The prevailing hypothesis is that carbofuranAfrica but estimated yield losses of upland rice between 15 prevents soil bacteria from consuming fertilizer The phytoshyand 30 from insects However losses in Thailand over a tonic effect is through greater availability of fertilizer to the4-yr period ranged from I to 13 averaging 5(106) plant Denitrifying bacteria are known to be more active inYield loss data however should be interpreted with aerobic than anaerobic soils therefore it may not be merecaution (184) The Philippine data were all derived from coincidence that the highest recorded yield losses (22-69)trials on farmers fields under farmers normal agronomic occurred in those trials with 12-30 kg basal Nha On themanagement practices except for Bukidnon and Capiz other hand contro with carbofuran granules may not be aswhere researchers used fertilizer Batangas farmers are good as a seed treatment against seed pests and perhapsatypical because ofhigh levels of fertilizer used The trials in yields would have been higher if a systemic seed treatmentBrazil were carried out at an experiment station but insecticide had been used conditions were similar to those of local farmers The 1979 and 1980 trials in Batangas had two full-

Carbofuran granules were used in every trial as a broad- protection treatments one using carbofuran granules andspectrum insecticide applied basally with the seed An the other using bendiocarb WP as a seed treatment Plotsobjection to using insecticides to measure yield losses isthat protected with bendiocarb yielded higher than those proshyinsecticidescandirectlyorindirectlystimulateplantgrowth tected with carbofuran granules The other trials in the

Table 4 Yield losses to insect pests and yield responses to insecticide in upland rice in the Philippines Brazil and Upper Volta

Yield (tha) Yield loss (5) Location Year Cultivar Protected

with Protected

without Check Total Vegetative Reproductive Ripening

Basal N (kgha) Reference

carbofuran carbofuran Protected No with carbofuran

carbofuran

Philippines Pili Camarines Sur Tanauan Batangas

Pangantucan Bukidnon

Dumarao t-apiz

BrazilGoias Minas Ge~ais

Upper Volta6

1975 1974 1974 1975 1975 1976 1977 1978

1979

1980

1979 1980 1979

1977 1978 1979

Bursiging puti Kinanda Dagge Kinanda Dagge Dagge Dagge Dagge UPL Ri5 Dagge UPL Ri5 Dagge UPL RiS UPL Ri5 UPL Ri5 UPL RiS

IAC47 IAC47 IAC47

14 33 11 36 35 31 30 27 37 26 40 30 40 42 46 35

19 21 09

--

-34 24 24 27 --

29 43 31 42 -

47 25

---

10 28 05 27 27 30 26 29 33 28 33 29 43 23 36 11

15 15 06

29 15 55 25 23

3 13 -7 11

-7 18

3 -7 45 22 69

24 28 35

--

21 -11 -20

4 -

-3

23 7

-2 -23 56

-

-

-__0

6 --

-

-23 14

5 0 8 2

22

-

--

0

-

--18 11

3 -10

4 10

7

--7 -

14 -

-1 6

10 -5

8 9 7

-

--

0 0 0 0 0 0 0 0 0 0 0

30 30 30

12 12 12

99 99

100 100 101 101 102 102 102 102 103 103 104 104 104

66 66 66

Farako-Ba

Mean

1977 1978

IRAT10 IRATIO

38 66

--

40 43

-5 35

21

-

10

--

--

--

- 242 243

Z

tJ

-w

24 IRPS No 123 January 1987

Philippines which had two treatments offering protection In the 9 Philippine trials where yield losses were measuredwith and without carbofuran granules relied on further by growth stage 10 of the yield loss (21) occurred at thecontrol with foliar spray and not on seed dressings The vegetative stage and 517 occurred at each of the reproshycarbofuran-protected plots had an 18yield los compared ductive and ripening stages Ibis result is similar to thewith only 10 with foliar sprayS This yield difference is pattern of yield loss in lowland rice measured by successiveprobably due to differences in protection against sown-seed treatments where the crop is unprotected at each growthpests (foliar spray do not control sown-seed pests) rather stage but proticted during the other growth stagesthan phytotonic effects became basal fertilizer wa used in However the insect complexes r sponsible for vegetativeonly 2 of the 10 comparisons stage yield losses in upland rice are entirely diftfrent than in

The recorded yield losses were not related to yield lowland rice The key vegetative stage pests in lowland ricepotential (Fig 3) One might expect higheryielding crops to arc aquatic whereas those in upland rice are sown--seedhave higher percentage of yield losses but this ws not tile (sown in soil) root and seedling pests This dift rence maycase One reason is that insect damage exacerbates plant explain why agricul torists have overlooked tile ilmportanccinjury from drought stress low yielding fields from d ought of upland rice insect pests Many reports of upland rice pestsstress would show disproportionate differences between in Asia ha Ve focused on major lowland species (46 108)insect-protected and -unprotected treatments Shoot and It isalso surprising that tei vegctative stage of upland rice root pests would be more responsible for losses from would record the highest yield losses becatuseseeral studiesdrought stress than sown-scd pests A second reason have shown that upland rice can readily recover from foliageconcerns seed pests which would have a greater impact on reinoval during that stage (117 152 165 188) Upland ricelow tillering low yielding varieties such as )aggc Low poscsses a hitgh obilitV to compensate for ca rly loss oftillering varieties nnot fill in the space created by removal foliage and will even be stinulated by foliage removal toof seeds oweer 1) R i 5 is high yielding because it tillersP1 produce a higher viel than plants ithout any foliageactively which can close the canopy to provide greater weed removed (63 192 231) [ie reason for this apparentcontrol discrepancy hovce is that yield loss at the vegetative

stage does not octUr frol leat arca loss but fron pestsYield loss () removing sown seed and feeding on devcloping tillers and roots -- damage that cannot be readily conpLtnsatCd for

70 - Measuring yield losses in upland rice cnvironrnents is further complicated by a report from Brazil that greater yield losses occurred in response to better insect control60 - =-09 as Insecticide seed treatments resuled in denser plant stands

(n=21) which in turn created a selectively more favorable environshyment for blast (69) This resuIt was parlticurla rlxexacerbated50 when cirbofuran granules were used in seed furrows Aside

ffrom a greater plant stand ca rbofuran iay have auigmented the N lecl and stimulated blast If so ca rbofuran should be40 - replaced with another insecticide or the seeding or fertilizerI rates decreased

Other methods to determine the impact of upland rice30 shy pests wvcrc to interview farners or to develop single speciesbull bull correlations of population levels and yield loss Feneira (66)2 NIM)st (75(1) Golas farmers reporteddid both in Brazil

20- insect losses in ufpland rice and 251 said those lossesS avcragcd 37(1 [his result is consistent with tie yield losses 0 0 measured by field trials in the same statc(Table4) For every

I0 4 of seedlings removC d by leaf-cutting ants a I 7yield loss is predicted Similarly Ferreira (66) found that ai

0 - average of 53 thrips per panicle before panicle initiation meant 3 times more unfillh d grains

-IO0 I 1 1 I I-101 0 2 3 4 5 6 7

Yield (tho) C)N [ROt IM FI IODS 3 Correlation between upland rice yield and yield lols caused by Upland rice insect pests may be controlled by culturalinsects measured in 21 trials in the Philippines Ivory Coast andBrazil genetic biological and chemical methods

IRPS No 123 January 1987 25

Cultural control dry season plowing will have little effect on them Plowing Crop husbandry to reduce upland insect pest populations when the soil is wet from the first monsoon rains will expose can be classified into practices effective in a single field and soil insects that tunnel close to the soil surface to predatory those effective only at the community level Cultural birds chickens dogs and even man (field crickets nole controls should come first in a pest management program crickets and white grubs a re delicacies in man tra ditional They have broad stable effects as pests have little pos- diets) sibility of overcoming them through selection of biotyps The greatest degree i1 inIscct control fron tillage comes (138) On the other hand some cultu ral controls work only from plowing soon after rice ha rvest when most soil insects with high labor or power inputs and connunity-wilt lie close to the soil surface anrd stern borers are in the rice methods require coordination of iany farniters stuble Whitegrubsare mature and tillage exposesthem to

Pantin time The luxury of being able to shift the time of predators Tirnelv control will prtec the crop following pla ntring upland rice is open onl hr la rniers in areas with a rice and lowr the populationIfor succeedinig years Plowingprolonged rainifall pa ttern lI copping intensity or underthe ice stulble helps it decompose killing sten borer rnechanied land preparatiot Gcnerally planlting as soon larvae inside as the rFiinv season begins will lower populatiois of 1110st [requent tillage destroys ait nests bit not termites insect pests Insect populationsarelo aftcra dlvseoll or Tillage and clean culture howes er rcliove food for winter fallow delayed plaiting lets them build tip on termites which then mtay attack a young rice crop Scarab alIternate hosts that gi vigorotsly with the first seasonial beetles also prefer to miposit in eceitly tilled fields Noshyratiris This first flush iof weds cros lItxuttriartlv with iceCult is dibbling faors the survival oftillace Clre Such rineralied N released during the dr season aid made stils i aels tnd insects thtt pttpate or lay eggs- in the soil

available to the plants with the rains It iliso igrtmitcally Zero tilla hio eerl conserves predators ofthe GIH (37) advanttgeous for the rice ciop to tap this trlLItlaI fertility nt it mcttoi lo lessen the impact o0 soil insects and grow vigorotsly to compete with wrus and to tolerate brotdcasting seed or planting in ftltrrows is prefc ired over pest infestation dihlirig in hills Crickets are large enotlgh to eat seedlings in

Larly plantngs terd to escape seedling maggolts Upland a hill atrd tillering normnally ca iriot fill in tlhe missing space rice in Bitangas is planted in Ma and ine itli the Iirst Oftcn dibbll rd is not well covered and iseasily found by rains Fairners are highly mtotiated to plant early tocscape foraging insets ioIdents and birds drlitght and typlotns ricar harct Arn Atrgtst planting Plan uit sii Incrcatsini sced density protects tile cropwould be evcely attacked by It riim orivzo At tti againrst seed and seelling pests especially ants particularly other Philippiie tipland sites Rea l Qteot antd (laelit if the crop will germinate and ermerge quickly M isainis Oriental seedlig mnaggots arnrige ric boe t se hr trftinll itI India rice intcrclopped witi cotton or farnes have a longer growing season and plnit 2 rin or pigeon pci had lower 6IIt and WBIlII poptrlations than mote after the ortset of the rainy season liaaiangas ftrmners rice alone (211) NfailC arid Urplald ir_ 2re coilltori in the plant ca rly becia use ofa shorter raiinfall period arnd the desi re Philippines he chatngC in pest status folii iniercropping is to lliarvet before the typhoor sea son peaks ltighly locatioll-specific (133) cmianarid the net elfc be either

Early plantings cornbind with early-iatiuring varictics nil bcnelficial ortdetriential (189) For example BIatangias provide higher crp tolerartce for white grubs Aestiatior farmers intercrip Itro in their ipland ricefields A liornshyterminates with the first rains but 5-6 moare ineeded for the nrii iotd-Ahis i (I iiiacius) which norlially feeds resting larvae to develop into beetles emerge and lay eggs on taro ill feed or miarbv rice plants Intercropping and for larvae to reach the danriigii g ird-instar stage probably would not afft0 seed pests A crluanion crop (134 In lataigas this normually iseiough tiite for harvest planted ith rice and haing gleater tolerarnce for root

It Brazil howecver delayed planting is a suggested dariage or being toxic nayact isi trapcrop to root-feeditig cultural control for the lesser cornstalk borer and insect pests Acial-fei i ig pests voultId be most affected by pests favored l drought (66 195) Cropping internsity in intrer ropping Shoot aphids leaf beetles sten borers and Brail is low and delayed plraing places the rice crop in the hopplers would be likely ca ntidates to note effcts On the niost stable rainfall period reducing likelihood old rought othlt halnd the corlipalior crop stch~ sitniaie Iiiv be more

In rilniy arca s rinmirsare roi atl to plant other crops benehited than rice from initecropping ( 137) before rice and althNgh it woultl be desirabh to avoid 11echi Weeuling luring the first month after clop infestations by planting rice cuatl lariners lack the labor cstablishent will force pcts such is anyvworris which and time to do so ol a low priority crp prefer grassy cetds ontor tle vorling rice Soil iampsecs will

77age Uplild rice Stils ae light textured and Call be mvnye to the rice crop Cleani citrltirc also lay lore territes plowed duiring the d rv seasor notl only to grin time for to attack I u cetincti inrice ceds i ay he le optinal planting with lie fitst Inilsoon riiains but also to desiccate solition 13) soil insects arid weeds such is ntsedge that foim rhizomes Irtlizr Itneatsing N Iciiliier oin hlwland rice has Htowever becta use most soil insects lie below die plow layer fAtwvored Irigher populains of pliithoppeiv Imlhoppers

26 IRPS No 123 January 1987

and leaffolde rs However because of the need to temper fertilizer use on upland rice o prevent blast fIrtilizer rates will be lower than is comonia r for lowlad rice causing relatixely less hopper and IaIfTfoide r population buildup However inupland rice areas without a history of blast such Is Batangas farmers apply high rates of N and conscqtrently the crop has high infcstation levels of WBPI]and RIF The fhrmrcrs leec insect buildup by splitting the fertili11r in0to two to three applicatiois bit do not applV fertilizer at the vegetati estage because this prodrces a tall crop which illeasily lodge during typhoons Greater crop fertility leads to greater insect survival (better nutrition) larger insects (to cat no re foliage) arid greater fec titd t (to lav nmre eggs) (128 145 171) Not all insect species respondequally hovr Increasing rates (if N l anrid K increased Dialraa s-acchralis and (iaeiurema flea beetle but reduced thrips (66) Zn rcdLuced deadheart density from the lesser cornIstalk borer

Fnldingi Although most Upland rice areas do nIot hawxcaccess to wa ter floodiig the fields is rRrrnriierdcd in areas of Brazil or root bug and white gib control ( 195)

Crop rotatioti Tillage before planting a crop after rc harvest will control stern boreis and other pests renain n the stubble arid will unearth soil pests Planting a non-gratriiaceous crop after rice is recommncrded for termite control (66) Crop rotation is less nttdcd in upland rice culture than in lowlands because atcrop-free period is assured even after harvest of crops following rice

t1ulchilt iryl plastic irli irig designed for weedcoitrol ii upland rice inJapan resulted in ntr highanri liy infestation of (ihilo sulytresvais (93 9-1 95) NIulched Upland rice grew more luxuriantlv and attracted more (Avipositing mothis than u lched rice Also the rice plant and stein borers imatoured 10rC quickly and a partialgeneratiori af vouing ista iNoverwintcred in the stubbleLotili--malitrt varieti Quickly maturing crops reduce the tmbCr of pCst generations that Calt build up Rapid crop establishment olan ca rl v-natu ring vaitvty if)pa 11icLa r will lessen white gib and stem borer damiage White grub larvae intle last iistar are highly destIruCtive so aii early-iiatri ng variety will escape serious root loss Stem borers build 1ip slowlyx in rice to ecormic abundant in a late-Maturing crop anrd caiuse whi tlieads Numibers of somespecies increase exponeitially with each succeeding genera-ti so a xarietyexhat atures I trowh earier thian another will hac less insect damage Early-naturing vaiicties however will not lessen seed pest daiage either ii the soil or oina standing crop

Strwhoi hAIJttam Inrscts tcd irig on thlie aerial portions of rice plats disperse from carly to late plantedfields AS iiiibCirs increase exponentially with eachinsect

gtscnrattm e eiglbrig f rmc~s1(uil upp wh rc generation pests build tIP where neighboring fihrTC e9r1sstagger their plantings II Batangas farmers plant uplandrice within I iro use 120-d s-arieties and plant at the start of the rainy season Ini Misarnis Oriental laricrs stagger their

plantings up to 3 mo apart and use a set of atieties that mature in 4-7 mo Insect pest danage particularly from stem borers is greater in Misarmis Oriental Also seedling maggots damage rice there because rainfall favors frequent planting of maic In Batangas rice escapes seedling maggots bCca usC of distinct drs season and synchron0us early planting with the onset of ains

Therefore combinations of cultural practices ea rlyplanting synchronous planing crop rotation and earlyshymaturing vaneties protect the rice crop against most insect pests

Ilmant resistance Beca tse upland rice has few specialized insect pests and many insect types attack the crop worldwide regional breeding and strong internatiornal cooperation are required

Sown-sced pests arc economically controlled by insectishycide seed treatment and should the relore receive lowxpriori ty inbreeding objectives

Root pests are normally cost ly to control with insecticides but finding resista t sonres mayvbe di fficult Tanaka (229) for exa -ple fi led to find varieties resistant to root aphids Selecting for large root bioniass perhaps should be the strategy for pests sLuch as Whlite grubs that renliove roots Fainter(174) howcvcr cites varietiesof crops tolerant ofor resistant t(i root pests Some sorghun varieties are high in cyanide so rice sorghun might incorporate broadshyspcctruri chemical resistance

Seedling vigor and drought tolerance are high prioritybreeding objectives so tupland rices should have large r root systernsand be able to tolerate ligherlevels of root loss Sonie root loss inolder plants is beneficial if it stimulates new root developnrut and therefore crihances intake of soil nutrients (53) The rice seedling inaggot rtiLotia spp may be

controlled thirough resistant aittics A hi rgc breeding effort on sorghniu has had fair success inideveloping arreties esrstait to Asoccata Rodani (251) A field trial comparing 10 upland ricc varieties showed differences aniong varieties va ring from II to 4(- damaged tilleis (215) Hovver resistarce to 1Iill-0r gtia species may rot cross ox-rti otier

Shiraki (210) reported that upland i-ice stens wcrc harder than lows-land rice stemsperhaps becaIusc (f a higher silicoricontent and thus were more resistant to yellow stern borer hc tortia Iow tillering (f uplaid iice xarneties prodtres a

higher percentage of infested tillers than would occor in higher tillering lowland iice under a siiilar egg density The corollary therefore is to select for high tilleriig upland rices

Stemii bor amcstheh main targes dobrecders inAfrica (571 204 )but nticl oifth e sc reen irig is dotie unde r lo wland co d t osrm s wih A h r~ o l pe i s n acorditions (11 96 218) As with lthrinrnwspecies mayhave to be dealt with independently however initial results show cross resistance between African and Asian species(92)

In Brazil screening Ior insect resistance focttses oil lou I pest problems Varieties tolerant of lasmolwalpus ligi-sellus and resistant to )iatraea saccharalis have been identified (70 153) Current work also emphasies 7iraca limativetntris and Thmialspis (=Iosis) flavouicta (66) These insect pests Ilt onlV citise IIigh yield losses tihey also ire difficult to control chemically ofr by other ilealls

ManV upland rice vaieties aic early maturing and are resistant to pests such as gall midge (I Xl ) Early mIturity is often a highly desirable trait to esuilpe pest buildup but it is not true resista nce bca usc ilthose va reties are plinted late they will be itsdamaged as ain early planted susc ible variety

P~erhaps tile cosiiiopolitall of all upland rice pestsimost are the sced pests (ripenling stagc) which pose a particular problem beta use scuds re ised IsIO for hitnit4ans Ayltoxic substatice introd tcld geticticllyilmay also be toxic to nal iniless it caln he placed old in thfe IdhuffSelectmi fo hairiness or lotng wns kill mIlct sthilrohleiiis dutilig threshing pa rtictfarl maitial threshinfl [hireshers ill complain of itcfimin atid itteascd (fust levels

Attempts itIRRI to fitid tcistatncc to the rice bug plorisa have ilotbeCnl prmitisilg lIIetIattonlids iorc

through the lenilna or palca hillhairs sedstVIor seeds wilh awns that deter bugs promote complaints from threshers Seed bugs liptocorisa and Stcnocoris enter the seed through the opening between tile lemna and palC but varieties with narrow openings are diifficult to mill

Breeding priorities should concentrate ott inotCI gruLtLIps that are most difficult to c nttrol by other meains Steil) horers appear to be the most widespread group that sigiiificantly dalag s rice aind is hard to manage by other Ilnca us

Biological control Very fe stutdic of mitatt~l eneties of rice pests in upland envirollnletits wee in tile wefouiid literature Therefore have not compiled a list of species Ve present a list of natural enemies iOtid in upland ricefields tuic latialln Batangas for some conitno iOldiar pest species ([itble 5)

Muich ofl thc lollowinle itfor ittiol cotlliCs rol Stutdies in the lowlatnds or roi crops otithcr thlin tice

Qu(ailtr o 11ll t11t( Nat trrtf lii hcoeits ight Ile used agaitist ile social insects tilt and termites Sollie predator specice live ill theirf nests n ilnic theirnllt ip-pearanIlCe aMRidbehavior IJtrouMdCing a mote aggressise ant species to displace pest ant codonies is a possibility (169) Patihogens as potential natural encnies have reccived little atteition (I13) Natural ncnuics asa group however hac not been toIl iporta it in Iegulating aintshown be and ternite numbers

The nocturnal behavior aid subteitmien habitat of lole crickets and field crickets protect them from many ntllral enemies There are only limited records of natural cnelics for these groups of pests some pathogens (87) but mainly

IRPS No 123 JattuIny I 271987

parasites - scelionids on eggsand sphecids on ilymphs and adults Sphecid wasps Larra carutmaria (SImitI hI L huzwuensis Rohwer and lsaltiuhta Willi ams speciali in mole crickets pa ralyzing them nd dragging the ninto thcir nests as food lot their voting ()tier sphCcid species Lins aurulenoa (Iabricius) lot maiac (Ashmead ) 1 lhshylssellaus (Smith) and Ml lal)oriosu (Snithf specialie in field clickets

White grUbsO the other hand hae Icti scriuns enuigh to have been extensively studied for cotttroi 1W nati al enemies( 112 140 141) Scoliid parasites C(ajsoMcispp have beetn intrtduced itto tilePhilippites for control of Ielxl ]udisiro la The rcsearchcts w Intttrdced themle during ill era claimed that the white gttionthreak as controlled (142) howevcr it isnormal for pest epidemics to subside and there was no diect evidence to indictte that tfe parasite was responsible A tmore promisitg titctllod might he to identify iso ltcttass prt~f tcartd apply hiactelaln fingal pathogens to tile soil Ihe lalattese beetle in North America has hcci coitrolled hv tite conmnleciall aailable hactcrial prcpa ration itacihlpu pav ofr milksdiscas (71) Spote prtyefott the soil itloted ilrtier reitiain actic fr Veas

Root aphid sardtnctly bugs readilslic tilib to predatots and parlsites once their tetndine ants are controlled The larvac and adtilts of a coccinellid beetle 1Scynms sp prey on llriadlouais in slash and burn upland rice Baits treated with insecticide may be used to control tending ants by killing their ourig (244)

Root-tkcditig blugs false wireoris wireorms and root wccils aic r ofsuch little ifilpoltatice that thenomttll role of liatural Cticniics hias tiot beei asscssed A possibility hoevrl tiglt he tit ItaIIitic niaCIMtodcs lintt1g11en Itese soil-itlhlfbitine mCtitodcs W5uhtfhave to be imiass Produiced

large miltibcr of niatltrll cnemics ircknown ft CltIoriS IC IpblCeii is thtt clttvortnlls ofonlie the crop soott alter land ptel)tritiOti a1tithe bcgit iniig of the raittv

Citsoln hfien tiatllral leniiiy populations aric lo Colltloil WOtuld iii ole mass prodtticing andirclcaslng key spccies

le egg stage is the fpirt (olthe Cedlitgt Ialot life cycle tI hich to f)ctis ftor tittlril elctit cottntrol hplraites o1 pleditors ttsit elleggs highly expoIsYCl I Sgg pailsites or predators stolld probahly e ittass andtbe produced released Littlh i toruialioll is a ailable ott what species to trV o ho to Iniss podLice thiCIi A etiloplhid wasp Icmihlaol sp effctiicly regushy

lates the leatf lcloalapolm2a spioanoiiitr heat andc rice antid should be collsctvcd

leaf beetles whCllose e am] eggs ate laid in the grutindhirO ould be motC difficutlt to control by niatulral enelV Mnll ipuhltiOn But species sUici IS c_gsthispid beetles ith

and larVle oilleascs ate vtilcratble to iparaites and pledators

Natural enemisc of tlirips otl rice are little known

28 IRPS No 123 January 1987

Table S Parasites recorded from upland rice insect pests in Tanauan Batangas Philippines 1977-84

Order Family Species Host stage

I lymttenoptera Braconidae Leaffolders Cnaphalocrocis Marasinia spy

Cardiochilesphilipplinensis Ashmead a Larva

Cotesia angustillasis(Gahan)Cotesia or cypris (Nixon)

Larva Larva

Cotesia or taeniaticornis(Nixon) Larva Cilonusinotakatae Munakata Chelonits SppAlacrocetrts philippinensis (Ashtmead)

Egg to larva Egg to larva Larva

Iclineumonidac Orgilts spp Isclutojppa hireator(Fabricius)

Larva Larva

Itoplectis naraigav(Ashmcad) Larva Tenelhtcha philippiensis (Ashmead) Larva Tenelchha stangli (Ashmead) a

irichomma cnaphalocrosis Uchida Larva Larva

Bethylidac Elasmidac

Goniozms or trianguli r Kieffer lasmtusalhopictus Crawford

Larva Larva

Diptera

Encyrtidac Chalcididac

Tachinidac

Elasmtus spp Copidosomopsisnacohiae (Fidy) Brach nteria evcarinata Gahan Biachyieria lasus (Walker) Ztgohothria ciliata (Wulp)

Larva Egg to larva Pupa Pupa Larva

Argyrophylx nigrotihialis(Blaranov) Larva

l-ymenoptera Braconidac Yellow stern borer Scirpopiaga incertulas

Braco chinensisSzeplegeti Larva CIthelotis spp Chelonus niunakatae Munakata Cotesi flaripes Cameron Stenoracon nicerilli (Bingham) a

Larva Egg to larva Larva Larva

Ichneumonidac Tropobracon schoenobii (Viereck) Amatroinorpiaaccepta metathoracica(Ashmiead)Frihorus sinicus (I lolmgren)

Larva Larva Larva

Ischnojoppaluteator (Fabricius) Larva Isotima nr dammeratiti Rohwer Larva Tenelucha philippinensis (Ashmead) a

Larva Tenehicha stangli (Ashmcad)Trichomma enaphalocrosis Uchida

Larva Lara

Chalcididac Eulophidac Trichogranmmatidae

Xanthopimnpla stemmator (Thunberg) Charops hrachYpterumn Gupta and Malteswary Brachtm eria sppTetrasticttus schoenohii Ferriere Trichorigranm chihunis Ishi

Larva to pupa Larva Pupa Egg Egg

Diptera

Scelionidac Pteromalidae Tachinidae

Tt ichograintnta aponiicuitt (Ashniead) Telenomits rowani (Gahan) a

Trichoinalopsis apattelctena (Crawford) Jeirhaea spp

Egg Fgg Egg pupa Larva

Z4gobothria ciliata (Wulp) Larva

Ilymenoptera lecneumonirlac Gol fringed borer (hilo auricilius

Trichomm enoptalocrosisUchida Larva

Ieioenoptera Braconidae

Aantholimlh steiniator (Thtinberg) a

Pink stet Ioret Sesamia inferens StoInacoil nicrilhi (Bingh am)

Loarva to pupa

larva

Diptera Ilyinenoptera

Armvtnyorms Mythimna separata Spodoptera mauritia acronyetoidesTachinidae Zvgohothriaatropiora( Robinea i -Desvoid y)Eulophidae It-phctrtiscitapadoe (Ashimead)

Larva Larva

(halcididae Braconidac

Brachpin ria spip (oesia spp

Larva larva

IHyntenoptera I)iptera

Rice skipper Ielop ida (halcididae Tachinidae

math ias aidgreen iortedci ciliarNIelan itis eda Brat lhincria sp iii matuqitatt Cameron Irgyrolhirlaxi trotlhiais(marumtv)a1

ismtne Larva Larva

Diptera Ilymenoptera

Tachnidae lelineutionidae

fuown se tilo per Mois frugalisIrgtrophrlax 1igrothiahiv 3 a rat ov) anthopimpa pmnctata (Iabricius)

Larva Larva to pupa

aMost dominant

IRPS No 123 January 1987 29

Normally predatory thrips prey on herbivorous thrips strepsiptera on both Spiders dominate tilenymphaladultNatural enemies of stem bugs --Scolinoplara and Tibraa predators Spittle bugs move to rice from pasture grasses - are in order of importance egg par~isites and predators and biocontrol would be successful only ifcarried out before fungal pathogens and nymphal adult parasites Egg tiley immigrate into ricefields parasites could either be introduced fron other areas or Biological control tactics for hemipterous seed bugs mass produced and released ould follow those outlined for stein bugs

Armyworms are normally lwld in hcck by the activities Coml)arison of environnewtns Natural enemies are of egg and larval parasites When these natural enemies fail perhaps faced with even greaterconstraints than insect pestsusually because of drought armyworns become epidemic to survive ind reproduce in upland environments TheyIt may be worthwhile therefore to release parasites during have adapted ani survived by having wide host ranges the the rainy months following a drought Virus diseases of ability to aestivate or hibernate over unfavorable seasonslarvae would beanotheravnue to exploreforarnyworms and the ability to disperse or by other mechanisms

Parasites have ben traditionallyconsidered for biolegical Comparing the most dominant and therefore the most control attempts against stein borers usually by rekasing adapted species of natural enemies collected in various exotic species( 166) This approach has not met with success environments in the Philippines we can understand how and new avenues should be explored The role of predators different rice environments are arid pathogens is little inderstood and deserves greater Anagqrms olptahiisa mymarid planthopper egg parasiteattention Egg parasitism rates are normally reasonably is cquallyadapted to all Philippine rice environments (Tablehigh but should be supplemented with effective egg 6) It is selective for planthoppers and even parasitizes eggspredators Orthopterans are potential egg predators of the n-iize planthopper Peregrinitsmaidis itt upland areas Metioche vittaticollis (Sta ) Anavipha nifiwnnis (Saus- Alavohis also parasitizes maize planthopper eggs but is sure) and Conocephahs hongipnni (de i-laan) feed on most prevalent on BPH in the irrigated wetlands (onashystem borer eggs etioche and Anaxil)ha specialize in eggs tocerus spp like A optalhilis on BPH and WBPI Iattacks with no hair such as Chilo spp The omnivorous (ono- the eggs of GLH in ell rie environments It is also found cephahsspecializes in eggs cov red with hairs it is a proven parasitizing the eggs ef the white rice licahopper (oimna egg predator of Scirlouphaga spp but has the discouraging spetra Oirosita naias a trichogrammatid egg parasitehabit of eating rice guin (65) attacks BIPH and WBPH in the wetlands and has no other

Grasshoppers have egg (scelionid) and nymphalad ult host A related species 0 aesopi specializes in G1-H in (nemestrinid tachinid and sarcophagid) v irasites Among wetland environments Dryinid strepsipt-an and pipunshypathogens protozoans have been recorded most frequently culid nymphal and adult parasites of planthoppers and Control of locust species by natural enemies could con- leafhoppers also specialize either in hopper species or centrate on their habitual breeding areas environments Among the dryinids Pseuhdgonatoipsflavi-

Leaffolders have rich complexes of natural enemies some feimur prefers the lowlands and BPH while P nuch has no adapted to upland rice (22 27) ranging from egg predators environmental prefirences but is adapted only to WBPH (gryllids coccinellids) to larval parasites (braconids ichncu- The main d ryinid species on GLH aptogonatop)usspp is monids) to larval prcdators (ants and carabids) to larval most dominant in rainfed lowlands and also attacks other pathogeiis (vira and fungal diseases) Efforts to augment leafhoppers within Amnrasca Cicadulinaand Bachaltuathese natural enemies should focus on the egg predators Strepsiptera are more dominant on plarithoppers than (rearing releasing)and viral or fungal pathogens (culturing lealhoppers The Elenchus species in the Philippines attack spraying) all species of planthoppers within the genera Sogawilla

Parasitization of the large conspicuous larvae of Sogatodels NilaparsataHarualia and Opiconsiva Elenshyskippers green horned caterpillars and other polyphagous cusrvasuntaisuiis adapted to the major rice environments Lepidoptera by tachinids and chalcids is normally low ELg but prefers BPH in rainfed arid WBPIH in wetland environshyparasites and predators are perhaps the key to their control ments Ilalictoph(utsmtnroeiand HVpectrus attack G LH

Gall midges are parasitized by pterormalids and platy- and COfimravcctra respectivelygasterids that aestivate within their larval host between Pipunculids are mainly parasitic oii lealhoppers Each seasons Shoot aphids are preyed upon by coccinellids species has a unique environmental preference Tomosshysyrphids and chrysopids varireIa oraetoraprefers Ne7 otetltix nigropictrs and

Planthoppers arid leafhoppers have egg parasites and Pilunwulhtsjavanensisattacks Dehoceplhahls spp which predators as well as nymphialad nlt parasitesand predators are more abundant in upland than irrigated lowland their rich complex of natural enemies includes fungal environments T suhvirescensattacks Nephotettix virescens pathogens The principal egg parasites are myrnarids and and prefers the lowlands Pilpuiulus nttillattls is an trichogramniatids Egg predators are norinally mirid plant upland rice parasite arid attacks only Nephotetix speciesbugs that suck out the yolk Nymphal ad nIt parasites are More than 17 recorded species of leaffolder parasites in dryinids on planthoppers pipunculids on lealhoppers and the Philippines attack species of C(naplwocrocis and

30 IRPS No 123 January 1987

Table 6 Dominant natural enemies of rice pests in upland rainfed lowland and irrigated lowland rice environments of the Philippines

Natural enemy

Host stage Species

Brown planthop-Parasite Egg

MymaridaeAnagrus optabilis(Perkins) Anagrusflaveolus Waterhouse

TrichogrammatidacOligosita naiasGirault

NymphAdult Dryinidac

Pseudogotatopuslaiifemttr Esaki et Hashimoto Elenchidae

Elenchus yasurnatsui Kifune and Hirashima

Environment

Upand Rainfed lowland Irrigated lowland

Ailaparata lugens

x x X x

x

x x

x X X Whitebacked planthoppers Sogatella furcifera and Sogatodes pusanusEgg

MymaridaeAnagrus optabilis(Perkins)

TrichogrammatidacOl~gositanaias Girault

NymphAdult Dryinidae

Pseudogonatopusnudus Perkins Elenchidae

Elenchus yasutatsui Kifune et Hirashima

Parasite Egg

MymaridaeGonatocerus spp

TrichogrammatidaeOligositaaesopi Girault

NymphAdult Pipunculidac

Tomosvaryella subviresccns Loew Tomosvaryella oryzaetora (Koizumi) Pipunclusmutillatus (de Meijere)

DryinidaeIlaplogonatopusspp

Egg Copidosomopsisnacoleiae (Eady)

Larva BraLonidae

Cardiochiles philipphnensisAshnead lchneumonidac

Trichiomnia cnaphalocrosisUchida Temelucha stangli (Ashmead)

Pupa Chalcididae

BrachymeriaexcarinataGahan Brachymerialasus (Walker) Brachymeria nr tachardiaegroup

Parasite Egg

Scelionidae Telenomus rowani (Gahan)

EulophidacTettasticusschoenobhi Ferriere

Larva Braconidae

Stenobraconnicevillei (Bingham) Iclhncurnonidac

Temelucha phiiippinensis(Ashinead)Temelucha stangli (Ashmead)

x x x

x x

X x X

x x Green leafhopper Nephotettix virescens

X x x

x x

X X X X

X

X Rice leaffolders CnaphalocrocisMarasmia

x X x

X

X X

x x x x x X x x x

Yellow stem borer Scirpophaga incertulas

X X x

x

x

x X XAinauromorphaaccepta metathoracica(Ashmead) x

continued on next page

IRPS No 123 January 1987 31

Table 6 cntinued

Natural enemy Environment

Host stage Species Upland Rainfed lowland Irrigated lowland

Larva-pupa Gold fringed borer Chilo attriciiuslchneumonidae

Xanthopimpla stemmator (Thunberg) x aPupa x

Pteromalidae Trichoinalopsis apanteloctena (Crawford) ax

Pink stem borerb Sesamnia inferensParasite

Larva Braconidae

Stenobracon nicevillei (Bingham) x b b Armyworms Mvthinna SpodopteraLarva

Tachinidac Zygobothriaarroptvora (Robineau-Desvoidy) x x

Braconidac Cotesia spp

X Skipper Pelopidas mathias and green horned caterpillar Melanitis leda ismeneF-gg

TrichogramrnatidacTrichogrannaspp x

Scelionidae Undetermined Scelionid

x Larva

Tachinidae Argyroph lax nitrotibialis (Baranov) X x

Pteromalidae Trichomnalopsis apanteloctena (Crawford)

X Predator

CoenagrionidaeAgriocnenis femina femina Brauer x xIschnura senegalensis tambur

Miridae CrtorhinsliidipennisReuter x x

Nabidac Stcnonabis tagalicus (Stal) x

VeliidaeAlicrovelia douglasi atrolineata (Bergroth) x

Coccinellidae Micraspis crocea (Mulsant) X x x

Vespidae Eumenes campaniformis (Fabricius) x

Formicidac Diaecamna spp xOdontoponera transversa (F Smith) x

Carabidae Chaenius spp OphioneaW- aJasciataSchmidt-Goebel

x

Ophionea ishii ishii Habu x x X

Gryllidav Anaxipta longipennis (Saussur-) x xMetioche ittaticollis (Sial) x x x

Tettigoniidae Conocephalus longipennis (de Haan) x x x

aNo rearing made on rainfed lowland bNo rearing made on rainfed lowland or Irrigated lowland

MrasmiaAmong the Laval parasites the braconid Cardio- lowlands and parasitizes oriental maize borer OsIriniachilesphilijpphensis is most adapted to an upland environ- furnacalis (Guenee) larvae The leaffolder pupal parasitesment It also parasitizes Hydelepta indicata a common not only occur equally in all environments but also haveleaffolder of legumes The ichneumonids Trichomma cna- wvide host ranges- parasitizing species of OsIrinia Marucaphalocrosis and Temehcha stangli occur more in the Hydelepta and Honiona - in maize and legumeslowlands T stangli is most adapted to irrigated lowlands The principal egg parasite of yeltow stem borer (YSB)and also parasitizes Chilo and Scirpophaga stem borer Telenomus rowani is widely adapted to all environmentslarvae T cnaphalocrosis is most prevalent in rainfed and has an unusual alternate host -Tabanus eggs (25)

32 IRPS No 123 January 1987

Tetrastictu schoenobii specializes only in YSB and occurs mainly in irrigated lowlands -- further proof that YSB is most adapted to the wetter environments

Larval parasites of YSB are highly environment-specific The braconid Stenohracon nicevillei occurs only in upland rice but also attacks the pink stem borer (PSB) Tieinchwa phiiiplPwnsis an ichneumonid occurs in upland rice and also parasitizes the larvae of (7hiu stqpresaisinatro-morpha accepta melaihoracica like 7 stangi is mnost adapted to the irrigated lowlands

The dark-headed stem borer(l)HSB) (ih pochit3sus was not collected in all rice environments and the gold fringed borer Chilo auricilius nd PSB were collected only in upland rice Xaantiholiplastenrnaloran ichneumonid larval parasite appears to be widely adapted whereas TrihomaslopisapanitI)cteiiaa pteromalid pupal parasite occurs mostly in wetlands and also parasitizes larvae of Chilo suppressais and Pelhuidas mathias

Armyworms Myit hinin and Spodujipera have two main larval parasites - a tachinid Zrgohothriaatropivora in rainfed environments and a braconid Cotesia spp in irrigated lowlands

The butterfly pests Pelopidas and 4elanitishave tricho-grammatid egg parasites found only in irrigated lowland environments

Two larval parasites show environmental preferences --Argrroph av nigrotihialis a tachinid in rainfed habitats and Trichonialopsisapantelkcnela in irrigated ricefields A limgrotibialisalsoattacks the sweet potato hornwormAgrius convolvtdi and T apanteloctena parasitizes DH S1B

Predators as a group show more distinct environmental preferences The aquatic damselflies Coenagrionidae are most abundant in the lowlands Cvriorhinus is also most adapted to the lowlands The nabid Stenonabis tagalicus occurs only in the uplands and isalso prevalent on legumes The coccinellid lady beetles wasps and ants are more prevalent in upland areas AlicrasIpis cit-ea larvae prey ona variety of aphids attacking legumes and maize Etunienes campanonia vespid mud wasp makes nests in trees and is therefore most adapted to the more botanically diverse uplands Soil-dwelling ants cannot tolerate flooding

Upland environments are habitats to arboreal carabid beetles Three species of Chlaeniusbeetles prey on leaffolder larvae and are perhaps more important than parasites (27)

The gryllids and tettigoniids are widely adapted egg predators not only on rice but also on maize and legumes They feed on eggs of most insects that are laid on leaves

The spidercommunity of upland rice environments isrich in species At one site (Batangas Philippines) 31 species have been recorded (23) Of the 176 spider species recorded in Philippine ricefields about one-half (82) occur in upland rice The spiderspecies of upland rice environments overlap more with those of rainfed lowland than with those of irrigated lowland rice (24) The same study showed that of the three environments irrigated rice has (ie greatest spider species diversity foiiowed by rainfed lowland and upland

However of the 10 most prevalent ricefield spiders 9 were abundant in upland rice showing wide adaptation All three spider guilds -- orb-web space-web and huting spiders shywere prevalent in upland ricefields There were however differences in environmental prelerences for some spider species

Of the orb-wcavers Tetragniathamanlihulatawas partishycila rlv abundant 7 japonica was low and Ltucatge dehcorata was absent in upland rice

As in other environments Aiypeiwnaifirtosana was the most dominant space-web spider however its relative numbers were lowerthan for lowland sites Among hunting spiders Ox|opes javanus and Lcosa leatcost fgla were more abundant and Lvcosa lpseudoanmnlata was less abundant in upland compared with lowland rice envishyroniments

Extensive rearing of rice insect pests in upland rainfed lowland and irrigated sites over an 8-yr period in the Philippines provided insight into the effectiveness of parasites as natural enemies (Table 7 8) Most of toe data are from large sample sizes taken over at least one crop which overcomes the pitfalls expressed by Van Driesche (238)

VSB egg parasitization was surprisingly similar across all environments Larval parasitization was highly dynamic among sites and even years within the same site but showed somewhat lowerlevels in uplandsthan in irrigated wetlands This would have been predicted for egg parasites as well since YSB is adapted to the lowlands and so must be its parasites This isevidence that Telenomus rowani is as good at dispersing as its main hosts Trowani clings onto the anal tufts of female YSB and parasitizes eggs as they are laid This phoresy may explain its wide adaptation The more specialized lowland YSB larval parasites therefore appear more effective than their upland counterparts with more alternate hosts

Low collection levels of PSB larvae especially in the wetlands make comparisons between environments difficult Low levels of larval parasifization were recorded in both rainfed environments Except in the Cagayan rainfed wetland site in 1980 low levels ef larval parasitization were evident at irrigated and rainfed sitesalike No differences by environment were evident Higher levels of parasitization occurred in Sarawak during a 1967 outbreak also from tachinids Rothschild (197) indicated that higher parasishytization occurred in upland rice areas

Skipper and green horned caterpillar larval parasitization rates were generally low and showed no environmental effect htighest activity occurred in Cagayan in 1980 (48 parasitization) Although larval parasitization levels of RLF were variable year to year activity appeared greater in the irrigated lowlands than in the uplands Again the highest levels were recorded in Cagayan in 1980 Cagayan has extensive grasslands and fallow around the rice areas and a relatively long rainfall period Rainfall in 1930 was good for crop growth and apparently for increase of larval parasites

IRPS No 123 January 1987 33

Table 7 Parasitization rates of rice planthoppers and green leafhopper in upland rainfed lowland and irrigated lowland environments Philipshypines 1976-840

Environment and site Year Egg parasitization () Nymphaladult parasitization ()

BPH W13P GLH BPII WBPH GLIH

UplandBatangas

Rainfed lowland

1976 1979 1980 1982

15 17 12

2(47)8(39)

4(51)

9(1800)14 (212)

12(304)

8 (101)6 (127)

9(116)

Iloilo

Cagayan

1976 1977 1978 1981 1980 1981 1982

0 (69)

11

44 48 40

-

-

46 55 44

0 (471)

11

52 56 49

0 (200)9 (214)

10 (1259)8(235)6 (900) 3(800)

-7 (462)

11 (270)3 (313)5 (900) 1 (800)

4 (600)10 (725)10 (362)12(870)5 (900) 3 (800)

Pangasinan

Irrigatedlowland

1984 1976 1978 1979 1982

13 (436) 13(379) 14 (457)

6(14)7 (197)

17(198) 11 (486) 18 (275)

3 (81)10 (274)3 (116)

16(871) 19(88)

28(211)9 (764)5 (647)7 (1204) 8(305)

Laguna 1977 11 7 21 1978 31 1979 14 4 4 28 1980 7 8 20 1981 1983

56 7(278)

26 16 (1650)

16 27(1921)

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R I Apostol unpublished 1lo1lo and Pangasinan 136 Cagayan BCanapi unpublished Liuna 175 I C Pantua unpublished 118 234 43 Carino and Shepard unpublished BPH = brown planthopper Nila parvara htgens W131Pi =whitebacked planthopper Sogatelia furcifera GLH = green leafhopper Nephotettrix virescens Figiires in parentheses are sample sizes

Tauher et al (230) reported that internal external and genetic factors influence the seasonal activity of parasitoids

Hopper egg parasites but not nymphaladult parasites were also parlicularly abundant at the Cagayan site possibly the egg parasites have more alternative hopper prey in the weedy areas than nymphaladult paisites

The seasonal dynamics of hopper egg parasites can be seen for upland rice in Batangas(Fig I 2) and for rainfed lowland lice in Pangasinan( 135)and Cagayan( 105) Levels of BPHWBPI-L and GLH egg parasitization were corn-parable in upland and irrigated lowland environments Except for some high rates of paraMsitization early in the season the parasitization rate tended to be steady in all envilonments However hopper nymphaladult parasi-tizution levels were more similar in the rainfed environments and lower than in irrigated rice

Overall parasites do not appear highly effective by themselves against upland or even lowland pests Among natural enemies parasites have attracted the most attention because they can be more readily assesscd

Predators and pathogens lend themselves better to management practices Predators the most important group of natural enemies in riccfields are difficult to quantify On the other hand pathogens appear less im-portant but can be readily cultured and disseminated

The natural enemy commulnity of upland rice is rich in species and differs significantly from that of lowland rice

These beneficial organisms must be conserved by applying insecticides judiciously particularly sprays Seed or soil placement of chemicals minimizes exposure to natural enemies The strategy to derive the greatest benefit from natural enemies isto allow the greatest number of beneficial species to thrive Then perhaps at least one species will be effective against each pest at any tine overcoming the variable abundance of each species within and between years

Programs to introduce exotic species or mass produce indigenous ones are ambitious and expensive because they require trained people on a sustained not ad hoc basis

Chemical control Insecticides offer rapid and efficient control of upland insect pests (190) but-should be used only after other control measures have been considered Insecticides are rarely used on upland rice because of cost (32) Upland rice yields are normally too low to justify the expenditure Also most upland rice is grown as a cash crop Spraying upland rice is more difficult than lowland rice because water is less accessible Government subsidy programs to provide pestishycides to tribal peoples for upland rice production have created problems of toxicity to humans because isolated people have had no experience to be able to handle pesticides safely

However ins-eticide use has beenjustified when a crop is

0Z

Table 8 Parasitization rates of lepidopterous pests in upland rainfed lowland and irrigated lowland environments Philippines 1976-84a

Parasitization (5)

Environment and site Year YSB GFB PSB Earcutting Swarming Skipper Greenborned RLF Brown larva larva caterpillar caterpillar caterpillar caterpillar larva semilooperEgg Larva larva larva larva larva larva

UplandBatangas 1977 - 0 (57) - 0 (63) - 0 (71) - 0 (106) 11(142) 2 (101)1978 43 (4697) 8(451) 5 (38) 3 (40) 6 (61) 2(58) 15 (57) 13 (129) 13 (1286) 4 (30)1979 59(4135) 12(320) 13 (11) 5(28) 9(45) 6(46) 7(23) 8(160) 9(1152) 8(26)

Rainfed lowland Iloilo 1976 - 0 (10) 0(10) 0 (7) 0 (19) 2 13) 15 (180) 5 (76)1978 0 (54) 0 (3) 0 (17) 9 (64) 4 (78) 3 (73) 0 (121) 0 (191)1979 64 (2350) 16 (84) 6 (17) 0(6) 18 (68) 25 (104)

1981 56 (1322) 18(304) 0(5) 12(51) 18(138)Pangasinan 1976 60 (7 7 )b 62(82) 0(111) 5 (39)

1978 58 (53)b 5(51) Cagayan 1979 59(713) 8 (101) 6 (34) 0(7)

1980 0(11) 4 (28) 15 (286)

62 ( 48(177) 61(140)1981 0(5) 12 (2) 13(45) 14 (92)

Irrigatedlowland Laguna 1974 62 8

1974-75 50 41 1975 43 72 1978 96 (623) 21(24) 33 (60)1979 83(2017) 18(110) 8(62) 11 (89) 22(400)1980 78 (1625) 20(75) 0 (16) 4 (72)1983 60

9 (80) 7 (270) 30

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R F Apostol unpublished Iloilo and Pangasnan 136 Cagayan B Canapi unpublished Laguna 175 P CPantua unpublished 118 234 143 Carino and Shepard unpublished YSB = yellow stem borer ScirpophagaincertulasGFB = gold fringed borer Chilo auricilius PSB = pink stem borerSesamia inferens earcutting caterpillar Alythimna separata swarming caterpillar Spcdoptera mauritia acronyctoides skipper Pelopidas mathias reen horned caterpillar Melanitls ledaismene RLF = rice leaffoldersCnaphalocrocismedinalis and Marasmiaspp brown semilooper Mocisfrugalis Figures in parentheses are sample sizes Egg masses

IRPS No 123 January 1987 35

Table 9 Effect of seeding rate and insecticide on plant density and Yield losses from insect pests in upland rice often areyield of Dagge upland rice Batangas Philippines 1976a comparable to those of lowland rainfed or irrigated rice

Seeding rate Insecticide Dosage Plant density Yield (0-Y30o) but usually from different pests The principal (kg aiha) seed treatment (kg aiha) (noIra-row) (tha) groups are soil pests feeding on sown seed and rootsat 14 DE followed by seedling pests These groups have largely gone

50 None - 35 a 20 a unnoticed leading researchers to conclude that stem borers50 Carbofuran F 10 38 ab 23 a and seed bugs are the most important pests (17) Yield loss 10 beCabofuranF100 None - 4551 c 23 a trials that pinpoint major pests need to includc seed

100 Dieldrin WP 10 52 cd 22 a treatments Granules applied in the furrow may not give150 None - 59 d 22 a adequate protection from many seed pests

aAv of 8 fields DE = days after crop emergence In a column means The floristic and pedological diversity of upland rice followed by a common letter are not significantly different at the environments is matched by the diversity of arthropods5level by D)MRT F = flowable Wi = wettable powder both pests and their natural enemies Irrigation has homoshy

genized lowland rice environments Flooded soils tend tothreatened by epidemics of locusts or armyworms New have the same properties and to eliminate soil pests The technology stressing efficiency has minimized insecticide spatial and temporal dominance of cultivated rice has cost benefited specialist pests

Seed treatment is normally inexpensive and can be Upland rice has no specialist arthropod species The economicallyjustified in -nanycases(66) Treating seed with monophagous pests on upland rice prefer the lowlands systemic insecticide (216) but not chlorinated hydrocarbons where rice evolved Rice in the uplands exists only because (247) is most effective in controlling seedling maggots of man and even though lowland rice predates upland riceFoliar sprays have to be repeated to obtain control and specialist species evolved Wheat maize and sorghum do often are washed off by frequent early season rains not have monophagous pests leading us to conclude that Granular insecticides are effective but required dosages are there is no evolutionary advantage for inssects to have too expensive to justify except under heavy infestation narrow host ranges in upland graminaceae(247) When directed against first-instar larvae banding Pests have adapted to the uplands not only by having granules in seed furrows (177) can be effective against white wider plant host ranges but also by having long life cyclesgrubs at low dosages (025 kg aii ha) (134) Seed treatments undergoing dormancy or being dispersive Each pest has a are not promising for white grub control (248) unique life history strategy suited to highly unpredictable

Seed treatment is also effective ainst ants and may be environments In agivenyear someinsect specics an better cheaper than increasing the seeding rate In Batangas exploit upland rice as a food source but no pest is a senious treating 50 kg seed ha was as effective as increasing the one every year on upland rice seeding rate to 100 kgha and using untreated seeds (Table The low yielding potential of upland rice means less 9) In Brazil seed treatments control termites and other soil quantity and lower quality of food for pests to exploit but insects including the lessercornstalk borer(66) Domiciano also less incentive for farmers to undertake control (58) however found seed treatments inconsistent in their The most effective control would be early and synchroshyeffect against high populations of lesser corn stalk borer nous planting of early-maturing upland rice varieties to

Baiting can be an inexpensive way of controlling ants prevent insect pest buildup Breeding for insect resistance seedling maggot flies mole crickets and field crickets Baits should initially focus on stem borers because they are verymade of locally available material and impregnated with difficult to control by other means Biocontrol efforts insecticide can be sparingly distributed taking adantage of should first stress conservation of natural enemies preshythe pests ability to disperse and encounter bait sites dators and pathogens rather than parasites Efficient

The new low-volume hand-held controlled-droplet chemical control includes seed treatment or baiting for seed sprayers may offer an advantage over high-volume knap- seedling and some root pests and using controlled dropletsack sprayes for controlling foliar pests Upland rice sprayers for spot treatments against foliar pests and seed canopies are more open than those in lowland rice and bugs when economic thresholds are reached better droplet penetration should result Controlled droplet applicators would be ideal for seedbugs and defoliators

CONCLUSION

Upland rice is attacked by a wider array of insect pests than is lowland rice mainly beca use of the addition of soil pests Generally population levels of lowland rice pests are lower in upland than lowland rice leading many people to conclude that insect pests are not important on upland rice

36 IRPS No 123 January 1987

REFERENCES CITED

1Abraham C C and K S R Mony 1978 Occurrence ofLeptocorisa acuta Fabr (Coreidae Hemiptera) as a pest of nutmeg trees J Bombay Nat ttistSoc 74553

2 Agyen-Sampong M 1975 HlierogrlphusdiaganensisKrauss (Orthop-tera Acrididae) a new pest of rice in Northern Ghana Ghana J Agric Sci 8249-253

3 Agyen-Samnpong M 1976 Varietal resistance and other inle-ratcd pest control components as factors of rice varietal improvement in West Africa Prcsented at the Second varietal improvement serinar 14-18 Sep 1976 Bouak Ivory Coast West Africa Rice Development Association Monrovia Liberia 18 p

4 Agyen-Sampong M 1977 Outbreak of Ihccrorrdnisorae in rice seedlings near Rokupr Sierra Leone lit Rice Res Newsl 2(6)16-17

5 Agyen-Sampong M 1980 Dirty panicles and rice yield reduction caused by hugs IntRice Res Newsl 5(l)11-12

6 Agyen-Sampong M 1982 The major pest problems of irrigated upland and mangrove swanip rice ecosystems ir the humid tropical Guinea savanna and Salel climate zones Pages 318-328 in Pro-ceedings training course cn concepts techniques and application of irtegrated pest management in rice inWest Africa 10-20 Jan 1982 Fendall Liberia West Africa Rice l)evelopioent Association Monrovia Liberia 505 p

7 Akibo-Betts V I and S A Raymundo 1978 Aphids as rice pests in Sierra Leone lit Rice Res Newsl 3(6)15-16

8 Akinsola F A 1975 Present status of different rice stemborers in parts of Nigeria iicL Entomo Newsl 328

9 Akinsola FA 1979 Ihe biology and ecology of rice stemborers in Nigeria Ph 1) thesis University ol lbadan Nigeria 217 p

10 Akinsola E A 1979 Guidelines of integrated management of insect pests of upland rice in West Africa Presented at the Seminar on integrated management of rice diseases and insect pests at Bobo-Dioulasso Upper Volta 17-22 Sep 1979 West Africa Rice Develop-ment Association Monrovia Liberia

II Akinsola IA 1984 Insect pests of upland rice in Africa Pages 301-305 in An overview of upland rice research Proceedings of the 1982 lIouak Iwsrv Coast upland rice workshop International Rice Research Institute 13O Box 933 Manila Philippines

12 Alam M S V TJohn and Kaung Zan 1985 Insect pests and diseases of rice in Africa Pages 67-82 in Rice improvement in Eastern Central and Southern Africa proceedings of the international rice workshop at Iusaka Zambia 9-19 April 1984

13 Alticri M A A van Schoonhovcn and J D Doll 1977 The ecological role of weeds in insect pest management systems a review illustiated with bean (Phaseohsvulgaris L)cropping systems PANS (Pest Artie News Summ) 23195-205

14 Anmante F 1967 The leaf-cutting ant Atia capiguaraa pest of pastures [in Portuguese] Biol6gico (Sao Paulo) 33113-120 (Rev Appl Entomol A56207)

15Andrade A C and D Puzzi 1951 Preliminary results of experi-merits on the control of Sculptcoris iastantus on sugarcane [in Portuguese] Bioulgico (Sao Paulo) 1744-49 (Rev Appl Entomol A41421422)

16 Appert J 1970 Maliarplia separaelha (rice stem borer) new observations of entomological problems of rice in Madagascar [in French] Agron Trop (Paris) 25329-334

17 Ariadeau Mand Z Harahap 1986 Relevant upland rice breeding objectives Pages 189-197 in Progress in upland rice research Proceedings of the 1985 Jakarta conference International Rice Research Institute 1O Box 933 Manila Philippines

18Asar S and Kisimoto 1975 Seasonal occurrence of the smaller brown planthopper in upland rice [in Japanese] Proc Kanto-Tosan Plant Prot Soc 2228

19 Bakker W 1974 Characterization and ecological aspects of rice yellow mottle virus in Kenya Meded Landbouwhogesh Wage-ningen 152 p

20 Balut F F 1970 White grubs attacking rice Oryza ativa L [in Portuguese] Biol6gico (So Paulo) 36321-322

21 Banerjee S N and L M Pramanik 1967 The lepidopterous stalk borers of rice and their life cycles in the tropics pages 103-124 in The major insect pests ofthe rice plant Proceedings ofa symposium at the International Rice Research Institute September 1964 The Johns Hopkins Press Baltimore Maryland USA

22 Barrion A T and J A Litsinger 1980 Ants a natural enemy of cliaiarac in dryland rice IntRice Res Newsl

5(4)22-23 23 Barrion A T and J A litsinger 1981 The spider fauna of

Philippine rice agroecosystem 1Dryland Philipp Entomnol 5139-166

24 Barrion A T and 1 A Iitsinger 1984 The spider fauna of Philippine rice agroccosystcms II Welland Philipp Eiitomol 611-37

25 Barrion A T1 and J A Litsinger 1984 lihanus (Diptera Tabanidae) eggs an alternative host of rice stei borer (SI3) egg parasite Telenoonis digns (Iliymenoptcra Scelionidae) lit Rice Res Newvsl 9(6)19

26 Barrion A T and J A Litsinger 1985 Identilication of rice leaffolders by wing markings Int Rice Res Newsl 10( 1)24

27 Barrion A T and J A Litsinger 1985 Chctiosspp (Coleoptera Carabidae) a leaffolder ILF)predator lit Rice Res Newsl 10(1)21

28 Bertels A 1970 Insect pests of rice and theircontrol [in Portuguese] Pelotas Brazil Instituto de Expcrimentacao Agropccuararias do Sul Circular 43 24 p

29 Bonzi S M 1982 Ctilo (flisilineus J de Joannis (Lepidoptera Pyralidae) stem borer of irrigated cereals in Upper Volta LioFrench] Agron Trop Riu Rizic Cult Vivrieres Trop 37207-209

30 Bourke T V T L Fenner J N LStihick G Baker EHassan D F OSullivan and C S li 1977 Insect pest survey for the year ending 30th June 1969 Papta New Guinea )epartment of Agrishyculture Stock and Fisheries Port Moresby 57 p

31 Bowden 1 1973 Migration of pests in the tropics Meded Rijksfac Landbouwwet Gent 38785-796

32 Breniere J 1973 The main rice parasites in West Africa and their control Pages 27-42 in Proceedings seminar on plant protection for the rice crop West Africa Rice Development Association Monrovia Liberia

33 Breniere J 1976 The principal insect pests of rice in West Africa and their control West Africa Rice Development Association Monrovia Liberia 52 p

34 Brown ES 1962 The African arinyworn Spoiolptera eXI)Ina (Walker) (Lepidoptera Noctuidae) a review of the literature Cornshymonwealth Institute of Entomology London UK57 p

35 Brown F B 1968 Upland rice in Latin America FAO-International Rice Commission 12th Session of the Working Party on Rice Production and Protection 2-14 Sept Peradeniya Sri lanka Food and Agriculture Organization Rome Italy 6 p

36 Buyckx F J 1 1962 Disease and insect pests of crops inCongo Rwanda and Burundi [iil French] Publications dc LInstilut National pour lestude Agrononiquedu Congo M Weissenbruck S A Brussels Belgium 708 p

37 Cariiio F 0 1981 Role of natural enentics inpopulation suppression and pest management of green rice lealhoppers Ph D thesis University ofthe Philippines at Los Bafitos PhilippinesLaguna 136 -

38 Carpente A J 1983 The development of rice cultivation and extension of vegetable productin in Zanzibar Final Report Vol II1 October 1983 Food and Agricultme Organization Ronte Italy

39 Carroll C P and S1 Risch 1983 Tropical annual cropping systems ant ecology Environ Manage 751-57

4(0 Cates R G 198 1 Host plant predictability and the feeding patterns of monophagous oligophagous and polyphagous insect herbivores Oecologia (Berl) 48319-326

IRPS No 123 January 1987 37

41 Cendcafia S M and F B Caora 1967 Insect pests of rice in the 62 Ene 1 C 1981 I)iiorldhol)ierusihilis Slater (licteroplera Philippines Pages 591-616 in The najor insect pests ofthe rice plant lygacidue Illissinac) a new rice pest front Nigeria lit Rice Res Proceedings of a symposium at the International Rice Research Newsl N3)15 Institute September 1964 The Johns Hopkins Press Baltimore 63 Fugenio T S 1965 rhe effect oi e-vCgetation on rice production Maryland USA 729 p and the possibility of overcoming it by leaf pruning MS thesis

42 Centro Ihternacional de Agricklittira Tropical 1984 Upland rice in University ol the Philippines at Los Baijos ILaguna Ihilippins 69 p Iatin Anerica Pages 93-119 hi An os ervies of uplaind rice research 04 Farrow R A 1975 The Alrican nmigratory locust ii is main Proceedings of the 1982 Boinakamp Ivory Coast upland rice workshop outbreak area of the middle Niger qlinlitatise studies of solitary International Rice Research Institute P0 Blox 933 Manila populations in relation to environmental factors Locusla No II Philippines 198 p

43 Chandra G 1979 Taxonomy ad hn riononlics ofthe imect parasites of 65 Fennal R G 1943 Food crop investigation Windward and rice Icalloppers and planthoppers in the Pihilippines and their Ieeward Islands First report Report onl work dunring the periodiiiportairce iii natural biol)gical cointrol Philipp Fntoltol 4119-139 April 1942-April 1943 Imperial College of Tropical Agriculture

44 Chang T T 1976 The originii colut ion enlIivation dissemination Trinidad 16 p (Rev of Appl Fintornol A3222-25) and diersificition of Asian and African rices -uphytica 25425-44 1 66 Ferreira F 1983 Integrated pest ritaageiret of upland rice [ill

45 Chatierjee P B 1979 Rice leaffolder attacks in India lit Rice Res Portuguese English summary] Pages 323-341 in Upland riceculttre Newsl 4(3)21 factors affecting production E Ferreira T Yamada and F-

46 Cheng Cbien-pan 19(4 Upland rice in Taiwan International Rice Malmolta eds Institutoda Potassae Foslato Institnto Internacional Cnmmission Wiorking Party oit Rice Production ard Protection I()th da Potassa Piracicaha Sao Paulo lraid Meeting 3-10 March 1964 Manila Philippines Food and Agricul- 67 Ferreira E arid F dl Silva Martins 1979 Occurrence of ture Organiaion Rome Italy 5 p Sogatohs orvziola (Muir) inupland rice in Goias Ilrail lit Rice

47 Chcrrett 1 M and 1) 1Peregrine 1976 A review of the status of Rcs Ncwssl 4(4)18 leaf-cutting ants and their control Ann Appl Biol 84124-128 68 Ferreira E and 1F da Silva Martiris 1984 Insect pestof rice ill

48 Chipeta F M 1974 Pests of rice in Malawi Pages 90-95 in Brazil and their control Jill Portugucsej FMIlRAtPA-CNPAF Proceedings Sith eastern Africa cereals research conference 10-15 Goiania Brmiil 67 p March 1974 DRB Mialda ed Ministry of Agriculture lilongwe 69 Ferreira FI F ca Silha Miartins S S Nero and 1 It P de Malawi Carvallio 1982 M tltiplc regression analysis todeterimine the effect of

49 Chu Yau-i 1979 The d atiaging period of the rice seedling fly insect populatiois ott upland rice yield [inIPortuguese Pesqui (Atlterigotiavxigu Stcin) (NItiscide Diptera) oil cori) at Fast Java Agropecu Bras 17671-675 Indonesia Plant Prot Bill 21403413 70 Ferreira F 1F da Silva Martins and F 1PZimmermann 1979

50 Conklin H C 1957 IHaninoo agriculture a report on aln integral Rice varietal resistance to tle lesser corn stalk borer [in Port uguese] systent of shi fti rig cult ivationi in the Pltilippincs Food and Agriculture Pesqui Agropccu Biras 14317-321 Organization Rome Italy 209 p 71 Fleming W F 1968 Biological control of lie hapanese beetle U S

51 Cramer H II 1967 Plant proection and world crop production Dep Agric RLs Sers [ech Bull 1383 78 p Pilaniensclhutz-Nachr 201-524 72 Food and Agriculture Organiation of the United Nations 1982

52 l)ani R C and Majunidar 1978 Prelininary observations on tie Country report surmimary Pages 27-37 in Report on the first incidence of root aphids on differeit rice cultnrcs Sci Cult 4488-89 FAOIUN)IPhail and regional training coiuirse in iniproved ctltural

53 Davidson R I 1979 lffects of root feeding on foliage yield Pages practices for upland rice 5-23 Oct 1981 Northern Region Agrculturd117-120 in T K Ci osb and R B Pottinger eds Proceedings of the Developririt Center Chiang Mai Food and Agricuilture Organiiashy2nd Australian conlerence on grassland invertebrate ecology Govt tion Rome Italy 37 n Printer Wellington New Zealand 73 Franco do Ariaral S 195 1 Residual action of insecticides in turfy

54 l)ean G 1W 1976 Rice insect pests inlaos Int Rice Res Newsl soil--the results of a field experiment [in Portuguese] Biol6gico (Sao 1(2)15 Paulo) 17209-214 (Rev Appl Entomol A41423)

55 I)ean G J W 1978 Insect pests of rice in Laos PANS 24(3)280-289 74 Freeman ) 1970 Report onithe Iban london School of -coroiics 56 )ingle H 1984 Behavior genes and lifc histories conplex adapta- Monographs on Social Anthropology No 41 Athlane Press

tions to uncertain environments Pages 169-194 in A new ecology London UK 317 p novel approaches to interactive systems P W Price C N Slobod- 75 Fuchs T W1 A Harding and 1W Smithlr 1979 Induction and chikoff and W S Gaud cdsl ohn Wiley New York USA termination oldiapause in the sugarcane borer Ann Flitonlol Soc

57 Dobelmann-JP and M Falais 1968 Upland rice cultivation ill tie Am 72271-274 North-western part of Madagascar in 1966-1967 [in French English 76 (hbriel B1) 1975 A review tf the major insect pests of some tpland summary] Trop Agron 2313-21 crops in tlc Philippines Pages 86-111 in Ad Htoc panel of experts on

58 Doniiciano N L 1979 Chemical control of tie lesser corn stalk pest disease arnd weed problems in some rainfed crops Bangkok (lolsonno~paloIgnowslhts) (Zeller 1848) (I-epid(ptera Ihycitid ae) Thailand Food aid Agriculture Organization Roire Italy

on rice in Ponta Gressa Parara 1977-1978 [in Portuguese] Pages 77 Garg D K and N K Shal 1983 White grub outbreak on rainfed 291-303 in Anais 1 Reuniao de tecnicos cmiirizicultura do estado de dryland rice in Uttar Pradeslh Int Rice Res Ncwsl 81(4)16-17 Sao Paulo-Campinas Cati Brazil 78 Garg I) K and IP Tandon 1982 Major insect pests of rice on hilly

59 Dunsmore 1 R 1970 Investigations on tlie varieties pests and tracts of Uttar Pradesh India lilt Rice Res Newsl 7(11)11-12diseases of upland rice (hill padi) in Sarawak Malaysia Int Rice 79 Garrity 1) P 1983 Upland rice research an international bibliograpy Comm Newsl 1929-35 1975-1982 Iternational Rice Research hnstitnte PO Box 933

60 Elias 1R1965 The new decline of rice seedlings is not caused by Manila Philippines 121 p parasites [in Portuguese] Biol6gico (Sao Paulo) 313-6 80 Ghauri M S K 1982 A new subspecies of Iinorh)rone bull

61 Empresa Brasiliera de Pesquisa Agropecuaria 1984 Upland rice itt cornlus Slater (Hemiptera Iygaeidae) front Papua New Guinea on Brazil Pages 121-134 it Av overview of upland rice research rice and carpet grass Ihll lEntoniol Res 72133-137 Proceedings ofthe 1982 Blouak Ivory Coast Upland rice workshop 81 Gonzales F 0 PArregoces R I ihernandef and OT Parada International Rice Research Institute PO Box 933 Manila 1983 Insect and imite pests of rice arid their control in Latin America Philippines [in Spanish] la Federacion Nacional de Arrocerts (Fedearroz)

Bogota Colombia 60 n

38 IRPS No 123 January 1987

82 Grist D H and R J A W Lever 1969 Pests of rice Longmans London UK520 p

83 Hale P R 1979 Two subterranean pests of upland rice in Papua New Guirnca IntRice Res Newsl 4(2)16-17

84 Hale P R and J T Hale 1975 Insect pests of rice in the New Guinea Islands Rice Entomol Newsl 34

85 Harris W V 1969 Termites as pests of crops and trees Common-wealth Institute of Entomology London UK 41 p

86 Hattori 1 1968 A method to distinguish between grass leaf roller (Ctaplha rocis medinalisGuenee) and the rice leaf roller (Susurtia exigua Butler) [in Japanese] Plant Prot (Japan) 23167-170

87 Henry I E and 1A Onsager 1984 Experimental control of the mormon cricket Iahrosshnph by NJsena locustae(Aficrospora Microsporidae) a protozoan parasite of grasshoppers (Ort Acri-didae) Entomophaga 27197-201

88 Hidaka 1 P Vungsilabulr and SKadkao 1974 Studies on ecology and control of the rice gall midge in Thailand Tech Bull 6 TropicalAgricultural Research Center Tokyo Japan 113 p

89 Hincklcy A 1) 1963 Ecology and control of rice planthoppers in Fiji Bull Entomrol Res 53467 481

90 Hirao J 1982 Ecology and chtInical control of the rice leaffolder Jpn Pestic Inf 4114-17

91 Ho DT 1984 Sten borer incifience in rice ecosystems in KenyaEast Africa Pages 307-320 in A overview of upland rice research Proceedings of the 1982 Bouac Ivory Coast upland rice workshop International Rice Research Institute PO Box 933 Manila Philippines

92 Ho )T J 1Njokah and J GKibuka 1983 Studies on rice sten borers in Kenya with emphasis on ifaliarpiaipiraellRag Insect Sci Appl 465-73

93 Ichihara I T Ito A NIatsuo and M Sawada 1971 Chemical control of the rice striped stem borer in upland rice grown in plastic mulch [iii Japanese] Proc Kanto-Tosan Plant Prot Soc 1882

94 IclhiharaIand A Matsuo 1969 Damage caused by rice striped stett borer to upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1686-87

95 Ichhara I atid A MIatsun11970 Conside ration oft lie extraordinary prevalence of rice striped stem borer in upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1778-79

96 Institut de Recherches Agrolnliqes Iropicale et des Cultures Vivrieres 1978 Insect resistance Pages 12-13 in Upland rice 1978 atnual report Bouak Ivory Coast 27 p

97 International Institute for Tropical Agriculture 1983 Aphids itt rice roots I ITA Res Briefs 4(1-3)4-5

98 International Institite fi Tropical Agriculture 1983 Hopperburn in rice caused by brown plathopper IITA Res Briefs 4(l-3)6-7

99 International Rice Research Institute 1975 Annual report for 1974 PO Box 933 Manila Philippines p 216

100 International Rice Restarch Institute 1976 Annual repiort for 1975 PO Box 933 Manila Philippines p 228

101 International Rice Research Institute 1979 Cropping systems program Component technology development and evaluation Page 404 i Annual report for 1978 11O Box 933 Manila Philippines

102 International Rice Research Institute Department of Entomology 1981) Insecticid evaluation 1979 Tables 73 74 PO Box 933Manila Philipes

103 International Rice Research Institute Department of Entonmology 1981 Insecticide evalomtion 1980 Table 81 POBox 933 Manila Philippines

104 International Rice Research Institute 1981 Report of a meeting of Asian rice-based cropping systems entomologists Eleventh cropping systems working group meeting 18-22 May 1981 PO Box 933 Manila Philippines p 132 133 144

105 International Rice Research Institute 1984 Cropping systems program Pages 368-373 in Annual report for 1983 PO Box 933 Manila Philippines

106 International Rice Research Institutc 1984 Upland rice in Asia Pages 45-68 in An overview of upland rice research Proceedings of the 1982 Bouakamp Ivory Coast upland rice workiiop loternational Rice Research Institute 11O Box 933 Manila Philippines

107 Irshad M 1977 Grasshoppers in rice in Pakistan lit Rice Res Newsl 2(3)8-9

108 Ito R and TOno 1964 Upland lice its culture and varieties in Japan lIternational Rice Cormmission loth Meeting of the Working Party on Rice Production and Protection 3-10 March 1964 Manila Philippines Food and Agriculture Organization Rome Italy 12 p

109 Jeffrey M I and V A Dyck 1983 Effect of moonlight on lightshytrap catches of brown planthopper lit Rice Res Newsl 8(6) 13

110 Jennings P R and T Alicia Pineda 1983 Management ofthe rice plnthoppers Sogatodsorivi(coIl in Latin America Presented at the meeting on stable plant resistance FAO and lIndocsian I)inectorate of Food Crops Protection Bali lndonesia 29 Novcniber-2 l)ccemshyber 9 p

IllJimenez Gomez J A 1979 Rice insect pests i lla ios ()riCntales lil Spanish] Pages 99-103 in Rice ciltivation Instituto (olotibiatto Agropecuario Compndio No 29

112 Jongeleen F JJ Soesanto and A Mockidjo 1979 Studies oti pathogens of two species of Scarabacidac froinIndonesia Z Angew Enlomnol 87412-417

113 Jouvenaz 1) PC S lofgren and W A Baiks 1981 Biological control of imported fire antsa review of cirrent knowledge Bull Entomol Soc Am 27203-208

114 Kainoh Y K Shiniiu S MarUani Y Taiaki 1980 lost finding behavior of tie rice bu lplotori+JchinvittjI)allas (liciniptera Corcidac) with special reference to diet pattern ol aggregation and feeding on the rice plant Appl Entomol Zool 15225-233

115 Kakkar K 0 P Sharma and P I Sharma 1977 Note on Ihteronyrchus liodresRedt (Coleopicra I)ynastidae) as asporadic pest of paddy and maie inHimachal Pradesh Corr Agric 156-57

116 Kalode M131 79 Major insect pest problems and their control in upland rice (Unpublishcd mineogriph CWI oJNaaiat lni Chandaratat (198 1)original not seen)

117 Kalshoven L i 98 I The pests of crops in I ldoniesia (translated by P A vatn der Laan) 1 T Ichtiar B3aru-vati Iloeve lakarta Indonesia 701 p

118 Kamal N Q 1981 Suppression of whitebacked planto ner Sogatella fircaifra (Horvath) aLId rice lealfolder (nhal -cis licdb lis (Guenee) populations by natural cnclmie hlI) tiesis Grcgorio Araneta University Foundation Victoncta Park Metto Manila Philippines 179 Ip

119 Karim A N M Reaul 1975 A species ofliealy bug Trioni i70s ceres new oii rice in the Philippines Rice Entoniol Newsl 247

120 Katanyukul Wand C Chandaratat 1981 Insect pestsof uplaid rice and their control Report firstFAOUNlFi Tliailand training course on improved cultural practices for tplad rice Chiang Mai Thailand 5-23 Oct 1981 Food and Agriculture Organiation Rome Italy 22 p

121 Kern F 1955 lhletwioa(I)uninemts) bhhit Biturm(Coleoptera) a new pest of sugarcane inVenezuela [inSpanish with Etnglish sunmary] Agron Trop (Maracay) 527-29 (Rev Appl Eitoniol A4572)

122 Khan K M and S Ghai 1974 White grubs ald their control in India Pesticides (1Bombay) 819-25

123 Kiritani K 1975 Integrated pest management of rice pests Pages 66-70 itt Proceedings platning workshop ott cooperative hIeld research in pest managementP SMotooka ed East-West Center Honololu Hawaii USA

124 Kisimoto R 1967 Ecology of insect vectors forecasting and chemical control Pages 243-255 in The virus diseases of tle rice plant Proceedings of a symposium atthe International Rice Research Institute April 1967 The Johns Hopkins Press altimore Maryshyland USA

125 Kisimoto R and V A Dyck 1976 Climate and rice insects Pages 367-391 in Climate and rice International Rice Research Institute PO Box 933 Manila Philippines

126 Kureha S S Kobayashi SAsari and T Ishihara 1977 Life history and control of the mole cricket Grrllotalpa africana Palisot de Beauvois [in Japanese English translation 7520 1RRI library] Proc Kanto-Tosan Plant Prot Soc 21192-197

127 Kushwaha K S 1981 The black beetle as a pest of rice in Haryana India Int Rice Res Newsl 6(4)14

128 Lawani S M 1982 A review of tile effects of various agronomic practices on cereal stem borers Trop Pest Manage 28266-276

129 Leuck D B 1966 Biology of the lesser cornstalk borer in South Georgia J Econ Entomol 59 797-801

130 lever R J A W 1955 Insect pests of the rice plant in Malaya Int Rice Comm Newsl 1612-21

131 Li Ching-Sing and Shin-Foon Chiu 1951 The study ofthe rice gall midge Iachydiplosis orvza Wood Mason [in Chinese with English summary] 1Taiwan Agric Res 21-13

132 Iitsinger J A 1984 Assesament of the need-based application for rice Proceedings TechnologyTransfer Workshop 15-16 March 1984 International Rice Research Institute 1PO Box 933 Manila Philippines 6 p

133 Litsinger J A 1984 Environmental diversity and insect pest abundance with reference to the Pacific Pages 285-307 in Proceedings sub-regional training course on methods of controlling diseases insects and other pests of plants in the South Pacific 4-20 Oct 1982 Vaini Kingdom of Tonga Nukualofa

134 Litsinger J A RF Apostol and M BObusan 1983 White grub Leucopholis irrorata (Coleoptera Scarabaeidae) pest status popula-tion dynamics and chemical conllol in arice-maize cropping pattern in the Philippines J Econ Entoinol 761133-1138

135 Litsinger J A C de la Cruz F Raymundo A T Barrion M D Lumaban J P Bandong M S Venugopal F Paragna and E Balete 1982 Insect pests and insecticide reponse of dry-seeded bunded rice Pages 179-193 in Report of workshop on cropping systems research in Asia International Rice Research Institute Los Bafos Philippine

136 Litsinger J A M 1) Luinaban J P Bandong A T Barrion W Katanyukul and N Panda 1978 Insect pest management in rainfcd lowland rice agroecosystms of tropical Asia International Rice Research Conference 17-21 April 1978 International Rice Research Institute Los Bafios Philippines

137 Litsinger J A and K Moody 1976 Integrated pest management in multiple cropping systems Pages 293-317 in R I Papendick P A Sanchez and G B Triplett eds Multiple cropping American Society of Agronomy Madison Wisconsin USA

138 Loevinsohn M E 1984 The ecology and control of rice pests in relation to the intensity and synchrony of cultivation Ph D thesis University of London 36) p

139 Loevinsohn M E J A Litsinger N Panda M A Austria R F Apostol C Andrion and J Bandong 1982 A basis for predicting rice insect pest abundance in cropping systems Paper presented at the 1982 International rice research conference on rice and cropping systems 19-23 April 1982 International Rice Research Institute Los Bafios Iaguna Philippines 25 p

140 Lopez A W 1930 Repot of the Entomology Department Pages 145-172 in Annual report of the Philippine Sugar Association 1929-30 Manila

141 Lopez AW 1931 Entomology Department Pages 227-266 in Annual report of the director or research 1930-31 Philippine Sugar Association Manila

142 Lopez A W 1932 Entomology Deparment Pages 252-279 in Annual report of the Philippine Sugar Association 1931-32 Manila

143 Lungibill P and G G Ainslee 1917 The lessercornstalk borer US Dep Agric Agric Res Serv Tech Bull 539 27 p

IRPS No 123 January 1987 39

144 McGuire J U Jr and BS Crandall 1967 Survey of insect pests and plant diseases ofselected food crops of Mexico Central America and Panama U SDepartment of Agriculture International Agriculshytural Development Service Agricultuial Researrh Service October 1967 11 p

145 McNeill Sand T R ESouthwood 1978 The role of nitrogen in the development of insectplant relations Pages 77-9 in lliochemical aspects of plant and animal coevolution J B Ilarbornu ed Academic Press London UK

146 Maebara H 1955 Spatial distribution of lastinstiarlanaeof purplish stem borer (Sesamia iqferens Walker) hibernating in an upland rice field Bull Kagoshima Univ Fac Agric 438-41

147 Mallick E H and N G Hajra 1977 Response of rice plant to defoliation treatments at different developmental stages Indian Agric 2151-56

148 Mansingh A 1971 Physiological classification of dormancies in insects Canl Entomol 103983-10)9

149 Martell C G 1974 Catalogue of plant feeding insects of Mexico [in Spanish] Fitofilo 27(69)1-176

150 Martinez 0 J 1982 Principal rice pests in Ilanos orientalis [in Spanish] P~ages 80-94 in Curso deact nalization de conocimientos en arroz Villaniencis Instituto (olonmbiano Agropecuario

151 Martins J F da Silva F Ferrcira A S Prabhu F 1 P Zimmermann 1980 Use of pesticides for the control of the main subterranean upland rice pests esqui Agropecu Bras 1553-u

152 Martins I F da Silva F Fe eira rod BdamSilveira Pinheiro I Q2 Damage simulation caused by defoliators of upland rice [in Portuguese with English summary] Pesqui Agropecu Bras 171113-1119

153 Martins 1F da Silva N(iyuen van Tan and 1Ida Silveira Pinheiro 1981 Stem borer resistance in upland rice associated with plant morphological characters [in Portuuce Iesqui Agropecu Bras 16187-192

154 Masaki S 1980 Sumnmer diapause Ann Rev Entomiol 251-25 155 Masuzawa T 11 Suwa and F Nakasoji 1983 l)ifkleences of

oviposition preferencc and survival rate of two skipper butterflies Parnara guttata and PIchlpidai iathiais (Iepidoptera HIesperiidac) on rice plant and cogon grass New Entomol 3247-56

156 Maurya D M and C P Vaish 1984 Upland rice in India Pages 69-92 in An overview oftpland rice research Proceedings ofthe 1982 Bouakamp Ivory Coast upland rice workshop International Rice Research Institute IO Box 933 Mnila Philippines

157 Menon M G R R N Katiyai S Kuar and S S Misra 1970 Termites attacking the paddy crop Sci Cult 36294-295

158 Miyahara Y and 1 Nishikubo 1967 Behavior of young nymphs of the red rice root aphid Iholahsipiun ri liabhdontinalisSasaki in relation to movement from the leaf to the roots of upland rice [in Japanese] Pioc Assoc Plant Prot Kyushu 1363-66

159 Mo ljoa Tjien 1952 White grub of upland rice (Holotrichiahelleri Brsk) [in Indonesian with English summary] Contrib Agric Res Stn Bogor No 13134 p

160 Mochida 0 and T Suryana 1979 The occurrence of the brown planthopper and ragged stunt on lowland and upland rice ill Central Java and logyakarta in tile late of January 1979 Entomnolgy Survey Report No 15 Central Research Institute of Agriculture (CRIA) Sukamandi Indonesia I February 1979 9 p

161Monet C 1979 Varietal resistance to upland rice pests [inI French] Pages 35-39 in Annual report 1977 Institut des Savanes Department des Cultures Vivrieres Bouak6 Ivory Coast

162 Monte 0 1942 A caterpillar ofrice-fields[in Portuguese] Biol6gico (Sao Paulo) 8161-163 (Rev Appl Entomol A31414)

163 Munroe I) D 1979 Pest management strategy for hill rice in four resettlement areas in Sarawak Malaysia Pages 89-102 i1 Guidelines for integrated control of rice insect pests Food and Agriculture Organization Plant Production and Protection Paper 14 Rome Italy 115 p

40 IRPS No 123 January 1987

164 Nakasuji F M Ishii and Jen-Zon Ito 1984 Rice skippers in Taiwan and their life histories ZZrC (Taipei) Tech Bull 82 10 p

165 Navas I) 1976 Fall armyworum in rice Pages 99-106 il Ecological animal control by habitat management Proc Tall Timbers Con-ference 28 Feb-I Mar 1974 Fall Timbers Res Stn Misc Publ 6 Tallahassee Horida USA

166 Nickel 1 1 1961 Biological control of rice stem borer A feasibility study Tcch Bull 2 International Rice Rcarch Institute PO Box 933 Manila Philippines IIl p

167 Nigan P M and R A Vernia 1985 Insect pests of uplaild rice in Uttar Pradesh Int Rice Res Newsl 10(6)22

168 Nilakhe S S 1985 -cological observations on spittle bugs with emphasis on their occirence in rice Pcsqtui Agropecu Bras 20407-414

169 Nixon G EJ 195 1 lie association of ats with aphids and coccids Commonwealth Iusti lcte of Entonology Iondon UK 36 p

170 Nurdin F an Ishak Manti 1985 Study oit seedling fly of upland rice Interlational Upland Rice Moitoring Tour Report 26 lebruary-2 MarLI Sukaranti West Sumatra

171 Oka 1 N 1979 Ciltural conttol of the brown planthopper Pages 357-369 in Brown planthopper threat to rice production in Asia International Rice Research Institutc 1p0 Box 933 Manila Philippines

172 Ohni M and A Vilnaii 1975 Rice insect pests of Italy biology of Sipha grcria (Kaltenbach) and its control [in Italian] Riso (Milan) 2459-71

173 Otanes F Q atnd 1 I Sison 194 1 Pests of rice Philipp J Agric 12211-259

174 Painter R I 1951 Insect icsistance in crop plants University Press of Kansits Lawrence Kansas USA 520 p

175 Pantua P C and 1A litsinger 1981 Arthropod abundance in continuous vs biannual rice cropping systeis Int Rice Res Ncwsl 6(2)20-21

176 Pathak M ) and V A l)yck 1975 Sttudics ott insect pests of upland rice Pages 186-191 in Major research in upland rice International Rice Research Institute PO Box 933 Ilanila Philippines

177 Patil B R B M IHasabe and 1) S Ajri 198 I Chemical control of white gritb lIditriihinhorraia Fabr intfesting groundntut paddy atnd sorghum Indian 1Entomol 43232-236

178 Pawar A 1) 1974 (ollection and identification of insect pests of rice rlerminal repirt lInternational Rice Research Institute PO Box 933 Manila Philippines 72 p (tnimco)

179 IerrajU A aid A S Reedy 1965 Observations ott diapausc itt tie steti borer ohrice Rice News Teller (lai-Mar)9-13

180 Pont A C 1972 A review of the Oriental species of theri ona Rondani (I)iptera Muscidac) of econtomic importance Pages 27-102 in Control of sorglium shoot fly M G lomtani and W R Young eds Oxford atd IH1 Ptthl Co New I)clhi India 324 p

181 Prasad S C 1PChand aid M 1PSingh 1983 Incidence of rice gall midge in the upland plateau region of Hihar Oryza 20235-236

182 Rai P S 198 0 Nisaga simpex damage to rice in the hill tracts of South India Int Rice kles News[ 5(I)12

183 Rathorc Y S 1969 Gonocplhahli dcprssunt Fabr as itpredator of (tilo zonwIlhis (Swinhoc) and other moths in field cage at Patnagar Uttar Pradesh Indian 1Entotnol 3127

184 Reed W 1983 Crop losses causcd by insect pists in the developing world Proceedings 10th Internatial Congress of Plant Protection lrighton UK 174-80

185 Reedy M V 1981 Ateactogasterinhens(Walker)a new root-grub pest of tipland paddy and inaize Curr Sci 501076

186 Reveno M A 1968 Catalogue of the insect pests of economic crops in Colombia [in Spaiish] Asociacion Latino Americana de Entomo-logia (AIAE) 129-33

187 Reyes G M 1957 Rice dwarf disease in the Philippines Food and Agriculture Organiiation Plant Prot Bull 617-19

188 Rice S E A A Grigarick and M 0 Way 1982 Effect of leaf and panicle feeding byarmyworm (Lepidoptera Noctuidae) larvae on rice grain yield Environ Entomol 7593-595

189 Risch S J 1) Andow and M A Altieri 1983 Agioccosystem diversity arnd pest control data tentative conclusions and nizw research dirctions Environ Entomol 12625-629

190 Rivera C T 1956 Three organic insecticides in thecontrol of insects affecting upland rice Philipp Agric 39465-472

191 Roffrey 1 1972 Locusta outbreaks in the Philippines Acrida 1177-188

192 Rojas R 1) 1941 The effect of pruning at varying ages upon the yield of Ramairio Phipp Agric 29851-860

193 Roonwal M i 1979 Ternite life and termite control in tropical South Asia Scientilic Pnbl odhpr India 139 p

194 Rosetto C1 F da Silva Martins N C Schmidt and L EAzzini 1978 Damage caused by pasturespittle bugs 4Dvoi)lavopictaandDJ schach) on rice lBragantia Nota 6 (Matcli)25-27

195 Rosztto C1- S S Neto I- Amantc I) Iink 1Garcia-Vieira D M de SoLza N V Bansa t and A M de Olivcira 1971 Rice pests of Brazil [in Portuguese] Pages 149-238 inrFA()-International Rice Commission 2nd meeting of the Rice Committee for the Americas 6-11 l)ecember 1971 Pelotas Rio Grande do Sul Brazil

196 Rothschild G H L 1969 Obseratiois on the armyworm Spodoptera matritia acronrctoiiles Gn (Lep Noctuidae) Bull Entomol Res 59143-160

197 Rothschild G 1- L 1974 Parasites of the armworm Spolopiera mauritiaacroncitoihes in Malaysia Fntornopliaga 19293-299

198 Sachan i N S K Gangwar and 1N Katujar 1980 Field crickets damaging cages in North Eastern IHill Region Indian J Entomol 42526-527

199 Sands 1) P A 1977 The biology and ecology of Lepocorisa (Hemiptera Alydidac) in Papua New Guinea Res Bull 18 Departshyment of Primary Industry Port Moresby Papua New Guinea 104 p

200 Sarawak 1968 titomology division Pages 97-106 in Report of the research branch of the Department of Altriculture 1966 Kuching (Rev AIppl Eintomol A56603)

201 Satpathy JM M S Das and K Naik 1977 Effect of multiple and mixed cropping on the incidence of scine important pestsI Entomol Res 178-85

202 Sauer IHF G 1939 Notes Ott Elasmpnlpualtsligniosellths Zeller (Lep Pyr) a serious pest ofcereals in Sao Paolo [in Portuguese] Arq Inst Biol 10199-206 (Rev Appl Entotnol A28378)

203 Satinders 1L A 1 SKing and C L Vargas S 1983 Crop pests of Central Anerica with t bibliography [in Spanish] Centro Agroshynotnico Tropical de Investigacion y Ensenanza Turrialba Costa Rica (CATIE) Tec Bol 9 90 p

204 Sauphanor B 1985 Some factors of upland rice tolerttce to stem borers in West Africa Inseu Sci Appl 6429434

205 Sen P atnd S Chakravorty 1976 Stentflics in Haritighata Nadia West Bengal India [it Rice Res Newsl 1(12) 15

206 Senapai B and 1M Satpathy 1983 The seedling fly Atherigona orrzae Malloch is a pest of rice in Orissa (India) J Entomol Res 777-79

207 Shhjahan M S Alatn M Ah nad and BN Islam 1983 A new scarabaeid beetle pest of dryland rice in Bangladesh Int Rice Res Ncwsl 8(I)16-17

208 Sharma G L S C Modgol and R C Gautam 1979 Effect of depth of submergence ott incidence of bacterial blight and white grub infestation in transplanted rice Int Rice Res Newsl 4(6)12-13

209 Sharma S 1985 Scarabaeid beetle outbreak itt upland rice Int Rice Res Newsl 1(1(l)24

210 Shiraki T 1917 The paddy borer Schoenobhits incertulas Walker [achaku Agric Exp Stn Formosa 256 p

211 Silva A de Brito and 11P Magalhaes 1981 Insect pests of rice [in Portuguese] Circular Tecnica No 22 EMBRAPA Centro de Pesquisa t ecuaria de Tropico Unido (CPAJU) Belem Brazil 14 p

212 Singh S A 1979 Emergence of rice leaf butterfly as a rice pest in Manipur India int Rice Res Newsl 4(4)16-17

213 SinghS A 1983 Changing trends of insect pestsof rice in Manipur Int Rice Res Newsl 8(f)12

214 Siwi S S and P H van Doesburg 1984 Lenovorisa Latreille in Indonesia (Heteroptera Coreidae Alydinae) Zool Meded (Leiden) 58117-129

215 Soejitno J 1977 Reaction of some upland rice varieties toseedling fly eltherikonaexigua Stein [in Indonesian] Siri Hana I Penyakit 630-35 Lembga I osat Fenclilian Pertanian Bogor Indonesia

216 Socnardi 1967 Insect pests of rice in Indonesia Pages675-683 in The major inse pests of thu rice plant Proceedings of a symposium at the Internatital Rice Rec Wi itt S pternber 1964 The Johns Hopkins Press Baltimore Maryland USA

217 Soto P E A T Petcz and I W Buddenhagen 1976 Survey of insect pests and diseases of rice in different ecological zones in Nigeria Rice Entomol Ncwsl 435-46

218 Soto 11E and Z Ziddiqi 1976 Screening for resistance to African rice insect pests Varietal improsetnent sCmi n r Bouak Ivory Coast 13-17 Sept 1976 West Africa Rice l)evelopment Association Monrovia Iiberia

219 Squire F A 1972 Entomological problems in Bolivia PANS (PestArtic News Sumr) 18249-268

220 Stapely 1 II 1978 The rice leaf roller in the Solomon Islands Int Rice Res Newsl 3(l)13

221 Suenaga H 1937 On aphids attacking the roots of rice Oyo-I)oubts Zasshi 9150-152 [in Japanese] (Rev Appl Entomol A25745)

222 Suenaga H 1952 On the seasonal abundance of three subterranean aphids on upland rice Bull Kyushu Agric Exp Sti 1249-262

223 Sutynso Yas and Dandi Soekaria 1964 Beberapa hana yang ttenperang tana man padi tanah kering [in Indonesian] Rapat kerja Padi Tanah Kering 6 sd 8 Juily 1964 Kaliurang 9 p

224 Tallamy I) W and R F Denno 1983 Alternative life history patterns in risky environments in example from lace bugs Pages 129-147 in Insect life history patterns R F l)cnio and If Dingle eds Spring-Verlag 225 p

225 Tamura 1 1939 Observations on some insect pests of field crops [in Japanese] 1 Plant Prot 26414-420 (Rev Appl EntomolA27-556)

226 Tamura and Suzuki 1964 Influences of the early-transplanting practice in rice culture upon development of the rice stem borer Chilo siql)ressalisWalker Bull Ilokuriku Natil Agric Exp Stn 761-94

227 Tan J P 1924 The rice root aphid Drowhi ihirsoia Philipp Agric 13277-288

228 Tanaka 1 1957 Taxonomy and distribution of some subterranean aphids injurious to upland rice in Japan with description of a new species [in Japanese English summary] Botyu-Kagaku 22168-176

229 Tanaka T 1961 The rice root aphids their ecology and control Spec [full College of Agriculture Utsunomiya University Japan F3 p

230 Tauber M IC A Tauber R Nechols and 1Obrycki 1983 Seasonal activity of parasitoids control by external internal and genetic factors Pages 87-108 itt I)iapause and life cycle strategies in insects V K Brown and I Ilodek eds )r W Junk The Hague Netherlands

231 Taylor W F 1972 Effects of artificial defoliation (simulating pest damage) on varieties of upland rice Exp Agric 879-83

232 feotia T P S and S Natud 1966 Biotoomics of the rice skipper Parnaraniathiis Fabricius (Lepidoptera Hesperiidac) Indian J Entomol 28181-186

233 Teran F 0 1971 Rice pests of Santa Cruz [in Spanish Ministerio de Agricultura Sita Cruz Bolivia Bull 45 8 p

234 Than Itun 1976 Population dynamics of the yellow rice borer Tryporza itcertulas (Walker) and its damage to the rice plant MS thesis University of the Philippines at Ios Bafios Laguna Philippines 92 p

235 Uichanco L B 1911 Pests Pages 55-74 it Rice Philipp Agric 11-200

IRPS No 123 January 1987 41

236 van der Laan P A 195 1 Life history of the rice seedling fly Contrib Gen Agric Res Stn Boor Indonesia 118 15 p

237 van der Laan 11A 1959 Correlation between rainfall in the dry season and the occurrence of the white rice borer (Scirpolhaga innotala WIk) in lava Entomol Fxp Appl 212-20

238 Van Driesche R G 1983 Meaning of percent parasitismin studies of insect parasitism Fnviron Entomol 121 I1- 122

239 Venugopal M S and A litsingcr 1984 Fffect ofcarbofuran on rice growth Prot Ecol 7313-317

240 Vietnam Directorate of Rural Affairs 1962 Annual work progress report on crop improvement program of rice sugarcane vegetable and field crops (lor the period from July 1961 to June 1962) (11) 299 p (Rev Appl Entomol A51526-527)

241 Wan M T K 1972 Observations on rice leaf- and planthoppers in Sarawak (Malaysian Borneo) Malay Agric 148308-335

242 West Africa Rice I)esclopnient Association 1978 Annual research report Vol 2 Statistics Monrovia Liberia

243 West Africa Rice Development Association 1979 Annual research report p47 Monrovia Liberia

244 Williams ) F and CS lofgren 1981 Fli I illy l-468 anew bait toxicant for control of the red imported lire at Fla Entomol 64472477

245 Williams I) 1 1970 The mealybugs OFlomoptera Coccoidea Iscudococcidac) of sugar-cane rice and sorghum Bull Entomol Res 0109-188

246 Williams D) 1 1982 Pulinaria icervi (Signoret) and its allies ott sugar cane and other grasses Btll Entoinol Res 72111-117

247 Wirjosuiardjo S 1976 Study ofchenmical cont rot of insect pests ott upland rice Imiu Pertanian (Jogjakarta) 1323-329

248 Wirjosuhardjo S and N Wirarta 1976 Chemical control of the white grub tIolorithiatlltriin upland rice in M uneng Irobolinggo Indonesia) [in Indonesian] Imut Pertanian (Jogjakarta) 1330-332

249 Yano K T Miyake and V F Eastop 1983 The biology and economic importance of rice aphids (Henuiptera Aphid idae) a review Bull Entomol Res 73539-566

250 Yelp D and P Rhodes 19(4 lhe success of drilled dryland rice iii the Fiji Islands International Rice Commission Working Party on Rice Production and Protection Manila Philippines 3-10 March1964 Food and Agriculture Organization Rome Italy 13 p

251 Young W R 1981 Fifty-five years of research on the sorghum shootfly Insect Sci Appi 23-9

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No 104 Faridpur acomputer-assisted instruction nmodel forrainfed lowland random coefficient model No 116 Consumer demand for rice grain quality in Thailand Indonesia andrice

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No 106 Rice grassy stunt virus 2 a new strain of rice grassy stunt in tire literature

Philippines No 118 IRRI-Korca collaborative project for the development of cold-No 107 Physical losses and quality deterioration in rice plstprodnCtion tolerant lines through anthei culture

systemis No 119 Problen soils as potenial areas for adverse soils-tolerant rice No 108 (olptblication ofIRRI materials a survey of translators and varieties in South and Ssuthcasi Asia

publishers No 120 Changes isinall-farm iLtireliing techii)hgy in Thailand and [he No 109 Classification of Philippine rainfall patterns Philippines No 110 Contributions of modern rice varieties to rutrition in Asia No 121 Landforms and modern rice varieties No Ill Changes in rice breeding in 10 Asian countries 1965-84 No 122 Yield stability and modern rice technology No 112 Design parameters affecting the performance of the IRRI-designed

axial-flow pump

Page 12: UPLANDRICE. INSECTl--1i M T -E!R.EOLOU t RTANCE ,AND …delphacid.s3.amazonaws.com › 6958.pdf · The significance of abiotic factors in upland rice insect ecology will be apparent

12 IRPS No 123 January 1987

Root weevils A few species of upland weevils (IHporneces Donacia Atactogaster) attack rice roots as larvae and feed on stems and leaves as adults Damage ispatchy The larvae are not adapted to submergence and are true upland soil weevils isopposed to the so-called water weevils They rarely become sufficiently abundant to cause damage Adults on rice foliage or the soil surface show that the roots are being attacked by the grubs Root weevils are highly polyphagous (185)

Root heeths Soil-inhabiting false wireworms Gono-cephalumt spp are reported in upland ricefields The adults and larvae normally feed on decaying organic matter but land preparation for upland rice removes these sources The larvae feed on seedling roots and nocturnal adults roam on the soil surface to cut off seedlings at ground level (82) This group of ground beetles is highly omnivorous adults even prey on stem borer larvae (183) True wireworms (Elate-ridae) feed on living roots but rice as a host has been reported Infrequently (117) In Latin America the highlypclyphagous larvae of Diabrotica rootworms are also recorded on rice

Root htus Upland rice seedlings can be attacked by the adults anI nlViphs of chinch bugs Blissusand Caenoblissus and bro n bug Scaptocoris which feed on the roots with theirsucking nouthparts (80 83) Three genera of burrower bugs A tms (eotonhus and Stiharopus -are common iil grasslands in the Philippines and may attack the first crop (rice or maize) planted after the dry season Chinch bugs are most abundant after a dry spell These insects are large and can kill seedlings or stunt and reduce tillering on older plants I hey prefer other hosts and rarely damage rice Adults can also feed oii developing rice grains (81)

(utworms Soil-dwelling noctuid larvae hide under-ground during the day to avoid predation by birds and bccone active at night above grour cutting off you ing plants at ground level (hence tile naiie cvtworn) arvaedrag severed plants into hurrows llie larvae have wide host ranges arid pupatc in the soillgrotis ipsilon occurs worldwide and appears to be the most common cutworm pest of upland environments but is more prevalent inlowland rice particularly in dry seedbeds

IFoliar pests Other groups of insects are adapted to upland environments because they pass one growth stage underground either as egg (in heinrietabolous species) or pupa (in holomnetabolous groups)

Rice seedlibig maggots AitJeriota flies occur in non-fhoodcd environments in Asia and Africa (180) Feeding of larvae one per iller causes deadhearts Flies as opposed to moths only secondarily evolved as pliytophagous pestsfroni a saprophagous Origin Athrnonalarvae secondarihv feed onidecaying tillers whereas larvae of stem borer moths actively Iced on living tillers to cause deadlicarts Seedling maggots will die if they cannot sever a tiller on which to feed

as it decays Plants die under heavy attack whereas lighter infestations cause stunting delayed maturity and ragged leaves (236)

Eggs are laid only on actively growing plants (49) aid damage occurs wiihin the first week after crop emergence(170) Larvae pupate more in stems than in soil Seedling maggot attack is highly seasonal (206) normally peaking 2-3 mo after the onset of the rainy season These pests possiblyaestivatc over the dry season Rice isonly one of their many graminaceous crop and weed hosts

The rice stem maggot (hlorops orzae is adapted to temperate upland and lowland rice growing regions of Asia In Japan and China upland rice grows in mountainous regions a habitat of this chloropid fly The larvae tunnel into tilestems The stem maggot is more damaging during the seedling stage larvac hibernate in grasses

File several dipterans reported as stem maggots in dryshyseeded lowland seedlings inIndia (205) could presumably attack upland rio

Leaf urinrs Icaf miners are common on wheat and maize but are rarely reported on upland rice (213) Three leaf miners are more prevalent on upland than lowland rice inthe Philippines Pvudonalomy a asicanh Vficataand lIAgromrza orzae attack at the seedling stage The larvae develop inside the tunnels they construct in the parenchyma tissue P Vpicata and Aorvzac prefer maizeEehtsine and other grasses to rice spicatacan become abundant on wheat

Leaf beetles Chrysomelid and coccinellid eaf bcetles scrape or otherwise remove leaf tissue as larvae or adults or both This damage accelerates desiccation of plants Tile larvae of sonic species are root feeders Flea beetles Epitrix and (lraetocnetna and rootworms l)iahroticaspp feed on rice foliage only as adults Their root-feeding larvae have hosts other than rice The adults and larvae of the hispa-like Oediolmlpa spp beetles of Brazil and (hmootriha (=EPfishyachra) beetle of Africa feed on upland rice leaves Oealioshypapa spp larvae are leaf miners (mootriha is also pshyvalent in lowland environments (11) Leaf beetles prefer vegetative rice (81)

The rice hispa Dicladipaarnigera is not reported on upland rice and prcers more aquatic habitats In Central Africa however I)cladispm viridicyaiea attacks uplandrice in the vegetative stage as well as lowland seedbeds 7richipasericea is reported on upland rice in West Africa and Madagascar (57)

The rice leaf beetle 1d1ea 01izae riiuech like Chioropsor|rzae is adapted to temperate regions of Ashi and is usually a pest on lowland rice but seldom oi inpland rice in the mountains (82) The adult hibernates on grasses

Armmworms Species of grassland-adapted SpoptloeraPlatysemta and A ithimna larvae create widespread epishydemics They can defoliate ricefields generally in patchesfrom early vegetative growth to harvest (41) Armywornms pupate inthe soil and highly favor upland rice (34) Adults

can disperse long distances to colonize even remote upland areas They become more abundant than their natural enemies particularly after favorable rains following a prolonged drought (196) The drought kills the armyworm and natural enemies (mainly parasites) but the armyworm can recolonize rapidly

The natural increase insoil fertility from mineralization of soil N over the drought period promotes luxuriant plant growth to foster a rapid buildup of these highly fecund species Weeding rice increases tilelikelihood of damage The parasites normally return to control the armyworms but not until after serious damage

Outbreaks on upland rice have been reported in Ghana (3) inzibar (38) Central Africa (36) lPanama (165) Brazil (28) Malaysia (196) and India (167) Defoliation may be severe often leaving only the base of stems Armyworm larvae hide during the day under litter or in soil cracks At night they ascend plants to feed Larvae pupate in the soil

Thrips Thrips frequent upland rice but damage is less than in the lowlands Thrips larvae and adults feed on leaf blades by rasping causing leaf rolling and stippling Thrips prefer vegetative stage rice and their survival increases during drought (heavy rain washes them from foliage) although drought stress usually outweighs thrips damage Drought and thrips cause leaf rolling in upland rice (235) but thrips have been overlooked perhaps becaLse of their small size Thrips are most numerous indry season irrigated rice The combination of favorable plant growth and lack of rainfall is ideal for thrips In contrast the lack of rainfall in upland rice hurts the crop and eventually the thrips whose numbers must decline

In [atin America Frankinilla rodeos attacks panicles before they emerge from the boot calusing sterile grains

Stem hug~s Black bugs are the most common group of rice stem-feeding -lemiptera in Asia The most frequently mentioned species ----Scotinophara coarctaia and Shrida - prefer aquatic habitats and therefore could not occur in upland rice as sometimes reported Upland species in the same gerurs - notably S tarsalis and S scotti -- are morphologically similar and remove sap from stems of grasses and upland rice The upland species are notably more abundant in upland rice planted near forests

In Latin America tile Tibraca ima-large pentatomid tiventris causes deadhearts and whitehcads on upland rice Both adults and nymphs remove sap from internodes of plants more than 3 wk old Tillers are killed by mechanical damage and by the entry of secondary microbial infection This damage is often confused with that of lepidopterous stem borers (233) Eggs are laid on the leaves and stems Nymphs are gregarious This species can also be found on irrigated rice Drought reduces population buildup

Stiem borers Lepidopterous and dipterous stem borers are widely recorded on upland rice but cultural practices and alternate hosts determine abtndance Except for Elasmolmll)uS Iinosellts numbers usually build up toward

IRPS No 123 January 1987 13

the end of the crop growth cycle Therefore late-maturing varieties staggered planting and lack offloodingfavorstem borer buildup Many stem borer species can lie dormant in the stUbbleafter harvest and theirsurvival is encouraged by panicle harvesting (leaving tall stems) and lack of tillage after harvest

Stem borers disperse and lay many eggs Many species are polyphagous however the monophagous MaliarpiasepashyratellaDiopsis Ivmgicornis Scirpophaga incertlas and S innotata can also be abundant on upland rice (hilo Suplressalisdevelops fasterand becomes la rger when reared on upland than on lowland rice (226)

The lesser corn stalk borer lasmopal tt Ittoselhs is semisubterranean and is perhaps the stern borer most adapted to upland rice(129) Highly polyphagous it infests maize peanut and cowpea It attacks seedling rice (one larva can kill tip to four plants before maximum tillering) (195) (hiv and Maliarphastem borers prefer older plants But Scirpophagaspp will attack young plants even in the nursery lasn opalls larvae tunnel into the stems at or below the soil level mausing dcadhcarts Larvae are not found in their turnnels inside sterns beca use vhei disturbed they retreat into cases made from soil particles bound by silk The cases areattached to the tunnel entrance (143) This behavior probably evolved as protection from natural enemies Larvae pupate in the soil The lesser corn stalk borer is more abundant during drought damage is often confused with that of drought (233) This species prefers sandy soils arid its distribution within a field may relate to soil texture (143)

Diatraeasaccharalis aside from the lesser corn stalk borer is frequently a stem borer pest in Latin America According to Teran (233) rice and maize are more preferred hosts than sugarcaine its namesake

Atcitonacrvsograpla is a less known upland rice stem borer in Asia with habits similar to Easinopall s (82) because it is not found in lowlands Acutra has grassy weed alternate hosts As a moth it is often confused with the dark-headed stern borer(hi polur(standstriped stern borer (hi sup)lpsrcais

Other common upland rice stem borers are also reported in the lowlands The dominant species in upland rice vary greatly with location In Uttar lPradesh India (78) and in Japan (123) the pink stem borer Sesamia htfirens is more prevalent hIKenya the principal rice stem borer is Chi partelus

(92) The lowland African sten borers Aialiarpha sepashyratellaDiopsis vngicornis and Sesamia calatnisnis are less abundant Oimupland than on lowland rice(217) but perhaps cause greater yield loss because plant injury is greater if combined with drought css (92) The stalk-eyed stern borer D)ivpsis is more prevalent in wet habitats (6 1I217)

In Central and West Africa Maliarpha and Sesamia calanistis are the dominant stern borers of upland rice (8 9 10 II) (lilo dihlfilineui and Chio partellus inhabit the

14 IRPS No 123 January 1987

upland savannas whereas Chilo zacconio prefers the lowlands (II) Aaliarpha is most abundant in the mangrove swamp habitats (91) coastal regions (16) or rain forest zones (II) of Africa and appears to be the homologue ofScirlpophaga incertulas The yellow stem borr can also be prevalent in upland rice (120 167) The sugarcane borer Eldanasaccharinais so far only a potential pest of uplandrice (91) despite Grist and evers (82) warning

Grasshotl)ers andI locitas Short-horned grasshoppers and locusts -- Locvta Patanrga Schistocerca - layeggs in the soil inhabit grasslands and develop into swarms to seriously damage upland rice from time to time (173) Largenumbers can destroy a field leaving only the stubble In low cmuoroer s giassnoppeirs an cut panicles (173) Farmers may not plant if locust swarms are imminent Prolonged dryweather followed by favorable rain favors development of large swarms (173) Usually breeding in di) grasslands theydisperse toattack crops including upland rice(191) Eggsaresusceptible to desiccation d uring drough t and toinundation during seasonal rains With favorable soil moisture over consecutive seasons natural enemies ca nnot prevent a largepopulation increase leading to migratory swarms (64)

Most grasshoppers on rice are adapted to uplands (107)but lowland rice grasshoppers iervgl(rlhusspp and Ora spp occur occasionally on upland rice (2 82) They lay eggson rice foliage Nymphs are scmiaquatic

The gryllid Lsctrtus coucinnus is a pest of lowland and deep water rice in Bangladesh and Thailand In the Philippines it damages upland rice grown in isolated pockets near lakes The gryllid nymphs and ad ults feed on the central portions of folrage leasing only the midrib and leaf nargins

Leal ler Pvralid moths whose larvae fold leaves to make a feeding shelter occur worldwide (217) but arc appareritly most common in Asia Leaffolde rsattacking rice in Asia are a complex of species (26) (Ctalhahcrcisinedinalis the best-known rice Ia flolde r in Asia may prefer grassy weeds over rice (86) It is a late colonizer of uplandrice U1arasmja eviua a lowland species (220) nreflers rice to grass specres (86) and colonizes upland rice during il early growth stages llarasnria patnalis is the most corn-rnonly encountered species in upland rice in the Philippines Perennial grasses are their alternate hosts and they can survive year-round in upland rice environments The leaf-folders 11arasmia ruralisand Alarasmia spp are least aburndant Dormancy is unknown in leaffolders Cnaplha-ocrocis medhnalis however is known as a migrant (90)Leaffolders remove photosynthetic tissue attack during the flag leaf stage is particularly injurious Popurlations are normally held in check by natural enemies but leaffolders become parlicularly abundant in conditions of high plantfertility and shade

Butterflies Skippers PelopidasInathiasand Borhofanta are more prevalent in African lowland than in uplandhabitats (6) but extensive studies in Japan and China

showed that Pelopidas mathias prefers an upland habitat (155) Parnaragutata seasonally migrates between lowland and upland areas and is equally abundant in both environshyments (164)

The green horned caterpillar Melanitis leda ismene is more abundant in lowlands (212) but our experience in the Philippines suggests that Pelopidas niathiasand 11elanifis leda ismene are often more abundant in uplands than in lowlands Bamboo iaunbuiva spp annually sustains P mathiaspopulations in the Philippine uplands Adults feed on nectar from flowers and migrate to upland rice during the late vegetative stage

iPi erlled h i-1 1d dill L tIC i 7-0 l(become numerous probably because of high egg predationEgg and larval parasitization are often low Butterflies can readily disperse long distances to seek renlote uplandricefields However they have low biotic potential and even it predation rates are low the rarely become abundant Upland environments with their diverse microhabitats can provide more favorable sites than lowland rice plains

tutterfly pests of rice have broad host ranges and become dormant Juring urnfavorable times of the year

IolhagrotslaidopwraThe larvae of many polyshyphagous moths are reported to defoliate upland rice tussock moths lsaispelnnatula Iawlia sUifita luproctisirgtwltaE mnorand P vathorrhoeaslug caterpillar

litotiahicolorhairy caterpillars Nisaga sinp vandA mata spp and wooly bears l)iacrisiaohliqtia and Crvatonotts gangis Many of these polyphgous larvac migrate and are transients in rice including HelhWhis arm iera and 1t zea highly pol phagous species (82)

Gall nidges The Asian rice gall midge Orseoliaorrzae (88 131) and its African counterpart 0 ortzivora(6 11 12 32217) prelerlowland to upland rice Gall midges in rainfed areas are prevalent only during tle rainy season and dispersepoorly Their alternate hosts (wild rices LeersiaIschaetnuan and Pasahmt)are mostlyaquatic grasses and are less likelyto be near upland rice areas In an area of mixed uplandfloating and rainfed lowland rice cultures in northern Thailand upland rice which is ilanted a month earlier was attacked firstbut subsequcent populations were much lower than in rainfed lowla id or floating rice Low numbers of gallmidges passed the dry season orr perennial weeds such as Paspahim (88)

Smot (hid(s Reports of aphids attacking upland rice plants are few (249) Without maize the maize leaf aphidRho alosihhunm maidis attacks upland rice in the Philipshypines The plum peach aphid IIrsteroneuraselariae was reported in Sierra Leerne toinfest leavesand tnriperied grain(7) Siphaglrctriaecaused transverse linrea r necrotic striae iir rice leaves in Italy (172) I-lea v rains held populations in check Rice is not a preferred host for shoot aphids

Leafhoper andI )anthol)pers Cicadellids and delshyphacidsdispersecasilyand readilycolonie upland ricefrom nearby lowland areas A study in Sarawak (241) showed

IRPS No 123 January 1987 15

similar species in upland and lowland rice Nephoiettix The principal vectors of rice virus diseases generally have species dominated The most prevalent species accounting narrow host ranges and alternate hosts in the grass family for 90-95 of the sweep net samples wre N virescens N are mostly annuals dying in the dr3 season Some virus umropictus Nilaparvia hgens Soatellafiircifera Cfana vectors such as BPH can disperse lontg distances but few do

sectraand Recilia dorsalisLowlands had higher popula- Th restriction of upland rice and weed hosts to the rainy tions Katanyukul and Chandaratat (120) monitored hopper season the small area planted to upland rice and the hopper numbers in upland and lowland fields of northern Thaiiand vectors having narrow host ranges all work against virus and found similar results A comparative study of plant- diseases in upland rice hoppers in Fiji showed more brown planthoppe BPH) and In Latin America Sogctdhs o(rzicola and S cuhanus whitebacked planthopper (WBPH) in wetland rice than in transmit hoja blanca virus disease but reports on upland lowland rice and hopperburn was prevalent in lowland rice are rare except in the favored upland areas of Colombia areas with standing water (89) Peru and Venezuela (35 42) S rricolaocurs in Brazil

Hopperburn in upland ricefields in Asia is rare (84) In on upland rice but hojaI blanca has not been reported (67) Batangas Philippines isolated fields of upland rice become S orzicola has a wide host ramgc (10) similar to WBPI- in hopperburned infrequently from WBPH A report from Fiji Asia WBPH however is not known to transmit any virus cites hopperburn from BPH and WBPH (250) In these two disease situations upland rice was grown on highly fertic soil and In Africa pale yellow mosaic virus is endemic to swampy N fertilizer was applied Hoppers respond to better nutrition lowland areas where rice ratoons year-round (19) and is Normally upland rice is grown on poor soil and receives no mechanically vectored mainly by hispid beetles A minor fertilizer vector a flea beetle Chaeiocnema spp is an upland rice

In Asia upland rice ras are normally not far from pest and upland ricetields near endemic areas occasionally lowland ricefields-- a source of hoppers Otherwise hopper contract the virus species with greater polyphagy than BPH or Nelhotettiv Spittle bugs Cercopid nymphs producc a protective virescens would be dominant frothy covering that looks like saliva In Latin America

In Africa leathoppers and planthoppers do not become spittle bugs damage upland rice In parts of Brazil where abundant on upland rice (6 II) probably because lowland upland ricefields are surrounded by pastures adults rice is not widely rlanted nearby The only reported migrating from grasses can damage rice severely and cause hopperburn as from Nilapanrata maeander in breeder complete hopperburn in a young rice crop within a week of plots receiving high N rates in Nigeria (98) Hoppers colonization (194 195) Dispersal to rice occurs even if however are a greter problem in Latin America where pastures are verdant and spittle bugs prefer pasture grasses large grassland areas breed polyphagous species (Gra- to rice (168) Nymphs are rarely abundant enough to phocephala spp Hortensia spp lEvitianus ohscurunervis damage rice Regular rainfall favors their development Balclutha spp and Draeculacephala spp) vhich then (233) disperse to cause hopperburn on seedling rice (68) Rosetto et al (195) reported Graphocep7hala spp and Hortensia spp Seed pests to be equally abundant on upland and lowland rice Alydid and pentatomid bugs are seed pests of upland rice

The leafliopper and planthopper epidemics in Asia that worldwide In Asia Leptocorisa sp prevail Their habits caused severe losses from hopperburn and vijus dicases make them particularly suitable for upland rice 1)thev feed over the past several decades occurred principally on on many grasses (and therefore can survive the earl y wet lowland rice Except in a few isolated cases (160) upland season before rice sets its grain) 2)adults are long-ied and rice has been spared from tungro grassy stunt and ragged mobile allowing them to find isolated plantings 0 rice stunt virus diseases vectorcd by green lcafhopper (GLH) (besides seed bugs only birds and rats appear to have shyand BPI] (123) Virus diseases have occurred in upland rice capability) and 3) adults aestivate in forested areas or where lowland rice was nearby Rice dwarf disease however sugarcane fields during the dry sca son when neither rice nor was first recorded on upland rice in the Philippines (187) grassy weeds are present (I199) Rice seed bugs concentrate Virus diseases can be perpetuated only on livir plants or on small-scale upland riceficlds because they can actively insect vectors and no living rice - plans tooned or search them out The prevalence of groves of trees characshyvolunteer - grows year-round in upland arcas excert in a teristic of upland environments ensures nearby aostivation few with 12 mo of rainfall sites keeping the seed bugs close by

The smaller brown planthopper Laodcelphax striatellu Leptocorisae are uncommon in lowland rice plains occurs in upland rice in Japan (18) It spreads black- because 1) their populations are diluted in a sea of rice streaked dwarf and stripe viruses between winter wheat and planted more or less at the same time (they ca n only feed on barley to lowland rice Early planting of lowland rice rice during the milk to hard dough stages) 2) they lack nurseries spreads these viruses but upland ice has remained aestivation sites and 3) weedy fallow areas are limited uninfected The lowands of Asia are dominated by L oratorius

16 IRPS No 123 January 1987

which lays its eggs hign on tile foliage (199) L acuta and L solonlonlesis oviposit at ground level the former on litter and the latter loose on the ground These egg laying habis explain theirtenvironmental prefcrencecs(214) In Japan the postdiapause adults of L chinensis migrate from moun-tainous areas to upland ricefields during floweiing (1 14)

Pentatornid bugs prevail in Latin America (8182 195) and in Africa pentatomid and coreid bugs arc equally cited (6 82) l)amage is often characterstic of the species (5)rcoris ventralis and Mnida spp are the common

pentatomid seed bugs of Philippine upland rice but they prefer lEchinochva spp grass seed to rice Vearaviritda occurs worldwide and hoften more abundant in the uplands(54) We saw few reports of seed bugs on upland rice but see(d bugs apparently opelrate cqually between by land and upland rice Good dispersal is necessary ior species that call feed for only 2-3 xon a crop and must first find those crops at ripening stages

Other pct species feed n flowers In Mato Grosso Brazil (194) and Santa Cruz Bolivia the panicle weevil Aeo-hariuia amllitarsis attacks rice during flowering (68 233)The larva e rice sten horers adutlts feed at the bases of spikelets caIusing empty grains Also inLatin America thecydnid -lkiituttatralusisa seed bug as anad tilt and nymph (194)

A number of beetles ecd on poilen - coccinellids lialrotia Aulac lhora and Monohpl-a (55) - but becatise riCe is self-pollinating and fcrtiliation occurs before pikelets open they pose no threat to yield

IST ECOLOGY

We now look at characteiistics of the life cycles of upland rice insect pests to lea rn hew each plays an adaptive role in upland rice ecology

Life history strategies Orzasativadoes not tolerate drodgh well so upland rice is highly seasonal normally grown in the wettest months of the year Usually rice is present for less than half of the year The rice-fiee fallow poses serious problems of survival to upland rice insects which have evolved att least four mechanisms to ovrcolne tle cvcl ic-l lack of a hostI) polyphagy 2) longevity 3) dormancy and 4) vagility (dispersal)

Poldt)agy lolyphagy is defined as having hosts of at least two botanical families oligophagy of more than one gents and ionophagy of only one ventus (40) - inour case Orvza Most wild rices however are illi tiC and theieiirc are normally far removed from upland rice hahitat othewise more would be alternate hosts for uplanJ iice insects Thte highly mornophagous rice pests shySciriol)haita hiceritlas Nilal)arvaiahiens Iand Nealo--tettiyvircsctws in Asia and Aalirpha secparatella and 1ioj)sis lngicornis inAfrica - occur on upland rice but

are more abundant in lowland culture They can specialize in rice because they are highly vagile and can attack all growth stages By specializing in rice monophagous rice pests can outcompete related but polyphagous speciesIntensive andI extensive lowland rice culture has favored monophagous species Riqwla alhinella (Cramer) the onlymonophagous species in Latin America is rarely reported on upland ice (I1)

Because of its limited temporal and spatial existence upland rice has not favored the evolution of specialized species Clhang(44)shows evidence that upland rice has only recently been cultured by man from the lowlands thus pests have had little time to fully adapt to the crop All but the lowland-adapted insect pest species have alternate planthosts to rice Root pests with limited mobility are highly polyphagous Pests that specialize in one growth stage eg secdling maggots seed bugs are oligophagous or polyphagous

Most alternat hosts of rice pests are annual grasseswhich also are seasonal Upla_rice is also a host but not the preferred one of the most highly polypliagous spcciessoil pests grass hoppers armvwarrs and Vearaviirielda

The d ry season poses igreat obstacle to ani upland riceinsects survival and only sonic perennial plant hosts are suitable food during the off-seasoin Plolyphagy allows upland rice insects to suLrx-ixe during thC rainy season when plants- both annual and perennial -areactivelygrowing and therefore more nutritious

Loiugevitr The ability of aiinsect to live 2-3 mo without undergoing dormancy helps species sich as root-feedingwhite grbs (as larvae) and secd-feeding bugs (as adtilt) to survive unfavorable periods White grubs arc not highly fecund and their life strategy is to improve survivorship of limited progeny Their subterranean habitat hides them from manv natural enemies lBtut root tissue is not highly nutritious so they ltilst eat great quantities and need a largeinsect biomass to digest it Other soil-inhabiting rootshyfeeding insects have short life cycles but are more fecund

Seed pests need greater longcvi vas adults to locate ahost at early grain dcclopment arnd give sedentary offspring time to develop Few groups of upland rice insects live longin active development Polyphagy and longevity only increase their ability to survive during the rainy seasonOther mechanisms are needed to survive the dry season

orantc In the tropics the nonrice season is the dry season while intemperate areas tile off-+season is winter Many upland rice insects undergo dormancy inlsummer (aestivation) or winter (hibernation) Dormancy is simplyinactivity during winters with prolonged temperatures below 10 C insects find shelter arnd cease movement Activity is resu med with warmer t empcera tu res A deep state of irnactivity termed diapauise occurs in sole species that prepare physiologically (havxing low metabolism and convcrting gonad s to fatty fuel reserves) for the unfavorable season (148) They enter diapaUse inanticipation of unshy

IRPS No 123 January 1987 17

favorable weather and resume activity after favorable Armyworms skippers and locusts actively travel hundreds weather begins usually in response to photoperiod or of kilometers in air fronts such as intertropical convergencethrough a delayed response to a stimulus such as heavy zones to descend onto upland rice fields far from their rainfall (237) Not all individuals of a population have the breeding grounds (31) Several generations can build ip on same threshold for environmental responses for entering or alternate hosts in tile grass family before rice is plantedterminating diapause -- this is seen as a mechanism to These large insects are active fliers Smaller insects such as prevent total mortality if a false cue appears such as a thrips aphids and planthoppers passively migrate long sudden warm spell in winter or rain in the middle of the dry distances by the wind season (154) Otherspecies ean readily travel tens of kilometers - stem

Probably more upland rice insects undL go dormancy borers leaffolders leafhoppes cutworms spittle bugs leaf than we realize Candidates are Atherigona spp root beetles seed bugs ants and termites aphids root weevils crickets leaf beetles thrips and seed The least dispersive species are white grabs mealybugSbugs Insects known to entervarious states of aestivation are seedling maggots crickets weevils and gall midgesleptocorisaspp adults (199) Maliarphascparatellalarvae Each species has evolved a unique set of attributes to (9) Scirpophaga innotata larvae (117) Chilo partelhtsv enable it to survive and adapt to the changing uplandlarvae (21) ScirpopThagainccetulas (179) white grub larvae environments Several evolutionary avenues lead to fitness (134) Patangasuccincia adults (6 82) and Schistocerca (Table 2) but there is no reason to believe one set of gregariaeggs (82) attributes is better than another (224)

Insects known to enter hibernation are LodeIphaDstriatellus nymphs (124) Chtio Tvlpressalis larivae (125) DroughtSesaia inferens larvae (146) but not S calavisis(2) Well-drained rainfed upland soils are subject to drought ifIDiaraea saccharalis larvae (75) Scirpophaga- tcerlttas rains do not fall within 2-3 wk Prolonged drought followedlarae(5 lacchrli s lae (75)saSlOhagasshoperseggs (82)and felo pidas athias pOpae (232) by favorable rains stimulates armyworm and locust outshybreaks but mealybtigs root aphids and thrips become

Lcpmotorisa oratorisand L acuta adults aggregate on numerous onlyafter dry spells for two reasons First heavy trees sugarcane or other shady moist sites L paluwanmsis rain normally kills soft-bodied foliage-feeding thrips and confined to Sulu and Palawan Philippines lives on the mealybugs and soil-inhabiting root aphids and root mealyshygrass Brachiariamutica (Forsk) Stapf in the absence of bugs Drought eliminates this source of mortality Second rice The adults can live for 5 mo When aestivating their the rice plant responds to drought by breaking down rate of metabolism drops and they do not feed They will proteins into soluble N compounds which enter the phloem however puncture plant tissues of their aestivation host in and are taken up by these sap-feeding insects (245) Greater search of water This behavior has led to errors of the host nutrition therefore leads to greater survivorship and range of rice bugs (I) Aestivation quickly terminates with fecundity rain probably meaning aestivation is not deep Small rice area

Some stem borers aestivate as mature larvae in rice Except in the highly mechanized large landholdings of stubble or in straw stacked as livestock feed However Latin America upland ricefieldstypically are patchy within moriality increases proportionally with the duration of the a highly diverse flora The small fields result in higher ratios dry season even if the stubble remains undisturbed If the of perimeter to area than with the typically larger lowland stubble is plowed for a crop following rice few stem borers ricefields Upland ricefields themselves may be intercroppedsurvive Rainfall terminates dormancy but it normally with cereals legumes or root crops Mixed intercroppingtakes several weeks for moths to develop and emerge with a wide variety of species also occ in tribal slash-andshy

Last-instar white grub larvae tunnel 1-2 m deep in the soil burn agriculture (50 74) to construct pupal chambers in which they aestivate The Upland ricefields tend to be small because of the labor larvae are very sensitive to dryness so pupal chambers are needed to clear land in slash-and-burn areas and to till land sealed to coierve body moisture Aestivation terminates to minimize weeds Rice requires more tillage than does with the first heavy rains but as in stem borer 3-4 wk pass maize because rice competes less well with weeds Uplandbefore adults develop rice is more a subsistence than a cash crop and small areas

Insects may respond differently in each region to cold (I ha) can feed a family temperature or drought Biotypes or loal populations may Small fields favor some insect pests such as some ant evolve (56) Local populations of Scirpophagaincertulasor species that can forage from more permanent field borders Patangasuccincta may have become adapted to prevailing Trees provide food and mating sites for white grubs (134) as conditions well as aestivation sites for LetTocorisa spp (199) Field

iagiliti The ability to disperse combined with high border grasses plus tIhose within a ricefield provide alternate fecundity orshort life cycles enable some upland rice pests to food for all but the highly monophagous upland rice better exploit the temporarily favorable upland habitat colonizers and a habitat for natural enemies

0 z

Table 2 Life history patterns of upland rice insects or insect groups

Host range Longevity Dormancy Vagility FertilityMonophagy Oligophagy Polyphagy Low Medium High Yes No Low Dispersive Migratory Low Medium High(Oryza spp) (grasses sedges) (angiosperms) (lt3 wk) (gt2 mo) (lt1 kin) (lt50 eggs) (gt500 eggs)

Ants xTermites x x x

Mole crickets x x x xx x xField crickets x xx

White grubs x x x xx xRoot aphids x x xxMealybugs x

x x x xxSeedling maggots x

x x x x xLeaf beetles

x xx xArmyworms x x xx x

Thrips x x xx xChilo spp x x xx xMaliarpha x x x x

xLocusts x x Xx x xGall midges x x xxiVephotettL iirescens x x xx x x xLeprocorisaspp x

x xLeaftolders x x x

Sogatellafurcifera x

xx x x x x x x

00

IRPS No 123 January 1Q87 19

All this appeals to favor insect pest buildup but the notwithstanding the release of stored nutrients in response contrary is true Loevinsohn (138) showed that insect pest to drought that temporarily benefits philoem feeders populations respond exponentially with the proportion of land devoted to rice up to about 75 of the area then the M POR ANCE I I P1A 1) RWI P1St S

rate of response declines In most upland rico areas tie rice crop occupies less than 50i of the area- therefore the Visits to upland ricefields normally reveal 1kw insect pests potential for population buildup is low This relationship Loevinsohn (138) and Locvinsohn ct al (139) compared may not hold for seed pests that can locate small ricefields insect abundance using annual light trap atchcs in upland

Another outcome of the highly diverse flora and phical rainfed lowland and irrigated lowland sites in the Philipshyenvironment of upland rice isthateach upland ricearca has pines Although only one upland site was studied data a unique composition of insect pests showed equal or fewer insects at the upland site than at the

For example in Tanauan Batangas Philippines the rainfcd lowland sites and fewer than at the irrigated sites main upland rice pests are rice leaffolder(RLF) armvworm (Table 3) M Lltiple regression analysis showed that the key Aythntnaseparata WBPH ants Soli psis gentitataand factors explaining insect abundance among a set of cropping white grub Leucopholis irrorata Sixty kilometers away in intensity variables were the number of rice crops grown per Real Quezon the main pests are seedling maggot Athe- year followed by area devoted to rice Table 3 shows that rigona orae flea beetle (Thacitcmnra hasalis thrips the upland rice site had only I rice crop perym rand that rice Stenchaetothrips hiurnmis and rice bug Leptocorisa was planted in only 20Vi of the area Fertiliter and oratorius In a third Philippine upland rice site in Claveria insecticide use had little bea ring on the trend The armters at Misamis Oriental Mindanao the main pests are ants the upland site applied 60 kg N ha higher than at most Solenopsis gemtinata seedling maggot Alieriona orvzae irrigated sites white grubs Holotrichia in)iUaiuoan t and Lucopholiv Insect occurrence at the upland site was omparable to irrorata stem borers Sesamia ifiArelr (hio auriciliusC that at the Cagayan rainfed lowland site parlicularly in 1982stpessallis Scirpophaga incertulas S innoiata and when drought prevented many farmers fron planting(only

4c ona chrrsurapella root aphids Tetraneura niriah- 40(of the rice area eventually was planted that year) M ore dunrinalisand Rhupaosiphunm riahbdminalis lcaffolders favorable weather occurred in 1980 and 1981 which Marasniapataltis and Chaphalocrocis Medinalis army- supported greater rice areas and consequently more insects worm MIthinna separata and rice bug Leltucorisa The most intensive site for insect pests was South Cotabato oratoriusThe great diversity of insect problems is typical Mindanao where farmers plant 23-25 rice crops per year fer upland rice and makes control efforts more difficult with irrigation The Zaragoa Nutva Ecija site also had particularly regarding breeding for insect resistance high pest abundance and at this site the fields were highly

asynchronous Asynchrony was the third most important Low yield potential Iactor in explaining rice insect abundance Most upland rice is grown on low-nutrient and mineral- Rice cropping intensity in upland areas is bound to toxic soils which when combined with eiratic rainfall remain low Only one rainid upland crop is possible per make upland rice a highly riskycrop for the farmerto invest year (Farmers in Claveria Misamis Oriental Mindanao in costly soil amendments and land management Often Philippines tried asecond upland ricecrop i 1984a ycarof upland rice isgrown far from markets making inputs even favorable rains but no harvest was possible b-cause of more expensive Farmers growing a subsistence crop hosi- drought blast and birds) Rice area isaso bound to remain tate to invest in costly inputs to raise yields low because of the high labor and power rcquirelmenlts to

Even having varieties that could double existing yield prepare the land (because of weeds)and because upland rice potential under present management levels would probably wvill not compete with lowland rice as a cash crop Some not prompt many farmers to use purchased insecticide upland rice sites with prolonged rainy seasons howeverare unless tile risk of crop failure were reduced such as it was planted asynchronously such as the Clacra site with irrigation systems in lowland rice The seasonal abundance of Upland rice insect pests

The Tanauan Batangas site is atypical for upland rice attracted toa light trap wasgrapied for crop year 1980-81 in The soils and rainfall pattern are favorable and because of Tanauan FBatangas (Fig I 2) W13P11 was he most its nearness to Manila farmers have cash resources ftm common rice hopper and its population peaked at the sales of vegetables to purchase tertilier They applied an midgrowth stage as it does in lowland rice D-Vac suction average of 60 kg Nha after particle initiation and obtained samples showed an earlier peak than that from a light trap 25-35 tha yields The high fertility isone reason why RLF indicating emigration W13PI usually emigrate after the armyworms and WBPH developed into large numbers vegetative stage Populations in 1980 reached levels that Farmers however used no insecticide on upland rice caused patches of slight yellowing in some fields WBPH although they did on vegetables probably cause hopperburn in Batangas They immigrate

The poor growing conditions fOr rice generally mean year-round as shown by low levels in the light trap BPH poor growth and low fecundity for insect pests fIeding on it however were scarce in the light trap and not collected with

Table 3 Abundance of rice insect pests and factors in rice cropping intensity of Philippine upland rainfed lowland and irrigated lowland locations where light traps were operated daily for atleast I yr -I

0Rainfed Rainfed lowland Irrigated lowlandupland

Pangasinan Cagayan lloilo Santa Maria Nueva Ecija South Cotabato Tanauan Caaringayan Caaringayan Bangag Bangag Bangag Buray

LagunaCale

Batangas Manaoag 1980-81 Santa Motiica 1982-83 Cabanatuan Zaragoza Avancena NaniamanSolana 1981-82 1982-83 Oton

1980-81 1979-80 Oton 1981-82 1981-82 Koronadal Koronadal1980-81 1979-80 1979-80 1983 1983

Insect (kerosene light trapNephotcttixcatch nolyr)virescens 650 1240 500 1800 330 210 3400 1600 160N nigropictus 210 - 360 18400 5500 2900- 140 90 20 - shy 1700 -Reciliadorsalis 710 - 3200 1700180 890 630 140 170 2400 800Nilaparratalugens 130 690 - - 4200 4600300 520 750 180 60 3300 1600 740 310Sogatellafiircifera 1700 31000 13000 8200310 210 3300 610 270 980 1600 6300 -Cyrrorhinus - 12000 6700600 540 3200 290 160 330 120 410Scirpophaga spp 50 410 620 - - 21000 12000150 180 220 270 30 1000Cnaphalocrocismedinalis 70 70 540 610 270 11000 6100200 140 100 6 60 250Chilo suppressalis 0 60 60 1900 300 200- - 30 20 0 6 3 160 -C auricilius - 0 00 - - 20 3 60 20 3 0Sesania inferens 0 - 80 60 3 80 10 51 -Cropping intensity _ -

Rice crops (noyr) 10 10 12 10 10 10 13Rice area (7c) 17 19 20 20 25 2320 85 85 60 70 40 85 85 90Fertilizer (kg Nha) 60 20 85 80 70 7020 0 0 0 30 30Insecticide (no of 40 60 40 300 04 04 0 03 300 13 13 30 50 40 33 33applicationscrop)

__________

IRPS No 123 January 1987 21

Light trapped Insects Drytnd __ 2(notrap-nght) Suctioned Insects (nom )

50 -)o 9 jO40- SOW ffurciWO Light trp

D-Vc suction

20 to 4

20

Egg porasitizahion (M) 75 60

300 SOOampReOftwciffr

45 30shy

15

-insects (notrop-night) 4 Nileporvata Iuen5

Egg parastization()

Light trapped insects (no trop-nght) Suchoned insects (no m 2 )

6 8 Cytorh~nuls hvidipennii 0

4 05 2

50 0 Insects (noltrop-night)40-ui Ipanh pe g

Moohase 1 Seasonal abundance measured by light trap

Rainfall(etod) and D-Vac suction sampler of whitebacked

to ~ hopper Nilaparvata lugens and lea ffolder60)-t Cnaphalocrocis mnedinalis andplthpe g20 L I II I I I parasites and mtrid predator Ortorhjntus livi-Apr May Jun Jul Aug Sep Oct Nov Dec JaIn Feb Mar dipennis set against cropping pattern daily1 9 80o 19 8 1rainfall and moon phase Tanauan Batangas

1980-81

the D- Vac Cyrtorhinus build up on the crop coterminous are collected in light traps during full moon (109)with WBPH buildup Cyrtorhinus disperses during the Because upland rice shares most of the lowland rice pestsrainy season RLF not readily collected in light traps was and those pest species are generally less impotant to uplandprevalent only during the rice crop rice than lowland rice there is a general belief that insect

Nephotettix virescens was more abundant than Nnigro- pests are less important onpicLus in the light trap collections and on the crop GLH upland than lowland rice Arcview of limited data on upland rice shows an averag of population peaked at the reproductive stage as did zigzag 10-21 yield loss from insect pests on a par with that ofleafhopper (ZLH) Crtorhinus tracked the populations of lowland uee determined by the same methodology (132) planthoppers and leafhoppers Both GLH species were (Table 4) McGuire and Crandall (144) made a similarcollected year-round in the light trap indicating immigra- estimate for Central America Cramer (51) howevertion from nearby irrigated rice areas Possibly because of the estimated 35 yieldlow populations loss in South America based on ainsect number did not correlate with similar pest complex - stem borers and seed bugs shymoon phase although reports indicate that more hoppers attacking irrigated rice in North America This estimate did

22 IRPS No 123 January 1987

Light trapped insects D~~nd (no trap-night )

St plht Suctined insects (nom 2

)

D-Voc suction 1520

1o

0 Lih trap 5 ~105 o5

00 Eggparositizotion()5040 NephOtgtha virneens

30 20

I010 A-Insects (notrop -night)

4 Nephatelh nigropccent i

2 o I vV^J - -- ^ VA____ f _

Light trapped insects (no trap-night) Suctioned insects (no rnE

10 -1208 ReCdO doRsol

6 15

4 10 052 iwxALAiamp 0

Light trapped insects (notrap-night) 2Suc oned nsects (nom )

8 CyrtorhmnusI6o nn]10 i0075

4 05

2 0 0

025

2 Seasonal abundance measured by light trap Roinfall (mmd) Moon phase and D-Vac suction sampler of green leafhoppersNephotettix virescens and N ngropicrus and I zigzag leafhopper Recilia dorsalis their egg 60

parasites and micid predator Cyrtorhinus livi-dipennis set against cropping pattern dailyrainfall and moon phase Tanauan Batangas1980-81

40

5 Lf Apr May Jun

1980 Jul Aug Sp Oct Nov I

-aMar

1981

not account for soil pests and was not based on field data in the absence of pests Carbofuran granules are phytotonicAkinsola (I1) cited mainly data from irrigated rice areas in to rice (239) The prevailing hypothesis is that carbofuranAfrica but estimated yield losses of upland rice between 15 prevents soil bacteria from consuming fertilizer The phytoshyand 30 from insects However losses in Thailand over a tonic effect is through greater availability of fertilizer to the4-yr period ranged from I to 13 averaging 5(106) plant Denitrifying bacteria are known to be more active inYield loss data however should be interpreted with aerobic than anaerobic soils therefore it may not be merecaution (184) The Philippine data were all derived from coincidence that the highest recorded yield losses (22-69)trials on farmers fields under farmers normal agronomic occurred in those trials with 12-30 kg basal Nha On themanagement practices except for Bukidnon and Capiz other hand contro with carbofuran granules may not be aswhere researchers used fertilizer Batangas farmers are good as a seed treatment against seed pests and perhapsatypical because ofhigh levels of fertilizer used The trials in yields would have been higher if a systemic seed treatmentBrazil were carried out at an experiment station but insecticide had been used conditions were similar to those of local farmers The 1979 and 1980 trials in Batangas had two full-

Carbofuran granules were used in every trial as a broad- protection treatments one using carbofuran granules andspectrum insecticide applied basally with the seed An the other using bendiocarb WP as a seed treatment Plotsobjection to using insecticides to measure yield losses isthat protected with bendiocarb yielded higher than those proshyinsecticidescandirectlyorindirectlystimulateplantgrowth tected with carbofuran granules The other trials in the

Table 4 Yield losses to insect pests and yield responses to insecticide in upland rice in the Philippines Brazil and Upper Volta

Yield (tha) Yield loss (5) Location Year Cultivar Protected

with Protected

without Check Total Vegetative Reproductive Ripening

Basal N (kgha) Reference

carbofuran carbofuran Protected No with carbofuran

carbofuran

Philippines Pili Camarines Sur Tanauan Batangas

Pangantucan Bukidnon

Dumarao t-apiz

BrazilGoias Minas Ge~ais

Upper Volta6

1975 1974 1974 1975 1975 1976 1977 1978

1979

1980

1979 1980 1979

1977 1978 1979

Bursiging puti Kinanda Dagge Kinanda Dagge Dagge Dagge Dagge UPL Ri5 Dagge UPL Ri5 Dagge UPL RiS UPL Ri5 UPL Ri5 UPL RiS

IAC47 IAC47 IAC47

14 33 11 36 35 31 30 27 37 26 40 30 40 42 46 35

19 21 09

--

-34 24 24 27 --

29 43 31 42 -

47 25

---

10 28 05 27 27 30 26 29 33 28 33 29 43 23 36 11

15 15 06

29 15 55 25 23

3 13 -7 11

-7 18

3 -7 45 22 69

24 28 35

--

21 -11 -20

4 -

-3

23 7

-2 -23 56

-

-

-__0

6 --

-

-23 14

5 0 8 2

22

-

--

0

-

--18 11

3 -10

4 10

7

--7 -

14 -

-1 6

10 -5

8 9 7

-

--

0 0 0 0 0 0 0 0 0 0 0

30 30 30

12 12 12

99 99

100 100 101 101 102 102 102 102 103 103 104 104 104

66 66 66

Farako-Ba

Mean

1977 1978

IRAT10 IRATIO

38 66

--

40 43

-5 35

21

-

10

--

--

--

- 242 243

Z

tJ

-w

24 IRPS No 123 January 1987

Philippines which had two treatments offering protection In the 9 Philippine trials where yield losses were measuredwith and without carbofuran granules relied on further by growth stage 10 of the yield loss (21) occurred at thecontrol with foliar spray and not on seed dressings The vegetative stage and 517 occurred at each of the reproshycarbofuran-protected plots had an 18yield los compared ductive and ripening stages Ibis result is similar to thewith only 10 with foliar sprayS This yield difference is pattern of yield loss in lowland rice measured by successiveprobably due to differences in protection against sown-seed treatments where the crop is unprotected at each growthpests (foliar spray do not control sown-seed pests) rather stage but proticted during the other growth stagesthan phytotonic effects became basal fertilizer wa used in However the insect complexes r sponsible for vegetativeonly 2 of the 10 comparisons stage yield losses in upland rice are entirely diftfrent than in

The recorded yield losses were not related to yield lowland rice The key vegetative stage pests in lowland ricepotential (Fig 3) One might expect higheryielding crops to arc aquatic whereas those in upland rice are sown--seedhave higher percentage of yield losses but this ws not tile (sown in soil) root and seedling pests This dift rence maycase One reason is that insect damage exacerbates plant explain why agricul torists have overlooked tile ilmportanccinjury from drought stress low yielding fields from d ought of upland rice insect pests Many reports of upland rice pestsstress would show disproportionate differences between in Asia ha Ve focused on major lowland species (46 108)insect-protected and -unprotected treatments Shoot and It isalso surprising that tei vegctative stage of upland rice root pests would be more responsible for losses from would record the highest yield losses becatuseseeral studiesdrought stress than sown-scd pests A second reason have shown that upland rice can readily recover from foliageconcerns seed pests which would have a greater impact on reinoval during that stage (117 152 165 188) Upland ricelow tillering low yielding varieties such as )aggc Low poscsses a hitgh obilitV to compensate for ca rly loss oftillering varieties nnot fill in the space created by removal foliage and will even be stinulated by foliage removal toof seeds oweer 1) R i 5 is high yielding because it tillersP1 produce a higher viel than plants ithout any foliageactively which can close the canopy to provide greater weed removed (63 192 231) [ie reason for this apparentcontrol discrepancy hovce is that yield loss at the vegetative

stage does not octUr frol leat arca loss but fron pestsYield loss () removing sown seed and feeding on devcloping tillers and roots -- damage that cannot be readily conpLtnsatCd for

70 - Measuring yield losses in upland rice cnvironrnents is further complicated by a report from Brazil that greater yield losses occurred in response to better insect control60 - =-09 as Insecticide seed treatments resuled in denser plant stands

(n=21) which in turn created a selectively more favorable environshyment for blast (69) This resuIt was parlticurla rlxexacerbated50 when cirbofuran granules were used in seed furrows Aside

ffrom a greater plant stand ca rbofuran iay have auigmented the N lecl and stimulated blast If so ca rbofuran should be40 - replaced with another insecticide or the seeding or fertilizerI rates decreased

Other methods to determine the impact of upland rice30 shy pests wvcrc to interview farners or to develop single speciesbull bull correlations of population levels and yield loss Feneira (66)2 NIM)st (75(1) Golas farmers reporteddid both in Brazil

20- insect losses in ufpland rice and 251 said those lossesS avcragcd 37(1 [his result is consistent with tie yield losses 0 0 measured by field trials in the same statc(Table4) For every

I0 4 of seedlings removC d by leaf-cutting ants a I 7yield loss is predicted Similarly Ferreira (66) found that ai

0 - average of 53 thrips per panicle before panicle initiation meant 3 times more unfillh d grains

-IO0 I 1 1 I I-101 0 2 3 4 5 6 7

Yield (tho) C)N [ROt IM FI IODS 3 Correlation between upland rice yield and yield lols caused by Upland rice insect pests may be controlled by culturalinsects measured in 21 trials in the Philippines Ivory Coast andBrazil genetic biological and chemical methods

IRPS No 123 January 1987 25

Cultural control dry season plowing will have little effect on them Plowing Crop husbandry to reduce upland insect pest populations when the soil is wet from the first monsoon rains will expose can be classified into practices effective in a single field and soil insects that tunnel close to the soil surface to predatory those effective only at the community level Cultural birds chickens dogs and even man (field crickets nole controls should come first in a pest management program crickets and white grubs a re delicacies in man tra ditional They have broad stable effects as pests have little pos- diets) sibility of overcoming them through selection of biotyps The greatest degree i1 inIscct control fron tillage comes (138) On the other hand some cultu ral controls work only from plowing soon after rice ha rvest when most soil insects with high labor or power inputs and connunity-wilt lie close to the soil surface anrd stern borers are in the rice methods require coordination of iany farniters stuble Whitegrubsare mature and tillage exposesthem to

Pantin time The luxury of being able to shift the time of predators Tirnelv control will prtec the crop following pla ntring upland rice is open onl hr la rniers in areas with a rice and lowr the populationIfor succeedinig years Plowingprolonged rainifall pa ttern lI copping intensity or underthe ice stulble helps it decompose killing sten borer rnechanied land preparatiot Gcnerally planlting as soon larvae inside as the rFiinv season begins will lower populatiois of 1110st [requent tillage destroys ait nests bit not termites insect pests Insect populationsarelo aftcra dlvseoll or Tillage and clean culture howes er rcliove food for winter fallow delayed plaiting lets them build tip on termites which then mtay attack a young rice crop Scarab alIternate hosts that gi vigorotsly with the first seasonial beetles also prefer to miposit in eceitly tilled fields Noshyratiris This first flush iof weds cros lItxuttriartlv with iceCult is dibbling faors the survival oftillace Clre Such rineralied N released during the dr season aid made stils i aels tnd insects thtt pttpate or lay eggs- in the soil

available to the plants with the rains It iliso igrtmitcally Zero tilla hio eerl conserves predators ofthe GIH (37) advanttgeous for the rice ciop to tap this trlLItlaI fertility nt it mcttoi lo lessen the impact o0 soil insects and grow vigorotsly to compete with wrus and to tolerate brotdcasting seed or planting in ftltrrows is prefc ired over pest infestation dihlirig in hills Crickets are large enotlgh to eat seedlings in

Larly plantngs terd to escape seedling maggolts Upland a hill atrd tillering normnally ca iriot fill in tlhe missing space rice in Bitangas is planted in Ma and ine itli the Iirst Oftcn dibbll rd is not well covered and iseasily found by rains Fairners are highly mtotiated to plant early tocscape foraging insets ioIdents and birds drlitght and typlotns ricar harct Arn Atrgtst planting Plan uit sii Incrcatsini sced density protects tile cropwould be evcely attacked by It riim orivzo At tti againrst seed and seelling pests especially ants particularly other Philippiie tipland sites Rea l Qteot antd (laelit if the crop will germinate and ermerge quickly M isainis Oriental seedlig mnaggots arnrige ric boe t se hr trftinll itI India rice intcrclopped witi cotton or farnes have a longer growing season and plnit 2 rin or pigeon pci had lower 6IIt and WBIlII poptrlations than mote after the ortset of the rainy season liaaiangas ftrmners rice alone (211) NfailC arid Urplald ir_ 2re coilltori in the plant ca rly becia use ofa shorter raiinfall period arnd the desi re Philippines he chatngC in pest status folii iniercropping is to lliarvet before the typhoor sea son peaks ltighly locatioll-specific (133) cmianarid the net elfc be either

Early plantings cornbind with early-iatiuring varictics nil bcnelficial ortdetriential (189) For example BIatangias provide higher crp tolerartce for white grubs Aestiatior farmers intercrip Itro in their ipland ricefields A liornshyterminates with the first rains but 5-6 moare ineeded for the nrii iotd-Ahis i (I iiiacius) which norlially feeds resting larvae to develop into beetles emerge and lay eggs on taro ill feed or miarbv rice plants Intercropping and for larvae to reach the danriigii g ird-instar stage probably would not afft0 seed pests A crluanion crop (134 In lataigas this normually iseiough tiite for harvest planted ith rice and haing gleater tolerarnce for root

It Brazil howecver delayed planting is a suggested dariage or being toxic nayact isi trapcrop to root-feeditig cultural control for the lesser cornstalk borer and insect pests Acial-fei i ig pests voultId be most affected by pests favored l drought (66 195) Cropping internsity in intrer ropping Shoot aphids leaf beetles sten borers and Brail is low and delayed plraing places the rice crop in the hopplers would be likely ca ntidates to note effcts On the niost stable rainfall period reducing likelihood old rought othlt halnd the corlipalior crop stch~ sitniaie Iiiv be more

In rilniy arca s rinmirsare roi atl to plant other crops benehited than rice from initecropping ( 137) before rice and althNgh it woultl be desirabh to avoid 11echi Weeuling luring the first month after clop infestations by planting rice cuatl lariners lack the labor cstablishent will force pcts such is anyvworris which and time to do so ol a low priority crp prefer grassy cetds ontor tle vorling rice Soil iampsecs will

77age Uplild rice Stils ae light textured and Call be mvnye to the rice crop Cleani citrltirc also lay lore territes plowed duiring the d rv seasor notl only to grin time for to attack I u cetincti inrice ceds i ay he le optinal planting with lie fitst Inilsoon riiains but also to desiccate solition 13) soil insects arid weeds such is ntsedge that foim rhizomes Irtlizr Itneatsing N Iciiliier oin hlwland rice has Htowever becta use most soil insects lie below die plow layer fAtwvored Irigher populains of pliithoppeiv Imlhoppers

26 IRPS No 123 January 1987

and leaffolde rs However because of the need to temper fertilizer use on upland rice o prevent blast fIrtilizer rates will be lower than is comonia r for lowlad rice causing relatixely less hopper and IaIfTfoide r population buildup However inupland rice areas without a history of blast such Is Batangas farmers apply high rates of N and conscqtrently the crop has high infcstation levels of WBPI]and RIF The fhrmrcrs leec insect buildup by splitting the fertili11r in0to two to three applicatiois bit do not applV fertilizer at the vegetati estage because this prodrces a tall crop which illeasily lodge during typhoons Greater crop fertility leads to greater insect survival (better nutrition) larger insects (to cat no re foliage) arid greater fec titd t (to lav nmre eggs) (128 145 171) Not all insect species respondequally hovr Increasing rates (if N l anrid K increased Dialraa s-acchralis and (iaeiurema flea beetle but reduced thrips (66) Zn rcdLuced deadheart density from the lesser cornIstalk borer

Fnldingi Although most Upland rice areas do nIot hawxcaccess to wa ter floodiig the fields is rRrrnriierdcd in areas of Brazil or root bug and white gib control ( 195)

Crop rotatioti Tillage before planting a crop after rc harvest will control stern boreis and other pests renain n the stubble arid will unearth soil pests Planting a non-gratriiaceous crop after rice is recommncrded for termite control (66) Crop rotation is less nttdcd in upland rice culture than in lowlands because atcrop-free period is assured even after harvest of crops following rice

t1ulchilt iryl plastic irli irig designed for weedcoitrol ii upland rice inJapan resulted in ntr highanri liy infestation of (ihilo sulytresvais (93 9-1 95) NIulched Upland rice grew more luxuriantlv and attracted more (Avipositing mothis than u lched rice Also the rice plant and stein borers imatoured 10rC quickly and a partialgeneratiori af vouing ista iNoverwintcred in the stubbleLotili--malitrt varieti Quickly maturing crops reduce the tmbCr of pCst generations that Calt build up Rapid crop establishment olan ca rl v-natu ring vaitvty if)pa 11icLa r will lessen white gib and stem borer damiage White grub larvae intle last iistar are highly destIruCtive so aii early-iiatri ng variety will escape serious root loss Stem borers build 1ip slowlyx in rice to ecormic abundant in a late-Maturing crop anrd caiuse whi tlieads Numibers of somespecies increase exponeitially with each succeeding genera-ti so a xarietyexhat atures I trowh earier thian another will hac less insect damage Early-naturing vaiicties however will not lessen seed pest daiage either ii the soil or oina standing crop

Strwhoi hAIJttam Inrscts tcd irig on thlie aerial portions of rice plats disperse from carly to late plantedfields AS iiiibCirs increase exponentially with eachinsect

gtscnrattm e eiglbrig f rmc~s1(uil upp wh rc generation pests build tIP where neighboring fihrTC e9r1sstagger their plantings II Batangas farmers plant uplandrice within I iro use 120-d s-arieties and plant at the start of the rainy season Ini Misarnis Oriental laricrs stagger their

plantings up to 3 mo apart and use a set of atieties that mature in 4-7 mo Insect pest danage particularly from stem borers is greater in Misarmis Oriental Also seedling maggots damage rice there because rainfall favors frequent planting of maic In Batangas rice escapes seedling maggots bCca usC of distinct drs season and synchron0us early planting with the onset of ains

Therefore combinations of cultural practices ea rlyplanting synchronous planing crop rotation and earlyshymaturing vaneties protect the rice crop against most insect pests

Ilmant resistance Beca tse upland rice has few specialized insect pests and many insect types attack the crop worldwide regional breeding and strong internatiornal cooperation are required

Sown-sced pests arc economically controlled by insectishycide seed treatment and should the relore receive lowxpriori ty inbreeding objectives

Root pests are normally cost ly to control with insecticides but finding resista t sonres mayvbe di fficult Tanaka (229) for exa -ple fi led to find varieties resistant to root aphids Selecting for large root bioniass perhaps should be the strategy for pests sLuch as Whlite grubs that renliove roots Fainter(174) howcvcr cites varietiesof crops tolerant ofor resistant t(i root pests Some sorghun varieties are high in cyanide so rice sorghun might incorporate broadshyspcctruri chemical resistance

Seedling vigor and drought tolerance are high prioritybreeding objectives so tupland rices should have large r root systernsand be able to tolerate ligherlevels of root loss Sonie root loss inolder plants is beneficial if it stimulates new root developnrut and therefore crihances intake of soil nutrients (53) The rice seedling inaggot rtiLotia spp may be

controlled thirough resistant aittics A hi rgc breeding effort on sorghniu has had fair success inideveloping arreties esrstait to Asoccata Rodani (251) A field trial comparing 10 upland ricc varieties showed differences aniong varieties va ring from II to 4(- damaged tilleis (215) Hovver resistarce to 1Iill-0r gtia species may rot cross ox-rti otier

Shiraki (210) reported that upland i-ice stens wcrc harder than lows-land rice stemsperhaps becaIusc (f a higher silicoricontent and thus were more resistant to yellow stern borer hc tortia Iow tillering (f uplaid iice xarneties prodtres a

higher percentage of infested tillers than would occor in higher tillering lowland iice under a siiilar egg density The corollary therefore is to select for high tilleriig upland rices

Stemii bor amcstheh main targes dobrecders inAfrica (571 204 )but nticl oifth e sc reen irig is dotie unde r lo wland co d t osrm s wih A h r~ o l pe i s n acorditions (11 96 218) As with lthrinrnwspecies mayhave to be dealt with independently however initial results show cross resistance between African and Asian species(92)

In Brazil screening Ior insect resistance focttses oil lou I pest problems Varieties tolerant of lasmolwalpus ligi-sellus and resistant to )iatraea saccharalis have been identified (70 153) Current work also emphasies 7iraca limativetntris and Thmialspis (=Iosis) flavouicta (66) These insect pests Ilt onlV citise IIigh yield losses tihey also ire difficult to control chemically ofr by other ilealls

ManV upland rice vaieties aic early maturing and are resistant to pests such as gall midge (I Xl ) Early mIturity is often a highly desirable trait to esuilpe pest buildup but it is not true resista nce bca usc ilthose va reties are plinted late they will be itsdamaged as ain early planted susc ible variety

P~erhaps tile cosiiiopolitall of all upland rice pestsimost are the sced pests (ripenling stagc) which pose a particular problem beta use scuds re ised IsIO for hitnit4ans Ayltoxic substatice introd tcld geticticllyilmay also be toxic to nal iniless it caln he placed old in thfe IdhuffSelectmi fo hairiness or lotng wns kill mIlct sthilrohleiiis dutilig threshing pa rtictfarl maitial threshinfl [hireshers ill complain of itcfimin atid itteascd (fust levels

Attempts itIRRI to fitid tcistatncc to the rice bug plorisa have ilotbeCnl prmitisilg lIIetIattonlids iorc

through the lenilna or palca hillhairs sedstVIor seeds wilh awns that deter bugs promote complaints from threshers Seed bugs liptocorisa and Stcnocoris enter the seed through the opening between tile lemna and palC but varieties with narrow openings are diifficult to mill

Breeding priorities should concentrate ott inotCI gruLtLIps that are most difficult to c nttrol by other meains Steil) horers appear to be the most widespread group that sigiiificantly dalag s rice aind is hard to manage by other Ilnca us

Biological control Very fe stutdic of mitatt~l eneties of rice pests in upland envirollnletits wee in tile wefouiid literature Therefore have not compiled a list of species Ve present a list of natural enemies iOtid in upland ricefields tuic latialln Batangas for some conitno iOldiar pest species ([itble 5)

Muich ofl thc lollowinle itfor ittiol cotlliCs rol Stutdies in the lowlatnds or roi crops otithcr thlin tice

Qu(ailtr o 11ll t11t( Nat trrtf lii hcoeits ight Ile used agaitist ile social insects tilt and termites Sollie predator specice live ill theirf nests n ilnic theirnllt ip-pearanIlCe aMRidbehavior IJtrouMdCing a mote aggressise ant species to displace pest ant codonies is a possibility (169) Patihogens as potential natural encnies have reccived little atteition (I13) Natural ncnuics asa group however hac not been toIl iporta it in Iegulating aintshown be and ternite numbers

The nocturnal behavior aid subteitmien habitat of lole crickets and field crickets protect them from many ntllral enemies There are only limited records of natural cnelics for these groups of pests some pathogens (87) but mainly

IRPS No 123 JattuIny I 271987

parasites - scelionids on eggsand sphecids on ilymphs and adults Sphecid wasps Larra carutmaria (SImitI hI L huzwuensis Rohwer and lsaltiuhta Willi ams speciali in mole crickets pa ralyzing them nd dragging the ninto thcir nests as food lot their voting ()tier sphCcid species Lins aurulenoa (Iabricius) lot maiac (Ashmead ) 1 lhshylssellaus (Smith) and Ml lal)oriosu (Snithf specialie in field clickets

White grUbsO the other hand hae Icti scriuns enuigh to have been extensively studied for cotttroi 1W nati al enemies( 112 140 141) Scoliid parasites C(ajsoMcispp have beetn intrtduced itto tilePhilippites for control of Ielxl ]udisiro la The rcsearchcts w Intttrdced themle during ill era claimed that the white gttionthreak as controlled (142) howevcr it isnormal for pest epidemics to subside and there was no diect evidence to indictte that tfe parasite was responsible A tmore promisitg titctllod might he to identify iso ltcttass prt~f tcartd apply hiactelaln fingal pathogens to tile soil Ihe lalattese beetle in North America has hcci coitrolled hv tite conmnleciall aailable hactcrial prcpa ration itacihlpu pav ofr milksdiscas (71) Spote prtyefott the soil itloted ilrtier reitiain actic fr Veas

Root aphid sardtnctly bugs readilslic tilib to predatots and parlsites once their tetndine ants are controlled The larvac and adtilts of a coccinellid beetle 1Scynms sp prey on llriadlouais in slash and burn upland rice Baits treated with insecticide may be used to control tending ants by killing their ourig (244)

Root-tkcditig blugs false wireoris wireorms and root wccils aic r ofsuch little ifilpoltatice that thenomttll role of liatural Cticniics hias tiot beei asscssed A possibility hoevrl tiglt he tit ItaIIitic niaCIMtodcs lintt1g11en Itese soil-itlhlfbitine mCtitodcs W5uhtfhave to be imiass Produiced

large miltibcr of niatltrll cnemics ircknown ft CltIoriS IC IpblCeii is thtt clttvortnlls ofonlie the crop soott alter land ptel)tritiOti a1tithe bcgit iniig of the raittv

Citsoln hfien tiatllral leniiiy populations aric lo Colltloil WOtuld iii ole mass prodtticing andirclcaslng key spccies

le egg stage is the fpirt (olthe Cedlitgt Ialot life cycle tI hich to f)ctis ftor tittlril elctit cottntrol hplraites o1 pleditors ttsit elleggs highly expoIsYCl I Sgg pailsites or predators stolld probahly e ittass andtbe produced released Littlh i toruialioll is a ailable ott what species to trV o ho to Iniss podLice thiCIi A etiloplhid wasp Icmihlaol sp effctiicly regushy

lates the leatf lcloalapolm2a spioanoiiitr heat andc rice antid should be collsctvcd

leaf beetles whCllose e am] eggs ate laid in the grutindhirO ould be motC difficutlt to control by niatulral enelV Mnll ipuhltiOn But species sUici IS c_gsthispid beetles ith

and larVle oilleascs ate vtilcratble to iparaites and pledators

Natural enemisc of tlirips otl rice are little known

28 IRPS No 123 January 1987

Table S Parasites recorded from upland rice insect pests in Tanauan Batangas Philippines 1977-84

Order Family Species Host stage

I lymttenoptera Braconidae Leaffolders Cnaphalocrocis Marasinia spy

Cardiochilesphilipplinensis Ashmead a Larva

Cotesia angustillasis(Gahan)Cotesia or cypris (Nixon)

Larva Larva

Cotesia or taeniaticornis(Nixon) Larva Cilonusinotakatae Munakata Chelonits SppAlacrocetrts philippinensis (Ashtmead)

Egg to larva Egg to larva Larva

Iclineumonidac Orgilts spp Isclutojppa hireator(Fabricius)

Larva Larva

Itoplectis naraigav(Ashmcad) Larva Tenelhtcha philippiensis (Ashmead) Larva Tenelchha stangli (Ashmead) a

irichomma cnaphalocrosis Uchida Larva Larva

Bethylidac Elasmidac

Goniozms or trianguli r Kieffer lasmtusalhopictus Crawford

Larva Larva

Diptera

Encyrtidac Chalcididac

Tachinidac

Elasmtus spp Copidosomopsisnacohiae (Fidy) Brach nteria evcarinata Gahan Biachyieria lasus (Walker) Ztgohothria ciliata (Wulp)

Larva Egg to larva Pupa Pupa Larva

Argyrophylx nigrotihialis(Blaranov) Larva

l-ymenoptera Braconidac Yellow stern borer Scirpopiaga incertulas

Braco chinensisSzeplegeti Larva CIthelotis spp Chelonus niunakatae Munakata Cotesi flaripes Cameron Stenoracon nicerilli (Bingham) a

Larva Egg to larva Larva Larva

Ichneumonidac Tropobracon schoenobii (Viereck) Amatroinorpiaaccepta metathoracica(Ashmiead)Frihorus sinicus (I lolmgren)

Larva Larva Larva

Ischnojoppaluteator (Fabricius) Larva Isotima nr dammeratiti Rohwer Larva Tenelucha philippinensis (Ashmead) a

Larva Tenehicha stangli (Ashmcad)Trichomma enaphalocrosis Uchida

Larva Lara

Chalcididac Eulophidac Trichogranmmatidae

Xanthopimnpla stemmator (Thunberg) Charops hrachYpterumn Gupta and Malteswary Brachtm eria sppTetrasticttus schoenohii Ferriere Trichorigranm chihunis Ishi

Larva to pupa Larva Pupa Egg Egg

Diptera

Scelionidac Pteromalidae Tachinidae

Tt ichograintnta aponiicuitt (Ashniead) Telenomits rowani (Gahan) a

Trichoinalopsis apattelctena (Crawford) Jeirhaea spp

Egg Fgg Egg pupa Larva

Z4gobothria ciliata (Wulp) Larva

Ilymenoptera lecneumonirlac Gol fringed borer (hilo auricilius

Trichomm enoptalocrosisUchida Larva

Ieioenoptera Braconidae

Aantholimlh steiniator (Thtinberg) a

Pink stet Ioret Sesamia inferens StoInacoil nicrilhi (Bingh am)

Loarva to pupa

larva

Diptera Ilyinenoptera

Armvtnyorms Mythimna separata Spodoptera mauritia acronyetoidesTachinidae Zvgohothriaatropiora( Robinea i -Desvoid y)Eulophidae It-phctrtiscitapadoe (Ashimead)

Larva Larva

(halcididae Braconidac

Brachpin ria spip (oesia spp

Larva larva

IHyntenoptera I)iptera

Rice skipper Ielop ida (halcididae Tachinidae

math ias aidgreen iortedci ciliarNIelan itis eda Brat lhincria sp iii matuqitatt Cameron Irgyrolhirlaxi trotlhiais(marumtv)a1

ismtne Larva Larva

Diptera Ilymenoptera

Tachnidae lelineutionidae

fuown se tilo per Mois frugalisIrgtrophrlax 1igrothiahiv 3 a rat ov) anthopimpa pmnctata (Iabricius)

Larva Larva to pupa

aMost dominant

IRPS No 123 January 1987 29

Normally predatory thrips prey on herbivorous thrips strepsiptera on both Spiders dominate tilenymphaladultNatural enemies of stem bugs --Scolinoplara and Tibraa predators Spittle bugs move to rice from pasture grasses - are in order of importance egg par~isites and predators and biocontrol would be successful only ifcarried out before fungal pathogens and nymphal adult parasites Egg tiley immigrate into ricefields parasites could either be introduced fron other areas or Biological control tactics for hemipterous seed bugs mass produced and released ould follow those outlined for stein bugs

Armyworms are normally lwld in hcck by the activities Coml)arison of environnewtns Natural enemies are of egg and larval parasites When these natural enemies fail perhaps faced with even greaterconstraints than insect pestsusually because of drought armyworns become epidemic to survive ind reproduce in upland environments TheyIt may be worthwhile therefore to release parasites during have adapted ani survived by having wide host ranges the the rainy months following a drought Virus diseases of ability to aestivate or hibernate over unfavorable seasonslarvae would beanotheravnue to exploreforarnyworms and the ability to disperse or by other mechanisms

Parasites have ben traditionallyconsidered for biolegical Comparing the most dominant and therefore the most control attempts against stein borers usually by rekasing adapted species of natural enemies collected in various exotic species( 166) This approach has not met with success environments in the Philippines we can understand how and new avenues should be explored The role of predators different rice environments are arid pathogens is little inderstood and deserves greater Anagqrms olptahiisa mymarid planthopper egg parasiteattention Egg parasitism rates are normally reasonably is cquallyadapted to all Philippine rice environments (Tablehigh but should be supplemented with effective egg 6) It is selective for planthoppers and even parasitizes eggspredators Orthopterans are potential egg predators of the n-iize planthopper Peregrinitsmaidis itt upland areas Metioche vittaticollis (Sta ) Anavipha nifiwnnis (Saus- Alavohis also parasitizes maize planthopper eggs but is sure) and Conocephahs hongipnni (de i-laan) feed on most prevalent on BPH in the irrigated wetlands (onashystem borer eggs etioche and Anaxil)ha specialize in eggs tocerus spp like A optalhilis on BPH and WBPI Iattacks with no hair such as Chilo spp The omnivorous (ono- the eggs of GLH in ell rie environments It is also found cephahsspecializes in eggs cov red with hairs it is a proven parasitizing the eggs ef the white rice licahopper (oimna egg predator of Scirlouphaga spp but has the discouraging spetra Oirosita naias a trichogrammatid egg parasitehabit of eating rice guin (65) attacks BIPH and WBPH in the wetlands and has no other

Grasshoppers have egg (scelionid) and nymphalad ult host A related species 0 aesopi specializes in G1-H in (nemestrinid tachinid and sarcophagid) v irasites Among wetland environments Dryinid strepsipt-an and pipunshypathogens protozoans have been recorded most frequently culid nymphal and adult parasites of planthoppers and Control of locust species by natural enemies could con- leafhoppers also specialize either in hopper species or centrate on their habitual breeding areas environments Among the dryinids Pseuhdgonatoipsflavi-

Leaffolders have rich complexes of natural enemies some feimur prefers the lowlands and BPH while P nuch has no adapted to upland rice (22 27) ranging from egg predators environmental prefirences but is adapted only to WBPH (gryllids coccinellids) to larval parasites (braconids ichncu- The main d ryinid species on GLH aptogonatop)usspp is monids) to larval prcdators (ants and carabids) to larval most dominant in rainfed lowlands and also attacks other pathogeiis (vira and fungal diseases) Efforts to augment leafhoppers within Amnrasca Cicadulinaand Bachaltuathese natural enemies should focus on the egg predators Strepsiptera are more dominant on plarithoppers than (rearing releasing)and viral or fungal pathogens (culturing lealhoppers The Elenchus species in the Philippines attack spraying) all species of planthoppers within the genera Sogawilla

Parasitization of the large conspicuous larvae of Sogatodels NilaparsataHarualia and Opiconsiva Elenshyskippers green horned caterpillars and other polyphagous cusrvasuntaisuiis adapted to the major rice environments Lepidoptera by tachinids and chalcids is normally low ELg but prefers BPH in rainfed arid WBPIH in wetland environshyparasites and predators are perhaps the key to their control ments Ilalictoph(utsmtnroeiand HVpectrus attack G LH

Gall midges are parasitized by pterormalids and platy- and COfimravcctra respectivelygasterids that aestivate within their larval host between Pipunculids are mainly parasitic oii lealhoppers Each seasons Shoot aphids are preyed upon by coccinellids species has a unique environmental preference Tomosshysyrphids and chrysopids varireIa oraetoraprefers Ne7 otetltix nigropictrs and

Planthoppers arid leafhoppers have egg parasites and Pilunwulhtsjavanensisattacks Dehoceplhahls spp which predators as well as nymphialad nlt parasitesand predators are more abundant in upland than irrigated lowland their rich complex of natural enemies includes fungal environments T suhvirescensattacks Nephotettix virescens pathogens The principal egg parasites are myrnarids and and prefers the lowlands Pilpuiulus nttillattls is an trichogramniatids Egg predators are norinally mirid plant upland rice parasite arid attacks only Nephotetix speciesbugs that suck out the yolk Nymphal ad nIt parasites are More than 17 recorded species of leaffolder parasites in dryinids on planthoppers pipunculids on lealhoppers and the Philippines attack species of C(naplwocrocis and

30 IRPS No 123 January 1987

Table 6 Dominant natural enemies of rice pests in upland rainfed lowland and irrigated lowland rice environments of the Philippines

Natural enemy

Host stage Species

Brown planthop-Parasite Egg

MymaridaeAnagrus optabilis(Perkins) Anagrusflaveolus Waterhouse

TrichogrammatidacOligosita naiasGirault

NymphAdult Dryinidac

Pseudogotatopuslaiifemttr Esaki et Hashimoto Elenchidae

Elenchus yasurnatsui Kifune and Hirashima

Environment

Upand Rainfed lowland Irrigated lowland

Ailaparata lugens

x x X x

x

x x

x X X Whitebacked planthoppers Sogatella furcifera and Sogatodes pusanusEgg

MymaridaeAnagrus optabilis(Perkins)

TrichogrammatidacOl~gositanaias Girault

NymphAdult Dryinidae

Pseudogonatopusnudus Perkins Elenchidae

Elenchus yasutatsui Kifune et Hirashima

Parasite Egg

MymaridaeGonatocerus spp

TrichogrammatidaeOligositaaesopi Girault

NymphAdult Pipunculidac

Tomosvaryella subviresccns Loew Tomosvaryella oryzaetora (Koizumi) Pipunclusmutillatus (de Meijere)

DryinidaeIlaplogonatopusspp

Egg Copidosomopsisnacoleiae (Eady)

Larva BraLonidae

Cardiochiles philipphnensisAshnead lchneumonidac

Trichiomnia cnaphalocrosisUchida Temelucha stangli (Ashmead)

Pupa Chalcididae

BrachymeriaexcarinataGahan Brachymerialasus (Walker) Brachymeria nr tachardiaegroup

Parasite Egg

Scelionidae Telenomus rowani (Gahan)

EulophidacTettasticusschoenobhi Ferriere

Larva Braconidae

Stenobraconnicevillei (Bingham) Iclhncurnonidac

Temelucha phiiippinensis(Ashinead)Temelucha stangli (Ashmead)

x x x

x x

X x X

x x Green leafhopper Nephotettix virescens

X x x

x x

X X X X

X

X Rice leaffolders CnaphalocrocisMarasmia

x X x

X

X X

x x x x x X x x x

Yellow stem borer Scirpophaga incertulas

X X x

x

x

x X XAinauromorphaaccepta metathoracica(Ashmead) x

continued on next page

IRPS No 123 January 1987 31

Table 6 cntinued

Natural enemy Environment

Host stage Species Upland Rainfed lowland Irrigated lowland

Larva-pupa Gold fringed borer Chilo attriciiuslchneumonidae

Xanthopimpla stemmator (Thunberg) x aPupa x

Pteromalidae Trichoinalopsis apanteloctena (Crawford) ax

Pink stem borerb Sesamnia inferensParasite

Larva Braconidae

Stenobracon nicevillei (Bingham) x b b Armyworms Mvthinna SpodopteraLarva

Tachinidac Zygobothriaarroptvora (Robineau-Desvoidy) x x

Braconidac Cotesia spp

X Skipper Pelopidas mathias and green horned caterpillar Melanitis leda ismeneF-gg

TrichogramrnatidacTrichogrannaspp x

Scelionidae Undetermined Scelionid

x Larva

Tachinidae Argyroph lax nitrotibialis (Baranov) X x

Pteromalidae Trichomnalopsis apanteloctena (Crawford)

X Predator

CoenagrionidaeAgriocnenis femina femina Brauer x xIschnura senegalensis tambur

Miridae CrtorhinsliidipennisReuter x x

Nabidac Stcnonabis tagalicus (Stal) x

VeliidaeAlicrovelia douglasi atrolineata (Bergroth) x

Coccinellidae Micraspis crocea (Mulsant) X x x

Vespidae Eumenes campaniformis (Fabricius) x

Formicidac Diaecamna spp xOdontoponera transversa (F Smith) x

Carabidae Chaenius spp OphioneaW- aJasciataSchmidt-Goebel

x

Ophionea ishii ishii Habu x x X

Gryllidav Anaxipta longipennis (Saussur-) x xMetioche ittaticollis (Sial) x x x

Tettigoniidae Conocephalus longipennis (de Haan) x x x

aNo rearing made on rainfed lowland bNo rearing made on rainfed lowland or Irrigated lowland

MrasmiaAmong the Laval parasites the braconid Cardio- lowlands and parasitizes oriental maize borer OsIriniachilesphilijpphensis is most adapted to an upland environ- furnacalis (Guenee) larvae The leaffolder pupal parasitesment It also parasitizes Hydelepta indicata a common not only occur equally in all environments but also haveleaffolder of legumes The ichneumonids Trichomma cna- wvide host ranges- parasitizing species of OsIrinia Marucaphalocrosis and Temehcha stangli occur more in the Hydelepta and Honiona - in maize and legumeslowlands T stangli is most adapted to irrigated lowlands The principal egg parasite of yeltow stem borer (YSB)and also parasitizes Chilo and Scirpophaga stem borer Telenomus rowani is widely adapted to all environmentslarvae T cnaphalocrosis is most prevalent in rainfed and has an unusual alternate host -Tabanus eggs (25)

32 IRPS No 123 January 1987

Tetrastictu schoenobii specializes only in YSB and occurs mainly in irrigated lowlands -- further proof that YSB is most adapted to the wetter environments

Larval parasites of YSB are highly environment-specific The braconid Stenohracon nicevillei occurs only in upland rice but also attacks the pink stem borer (PSB) Tieinchwa phiiiplPwnsis an ichneumonid occurs in upland rice and also parasitizes the larvae of (7hiu stqpresaisinatro-morpha accepta melaihoracica like 7 stangi is mnost adapted to the irrigated lowlands

The dark-headed stem borer(l)HSB) (ih pochit3sus was not collected in all rice environments and the gold fringed borer Chilo auricilius nd PSB were collected only in upland rice Xaantiholiplastenrnaloran ichneumonid larval parasite appears to be widely adapted whereas TrihomaslopisapanitI)cteiiaa pteromalid pupal parasite occurs mostly in wetlands and also parasitizes larvae of Chilo suppressais and Pelhuidas mathias

Armyworms Myit hinin and Spodujipera have two main larval parasites - a tachinid Zrgohothriaatropivora in rainfed environments and a braconid Cotesia spp in irrigated lowlands

The butterfly pests Pelopidas and 4elanitishave tricho-grammatid egg parasites found only in irrigated lowland environments

Two larval parasites show environmental preferences --Argrroph av nigrotihialis a tachinid in rainfed habitats and Trichonialopsisapantelkcnela in irrigated ricefields A limgrotibialisalsoattacks the sweet potato hornwormAgrius convolvtdi and T apanteloctena parasitizes DH S1B

Predators as a group show more distinct environmental preferences The aquatic damselflies Coenagrionidae are most abundant in the lowlands Cvriorhinus is also most adapted to the lowlands The nabid Stenonabis tagalicus occurs only in the uplands and isalso prevalent on legumes The coccinellid lady beetles wasps and ants are more prevalent in upland areas AlicrasIpis cit-ea larvae prey ona variety of aphids attacking legumes and maize Etunienes campanonia vespid mud wasp makes nests in trees and is therefore most adapted to the more botanically diverse uplands Soil-dwelling ants cannot tolerate flooding

Upland environments are habitats to arboreal carabid beetles Three species of Chlaeniusbeetles prey on leaffolder larvae and are perhaps more important than parasites (27)

The gryllids and tettigoniids are widely adapted egg predators not only on rice but also on maize and legumes They feed on eggs of most insects that are laid on leaves

The spidercommunity of upland rice environments isrich in species At one site (Batangas Philippines) 31 species have been recorded (23) Of the 176 spider species recorded in Philippine ricefields about one-half (82) occur in upland rice The spiderspecies of upland rice environments overlap more with those of rainfed lowland than with those of irrigated lowland rice (24) The same study showed that of the three environments irrigated rice has (ie greatest spider species diversity foiiowed by rainfed lowland and upland

However of the 10 most prevalent ricefield spiders 9 were abundant in upland rice showing wide adaptation All three spider guilds -- orb-web space-web and huting spiders shywere prevalent in upland ricefields There were however differences in environmental prelerences for some spider species

Of the orb-wcavers Tetragniathamanlihulatawas partishycila rlv abundant 7 japonica was low and Ltucatge dehcorata was absent in upland rice

As in other environments Aiypeiwnaifirtosana was the most dominant space-web spider however its relative numbers were lowerthan for lowland sites Among hunting spiders Ox|opes javanus and Lcosa leatcost fgla were more abundant and Lvcosa lpseudoanmnlata was less abundant in upland compared with lowland rice envishyroniments

Extensive rearing of rice insect pests in upland rainfed lowland and irrigated sites over an 8-yr period in the Philippines provided insight into the effectiveness of parasites as natural enemies (Table 7 8) Most of toe data are from large sample sizes taken over at least one crop which overcomes the pitfalls expressed by Van Driesche (238)

VSB egg parasitization was surprisingly similar across all environments Larval parasitization was highly dynamic among sites and even years within the same site but showed somewhat lowerlevels in uplandsthan in irrigated wetlands This would have been predicted for egg parasites as well since YSB is adapted to the lowlands and so must be its parasites This isevidence that Telenomus rowani is as good at dispersing as its main hosts Trowani clings onto the anal tufts of female YSB and parasitizes eggs as they are laid This phoresy may explain its wide adaptation The more specialized lowland YSB larval parasites therefore appear more effective than their upland counterparts with more alternate hosts

Low collection levels of PSB larvae especially in the wetlands make comparisons between environments difficult Low levels of larval parasifization were recorded in both rainfed environments Except in the Cagayan rainfed wetland site in 1980 low levels ef larval parasitization were evident at irrigated and rainfed sitesalike No differences by environment were evident Higher levels of parasitization occurred in Sarawak during a 1967 outbreak also from tachinids Rothschild (197) indicated that higher parasishytization occurred in upland rice areas

Skipper and green horned caterpillar larval parasitization rates were generally low and showed no environmental effect htighest activity occurred in Cagayan in 1980 (48 parasitization) Although larval parasitization levels of RLF were variable year to year activity appeared greater in the irrigated lowlands than in the uplands Again the highest levels were recorded in Cagayan in 1980 Cagayan has extensive grasslands and fallow around the rice areas and a relatively long rainfall period Rainfall in 1930 was good for crop growth and apparently for increase of larval parasites

IRPS No 123 January 1987 33

Table 7 Parasitization rates of rice planthoppers and green leafhopper in upland rainfed lowland and irrigated lowland environments Philipshypines 1976-840

Environment and site Year Egg parasitization () Nymphaladult parasitization ()

BPH W13P GLH BPII WBPH GLIH

UplandBatangas

Rainfed lowland

1976 1979 1980 1982

15 17 12

2(47)8(39)

4(51)

9(1800)14 (212)

12(304)

8 (101)6 (127)

9(116)

Iloilo

Cagayan

1976 1977 1978 1981 1980 1981 1982

0 (69)

11

44 48 40

-

-

46 55 44

0 (471)

11

52 56 49

0 (200)9 (214)

10 (1259)8(235)6 (900) 3(800)

-7 (462)

11 (270)3 (313)5 (900) 1 (800)

4 (600)10 (725)10 (362)12(870)5 (900) 3 (800)

Pangasinan

Irrigatedlowland

1984 1976 1978 1979 1982

13 (436) 13(379) 14 (457)

6(14)7 (197)

17(198) 11 (486) 18 (275)

3 (81)10 (274)3 (116)

16(871) 19(88)

28(211)9 (764)5 (647)7 (1204) 8(305)

Laguna 1977 11 7 21 1978 31 1979 14 4 4 28 1980 7 8 20 1981 1983

56 7(278)

26 16 (1650)

16 27(1921)

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R I Apostol unpublished 1lo1lo and Pangasinan 136 Cagayan BCanapi unpublished Liuna 175 I C Pantua unpublished 118 234 43 Carino and Shepard unpublished BPH = brown planthopper Nila parvara htgens W131Pi =whitebacked planthopper Sogatelia furcifera GLH = green leafhopper Nephotettrix virescens Figiires in parentheses are sample sizes

Tauher et al (230) reported that internal external and genetic factors influence the seasonal activity of parasitoids

Hopper egg parasites but not nymphaladult parasites were also parlicularly abundant at the Cagayan site possibly the egg parasites have more alternative hopper prey in the weedy areas than nymphaladult paisites

The seasonal dynamics of hopper egg parasites can be seen for upland rice in Batangas(Fig I 2) and for rainfed lowland lice in Pangasinan( 135)and Cagayan( 105) Levels of BPHWBPI-L and GLH egg parasitization were corn-parable in upland and irrigated lowland environments Except for some high rates of paraMsitization early in the season the parasitization rate tended to be steady in all envilonments However hopper nymphaladult parasi-tizution levels were more similar in the rainfed environments and lower than in irrigated rice

Overall parasites do not appear highly effective by themselves against upland or even lowland pests Among natural enemies parasites have attracted the most attention because they can be more readily assesscd

Predators and pathogens lend themselves better to management practices Predators the most important group of natural enemies in riccfields are difficult to quantify On the other hand pathogens appear less im-portant but can be readily cultured and disseminated

The natural enemy commulnity of upland rice is rich in species and differs significantly from that of lowland rice

These beneficial organisms must be conserved by applying insecticides judiciously particularly sprays Seed or soil placement of chemicals minimizes exposure to natural enemies The strategy to derive the greatest benefit from natural enemies isto allow the greatest number of beneficial species to thrive Then perhaps at least one species will be effective against each pest at any tine overcoming the variable abundance of each species within and between years

Programs to introduce exotic species or mass produce indigenous ones are ambitious and expensive because they require trained people on a sustained not ad hoc basis

Chemical control Insecticides offer rapid and efficient control of upland insect pests (190) but-should be used only after other control measures have been considered Insecticides are rarely used on upland rice because of cost (32) Upland rice yields are normally too low to justify the expenditure Also most upland rice is grown as a cash crop Spraying upland rice is more difficult than lowland rice because water is less accessible Government subsidy programs to provide pestishycides to tribal peoples for upland rice production have created problems of toxicity to humans because isolated people have had no experience to be able to handle pesticides safely

However ins-eticide use has beenjustified when a crop is

0Z

Table 8 Parasitization rates of lepidopterous pests in upland rainfed lowland and irrigated lowland environments Philippines 1976-84a

Parasitization (5)

Environment and site Year YSB GFB PSB Earcutting Swarming Skipper Greenborned RLF Brown larva larva caterpillar caterpillar caterpillar caterpillar larva semilooperEgg Larva larva larva larva larva larva

UplandBatangas 1977 - 0 (57) - 0 (63) - 0 (71) - 0 (106) 11(142) 2 (101)1978 43 (4697) 8(451) 5 (38) 3 (40) 6 (61) 2(58) 15 (57) 13 (129) 13 (1286) 4 (30)1979 59(4135) 12(320) 13 (11) 5(28) 9(45) 6(46) 7(23) 8(160) 9(1152) 8(26)

Rainfed lowland Iloilo 1976 - 0 (10) 0(10) 0 (7) 0 (19) 2 13) 15 (180) 5 (76)1978 0 (54) 0 (3) 0 (17) 9 (64) 4 (78) 3 (73) 0 (121) 0 (191)1979 64 (2350) 16 (84) 6 (17) 0(6) 18 (68) 25 (104)

1981 56 (1322) 18(304) 0(5) 12(51) 18(138)Pangasinan 1976 60 (7 7 )b 62(82) 0(111) 5 (39)

1978 58 (53)b 5(51) Cagayan 1979 59(713) 8 (101) 6 (34) 0(7)

1980 0(11) 4 (28) 15 (286)

62 ( 48(177) 61(140)1981 0(5) 12 (2) 13(45) 14 (92)

Irrigatedlowland Laguna 1974 62 8

1974-75 50 41 1975 43 72 1978 96 (623) 21(24) 33 (60)1979 83(2017) 18(110) 8(62) 11 (89) 22(400)1980 78 (1625) 20(75) 0 (16) 4 (72)1983 60

9 (80) 7 (270) 30

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R F Apostol unpublished Iloilo and Pangasnan 136 Cagayan B Canapi unpublished Laguna 175 P CPantua unpublished 118 234 143 Carino and Shepard unpublished YSB = yellow stem borer ScirpophagaincertulasGFB = gold fringed borer Chilo auricilius PSB = pink stem borerSesamia inferens earcutting caterpillar Alythimna separata swarming caterpillar Spcdoptera mauritia acronyctoides skipper Pelopidas mathias reen horned caterpillar Melanitls ledaismene RLF = rice leaffoldersCnaphalocrocismedinalis and Marasmiaspp brown semilooper Mocisfrugalis Figures in parentheses are sample sizes Egg masses

IRPS No 123 January 1987 35

Table 9 Effect of seeding rate and insecticide on plant density and Yield losses from insect pests in upland rice often areyield of Dagge upland rice Batangas Philippines 1976a comparable to those of lowland rainfed or irrigated rice

Seeding rate Insecticide Dosage Plant density Yield (0-Y30o) but usually from different pests The principal (kg aiha) seed treatment (kg aiha) (noIra-row) (tha) groups are soil pests feeding on sown seed and rootsat 14 DE followed by seedling pests These groups have largely gone

50 None - 35 a 20 a unnoticed leading researchers to conclude that stem borers50 Carbofuran F 10 38 ab 23 a and seed bugs are the most important pests (17) Yield loss 10 beCabofuranF100 None - 4551 c 23 a trials that pinpoint major pests need to includc seed

100 Dieldrin WP 10 52 cd 22 a treatments Granules applied in the furrow may not give150 None - 59 d 22 a adequate protection from many seed pests

aAv of 8 fields DE = days after crop emergence In a column means The floristic and pedological diversity of upland rice followed by a common letter are not significantly different at the environments is matched by the diversity of arthropods5level by D)MRT F = flowable Wi = wettable powder both pests and their natural enemies Irrigation has homoshy

genized lowland rice environments Flooded soils tend tothreatened by epidemics of locusts or armyworms New have the same properties and to eliminate soil pests The technology stressing efficiency has minimized insecticide spatial and temporal dominance of cultivated rice has cost benefited specialist pests

Seed treatment is normally inexpensive and can be Upland rice has no specialist arthropod species The economicallyjustified in -nanycases(66) Treating seed with monophagous pests on upland rice prefer the lowlands systemic insecticide (216) but not chlorinated hydrocarbons where rice evolved Rice in the uplands exists only because (247) is most effective in controlling seedling maggots of man and even though lowland rice predates upland riceFoliar sprays have to be repeated to obtain control and specialist species evolved Wheat maize and sorghum do often are washed off by frequent early season rains not have monophagous pests leading us to conclude that Granular insecticides are effective but required dosages are there is no evolutionary advantage for inssects to have too expensive to justify except under heavy infestation narrow host ranges in upland graminaceae(247) When directed against first-instar larvae banding Pests have adapted to the uplands not only by having granules in seed furrows (177) can be effective against white wider plant host ranges but also by having long life cyclesgrubs at low dosages (025 kg aii ha) (134) Seed treatments undergoing dormancy or being dispersive Each pest has a are not promising for white grub control (248) unique life history strategy suited to highly unpredictable

Seed treatment is also effective ainst ants and may be environments In agivenyear someinsect specics an better cheaper than increasing the seeding rate In Batangas exploit upland rice as a food source but no pest is a senious treating 50 kg seed ha was as effective as increasing the one every year on upland rice seeding rate to 100 kgha and using untreated seeds (Table The low yielding potential of upland rice means less 9) In Brazil seed treatments control termites and other soil quantity and lower quality of food for pests to exploit but insects including the lessercornstalk borer(66) Domiciano also less incentive for farmers to undertake control (58) however found seed treatments inconsistent in their The most effective control would be early and synchroshyeffect against high populations of lesser corn stalk borer nous planting of early-maturing upland rice varieties to

Baiting can be an inexpensive way of controlling ants prevent insect pest buildup Breeding for insect resistance seedling maggot flies mole crickets and field crickets Baits should initially focus on stem borers because they are verymade of locally available material and impregnated with difficult to control by other means Biocontrol efforts insecticide can be sparingly distributed taking adantage of should first stress conservation of natural enemies preshythe pests ability to disperse and encounter bait sites dators and pathogens rather than parasites Efficient

The new low-volume hand-held controlled-droplet chemical control includes seed treatment or baiting for seed sprayers may offer an advantage over high-volume knap- seedling and some root pests and using controlled dropletsack sprayes for controlling foliar pests Upland rice sprayers for spot treatments against foliar pests and seed canopies are more open than those in lowland rice and bugs when economic thresholds are reached better droplet penetration should result Controlled droplet applicators would be ideal for seedbugs and defoliators

CONCLUSION

Upland rice is attacked by a wider array of insect pests than is lowland rice mainly beca use of the addition of soil pests Generally population levels of lowland rice pests are lower in upland than lowland rice leading many people to conclude that insect pests are not important on upland rice

36 IRPS No 123 January 1987

REFERENCES CITED

1Abraham C C and K S R Mony 1978 Occurrence ofLeptocorisa acuta Fabr (Coreidae Hemiptera) as a pest of nutmeg trees J Bombay Nat ttistSoc 74553

2 Agyen-Sampong M 1975 HlierogrlphusdiaganensisKrauss (Orthop-tera Acrididae) a new pest of rice in Northern Ghana Ghana J Agric Sci 8249-253

3 Agyen-Samnpong M 1976 Varietal resistance and other inle-ratcd pest control components as factors of rice varietal improvement in West Africa Prcsented at the Second varietal improvement serinar 14-18 Sep 1976 Bouak Ivory Coast West Africa Rice Development Association Monrovia Liberia 18 p

4 Agyen-Sampong M 1977 Outbreak of Ihccrorrdnisorae in rice seedlings near Rokupr Sierra Leone lit Rice Res Newsl 2(6)16-17

5 Agyen-Sampong M 1980 Dirty panicles and rice yield reduction caused by hugs IntRice Res Newsl 5(l)11-12

6 Agyen-Sampong M 1982 The major pest problems of irrigated upland and mangrove swanip rice ecosystems ir the humid tropical Guinea savanna and Salel climate zones Pages 318-328 in Pro-ceedings training course cn concepts techniques and application of irtegrated pest management in rice inWest Africa 10-20 Jan 1982 Fendall Liberia West Africa Rice l)evelopioent Association Monrovia Liberia 505 p

7 Akibo-Betts V I and S A Raymundo 1978 Aphids as rice pests in Sierra Leone lit Rice Res Newsl 3(6)15-16

8 Akinsola F A 1975 Present status of different rice stemborers in parts of Nigeria iicL Entomo Newsl 328

9 Akinsola FA 1979 Ihe biology and ecology of rice stemborers in Nigeria Ph 1) thesis University ol lbadan Nigeria 217 p

10 Akinsola E A 1979 Guidelines of integrated management of insect pests of upland rice in West Africa Presented at the Seminar on integrated management of rice diseases and insect pests at Bobo-Dioulasso Upper Volta 17-22 Sep 1979 West Africa Rice Develop-ment Association Monrovia Liberia

II Akinsola IA 1984 Insect pests of upland rice in Africa Pages 301-305 in An overview of upland rice research Proceedings of the 1982 lIouak Iwsrv Coast upland rice workshop International Rice Research Institute 13O Box 933 Manila Philippines

12 Alam M S V TJohn and Kaung Zan 1985 Insect pests and diseases of rice in Africa Pages 67-82 in Rice improvement in Eastern Central and Southern Africa proceedings of the international rice workshop at Iusaka Zambia 9-19 April 1984

13 Alticri M A A van Schoonhovcn and J D Doll 1977 The ecological role of weeds in insect pest management systems a review illustiated with bean (Phaseohsvulgaris L)cropping systems PANS (Pest Artie News Summ) 23195-205

14 Anmante F 1967 The leaf-cutting ant Atia capiguaraa pest of pastures [in Portuguese] Biol6gico (Sao Paulo) 33113-120 (Rev Appl Entomol A56207)

15Andrade A C and D Puzzi 1951 Preliminary results of experi-merits on the control of Sculptcoris iastantus on sugarcane [in Portuguese] Bioulgico (Sao Paulo) 1744-49 (Rev Appl Entomol A41421422)

16 Appert J 1970 Maliarplia separaelha (rice stem borer) new observations of entomological problems of rice in Madagascar [in French] Agron Trop (Paris) 25329-334

17 Ariadeau Mand Z Harahap 1986 Relevant upland rice breeding objectives Pages 189-197 in Progress in upland rice research Proceedings of the 1985 Jakarta conference International Rice Research Institute 1O Box 933 Manila Philippines

18Asar S and Kisimoto 1975 Seasonal occurrence of the smaller brown planthopper in upland rice [in Japanese] Proc Kanto-Tosan Plant Prot Soc 2228

19 Bakker W 1974 Characterization and ecological aspects of rice yellow mottle virus in Kenya Meded Landbouwhogesh Wage-ningen 152 p

20 Balut F F 1970 White grubs attacking rice Oryza ativa L [in Portuguese] Biol6gico (So Paulo) 36321-322

21 Banerjee S N and L M Pramanik 1967 The lepidopterous stalk borers of rice and their life cycles in the tropics pages 103-124 in The major insect pests ofthe rice plant Proceedings ofa symposium at the International Rice Research Institute September 1964 The Johns Hopkins Press Baltimore Maryland USA

22 Barrion A T and J A Litsinger 1980 Ants a natural enemy of cliaiarac in dryland rice IntRice Res Newsl

5(4)22-23 23 Barrion A T and J A litsinger 1981 The spider fauna of

Philippine rice agroecosystem 1Dryland Philipp Entomnol 5139-166

24 Barrion A T and 1 A Iitsinger 1984 The spider fauna of Philippine rice agroccosystcms II Welland Philipp Eiitomol 611-37

25 Barrion A T1 and J A Litsinger 1984 lihanus (Diptera Tabanidae) eggs an alternative host of rice stei borer (SI3) egg parasite Telenoonis digns (Iliymenoptcra Scelionidae) lit Rice Res Newvsl 9(6)19

26 Barrion A T and J A Litsinger 1985 Identilication of rice leaffolders by wing markings Int Rice Res Newsl 10( 1)24

27 Barrion A T and J A Litsinger 1985 Chctiosspp (Coleoptera Carabidae) a leaffolder ILF)predator lit Rice Res Newsl 10(1)21

28 Bertels A 1970 Insect pests of rice and theircontrol [in Portuguese] Pelotas Brazil Instituto de Expcrimentacao Agropccuararias do Sul Circular 43 24 p

29 Bonzi S M 1982 Ctilo (flisilineus J de Joannis (Lepidoptera Pyralidae) stem borer of irrigated cereals in Upper Volta LioFrench] Agron Trop Riu Rizic Cult Vivrieres Trop 37207-209

30 Bourke T V T L Fenner J N LStihick G Baker EHassan D F OSullivan and C S li 1977 Insect pest survey for the year ending 30th June 1969 Papta New Guinea )epartment of Agrishyculture Stock and Fisheries Port Moresby 57 p

31 Bowden 1 1973 Migration of pests in the tropics Meded Rijksfac Landbouwwet Gent 38785-796

32 Breniere J 1973 The main rice parasites in West Africa and their control Pages 27-42 in Proceedings seminar on plant protection for the rice crop West Africa Rice Development Association Monrovia Liberia

33 Breniere J 1976 The principal insect pests of rice in West Africa and their control West Africa Rice Development Association Monrovia Liberia 52 p

34 Brown ES 1962 The African arinyworn Spoiolptera eXI)Ina (Walker) (Lepidoptera Noctuidae) a review of the literature Cornshymonwealth Institute of Entomology London UK57 p

35 Brown F B 1968 Upland rice in Latin America FAO-International Rice Commission 12th Session of the Working Party on Rice Production and Protection 2-14 Sept Peradeniya Sri lanka Food and Agriculture Organization Rome Italy 6 p

36 Buyckx F J 1 1962 Disease and insect pests of crops inCongo Rwanda and Burundi [iil French] Publications dc LInstilut National pour lestude Agrononiquedu Congo M Weissenbruck S A Brussels Belgium 708 p

37 Cariiio F 0 1981 Role of natural enentics inpopulation suppression and pest management of green rice lealhoppers Ph D thesis University ofthe Philippines at Los Bafitos PhilippinesLaguna 136 -

38 Carpente A J 1983 The development of rice cultivation and extension of vegetable productin in Zanzibar Final Report Vol II1 October 1983 Food and Agricultme Organization Ronte Italy

39 Carroll C P and S1 Risch 1983 Tropical annual cropping systems ant ecology Environ Manage 751-57

4(0 Cates R G 198 1 Host plant predictability and the feeding patterns of monophagous oligophagous and polyphagous insect herbivores Oecologia (Berl) 48319-326

IRPS No 123 January 1987 37

41 Cendcafia S M and F B Caora 1967 Insect pests of rice in the 62 Ene 1 C 1981 I)iiorldhol)ierusihilis Slater (licteroplera Philippines Pages 591-616 in The najor insect pests ofthe rice plant lygacidue Illissinac) a new rice pest front Nigeria lit Rice Res Proceedings of a symposium at the International Rice Research Newsl N3)15 Institute September 1964 The Johns Hopkins Press Baltimore 63 Fugenio T S 1965 rhe effect oi e-vCgetation on rice production Maryland USA 729 p and the possibility of overcoming it by leaf pruning MS thesis

42 Centro Ihternacional de Agricklittira Tropical 1984 Upland rice in University ol the Philippines at Los Baijos ILaguna Ihilippins 69 p Iatin Anerica Pages 93-119 hi An os ervies of uplaind rice research 04 Farrow R A 1975 The Alrican nmigratory locust ii is main Proceedings of the 1982 Boinakamp Ivory Coast upland rice workshop outbreak area of the middle Niger qlinlitatise studies of solitary International Rice Research Institute P0 Blox 933 Manila populations in relation to environmental factors Locusla No II Philippines 198 p

43 Chandra G 1979 Taxonomy ad hn riononlics ofthe imect parasites of 65 Fennal R G 1943 Food crop investigation Windward and rice Icalloppers and planthoppers in the Pihilippines and their Ieeward Islands First report Report onl work dunring the periodiiiportairce iii natural biol)gical cointrol Philipp Fntoltol 4119-139 April 1942-April 1943 Imperial College of Tropical Agriculture

44 Chang T T 1976 The originii colut ion enlIivation dissemination Trinidad 16 p (Rev of Appl Fintornol A3222-25) and diersificition of Asian and African rices -uphytica 25425-44 1 66 Ferreira F 1983 Integrated pest ritaageiret of upland rice [ill

45 Chatierjee P B 1979 Rice leaffolder attacks in India lit Rice Res Portuguese English summary] Pages 323-341 in Upland riceculttre Newsl 4(3)21 factors affecting production E Ferreira T Yamada and F-

46 Cheng Cbien-pan 19(4 Upland rice in Taiwan International Rice Malmolta eds Institutoda Potassae Foslato Institnto Internacional Cnmmission Wiorking Party oit Rice Production ard Protection I()th da Potassa Piracicaha Sao Paulo lraid Meeting 3-10 March 1964 Manila Philippines Food and Agricul- 67 Ferreira E arid F dl Silva Martins 1979 Occurrence of ture Organiaion Rome Italy 5 p Sogatohs orvziola (Muir) inupland rice in Goias Ilrail lit Rice

47 Chcrrett 1 M and 1) 1Peregrine 1976 A review of the status of Rcs Ncwssl 4(4)18 leaf-cutting ants and their control Ann Appl Biol 84124-128 68 Ferreira E and 1F da Silva Martiris 1984 Insect pestof rice ill

48 Chipeta F M 1974 Pests of rice in Malawi Pages 90-95 in Brazil and their control Jill Portugucsej FMIlRAtPA-CNPAF Proceedings Sith eastern Africa cereals research conference 10-15 Goiania Brmiil 67 p March 1974 DRB Mialda ed Ministry of Agriculture lilongwe 69 Ferreira FI F ca Silha Miartins S S Nero and 1 It P de Malawi Carvallio 1982 M tltiplc regression analysis todeterimine the effect of

49 Chu Yau-i 1979 The d atiaging period of the rice seedling fly insect populatiois ott upland rice yield [inIPortuguese Pesqui (Atlterigotiavxigu Stcin) (NItiscide Diptera) oil cori) at Fast Java Agropecu Bras 17671-675 Indonesia Plant Prot Bill 21403413 70 Ferreira F 1F da Silva Martins and F 1PZimmermann 1979

50 Conklin H C 1957 IHaninoo agriculture a report on aln integral Rice varietal resistance to tle lesser corn stalk borer [in Port uguese] systent of shi fti rig cult ivationi in the Pltilippincs Food and Agriculture Pesqui Agropccu Biras 14317-321 Organization Rome Italy 209 p 71 Fleming W F 1968 Biological control of lie hapanese beetle U S

51 Cramer H II 1967 Plant proection and world crop production Dep Agric RLs Sers [ech Bull 1383 78 p Pilaniensclhutz-Nachr 201-524 72 Food and Agriculture Organiation of the United Nations 1982

52 l)ani R C and Majunidar 1978 Prelininary observations on tie Country report surmimary Pages 27-37 in Report on the first incidence of root aphids on differeit rice cultnrcs Sci Cult 4488-89 FAOIUN)IPhail and regional training coiuirse in iniproved ctltural

53 Davidson R I 1979 lffects of root feeding on foliage yield Pages practices for upland rice 5-23 Oct 1981 Northern Region Agrculturd117-120 in T K Ci osb and R B Pottinger eds Proceedings of the Developririt Center Chiang Mai Food and Agricuilture Organiiashy2nd Australian conlerence on grassland invertebrate ecology Govt tion Rome Italy 37 n Printer Wellington New Zealand 73 Franco do Ariaral S 195 1 Residual action of insecticides in turfy

54 l)ean G 1W 1976 Rice insect pests inlaos Int Rice Res Newsl soil--the results of a field experiment [in Portuguese] Biol6gico (Sao 1(2)15 Paulo) 17209-214 (Rev Appl Entomol A41423)

55 I)ean G J W 1978 Insect pests of rice in Laos PANS 24(3)280-289 74 Freeman ) 1970 Report onithe Iban london School of -coroiics 56 )ingle H 1984 Behavior genes and lifc histories conplex adapta- Monographs on Social Anthropology No 41 Athlane Press

tions to uncertain environments Pages 169-194 in A new ecology London UK 317 p novel approaches to interactive systems P W Price C N Slobod- 75 Fuchs T W1 A Harding and 1W Smithlr 1979 Induction and chikoff and W S Gaud cdsl ohn Wiley New York USA termination oldiapause in the sugarcane borer Ann Flitonlol Soc

57 Dobelmann-JP and M Falais 1968 Upland rice cultivation ill tie Am 72271-274 North-western part of Madagascar in 1966-1967 [in French English 76 (hbriel B1) 1975 A review tf the major insect pests of some tpland summary] Trop Agron 2313-21 crops in tlc Philippines Pages 86-111 in Ad Htoc panel of experts on

58 Doniiciano N L 1979 Chemical control of tie lesser corn stalk pest disease arnd weed problems in some rainfed crops Bangkok (lolsonno~paloIgnowslhts) (Zeller 1848) (I-epid(ptera Ihycitid ae) Thailand Food aid Agriculture Organization Roire Italy

on rice in Ponta Gressa Parara 1977-1978 [in Portuguese] Pages 77 Garg D K and N K Shal 1983 White grub outbreak on rainfed 291-303 in Anais 1 Reuniao de tecnicos cmiirizicultura do estado de dryland rice in Uttar Pradeslh Int Rice Res Ncwsl 81(4)16-17 Sao Paulo-Campinas Cati Brazil 78 Garg I) K and IP Tandon 1982 Major insect pests of rice on hilly

59 Dunsmore 1 R 1970 Investigations on tlie varieties pests and tracts of Uttar Pradesh India lilt Rice Res Newsl 7(11)11-12diseases of upland rice (hill padi) in Sarawak Malaysia Int Rice 79 Garrity 1) P 1983 Upland rice research an international bibliograpy Comm Newsl 1929-35 1975-1982 Iternational Rice Research hnstitnte PO Box 933

60 Elias 1R1965 The new decline of rice seedlings is not caused by Manila Philippines 121 p parasites [in Portuguese] Biol6gico (Sao Paulo) 313-6 80 Ghauri M S K 1982 A new subspecies of Iinorh)rone bull

61 Empresa Brasiliera de Pesquisa Agropecuaria 1984 Upland rice itt cornlus Slater (Hemiptera Iygaeidae) front Papua New Guinea on Brazil Pages 121-134 it Av overview of upland rice research rice and carpet grass Ihll lEntoniol Res 72133-137 Proceedings ofthe 1982 Blouak Ivory Coast Upland rice workshop 81 Gonzales F 0 PArregoces R I ihernandef and OT Parada International Rice Research Institute PO Box 933 Manila 1983 Insect and imite pests of rice arid their control in Latin America Philippines [in Spanish] la Federacion Nacional de Arrocerts (Fedearroz)

Bogota Colombia 60 n

38 IRPS No 123 January 1987

82 Grist D H and R J A W Lever 1969 Pests of rice Longmans London UK520 p

83 Hale P R 1979 Two subterranean pests of upland rice in Papua New Guirnca IntRice Res Newsl 4(2)16-17

84 Hale P R and J T Hale 1975 Insect pests of rice in the New Guinea Islands Rice Entomol Newsl 34

85 Harris W V 1969 Termites as pests of crops and trees Common-wealth Institute of Entomology London UK 41 p

86 Hattori 1 1968 A method to distinguish between grass leaf roller (Ctaplha rocis medinalisGuenee) and the rice leaf roller (Susurtia exigua Butler) [in Japanese] Plant Prot (Japan) 23167-170

87 Henry I E and 1A Onsager 1984 Experimental control of the mormon cricket Iahrosshnph by NJsena locustae(Aficrospora Microsporidae) a protozoan parasite of grasshoppers (Ort Acri-didae) Entomophaga 27197-201

88 Hidaka 1 P Vungsilabulr and SKadkao 1974 Studies on ecology and control of the rice gall midge in Thailand Tech Bull 6 TropicalAgricultural Research Center Tokyo Japan 113 p

89 Hincklcy A 1) 1963 Ecology and control of rice planthoppers in Fiji Bull Entomrol Res 53467 481

90 Hirao J 1982 Ecology and chtInical control of the rice leaffolder Jpn Pestic Inf 4114-17

91 Ho DT 1984 Sten borer incifience in rice ecosystems in KenyaEast Africa Pages 307-320 in A overview of upland rice research Proceedings of the 1982 Bouac Ivory Coast upland rice workshop International Rice Research Institute PO Box 933 Manila Philippines

92 Ho )T J 1Njokah and J GKibuka 1983 Studies on rice sten borers in Kenya with emphasis on ifaliarpiaipiraellRag Insect Sci Appl 465-73

93 Ichihara I T Ito A NIatsuo and M Sawada 1971 Chemical control of the rice striped stem borer in upland rice grown in plastic mulch [iii Japanese] Proc Kanto-Tosan Plant Prot Soc 1882

94 IclhiharaIand A Matsuo 1969 Damage caused by rice striped stett borer to upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1686-87

95 Ichhara I atid A MIatsun11970 Conside ration oft lie extraordinary prevalence of rice striped stem borer in upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1778-79

96 Institut de Recherches Agrolnliqes Iropicale et des Cultures Vivrieres 1978 Insect resistance Pages 12-13 in Upland rice 1978 atnual report Bouak Ivory Coast 27 p

97 International Institute for Tropical Agriculture 1983 Aphids itt rice roots I ITA Res Briefs 4(1-3)4-5

98 International Institite fi Tropical Agriculture 1983 Hopperburn in rice caused by brown plathopper IITA Res Briefs 4(l-3)6-7

99 International Rice Research Institute 1975 Annual report for 1974 PO Box 933 Manila Philippines p 216

100 International Rice Restarch Institute 1976 Annual repiort for 1975 PO Box 933 Manila Philippines p 228

101 International Rice Research Institute 1979 Cropping systems program Component technology development and evaluation Page 404 i Annual report for 1978 11O Box 933 Manila Philippines

102 International Rice Research Institute Department of Entomology 1981) Insecticid evaluation 1979 Tables 73 74 PO Box 933Manila Philipes

103 International Rice Research Institute Department of Entonmology 1981 Insecticide evalomtion 1980 Table 81 POBox 933 Manila Philippines

104 International Rice Research Institute 1981 Report of a meeting of Asian rice-based cropping systems entomologists Eleventh cropping systems working group meeting 18-22 May 1981 PO Box 933 Manila Philippines p 132 133 144

105 International Rice Research Institute 1984 Cropping systems program Pages 368-373 in Annual report for 1983 PO Box 933 Manila Philippines

106 International Rice Research Institutc 1984 Upland rice in Asia Pages 45-68 in An overview of upland rice research Proceedings of the 1982 Bouakamp Ivory Coast upland rice workiiop loternational Rice Research Institute 11O Box 933 Manila Philippines

107 Irshad M 1977 Grasshoppers in rice in Pakistan lit Rice Res Newsl 2(3)8-9

108 Ito R and TOno 1964 Upland lice its culture and varieties in Japan lIternational Rice Cormmission loth Meeting of the Working Party on Rice Production and Protection 3-10 March 1964 Manila Philippines Food and Agriculture Organization Rome Italy 12 p

109 Jeffrey M I and V A Dyck 1983 Effect of moonlight on lightshytrap catches of brown planthopper lit Rice Res Newsl 8(6) 13

110 Jennings P R and T Alicia Pineda 1983 Management ofthe rice plnthoppers Sogatodsorivi(coIl in Latin America Presented at the meeting on stable plant resistance FAO and lIndocsian I)inectorate of Food Crops Protection Bali lndonesia 29 Novcniber-2 l)ccemshyber 9 p

IllJimenez Gomez J A 1979 Rice insect pests i lla ios ()riCntales lil Spanish] Pages 99-103 in Rice ciltivation Instituto (olotibiatto Agropecuario Compndio No 29

112 Jongeleen F JJ Soesanto and A Mockidjo 1979 Studies oti pathogens of two species of Scarabacidac froinIndonesia Z Angew Enlomnol 87412-417

113 Jouvenaz 1) PC S lofgren and W A Baiks 1981 Biological control of imported fire antsa review of cirrent knowledge Bull Entomol Soc Am 27203-208

114 Kainoh Y K Shiniiu S MarUani Y Taiaki 1980 lost finding behavior of tie rice bu lplotori+JchinvittjI)allas (liciniptera Corcidac) with special reference to diet pattern ol aggregation and feeding on the rice plant Appl Entomol Zool 15225-233

115 Kakkar K 0 P Sharma and P I Sharma 1977 Note on Ihteronyrchus liodresRedt (Coleopicra I)ynastidae) as asporadic pest of paddy and maie inHimachal Pradesh Corr Agric 156-57

116 Kalode M131 79 Major insect pest problems and their control in upland rice (Unpublishcd mineogriph CWI oJNaaiat lni Chandaratat (198 1)original not seen)

117 Kalshoven L i 98 I The pests of crops in I ldoniesia (translated by P A vatn der Laan) 1 T Ichtiar B3aru-vati Iloeve lakarta Indonesia 701 p

118 Kamal N Q 1981 Suppression of whitebacked planto ner Sogatella fircaifra (Horvath) aLId rice lealfolder (nhal -cis licdb lis (Guenee) populations by natural cnclmie hlI) tiesis Grcgorio Araneta University Foundation Victoncta Park Metto Manila Philippines 179 Ip

119 Karim A N M Reaul 1975 A species ofliealy bug Trioni i70s ceres new oii rice in the Philippines Rice Entoniol Newsl 247

120 Katanyukul Wand C Chandaratat 1981 Insect pestsof uplaid rice and their control Report firstFAOUNlFi Tliailand training course on improved cultural practices for tplad rice Chiang Mai Thailand 5-23 Oct 1981 Food and Agriculture Organiation Rome Italy 22 p

121 Kern F 1955 lhletwioa(I)uninemts) bhhit Biturm(Coleoptera) a new pest of sugarcane inVenezuela [inSpanish with Etnglish sunmary] Agron Trop (Maracay) 527-29 (Rev Appl Eitoniol A4572)

122 Khan K M and S Ghai 1974 White grubs ald their control in India Pesticides (1Bombay) 819-25

123 Kiritani K 1975 Integrated pest management of rice pests Pages 66-70 itt Proceedings platning workshop ott cooperative hIeld research in pest managementP SMotooka ed East-West Center Honololu Hawaii USA

124 Kisimoto R 1967 Ecology of insect vectors forecasting and chemical control Pages 243-255 in The virus diseases of tle rice plant Proceedings of a symposium atthe International Rice Research Institute April 1967 The Johns Hopkins Press altimore Maryshyland USA

125 Kisimoto R and V A Dyck 1976 Climate and rice insects Pages 367-391 in Climate and rice International Rice Research Institute PO Box 933 Manila Philippines

126 Kureha S S Kobayashi SAsari and T Ishihara 1977 Life history and control of the mole cricket Grrllotalpa africana Palisot de Beauvois [in Japanese English translation 7520 1RRI library] Proc Kanto-Tosan Plant Prot Soc 21192-197

127 Kushwaha K S 1981 The black beetle as a pest of rice in Haryana India Int Rice Res Newsl 6(4)14

128 Lawani S M 1982 A review of tile effects of various agronomic practices on cereal stem borers Trop Pest Manage 28266-276

129 Leuck D B 1966 Biology of the lesser cornstalk borer in South Georgia J Econ Entomol 59 797-801

130 lever R J A W 1955 Insect pests of the rice plant in Malaya Int Rice Comm Newsl 1612-21

131 Li Ching-Sing and Shin-Foon Chiu 1951 The study ofthe rice gall midge Iachydiplosis orvza Wood Mason [in Chinese with English summary] 1Taiwan Agric Res 21-13

132 Iitsinger J A 1984 Assesament of the need-based application for rice Proceedings TechnologyTransfer Workshop 15-16 March 1984 International Rice Research Institute 1PO Box 933 Manila Philippines 6 p

133 Litsinger J A 1984 Environmental diversity and insect pest abundance with reference to the Pacific Pages 285-307 in Proceedings sub-regional training course on methods of controlling diseases insects and other pests of plants in the South Pacific 4-20 Oct 1982 Vaini Kingdom of Tonga Nukualofa

134 Litsinger J A RF Apostol and M BObusan 1983 White grub Leucopholis irrorata (Coleoptera Scarabaeidae) pest status popula-tion dynamics and chemical conllol in arice-maize cropping pattern in the Philippines J Econ Entoinol 761133-1138

135 Litsinger J A C de la Cruz F Raymundo A T Barrion M D Lumaban J P Bandong M S Venugopal F Paragna and E Balete 1982 Insect pests and insecticide reponse of dry-seeded bunded rice Pages 179-193 in Report of workshop on cropping systems research in Asia International Rice Research Institute Los Bafos Philippine

136 Litsinger J A M 1) Luinaban J P Bandong A T Barrion W Katanyukul and N Panda 1978 Insect pest management in rainfcd lowland rice agroecosystms of tropical Asia International Rice Research Conference 17-21 April 1978 International Rice Research Institute Los Bafios Philippines

137 Litsinger J A and K Moody 1976 Integrated pest management in multiple cropping systems Pages 293-317 in R I Papendick P A Sanchez and G B Triplett eds Multiple cropping American Society of Agronomy Madison Wisconsin USA

138 Loevinsohn M E 1984 The ecology and control of rice pests in relation to the intensity and synchrony of cultivation Ph D thesis University of London 36) p

139 Loevinsohn M E J A Litsinger N Panda M A Austria R F Apostol C Andrion and J Bandong 1982 A basis for predicting rice insect pest abundance in cropping systems Paper presented at the 1982 International rice research conference on rice and cropping systems 19-23 April 1982 International Rice Research Institute Los Bafios Iaguna Philippines 25 p

140 Lopez A W 1930 Repot of the Entomology Department Pages 145-172 in Annual report of the Philippine Sugar Association 1929-30 Manila

141 Lopez AW 1931 Entomology Department Pages 227-266 in Annual report of the director or research 1930-31 Philippine Sugar Association Manila

142 Lopez A W 1932 Entomology Deparment Pages 252-279 in Annual report of the Philippine Sugar Association 1931-32 Manila

143 Lungibill P and G G Ainslee 1917 The lessercornstalk borer US Dep Agric Agric Res Serv Tech Bull 539 27 p

IRPS No 123 January 1987 39

144 McGuire J U Jr and BS Crandall 1967 Survey of insect pests and plant diseases ofselected food crops of Mexico Central America and Panama U SDepartment of Agriculture International Agriculshytural Development Service Agricultuial Researrh Service October 1967 11 p

145 McNeill Sand T R ESouthwood 1978 The role of nitrogen in the development of insectplant relations Pages 77-9 in lliochemical aspects of plant and animal coevolution J B Ilarbornu ed Academic Press London UK

146 Maebara H 1955 Spatial distribution of lastinstiarlanaeof purplish stem borer (Sesamia iqferens Walker) hibernating in an upland rice field Bull Kagoshima Univ Fac Agric 438-41

147 Mallick E H and N G Hajra 1977 Response of rice plant to defoliation treatments at different developmental stages Indian Agric 2151-56

148 Mansingh A 1971 Physiological classification of dormancies in insects Canl Entomol 103983-10)9

149 Martell C G 1974 Catalogue of plant feeding insects of Mexico [in Spanish] Fitofilo 27(69)1-176

150 Martinez 0 J 1982 Principal rice pests in Ilanos orientalis [in Spanish] P~ages 80-94 in Curso deact nalization de conocimientos en arroz Villaniencis Instituto (olonmbiano Agropecuario

151 Martins J F da Silva F Ferrcira A S Prabhu F 1 P Zimmermann 1980 Use of pesticides for the control of the main subterranean upland rice pests esqui Agropecu Bras 1553-u

152 Martins I F da Silva F Fe eira rod BdamSilveira Pinheiro I Q2 Damage simulation caused by defoliators of upland rice [in Portuguese with English summary] Pesqui Agropecu Bras 171113-1119

153 Martins 1F da Silva N(iyuen van Tan and 1Ida Silveira Pinheiro 1981 Stem borer resistance in upland rice associated with plant morphological characters [in Portuuce Iesqui Agropecu Bras 16187-192

154 Masaki S 1980 Sumnmer diapause Ann Rev Entomiol 251-25 155 Masuzawa T 11 Suwa and F Nakasoji 1983 l)ifkleences of

oviposition preferencc and survival rate of two skipper butterflies Parnara guttata and PIchlpidai iathiais (Iepidoptera HIesperiidac) on rice plant and cogon grass New Entomol 3247-56

156 Maurya D M and C P Vaish 1984 Upland rice in India Pages 69-92 in An overview oftpland rice research Proceedings ofthe 1982 Bouakamp Ivory Coast upland rice workshop International Rice Research Institute IO Box 933 Mnila Philippines

157 Menon M G R R N Katiyai S Kuar and S S Misra 1970 Termites attacking the paddy crop Sci Cult 36294-295

158 Miyahara Y and 1 Nishikubo 1967 Behavior of young nymphs of the red rice root aphid Iholahsipiun ri liabhdontinalisSasaki in relation to movement from the leaf to the roots of upland rice [in Japanese] Pioc Assoc Plant Prot Kyushu 1363-66

159 Mo ljoa Tjien 1952 White grub of upland rice (Holotrichiahelleri Brsk) [in Indonesian with English summary] Contrib Agric Res Stn Bogor No 13134 p

160 Mochida 0 and T Suryana 1979 The occurrence of the brown planthopper and ragged stunt on lowland and upland rice ill Central Java and logyakarta in tile late of January 1979 Entomnolgy Survey Report No 15 Central Research Institute of Agriculture (CRIA) Sukamandi Indonesia I February 1979 9 p

161Monet C 1979 Varietal resistance to upland rice pests [inI French] Pages 35-39 in Annual report 1977 Institut des Savanes Department des Cultures Vivrieres Bouak6 Ivory Coast

162 Monte 0 1942 A caterpillar ofrice-fields[in Portuguese] Biol6gico (Sao Paulo) 8161-163 (Rev Appl Entomol A31414)

163 Munroe I) D 1979 Pest management strategy for hill rice in four resettlement areas in Sarawak Malaysia Pages 89-102 i1 Guidelines for integrated control of rice insect pests Food and Agriculture Organization Plant Production and Protection Paper 14 Rome Italy 115 p

40 IRPS No 123 January 1987

164 Nakasuji F M Ishii and Jen-Zon Ito 1984 Rice skippers in Taiwan and their life histories ZZrC (Taipei) Tech Bull 82 10 p

165 Navas I) 1976 Fall armyworum in rice Pages 99-106 il Ecological animal control by habitat management Proc Tall Timbers Con-ference 28 Feb-I Mar 1974 Fall Timbers Res Stn Misc Publ 6 Tallahassee Horida USA

166 Nickel 1 1 1961 Biological control of rice stem borer A feasibility study Tcch Bull 2 International Rice Rcarch Institute PO Box 933 Manila Philippines IIl p

167 Nigan P M and R A Vernia 1985 Insect pests of uplaild rice in Uttar Pradesh Int Rice Res Newsl 10(6)22

168 Nilakhe S S 1985 -cological observations on spittle bugs with emphasis on their occirence in rice Pcsqtui Agropecu Bras 20407-414

169 Nixon G EJ 195 1 lie association of ats with aphids and coccids Commonwealth Iusti lcte of Entonology Iondon UK 36 p

170 Nurdin F an Ishak Manti 1985 Study oit seedling fly of upland rice Interlational Upland Rice Moitoring Tour Report 26 lebruary-2 MarLI Sukaranti West Sumatra

171 Oka 1 N 1979 Ciltural conttol of the brown planthopper Pages 357-369 in Brown planthopper threat to rice production in Asia International Rice Research Institutc 1p0 Box 933 Manila Philippines

172 Ohni M and A Vilnaii 1975 Rice insect pests of Italy biology of Sipha grcria (Kaltenbach) and its control [in Italian] Riso (Milan) 2459-71

173 Otanes F Q atnd 1 I Sison 194 1 Pests of rice Philipp J Agric 12211-259

174 Painter R I 1951 Insect icsistance in crop plants University Press of Kansits Lawrence Kansas USA 520 p

175 Pantua P C and 1A litsinger 1981 Arthropod abundance in continuous vs biannual rice cropping systeis Int Rice Res Ncwsl 6(2)20-21

176 Pathak M ) and V A l)yck 1975 Sttudics ott insect pests of upland rice Pages 186-191 in Major research in upland rice International Rice Research Institute PO Box 933 Ilanila Philippines

177 Patil B R B M IHasabe and 1) S Ajri 198 I Chemical control of white gritb lIditriihinhorraia Fabr intfesting groundntut paddy atnd sorghum Indian 1Entomol 43232-236

178 Pawar A 1) 1974 (ollection and identification of insect pests of rice rlerminal repirt lInternational Rice Research Institute PO Box 933 Manila Philippines 72 p (tnimco)

179 IerrajU A aid A S Reedy 1965 Observations ott diapausc itt tie steti borer ohrice Rice News Teller (lai-Mar)9-13

180 Pont A C 1972 A review of the Oriental species of theri ona Rondani (I)iptera Muscidac) of econtomic importance Pages 27-102 in Control of sorglium shoot fly M G lomtani and W R Young eds Oxford atd IH1 Ptthl Co New I)clhi India 324 p

181 Prasad S C 1PChand aid M 1PSingh 1983 Incidence of rice gall midge in the upland plateau region of Hihar Oryza 20235-236

182 Rai P S 198 0 Nisaga simpex damage to rice in the hill tracts of South India Int Rice kles News[ 5(I)12

183 Rathorc Y S 1969 Gonocplhahli dcprssunt Fabr as itpredator of (tilo zonwIlhis (Swinhoc) and other moths in field cage at Patnagar Uttar Pradesh Indian 1Entotnol 3127

184 Reed W 1983 Crop losses causcd by insect pists in the developing world Proceedings 10th Internatial Congress of Plant Protection lrighton UK 174-80

185 Reedy M V 1981 Ateactogasterinhens(Walker)a new root-grub pest of tipland paddy and inaize Curr Sci 501076

186 Reveno M A 1968 Catalogue of the insect pests of economic crops in Colombia [in Spaiish] Asociacion Latino Americana de Entomo-logia (AIAE) 129-33

187 Reyes G M 1957 Rice dwarf disease in the Philippines Food and Agriculture Organiiation Plant Prot Bull 617-19

188 Rice S E A A Grigarick and M 0 Way 1982 Effect of leaf and panicle feeding byarmyworm (Lepidoptera Noctuidae) larvae on rice grain yield Environ Entomol 7593-595

189 Risch S J 1) Andow and M A Altieri 1983 Agioccosystem diversity arnd pest control data tentative conclusions and nizw research dirctions Environ Entomol 12625-629

190 Rivera C T 1956 Three organic insecticides in thecontrol of insects affecting upland rice Philipp Agric 39465-472

191 Roffrey 1 1972 Locusta outbreaks in the Philippines Acrida 1177-188

192 Rojas R 1) 1941 The effect of pruning at varying ages upon the yield of Ramairio Phipp Agric 29851-860

193 Roonwal M i 1979 Ternite life and termite control in tropical South Asia Scientilic Pnbl odhpr India 139 p

194 Rosetto C1 F da Silva Martins N C Schmidt and L EAzzini 1978 Damage caused by pasturespittle bugs 4Dvoi)lavopictaandDJ schach) on rice lBragantia Nota 6 (Matcli)25-27

195 Rosztto C1- S S Neto I- Amantc I) Iink 1Garcia-Vieira D M de SoLza N V Bansa t and A M de Olivcira 1971 Rice pests of Brazil [in Portuguese] Pages 149-238 inrFA()-International Rice Commission 2nd meeting of the Rice Committee for the Americas 6-11 l)ecember 1971 Pelotas Rio Grande do Sul Brazil

196 Rothschild G H L 1969 Obseratiois on the armyworm Spodoptera matritia acronrctoiiles Gn (Lep Noctuidae) Bull Entomol Res 59143-160

197 Rothschild G 1- L 1974 Parasites of the armworm Spolopiera mauritiaacroncitoihes in Malaysia Fntornopliaga 19293-299

198 Sachan i N S K Gangwar and 1N Katujar 1980 Field crickets damaging cages in North Eastern IHill Region Indian J Entomol 42526-527

199 Sands 1) P A 1977 The biology and ecology of Lepocorisa (Hemiptera Alydidac) in Papua New Guinea Res Bull 18 Departshyment of Primary Industry Port Moresby Papua New Guinea 104 p

200 Sarawak 1968 titomology division Pages 97-106 in Report of the research branch of the Department of Altriculture 1966 Kuching (Rev AIppl Eintomol A56603)

201 Satpathy JM M S Das and K Naik 1977 Effect of multiple and mixed cropping on the incidence of scine important pestsI Entomol Res 178-85

202 Sauer IHF G 1939 Notes Ott Elasmpnlpualtsligniosellths Zeller (Lep Pyr) a serious pest ofcereals in Sao Paolo [in Portuguese] Arq Inst Biol 10199-206 (Rev Appl Entotnol A28378)

203 Satinders 1L A 1 SKing and C L Vargas S 1983 Crop pests of Central Anerica with t bibliography [in Spanish] Centro Agroshynotnico Tropical de Investigacion y Ensenanza Turrialba Costa Rica (CATIE) Tec Bol 9 90 p

204 Sauphanor B 1985 Some factors of upland rice tolerttce to stem borers in West Africa Inseu Sci Appl 6429434

205 Sen P atnd S Chakravorty 1976 Stentflics in Haritighata Nadia West Bengal India [it Rice Res Newsl 1(12) 15

206 Senapai B and 1M Satpathy 1983 The seedling fly Atherigona orrzae Malloch is a pest of rice in Orissa (India) J Entomol Res 777-79

207 Shhjahan M S Alatn M Ah nad and BN Islam 1983 A new scarabaeid beetle pest of dryland rice in Bangladesh Int Rice Res Ncwsl 8(I)16-17

208 Sharma G L S C Modgol and R C Gautam 1979 Effect of depth of submergence ott incidence of bacterial blight and white grub infestation in transplanted rice Int Rice Res Newsl 4(6)12-13

209 Sharma S 1985 Scarabaeid beetle outbreak itt upland rice Int Rice Res Newsl 1(1(l)24

210 Shiraki T 1917 The paddy borer Schoenobhits incertulas Walker [achaku Agric Exp Stn Formosa 256 p

211 Silva A de Brito and 11P Magalhaes 1981 Insect pests of rice [in Portuguese] Circular Tecnica No 22 EMBRAPA Centro de Pesquisa t ecuaria de Tropico Unido (CPAJU) Belem Brazil 14 p

212 Singh S A 1979 Emergence of rice leaf butterfly as a rice pest in Manipur India int Rice Res Newsl 4(4)16-17

213 SinghS A 1983 Changing trends of insect pestsof rice in Manipur Int Rice Res Newsl 8(f)12

214 Siwi S S and P H van Doesburg 1984 Lenovorisa Latreille in Indonesia (Heteroptera Coreidae Alydinae) Zool Meded (Leiden) 58117-129

215 Soejitno J 1977 Reaction of some upland rice varieties toseedling fly eltherikonaexigua Stein [in Indonesian] Siri Hana I Penyakit 630-35 Lembga I osat Fenclilian Pertanian Bogor Indonesia

216 Socnardi 1967 Insect pests of rice in Indonesia Pages675-683 in The major inse pests of thu rice plant Proceedings of a symposium at the Internatital Rice Rec Wi itt S pternber 1964 The Johns Hopkins Press Baltimore Maryland USA

217 Soto P E A T Petcz and I W Buddenhagen 1976 Survey of insect pests and diseases of rice in different ecological zones in Nigeria Rice Entomol Ncwsl 435-46

218 Soto 11E and Z Ziddiqi 1976 Screening for resistance to African rice insect pests Varietal improsetnent sCmi n r Bouak Ivory Coast 13-17 Sept 1976 West Africa Rice l)evelopment Association Monrovia Iiberia

219 Squire F A 1972 Entomological problems in Bolivia PANS (PestArtic News Sumr) 18249-268

220 Stapely 1 II 1978 The rice leaf roller in the Solomon Islands Int Rice Res Newsl 3(l)13

221 Suenaga H 1937 On aphids attacking the roots of rice Oyo-I)oubts Zasshi 9150-152 [in Japanese] (Rev Appl Entomol A25745)

222 Suenaga H 1952 On the seasonal abundance of three subterranean aphids on upland rice Bull Kyushu Agric Exp Sti 1249-262

223 Sutynso Yas and Dandi Soekaria 1964 Beberapa hana yang ttenperang tana man padi tanah kering [in Indonesian] Rapat kerja Padi Tanah Kering 6 sd 8 Juily 1964 Kaliurang 9 p

224 Tallamy I) W and R F Denno 1983 Alternative life history patterns in risky environments in example from lace bugs Pages 129-147 in Insect life history patterns R F l)cnio and If Dingle eds Spring-Verlag 225 p

225 Tamura 1 1939 Observations on some insect pests of field crops [in Japanese] 1 Plant Prot 26414-420 (Rev Appl EntomolA27-556)

226 Tamura and Suzuki 1964 Influences of the early-transplanting practice in rice culture upon development of the rice stem borer Chilo siql)ressalisWalker Bull Ilokuriku Natil Agric Exp Stn 761-94

227 Tan J P 1924 The rice root aphid Drowhi ihirsoia Philipp Agric 13277-288

228 Tanaka 1 1957 Taxonomy and distribution of some subterranean aphids injurious to upland rice in Japan with description of a new species [in Japanese English summary] Botyu-Kagaku 22168-176

229 Tanaka T 1961 The rice root aphids their ecology and control Spec [full College of Agriculture Utsunomiya University Japan F3 p

230 Tauber M IC A Tauber R Nechols and 1Obrycki 1983 Seasonal activity of parasitoids control by external internal and genetic factors Pages 87-108 itt I)iapause and life cycle strategies in insects V K Brown and I Ilodek eds )r W Junk The Hague Netherlands

231 Taylor W F 1972 Effects of artificial defoliation (simulating pest damage) on varieties of upland rice Exp Agric 879-83

232 feotia T P S and S Natud 1966 Biotoomics of the rice skipper Parnaraniathiis Fabricius (Lepidoptera Hesperiidac) Indian J Entomol 28181-186

233 Teran F 0 1971 Rice pests of Santa Cruz [in Spanish Ministerio de Agricultura Sita Cruz Bolivia Bull 45 8 p

234 Than Itun 1976 Population dynamics of the yellow rice borer Tryporza itcertulas (Walker) and its damage to the rice plant MS thesis University of the Philippines at Ios Bafios Laguna Philippines 92 p

235 Uichanco L B 1911 Pests Pages 55-74 it Rice Philipp Agric 11-200

IRPS No 123 January 1987 41

236 van der Laan P A 195 1 Life history of the rice seedling fly Contrib Gen Agric Res Stn Boor Indonesia 118 15 p

237 van der Laan 11A 1959 Correlation between rainfall in the dry season and the occurrence of the white rice borer (Scirpolhaga innotala WIk) in lava Entomol Fxp Appl 212-20

238 Van Driesche R G 1983 Meaning of percent parasitismin studies of insect parasitism Fnviron Entomol 121 I1- 122

239 Venugopal M S and A litsingcr 1984 Fffect ofcarbofuran on rice growth Prot Ecol 7313-317

240 Vietnam Directorate of Rural Affairs 1962 Annual work progress report on crop improvement program of rice sugarcane vegetable and field crops (lor the period from July 1961 to June 1962) (11) 299 p (Rev Appl Entomol A51526-527)

241 Wan M T K 1972 Observations on rice leaf- and planthoppers in Sarawak (Malaysian Borneo) Malay Agric 148308-335

242 West Africa Rice I)esclopnient Association 1978 Annual research report Vol 2 Statistics Monrovia Liberia

243 West Africa Rice Development Association 1979 Annual research report p47 Monrovia Liberia

244 Williams ) F and CS lofgren 1981 Fli I illy l-468 anew bait toxicant for control of the red imported lire at Fla Entomol 64472477

245 Williams I) 1 1970 The mealybugs OFlomoptera Coccoidea Iscudococcidac) of sugar-cane rice and sorghum Bull Entomol Res 0109-188

246 Williams D) 1 1982 Pulinaria icervi (Signoret) and its allies ott sugar cane and other grasses Btll Entoinol Res 72111-117

247 Wirjosuiardjo S 1976 Study ofchenmical cont rot of insect pests ott upland rice Imiu Pertanian (Jogjakarta) 1323-329

248 Wirjosuhardjo S and N Wirarta 1976 Chemical control of the white grub tIolorithiatlltriin upland rice in M uneng Irobolinggo Indonesia) [in Indonesian] Imut Pertanian (Jogjakarta) 1330-332

249 Yano K T Miyake and V F Eastop 1983 The biology and economic importance of rice aphids (Henuiptera Aphid idae) a review Bull Entomol Res 73539-566

250 Yelp D and P Rhodes 19(4 lhe success of drilled dryland rice iii the Fiji Islands International Rice Commission Working Party on Rice Production and Protection Manila Philippines 3-10 March1964 Food and Agriculture Organization Rome Italy 13 p

251 Young W R 1981 Fifty-five years of research on the sorghum shootfly Insect Sci Appi 23-9

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ISSN 05-3862Other papers in this series TITLES OF NUMBERS 1-45 ARE LISTED ON THE LAST PAGE OF NO46 THOSE OF NUMBERS 46-70 ARE ON THE LAST PAGE OF NO 71-80AND THOSE OF NUMBERS 71-1O0ARE ON

THE LAST PAGE OF NO 101-122

No 101The economics of hybrid rice production in (hina No 113 Boron toxicity in rice No 102 Rice ratooning No 114 Energy analysis rice production systems and rice research No 103 Growth and development of the (Jeep water rice plant No 115 Production risk and optimal fertilizer rates an application of the

No 104 Faridpur acomputer-assisted instruction nmodel forrainfed lowland random coefficient model No 116 Consumer demand for rice grain quality in Thailand Indonesia andrice

No 105 A readingand listening comprehcnsion test inEnglish for non native the Philippines speakers applying for tiraining at IRIRI No 117 Morphological changes in rice panicle development a review of

No 106 Rice grassy stunt virus 2 a new strain of rice grassy stunt in tire literature

Philippines No 118 IRRI-Korca collaborative project for the development of cold-No 107 Physical losses and quality deterioration in rice plstprodnCtion tolerant lines through anthei culture

systemis No 119 Problen soils as potenial areas for adverse soils-tolerant rice No 108 (olptblication ofIRRI materials a survey of translators and varieties in South and Ssuthcasi Asia

publishers No 120 Changes isinall-farm iLtireliing techii)hgy in Thailand and [he No 109 Classification of Philippine rainfall patterns Philippines No 110 Contributions of modern rice varieties to rutrition in Asia No 121 Landforms and modern rice varieties No Ill Changes in rice breeding in 10 Asian countries 1965-84 No 122 Yield stability and modern rice technology No 112 Design parameters affecting the performance of the IRRI-designed

axial-flow pump

Page 13: UPLANDRICE. INSECTl--1i M T -E!R.EOLOU t RTANCE ,AND …delphacid.s3.amazonaws.com › 6958.pdf · The significance of abiotic factors in upland rice insect ecology will be apparent

can disperse long distances to colonize even remote upland areas They become more abundant than their natural enemies particularly after favorable rains following a prolonged drought (196) The drought kills the armyworm and natural enemies (mainly parasites) but the armyworm can recolonize rapidly

The natural increase insoil fertility from mineralization of soil N over the drought period promotes luxuriant plant growth to foster a rapid buildup of these highly fecund species Weeding rice increases tilelikelihood of damage The parasites normally return to control the armyworms but not until after serious damage

Outbreaks on upland rice have been reported in Ghana (3) inzibar (38) Central Africa (36) lPanama (165) Brazil (28) Malaysia (196) and India (167) Defoliation may be severe often leaving only the base of stems Armyworm larvae hide during the day under litter or in soil cracks At night they ascend plants to feed Larvae pupate in the soil

Thrips Thrips frequent upland rice but damage is less than in the lowlands Thrips larvae and adults feed on leaf blades by rasping causing leaf rolling and stippling Thrips prefer vegetative stage rice and their survival increases during drought (heavy rain washes them from foliage) although drought stress usually outweighs thrips damage Drought and thrips cause leaf rolling in upland rice (235) but thrips have been overlooked perhaps becaLse of their small size Thrips are most numerous indry season irrigated rice The combination of favorable plant growth and lack of rainfall is ideal for thrips In contrast the lack of rainfall in upland rice hurts the crop and eventually the thrips whose numbers must decline

In [atin America Frankinilla rodeos attacks panicles before they emerge from the boot calusing sterile grains

Stem hug~s Black bugs are the most common group of rice stem-feeding -lemiptera in Asia The most frequently mentioned species ----Scotinophara coarctaia and Shrida - prefer aquatic habitats and therefore could not occur in upland rice as sometimes reported Upland species in the same gerurs - notably S tarsalis and S scotti -- are morphologically similar and remove sap from stems of grasses and upland rice The upland species are notably more abundant in upland rice planted near forests

In Latin America tile Tibraca ima-large pentatomid tiventris causes deadhearts and whitehcads on upland rice Both adults and nymphs remove sap from internodes of plants more than 3 wk old Tillers are killed by mechanical damage and by the entry of secondary microbial infection This damage is often confused with that of lepidopterous stem borers (233) Eggs are laid on the leaves and stems Nymphs are gregarious This species can also be found on irrigated rice Drought reduces population buildup

Stiem borers Lepidopterous and dipterous stem borers are widely recorded on upland rice but cultural practices and alternate hosts determine abtndance Except for Elasmolmll)uS Iinosellts numbers usually build up toward

IRPS No 123 January 1987 13

the end of the crop growth cycle Therefore late-maturing varieties staggered planting and lack offloodingfavorstem borer buildup Many stem borer species can lie dormant in the stUbbleafter harvest and theirsurvival is encouraged by panicle harvesting (leaving tall stems) and lack of tillage after harvest

Stem borers disperse and lay many eggs Many species are polyphagous however the monophagous MaliarpiasepashyratellaDiopsis Ivmgicornis Scirpophaga incertlas and S innotata can also be abundant on upland rice (hilo Suplressalisdevelops fasterand becomes la rger when reared on upland than on lowland rice (226)

The lesser corn stalk borer lasmopal tt Ittoselhs is semisubterranean and is perhaps the stern borer most adapted to upland rice(129) Highly polyphagous it infests maize peanut and cowpea It attacks seedling rice (one larva can kill tip to four plants before maximum tillering) (195) (hiv and Maliarphastem borers prefer older plants But Scirpophagaspp will attack young plants even in the nursery lasn opalls larvae tunnel into the stems at or below the soil level mausing dcadhcarts Larvae are not found in their turnnels inside sterns beca use vhei disturbed they retreat into cases made from soil particles bound by silk The cases areattached to the tunnel entrance (143) This behavior probably evolved as protection from natural enemies Larvae pupate in the soil The lesser corn stalk borer is more abundant during drought damage is often confused with that of drought (233) This species prefers sandy soils arid its distribution within a field may relate to soil texture (143)

Diatraeasaccharalis aside from the lesser corn stalk borer is frequently a stem borer pest in Latin America According to Teran (233) rice and maize are more preferred hosts than sugarcaine its namesake

Atcitonacrvsograpla is a less known upland rice stem borer in Asia with habits similar to Easinopall s (82) because it is not found in lowlands Acutra has grassy weed alternate hosts As a moth it is often confused with the dark-headed stern borer(hi polur(standstriped stern borer (hi sup)lpsrcais

Other common upland rice stem borers are also reported in the lowlands The dominant species in upland rice vary greatly with location In Uttar lPradesh India (78) and in Japan (123) the pink stem borer Sesamia htfirens is more prevalent hIKenya the principal rice stem borer is Chi partelus

(92) The lowland African sten borers Aialiarpha sepashyratellaDiopsis vngicornis and Sesamia calatnisnis are less abundant Oimupland than on lowland rice(217) but perhaps cause greater yield loss because plant injury is greater if combined with drought css (92) The stalk-eyed stern borer D)ivpsis is more prevalent in wet habitats (6 1I217)

In Central and West Africa Maliarpha and Sesamia calanistis are the dominant stern borers of upland rice (8 9 10 II) (lilo dihlfilineui and Chio partellus inhabit the

14 IRPS No 123 January 1987

upland savannas whereas Chilo zacconio prefers the lowlands (II) Aaliarpha is most abundant in the mangrove swamp habitats (91) coastal regions (16) or rain forest zones (II) of Africa and appears to be the homologue ofScirlpophaga incertulas The yellow stem borr can also be prevalent in upland rice (120 167) The sugarcane borer Eldanasaccharinais so far only a potential pest of uplandrice (91) despite Grist and evers (82) warning

Grasshotl)ers andI locitas Short-horned grasshoppers and locusts -- Locvta Patanrga Schistocerca - layeggs in the soil inhabit grasslands and develop into swarms to seriously damage upland rice from time to time (173) Largenumbers can destroy a field leaving only the stubble In low cmuoroer s giassnoppeirs an cut panicles (173) Farmers may not plant if locust swarms are imminent Prolonged dryweather followed by favorable rain favors development of large swarms (173) Usually breeding in di) grasslands theydisperse toattack crops including upland rice(191) Eggsaresusceptible to desiccation d uring drough t and toinundation during seasonal rains With favorable soil moisture over consecutive seasons natural enemies ca nnot prevent a largepopulation increase leading to migratory swarms (64)

Most grasshoppers on rice are adapted to uplands (107)but lowland rice grasshoppers iervgl(rlhusspp and Ora spp occur occasionally on upland rice (2 82) They lay eggson rice foliage Nymphs are scmiaquatic

The gryllid Lsctrtus coucinnus is a pest of lowland and deep water rice in Bangladesh and Thailand In the Philippines it damages upland rice grown in isolated pockets near lakes The gryllid nymphs and ad ults feed on the central portions of folrage leasing only the midrib and leaf nargins

Leal ler Pvralid moths whose larvae fold leaves to make a feeding shelter occur worldwide (217) but arc appareritly most common in Asia Leaffolde rsattacking rice in Asia are a complex of species (26) (Ctalhahcrcisinedinalis the best-known rice Ia flolde r in Asia may prefer grassy weeds over rice (86) It is a late colonizer of uplandrice U1arasmja eviua a lowland species (220) nreflers rice to grass specres (86) and colonizes upland rice during il early growth stages llarasnria patnalis is the most corn-rnonly encountered species in upland rice in the Philippines Perennial grasses are their alternate hosts and they can survive year-round in upland rice environments The leaf-folders 11arasmia ruralisand Alarasmia spp are least aburndant Dormancy is unknown in leaffolders Cnaplha-ocrocis medhnalis however is known as a migrant (90)Leaffolders remove photosynthetic tissue attack during the flag leaf stage is particularly injurious Popurlations are normally held in check by natural enemies but leaffolders become parlicularly abundant in conditions of high plantfertility and shade

Butterflies Skippers PelopidasInathiasand Borhofanta are more prevalent in African lowland than in uplandhabitats (6) but extensive studies in Japan and China

showed that Pelopidas mathias prefers an upland habitat (155) Parnaragutata seasonally migrates between lowland and upland areas and is equally abundant in both environshyments (164)

The green horned caterpillar Melanitis leda ismene is more abundant in lowlands (212) but our experience in the Philippines suggests that Pelopidas niathiasand 11elanifis leda ismene are often more abundant in uplands than in lowlands Bamboo iaunbuiva spp annually sustains P mathiaspopulations in the Philippine uplands Adults feed on nectar from flowers and migrate to upland rice during the late vegetative stage

iPi erlled h i-1 1d dill L tIC i 7-0 l(become numerous probably because of high egg predationEgg and larval parasitization are often low Butterflies can readily disperse long distances to seek renlote uplandricefields However they have low biotic potential and even it predation rates are low the rarely become abundant Upland environments with their diverse microhabitats can provide more favorable sites than lowland rice plains

tutterfly pests of rice have broad host ranges and become dormant Juring urnfavorable times of the year

IolhagrotslaidopwraThe larvae of many polyshyphagous moths are reported to defoliate upland rice tussock moths lsaispelnnatula Iawlia sUifita luproctisirgtwltaE mnorand P vathorrhoeaslug caterpillar

litotiahicolorhairy caterpillars Nisaga sinp vandA mata spp and wooly bears l)iacrisiaohliqtia and Crvatonotts gangis Many of these polyphgous larvac migrate and are transients in rice including HelhWhis arm iera and 1t zea highly pol phagous species (82)

Gall nidges The Asian rice gall midge Orseoliaorrzae (88 131) and its African counterpart 0 ortzivora(6 11 12 32217) prelerlowland to upland rice Gall midges in rainfed areas are prevalent only during tle rainy season and dispersepoorly Their alternate hosts (wild rices LeersiaIschaetnuan and Pasahmt)are mostlyaquatic grasses and are less likelyto be near upland rice areas In an area of mixed uplandfloating and rainfed lowland rice cultures in northern Thailand upland rice which is ilanted a month earlier was attacked firstbut subsequcent populations were much lower than in rainfed lowla id or floating rice Low numbers of gallmidges passed the dry season orr perennial weeds such as Paspahim (88)

Smot (hid(s Reports of aphids attacking upland rice plants are few (249) Without maize the maize leaf aphidRho alosihhunm maidis attacks upland rice in the Philipshypines The plum peach aphid IIrsteroneuraselariae was reported in Sierra Leerne toinfest leavesand tnriperied grain(7) Siphaglrctriaecaused transverse linrea r necrotic striae iir rice leaves in Italy (172) I-lea v rains held populations in check Rice is not a preferred host for shoot aphids

Leafhoper andI )anthol)pers Cicadellids and delshyphacidsdispersecasilyand readilycolonie upland ricefrom nearby lowland areas A study in Sarawak (241) showed

IRPS No 123 January 1987 15

similar species in upland and lowland rice Nephoiettix The principal vectors of rice virus diseases generally have species dominated The most prevalent species accounting narrow host ranges and alternate hosts in the grass family for 90-95 of the sweep net samples wre N virescens N are mostly annuals dying in the dr3 season Some virus umropictus Nilaparvia hgens Soatellafiircifera Cfana vectors such as BPH can disperse lontg distances but few do

sectraand Recilia dorsalisLowlands had higher popula- Th restriction of upland rice and weed hosts to the rainy tions Katanyukul and Chandaratat (120) monitored hopper season the small area planted to upland rice and the hopper numbers in upland and lowland fields of northern Thaiiand vectors having narrow host ranges all work against virus and found similar results A comparative study of plant- diseases in upland rice hoppers in Fiji showed more brown planthoppe BPH) and In Latin America Sogctdhs o(rzicola and S cuhanus whitebacked planthopper (WBPH) in wetland rice than in transmit hoja blanca virus disease but reports on upland lowland rice and hopperburn was prevalent in lowland rice are rare except in the favored upland areas of Colombia areas with standing water (89) Peru and Venezuela (35 42) S rricolaocurs in Brazil

Hopperburn in upland ricefields in Asia is rare (84) In on upland rice but hojaI blanca has not been reported (67) Batangas Philippines isolated fields of upland rice become S orzicola has a wide host ramgc (10) similar to WBPI- in hopperburned infrequently from WBPH A report from Fiji Asia WBPH however is not known to transmit any virus cites hopperburn from BPH and WBPH (250) In these two disease situations upland rice was grown on highly fertic soil and In Africa pale yellow mosaic virus is endemic to swampy N fertilizer was applied Hoppers respond to better nutrition lowland areas where rice ratoons year-round (19) and is Normally upland rice is grown on poor soil and receives no mechanically vectored mainly by hispid beetles A minor fertilizer vector a flea beetle Chaeiocnema spp is an upland rice

In Asia upland rice ras are normally not far from pest and upland ricetields near endemic areas occasionally lowland ricefields-- a source of hoppers Otherwise hopper contract the virus species with greater polyphagy than BPH or Nelhotettiv Spittle bugs Cercopid nymphs producc a protective virescens would be dominant frothy covering that looks like saliva In Latin America

In Africa leathoppers and planthoppers do not become spittle bugs damage upland rice In parts of Brazil where abundant on upland rice (6 II) probably because lowland upland ricefields are surrounded by pastures adults rice is not widely rlanted nearby The only reported migrating from grasses can damage rice severely and cause hopperburn as from Nilapanrata maeander in breeder complete hopperburn in a young rice crop within a week of plots receiving high N rates in Nigeria (98) Hoppers colonization (194 195) Dispersal to rice occurs even if however are a greter problem in Latin America where pastures are verdant and spittle bugs prefer pasture grasses large grassland areas breed polyphagous species (Gra- to rice (168) Nymphs are rarely abundant enough to phocephala spp Hortensia spp lEvitianus ohscurunervis damage rice Regular rainfall favors their development Balclutha spp and Draeculacephala spp) vhich then (233) disperse to cause hopperburn on seedling rice (68) Rosetto et al (195) reported Graphocep7hala spp and Hortensia spp Seed pests to be equally abundant on upland and lowland rice Alydid and pentatomid bugs are seed pests of upland rice

The leafliopper and planthopper epidemics in Asia that worldwide In Asia Leptocorisa sp prevail Their habits caused severe losses from hopperburn and vijus dicases make them particularly suitable for upland rice 1)thev feed over the past several decades occurred principally on on many grasses (and therefore can survive the earl y wet lowland rice Except in a few isolated cases (160) upland season before rice sets its grain) 2)adults are long-ied and rice has been spared from tungro grassy stunt and ragged mobile allowing them to find isolated plantings 0 rice stunt virus diseases vectorcd by green lcafhopper (GLH) (besides seed bugs only birds and rats appear to have shyand BPI] (123) Virus diseases have occurred in upland rice capability) and 3) adults aestivate in forested areas or where lowland rice was nearby Rice dwarf disease however sugarcane fields during the dry sca son when neither rice nor was first recorded on upland rice in the Philippines (187) grassy weeds are present (I199) Rice seed bugs concentrate Virus diseases can be perpetuated only on livir plants or on small-scale upland riceficlds because they can actively insect vectors and no living rice - plans tooned or search them out The prevalence of groves of trees characshyvolunteer - grows year-round in upland arcas excert in a teristic of upland environments ensures nearby aostivation few with 12 mo of rainfall sites keeping the seed bugs close by

The smaller brown planthopper Laodcelphax striatellu Leptocorisae are uncommon in lowland rice plains occurs in upland rice in Japan (18) It spreads black- because 1) their populations are diluted in a sea of rice streaked dwarf and stripe viruses between winter wheat and planted more or less at the same time (they ca n only feed on barley to lowland rice Early planting of lowland rice rice during the milk to hard dough stages) 2) they lack nurseries spreads these viruses but upland ice has remained aestivation sites and 3) weedy fallow areas are limited uninfected The lowands of Asia are dominated by L oratorius

16 IRPS No 123 January 1987

which lays its eggs hign on tile foliage (199) L acuta and L solonlonlesis oviposit at ground level the former on litter and the latter loose on the ground These egg laying habis explain theirtenvironmental prefcrencecs(214) In Japan the postdiapause adults of L chinensis migrate from moun-tainous areas to upland ricefields during floweiing (1 14)

Pentatornid bugs prevail in Latin America (8182 195) and in Africa pentatomid and coreid bugs arc equally cited (6 82) l)amage is often characterstic of the species (5)rcoris ventralis and Mnida spp are the common

pentatomid seed bugs of Philippine upland rice but they prefer lEchinochva spp grass seed to rice Vearaviritda occurs worldwide and hoften more abundant in the uplands(54) We saw few reports of seed bugs on upland rice but see(d bugs apparently opelrate cqually between by land and upland rice Good dispersal is necessary ior species that call feed for only 2-3 xon a crop and must first find those crops at ripening stages

Other pct species feed n flowers In Mato Grosso Brazil (194) and Santa Cruz Bolivia the panicle weevil Aeo-hariuia amllitarsis attacks rice during flowering (68 233)The larva e rice sten horers adutlts feed at the bases of spikelets caIusing empty grains Also inLatin America thecydnid -lkiituttatralusisa seed bug as anad tilt and nymph (194)

A number of beetles ecd on poilen - coccinellids lialrotia Aulac lhora and Monohpl-a (55) - but becatise riCe is self-pollinating and fcrtiliation occurs before pikelets open they pose no threat to yield

IST ECOLOGY

We now look at characteiistics of the life cycles of upland rice insect pests to lea rn hew each plays an adaptive role in upland rice ecology

Life history strategies Orzasativadoes not tolerate drodgh well so upland rice is highly seasonal normally grown in the wettest months of the year Usually rice is present for less than half of the year The rice-fiee fallow poses serious problems of survival to upland rice insects which have evolved att least four mechanisms to ovrcolne tle cvcl ic-l lack of a hostI) polyphagy 2) longevity 3) dormancy and 4) vagility (dispersal)

Poldt)agy lolyphagy is defined as having hosts of at least two botanical families oligophagy of more than one gents and ionophagy of only one ventus (40) - inour case Orvza Most wild rices however are illi tiC and theieiirc are normally far removed from upland rice hahitat othewise more would be alternate hosts for uplanJ iice insects Thte highly mornophagous rice pests shySciriol)haita hiceritlas Nilal)arvaiahiens Iand Nealo--tettiyvircsctws in Asia and Aalirpha secparatella and 1ioj)sis lngicornis inAfrica - occur on upland rice but

are more abundant in lowland culture They can specialize in rice because they are highly vagile and can attack all growth stages By specializing in rice monophagous rice pests can outcompete related but polyphagous speciesIntensive andI extensive lowland rice culture has favored monophagous species Riqwla alhinella (Cramer) the onlymonophagous species in Latin America is rarely reported on upland ice (I1)

Because of its limited temporal and spatial existence upland rice has not favored the evolution of specialized species Clhang(44)shows evidence that upland rice has only recently been cultured by man from the lowlands thus pests have had little time to fully adapt to the crop All but the lowland-adapted insect pest species have alternate planthosts to rice Root pests with limited mobility are highly polyphagous Pests that specialize in one growth stage eg secdling maggots seed bugs are oligophagous or polyphagous

Most alternat hosts of rice pests are annual grasseswhich also are seasonal Upla_rice is also a host but not the preferred one of the most highly polypliagous spcciessoil pests grass hoppers armvwarrs and Vearaviirielda

The d ry season poses igreat obstacle to ani upland riceinsects survival and only sonic perennial plant hosts are suitable food during the off-seasoin Plolyphagy allows upland rice insects to suLrx-ixe during thC rainy season when plants- both annual and perennial -areactivelygrowing and therefore more nutritious

Loiugevitr The ability of aiinsect to live 2-3 mo without undergoing dormancy helps species sich as root-feedingwhite grbs (as larvae) and secd-feeding bugs (as adtilt) to survive unfavorable periods White grubs arc not highly fecund and their life strategy is to improve survivorship of limited progeny Their subterranean habitat hides them from manv natural enemies lBtut root tissue is not highly nutritious so they ltilst eat great quantities and need a largeinsect biomass to digest it Other soil-inhabiting rootshyfeeding insects have short life cycles but are more fecund

Seed pests need greater longcvi vas adults to locate ahost at early grain dcclopment arnd give sedentary offspring time to develop Few groups of upland rice insects live longin active development Polyphagy and longevity only increase their ability to survive during the rainy seasonOther mechanisms are needed to survive the dry season

orantc In the tropics the nonrice season is the dry season while intemperate areas tile off-+season is winter Many upland rice insects undergo dormancy inlsummer (aestivation) or winter (hibernation) Dormancy is simplyinactivity during winters with prolonged temperatures below 10 C insects find shelter arnd cease movement Activity is resu med with warmer t empcera tu res A deep state of irnactivity termed diapauise occurs in sole species that prepare physiologically (havxing low metabolism and convcrting gonad s to fatty fuel reserves) for the unfavorable season (148) They enter diapaUse inanticipation of unshy

IRPS No 123 January 1987 17

favorable weather and resume activity after favorable Armyworms skippers and locusts actively travel hundreds weather begins usually in response to photoperiod or of kilometers in air fronts such as intertropical convergencethrough a delayed response to a stimulus such as heavy zones to descend onto upland rice fields far from their rainfall (237) Not all individuals of a population have the breeding grounds (31) Several generations can build ip on same threshold for environmental responses for entering or alternate hosts in tile grass family before rice is plantedterminating diapause -- this is seen as a mechanism to These large insects are active fliers Smaller insects such as prevent total mortality if a false cue appears such as a thrips aphids and planthoppers passively migrate long sudden warm spell in winter or rain in the middle of the dry distances by the wind season (154) Otherspecies ean readily travel tens of kilometers - stem

Probably more upland rice insects undL go dormancy borers leaffolders leafhoppes cutworms spittle bugs leaf than we realize Candidates are Atherigona spp root beetles seed bugs ants and termites aphids root weevils crickets leaf beetles thrips and seed The least dispersive species are white grabs mealybugSbugs Insects known to entervarious states of aestivation are seedling maggots crickets weevils and gall midgesleptocorisaspp adults (199) Maliarphascparatellalarvae Each species has evolved a unique set of attributes to (9) Scirpophaga innotata larvae (117) Chilo partelhtsv enable it to survive and adapt to the changing uplandlarvae (21) ScirpopThagainccetulas (179) white grub larvae environments Several evolutionary avenues lead to fitness (134) Patangasuccincia adults (6 82) and Schistocerca (Table 2) but there is no reason to believe one set of gregariaeggs (82) attributes is better than another (224)

Insects known to enter hibernation are LodeIphaDstriatellus nymphs (124) Chtio Tvlpressalis larivae (125) DroughtSesaia inferens larvae (146) but not S calavisis(2) Well-drained rainfed upland soils are subject to drought ifIDiaraea saccharalis larvae (75) Scirpophaga- tcerlttas rains do not fall within 2-3 wk Prolonged drought followedlarae(5 lacchrli s lae (75)saSlOhagasshoperseggs (82)and felo pidas athias pOpae (232) by favorable rains stimulates armyworm and locust outshybreaks but mealybtigs root aphids and thrips become

Lcpmotorisa oratorisand L acuta adults aggregate on numerous onlyafter dry spells for two reasons First heavy trees sugarcane or other shady moist sites L paluwanmsis rain normally kills soft-bodied foliage-feeding thrips and confined to Sulu and Palawan Philippines lives on the mealybugs and soil-inhabiting root aphids and root mealyshygrass Brachiariamutica (Forsk) Stapf in the absence of bugs Drought eliminates this source of mortality Second rice The adults can live for 5 mo When aestivating their the rice plant responds to drought by breaking down rate of metabolism drops and they do not feed They will proteins into soluble N compounds which enter the phloem however puncture plant tissues of their aestivation host in and are taken up by these sap-feeding insects (245) Greater search of water This behavior has led to errors of the host nutrition therefore leads to greater survivorship and range of rice bugs (I) Aestivation quickly terminates with fecundity rain probably meaning aestivation is not deep Small rice area

Some stem borers aestivate as mature larvae in rice Except in the highly mechanized large landholdings of stubble or in straw stacked as livestock feed However Latin America upland ricefieldstypically are patchy within moriality increases proportionally with the duration of the a highly diverse flora The small fields result in higher ratios dry season even if the stubble remains undisturbed If the of perimeter to area than with the typically larger lowland stubble is plowed for a crop following rice few stem borers ricefields Upland ricefields themselves may be intercroppedsurvive Rainfall terminates dormancy but it normally with cereals legumes or root crops Mixed intercroppingtakes several weeks for moths to develop and emerge with a wide variety of species also occ in tribal slash-andshy

Last-instar white grub larvae tunnel 1-2 m deep in the soil burn agriculture (50 74) to construct pupal chambers in which they aestivate The Upland ricefields tend to be small because of the labor larvae are very sensitive to dryness so pupal chambers are needed to clear land in slash-and-burn areas and to till land sealed to coierve body moisture Aestivation terminates to minimize weeds Rice requires more tillage than does with the first heavy rains but as in stem borer 3-4 wk pass maize because rice competes less well with weeds Uplandbefore adults develop rice is more a subsistence than a cash crop and small areas

Insects may respond differently in each region to cold (I ha) can feed a family temperature or drought Biotypes or loal populations may Small fields favor some insect pests such as some ant evolve (56) Local populations of Scirpophagaincertulasor species that can forage from more permanent field borders Patangasuccincta may have become adapted to prevailing Trees provide food and mating sites for white grubs (134) as conditions well as aestivation sites for LetTocorisa spp (199) Field

iagiliti The ability to disperse combined with high border grasses plus tIhose within a ricefield provide alternate fecundity orshort life cycles enable some upland rice pests to food for all but the highly monophagous upland rice better exploit the temporarily favorable upland habitat colonizers and a habitat for natural enemies

0 z

Table 2 Life history patterns of upland rice insects or insect groups

Host range Longevity Dormancy Vagility FertilityMonophagy Oligophagy Polyphagy Low Medium High Yes No Low Dispersive Migratory Low Medium High(Oryza spp) (grasses sedges) (angiosperms) (lt3 wk) (gt2 mo) (lt1 kin) (lt50 eggs) (gt500 eggs)

Ants xTermites x x x

Mole crickets x x x xx x xField crickets x xx

White grubs x x x xx xRoot aphids x x xxMealybugs x

x x x xxSeedling maggots x

x x x x xLeaf beetles

x xx xArmyworms x x xx x

Thrips x x xx xChilo spp x x xx xMaliarpha x x x x

xLocusts x x Xx x xGall midges x x xxiVephotettL iirescens x x xx x x xLeprocorisaspp x

x xLeaftolders x x x

Sogatellafurcifera x

xx x x x x x x

00

IRPS No 123 January 1Q87 19

All this appeals to favor insect pest buildup but the notwithstanding the release of stored nutrients in response contrary is true Loevinsohn (138) showed that insect pest to drought that temporarily benefits philoem feeders populations respond exponentially with the proportion of land devoted to rice up to about 75 of the area then the M POR ANCE I I P1A 1) RWI P1St S

rate of response declines In most upland rico areas tie rice crop occupies less than 50i of the area- therefore the Visits to upland ricefields normally reveal 1kw insect pests potential for population buildup is low This relationship Loevinsohn (138) and Locvinsohn ct al (139) compared may not hold for seed pests that can locate small ricefields insect abundance using annual light trap atchcs in upland

Another outcome of the highly diverse flora and phical rainfed lowland and irrigated lowland sites in the Philipshyenvironment of upland rice isthateach upland ricearca has pines Although only one upland site was studied data a unique composition of insect pests showed equal or fewer insects at the upland site than at the

For example in Tanauan Batangas Philippines the rainfcd lowland sites and fewer than at the irrigated sites main upland rice pests are rice leaffolder(RLF) armvworm (Table 3) M Lltiple regression analysis showed that the key Aythntnaseparata WBPH ants Soli psis gentitataand factors explaining insect abundance among a set of cropping white grub Leucopholis irrorata Sixty kilometers away in intensity variables were the number of rice crops grown per Real Quezon the main pests are seedling maggot Athe- year followed by area devoted to rice Table 3 shows that rigona orae flea beetle (Thacitcmnra hasalis thrips the upland rice site had only I rice crop perym rand that rice Stenchaetothrips hiurnmis and rice bug Leptocorisa was planted in only 20Vi of the area Fertiliter and oratorius In a third Philippine upland rice site in Claveria insecticide use had little bea ring on the trend The armters at Misamis Oriental Mindanao the main pests are ants the upland site applied 60 kg N ha higher than at most Solenopsis gemtinata seedling maggot Alieriona orvzae irrigated sites white grubs Holotrichia in)iUaiuoan t and Lucopholiv Insect occurrence at the upland site was omparable to irrorata stem borers Sesamia ifiArelr (hio auriciliusC that at the Cagayan rainfed lowland site parlicularly in 1982stpessallis Scirpophaga incertulas S innoiata and when drought prevented many farmers fron planting(only

4c ona chrrsurapella root aphids Tetraneura niriah- 40(of the rice area eventually was planted that year) M ore dunrinalisand Rhupaosiphunm riahbdminalis lcaffolders favorable weather occurred in 1980 and 1981 which Marasniapataltis and Chaphalocrocis Medinalis army- supported greater rice areas and consequently more insects worm MIthinna separata and rice bug Leltucorisa The most intensive site for insect pests was South Cotabato oratoriusThe great diversity of insect problems is typical Mindanao where farmers plant 23-25 rice crops per year fer upland rice and makes control efforts more difficult with irrigation The Zaragoa Nutva Ecija site also had particularly regarding breeding for insect resistance high pest abundance and at this site the fields were highly

asynchronous Asynchrony was the third most important Low yield potential Iactor in explaining rice insect abundance Most upland rice is grown on low-nutrient and mineral- Rice cropping intensity in upland areas is bound to toxic soils which when combined with eiratic rainfall remain low Only one rainid upland crop is possible per make upland rice a highly riskycrop for the farmerto invest year (Farmers in Claveria Misamis Oriental Mindanao in costly soil amendments and land management Often Philippines tried asecond upland ricecrop i 1984a ycarof upland rice isgrown far from markets making inputs even favorable rains but no harvest was possible b-cause of more expensive Farmers growing a subsistence crop hosi- drought blast and birds) Rice area isaso bound to remain tate to invest in costly inputs to raise yields low because of the high labor and power rcquirelmenlts to

Even having varieties that could double existing yield prepare the land (because of weeds)and because upland rice potential under present management levels would probably wvill not compete with lowland rice as a cash crop Some not prompt many farmers to use purchased insecticide upland rice sites with prolonged rainy seasons howeverare unless tile risk of crop failure were reduced such as it was planted asynchronously such as the Clacra site with irrigation systems in lowland rice The seasonal abundance of Upland rice insect pests

The Tanauan Batangas site is atypical for upland rice attracted toa light trap wasgrapied for crop year 1980-81 in The soils and rainfall pattern are favorable and because of Tanauan FBatangas (Fig I 2) W13P11 was he most its nearness to Manila farmers have cash resources ftm common rice hopper and its population peaked at the sales of vegetables to purchase tertilier They applied an midgrowth stage as it does in lowland rice D-Vac suction average of 60 kg Nha after particle initiation and obtained samples showed an earlier peak than that from a light trap 25-35 tha yields The high fertility isone reason why RLF indicating emigration W13PI usually emigrate after the armyworms and WBPH developed into large numbers vegetative stage Populations in 1980 reached levels that Farmers however used no insecticide on upland rice caused patches of slight yellowing in some fields WBPH although they did on vegetables probably cause hopperburn in Batangas They immigrate

The poor growing conditions fOr rice generally mean year-round as shown by low levels in the light trap BPH poor growth and low fecundity for insect pests fIeding on it however were scarce in the light trap and not collected with

Table 3 Abundance of rice insect pests and factors in rice cropping intensity of Philippine upland rainfed lowland and irrigated lowland locations where light traps were operated daily for atleast I yr -I

0Rainfed Rainfed lowland Irrigated lowlandupland

Pangasinan Cagayan lloilo Santa Maria Nueva Ecija South Cotabato Tanauan Caaringayan Caaringayan Bangag Bangag Bangag Buray

LagunaCale

Batangas Manaoag 1980-81 Santa Motiica 1982-83 Cabanatuan Zaragoza Avancena NaniamanSolana 1981-82 1982-83 Oton

1980-81 1979-80 Oton 1981-82 1981-82 Koronadal Koronadal1980-81 1979-80 1979-80 1983 1983

Insect (kerosene light trapNephotcttixcatch nolyr)virescens 650 1240 500 1800 330 210 3400 1600 160N nigropictus 210 - 360 18400 5500 2900- 140 90 20 - shy 1700 -Reciliadorsalis 710 - 3200 1700180 890 630 140 170 2400 800Nilaparratalugens 130 690 - - 4200 4600300 520 750 180 60 3300 1600 740 310Sogatellafiircifera 1700 31000 13000 8200310 210 3300 610 270 980 1600 6300 -Cyrrorhinus - 12000 6700600 540 3200 290 160 330 120 410Scirpophaga spp 50 410 620 - - 21000 12000150 180 220 270 30 1000Cnaphalocrocismedinalis 70 70 540 610 270 11000 6100200 140 100 6 60 250Chilo suppressalis 0 60 60 1900 300 200- - 30 20 0 6 3 160 -C auricilius - 0 00 - - 20 3 60 20 3 0Sesania inferens 0 - 80 60 3 80 10 51 -Cropping intensity _ -

Rice crops (noyr) 10 10 12 10 10 10 13Rice area (7c) 17 19 20 20 25 2320 85 85 60 70 40 85 85 90Fertilizer (kg Nha) 60 20 85 80 70 7020 0 0 0 30 30Insecticide (no of 40 60 40 300 04 04 0 03 300 13 13 30 50 40 33 33applicationscrop)

__________

IRPS No 123 January 1987 21

Light trapped Insects Drytnd __ 2(notrap-nght) Suctioned Insects (nom )

50 -)o 9 jO40- SOW ffurciWO Light trp

D-Vc suction

20 to 4

20

Egg porasitizahion (M) 75 60

300 SOOampReOftwciffr

45 30shy

15

-insects (notrop-night) 4 Nileporvata Iuen5

Egg parastization()

Light trapped insects (no trop-nght) Suchoned insects (no m 2 )

6 8 Cytorh~nuls hvidipennii 0

4 05 2

50 0 Insects (noltrop-night)40-ui Ipanh pe g

Moohase 1 Seasonal abundance measured by light trap

Rainfall(etod) and D-Vac suction sampler of whitebacked

to ~ hopper Nilaparvata lugens and lea ffolder60)-t Cnaphalocrocis mnedinalis andplthpe g20 L I II I I I parasites and mtrid predator Ortorhjntus livi-Apr May Jun Jul Aug Sep Oct Nov Dec JaIn Feb Mar dipennis set against cropping pattern daily1 9 80o 19 8 1rainfall and moon phase Tanauan Batangas

1980-81

the D- Vac Cyrtorhinus build up on the crop coterminous are collected in light traps during full moon (109)with WBPH buildup Cyrtorhinus disperses during the Because upland rice shares most of the lowland rice pestsrainy season RLF not readily collected in light traps was and those pest species are generally less impotant to uplandprevalent only during the rice crop rice than lowland rice there is a general belief that insect

Nephotettix virescens was more abundant than Nnigro- pests are less important onpicLus in the light trap collections and on the crop GLH upland than lowland rice Arcview of limited data on upland rice shows an averag of population peaked at the reproductive stage as did zigzag 10-21 yield loss from insect pests on a par with that ofleafhopper (ZLH) Crtorhinus tracked the populations of lowland uee determined by the same methodology (132) planthoppers and leafhoppers Both GLH species were (Table 4) McGuire and Crandall (144) made a similarcollected year-round in the light trap indicating immigra- estimate for Central America Cramer (51) howevertion from nearby irrigated rice areas Possibly because of the estimated 35 yieldlow populations loss in South America based on ainsect number did not correlate with similar pest complex - stem borers and seed bugs shymoon phase although reports indicate that more hoppers attacking irrigated rice in North America This estimate did

22 IRPS No 123 January 1987

Light trapped insects D~~nd (no trap-night )

St plht Suctined insects (nom 2

)

D-Voc suction 1520

1o

0 Lih trap 5 ~105 o5

00 Eggparositizotion()5040 NephOtgtha virneens

30 20

I010 A-Insects (notrop -night)

4 Nephatelh nigropccent i

2 o I vV^J - -- ^ VA____ f _

Light trapped insects (no trap-night) Suctioned insects (no rnE

10 -1208 ReCdO doRsol

6 15

4 10 052 iwxALAiamp 0

Light trapped insects (notrap-night) 2Suc oned nsects (nom )

8 CyrtorhmnusI6o nn]10 i0075

4 05

2 0 0

025

2 Seasonal abundance measured by light trap Roinfall (mmd) Moon phase and D-Vac suction sampler of green leafhoppersNephotettix virescens and N ngropicrus and I zigzag leafhopper Recilia dorsalis their egg 60

parasites and micid predator Cyrtorhinus livi-dipennis set against cropping pattern dailyrainfall and moon phase Tanauan Batangas1980-81

40

5 Lf Apr May Jun

1980 Jul Aug Sp Oct Nov I

-aMar

1981

not account for soil pests and was not based on field data in the absence of pests Carbofuran granules are phytotonicAkinsola (I1) cited mainly data from irrigated rice areas in to rice (239) The prevailing hypothesis is that carbofuranAfrica but estimated yield losses of upland rice between 15 prevents soil bacteria from consuming fertilizer The phytoshyand 30 from insects However losses in Thailand over a tonic effect is through greater availability of fertilizer to the4-yr period ranged from I to 13 averaging 5(106) plant Denitrifying bacteria are known to be more active inYield loss data however should be interpreted with aerobic than anaerobic soils therefore it may not be merecaution (184) The Philippine data were all derived from coincidence that the highest recorded yield losses (22-69)trials on farmers fields under farmers normal agronomic occurred in those trials with 12-30 kg basal Nha On themanagement practices except for Bukidnon and Capiz other hand contro with carbofuran granules may not be aswhere researchers used fertilizer Batangas farmers are good as a seed treatment against seed pests and perhapsatypical because ofhigh levels of fertilizer used The trials in yields would have been higher if a systemic seed treatmentBrazil were carried out at an experiment station but insecticide had been used conditions were similar to those of local farmers The 1979 and 1980 trials in Batangas had two full-

Carbofuran granules were used in every trial as a broad- protection treatments one using carbofuran granules andspectrum insecticide applied basally with the seed An the other using bendiocarb WP as a seed treatment Plotsobjection to using insecticides to measure yield losses isthat protected with bendiocarb yielded higher than those proshyinsecticidescandirectlyorindirectlystimulateplantgrowth tected with carbofuran granules The other trials in the

Table 4 Yield losses to insect pests and yield responses to insecticide in upland rice in the Philippines Brazil and Upper Volta

Yield (tha) Yield loss (5) Location Year Cultivar Protected

with Protected

without Check Total Vegetative Reproductive Ripening

Basal N (kgha) Reference

carbofuran carbofuran Protected No with carbofuran

carbofuran

Philippines Pili Camarines Sur Tanauan Batangas

Pangantucan Bukidnon

Dumarao t-apiz

BrazilGoias Minas Ge~ais

Upper Volta6

1975 1974 1974 1975 1975 1976 1977 1978

1979

1980

1979 1980 1979

1977 1978 1979

Bursiging puti Kinanda Dagge Kinanda Dagge Dagge Dagge Dagge UPL Ri5 Dagge UPL Ri5 Dagge UPL RiS UPL Ri5 UPL Ri5 UPL RiS

IAC47 IAC47 IAC47

14 33 11 36 35 31 30 27 37 26 40 30 40 42 46 35

19 21 09

--

-34 24 24 27 --

29 43 31 42 -

47 25

---

10 28 05 27 27 30 26 29 33 28 33 29 43 23 36 11

15 15 06

29 15 55 25 23

3 13 -7 11

-7 18

3 -7 45 22 69

24 28 35

--

21 -11 -20

4 -

-3

23 7

-2 -23 56

-

-

-__0

6 --

-

-23 14

5 0 8 2

22

-

--

0

-

--18 11

3 -10

4 10

7

--7 -

14 -

-1 6

10 -5

8 9 7

-

--

0 0 0 0 0 0 0 0 0 0 0

30 30 30

12 12 12

99 99

100 100 101 101 102 102 102 102 103 103 104 104 104

66 66 66

Farako-Ba

Mean

1977 1978

IRAT10 IRATIO

38 66

--

40 43

-5 35

21

-

10

--

--

--

- 242 243

Z

tJ

-w

24 IRPS No 123 January 1987

Philippines which had two treatments offering protection In the 9 Philippine trials where yield losses were measuredwith and without carbofuran granules relied on further by growth stage 10 of the yield loss (21) occurred at thecontrol with foliar spray and not on seed dressings The vegetative stage and 517 occurred at each of the reproshycarbofuran-protected plots had an 18yield los compared ductive and ripening stages Ibis result is similar to thewith only 10 with foliar sprayS This yield difference is pattern of yield loss in lowland rice measured by successiveprobably due to differences in protection against sown-seed treatments where the crop is unprotected at each growthpests (foliar spray do not control sown-seed pests) rather stage but proticted during the other growth stagesthan phytotonic effects became basal fertilizer wa used in However the insect complexes r sponsible for vegetativeonly 2 of the 10 comparisons stage yield losses in upland rice are entirely diftfrent than in

The recorded yield losses were not related to yield lowland rice The key vegetative stage pests in lowland ricepotential (Fig 3) One might expect higheryielding crops to arc aquatic whereas those in upland rice are sown--seedhave higher percentage of yield losses but this ws not tile (sown in soil) root and seedling pests This dift rence maycase One reason is that insect damage exacerbates plant explain why agricul torists have overlooked tile ilmportanccinjury from drought stress low yielding fields from d ought of upland rice insect pests Many reports of upland rice pestsstress would show disproportionate differences between in Asia ha Ve focused on major lowland species (46 108)insect-protected and -unprotected treatments Shoot and It isalso surprising that tei vegctative stage of upland rice root pests would be more responsible for losses from would record the highest yield losses becatuseseeral studiesdrought stress than sown-scd pests A second reason have shown that upland rice can readily recover from foliageconcerns seed pests which would have a greater impact on reinoval during that stage (117 152 165 188) Upland ricelow tillering low yielding varieties such as )aggc Low poscsses a hitgh obilitV to compensate for ca rly loss oftillering varieties nnot fill in the space created by removal foliage and will even be stinulated by foliage removal toof seeds oweer 1) R i 5 is high yielding because it tillersP1 produce a higher viel than plants ithout any foliageactively which can close the canopy to provide greater weed removed (63 192 231) [ie reason for this apparentcontrol discrepancy hovce is that yield loss at the vegetative

stage does not octUr frol leat arca loss but fron pestsYield loss () removing sown seed and feeding on devcloping tillers and roots -- damage that cannot be readily conpLtnsatCd for

70 - Measuring yield losses in upland rice cnvironrnents is further complicated by a report from Brazil that greater yield losses occurred in response to better insect control60 - =-09 as Insecticide seed treatments resuled in denser plant stands

(n=21) which in turn created a selectively more favorable environshyment for blast (69) This resuIt was parlticurla rlxexacerbated50 when cirbofuran granules were used in seed furrows Aside

ffrom a greater plant stand ca rbofuran iay have auigmented the N lecl and stimulated blast If so ca rbofuran should be40 - replaced with another insecticide or the seeding or fertilizerI rates decreased

Other methods to determine the impact of upland rice30 shy pests wvcrc to interview farners or to develop single speciesbull bull correlations of population levels and yield loss Feneira (66)2 NIM)st (75(1) Golas farmers reporteddid both in Brazil

20- insect losses in ufpland rice and 251 said those lossesS avcragcd 37(1 [his result is consistent with tie yield losses 0 0 measured by field trials in the same statc(Table4) For every

I0 4 of seedlings removC d by leaf-cutting ants a I 7yield loss is predicted Similarly Ferreira (66) found that ai

0 - average of 53 thrips per panicle before panicle initiation meant 3 times more unfillh d grains

-IO0 I 1 1 I I-101 0 2 3 4 5 6 7

Yield (tho) C)N [ROt IM FI IODS 3 Correlation between upland rice yield and yield lols caused by Upland rice insect pests may be controlled by culturalinsects measured in 21 trials in the Philippines Ivory Coast andBrazil genetic biological and chemical methods

IRPS No 123 January 1987 25

Cultural control dry season plowing will have little effect on them Plowing Crop husbandry to reduce upland insect pest populations when the soil is wet from the first monsoon rains will expose can be classified into practices effective in a single field and soil insects that tunnel close to the soil surface to predatory those effective only at the community level Cultural birds chickens dogs and even man (field crickets nole controls should come first in a pest management program crickets and white grubs a re delicacies in man tra ditional They have broad stable effects as pests have little pos- diets) sibility of overcoming them through selection of biotyps The greatest degree i1 inIscct control fron tillage comes (138) On the other hand some cultu ral controls work only from plowing soon after rice ha rvest when most soil insects with high labor or power inputs and connunity-wilt lie close to the soil surface anrd stern borers are in the rice methods require coordination of iany farniters stuble Whitegrubsare mature and tillage exposesthem to

Pantin time The luxury of being able to shift the time of predators Tirnelv control will prtec the crop following pla ntring upland rice is open onl hr la rniers in areas with a rice and lowr the populationIfor succeedinig years Plowingprolonged rainifall pa ttern lI copping intensity or underthe ice stulble helps it decompose killing sten borer rnechanied land preparatiot Gcnerally planlting as soon larvae inside as the rFiinv season begins will lower populatiois of 1110st [requent tillage destroys ait nests bit not termites insect pests Insect populationsarelo aftcra dlvseoll or Tillage and clean culture howes er rcliove food for winter fallow delayed plaiting lets them build tip on termites which then mtay attack a young rice crop Scarab alIternate hosts that gi vigorotsly with the first seasonial beetles also prefer to miposit in eceitly tilled fields Noshyratiris This first flush iof weds cros lItxuttriartlv with iceCult is dibbling faors the survival oftillace Clre Such rineralied N released during the dr season aid made stils i aels tnd insects thtt pttpate or lay eggs- in the soil

available to the plants with the rains It iliso igrtmitcally Zero tilla hio eerl conserves predators ofthe GIH (37) advanttgeous for the rice ciop to tap this trlLItlaI fertility nt it mcttoi lo lessen the impact o0 soil insects and grow vigorotsly to compete with wrus and to tolerate brotdcasting seed or planting in ftltrrows is prefc ired over pest infestation dihlirig in hills Crickets are large enotlgh to eat seedlings in

Larly plantngs terd to escape seedling maggolts Upland a hill atrd tillering normnally ca iriot fill in tlhe missing space rice in Bitangas is planted in Ma and ine itli the Iirst Oftcn dibbll rd is not well covered and iseasily found by rains Fairners are highly mtotiated to plant early tocscape foraging insets ioIdents and birds drlitght and typlotns ricar harct Arn Atrgtst planting Plan uit sii Incrcatsini sced density protects tile cropwould be evcely attacked by It riim orivzo At tti againrst seed and seelling pests especially ants particularly other Philippiie tipland sites Rea l Qteot antd (laelit if the crop will germinate and ermerge quickly M isainis Oriental seedlig mnaggots arnrige ric boe t se hr trftinll itI India rice intcrclopped witi cotton or farnes have a longer growing season and plnit 2 rin or pigeon pci had lower 6IIt and WBIlII poptrlations than mote after the ortset of the rainy season liaaiangas ftrmners rice alone (211) NfailC arid Urplald ir_ 2re coilltori in the plant ca rly becia use ofa shorter raiinfall period arnd the desi re Philippines he chatngC in pest status folii iniercropping is to lliarvet before the typhoor sea son peaks ltighly locatioll-specific (133) cmianarid the net elfc be either

Early plantings cornbind with early-iatiuring varictics nil bcnelficial ortdetriential (189) For example BIatangias provide higher crp tolerartce for white grubs Aestiatior farmers intercrip Itro in their ipland ricefields A liornshyterminates with the first rains but 5-6 moare ineeded for the nrii iotd-Ahis i (I iiiacius) which norlially feeds resting larvae to develop into beetles emerge and lay eggs on taro ill feed or miarbv rice plants Intercropping and for larvae to reach the danriigii g ird-instar stage probably would not afft0 seed pests A crluanion crop (134 In lataigas this normually iseiough tiite for harvest planted ith rice and haing gleater tolerarnce for root

It Brazil howecver delayed planting is a suggested dariage or being toxic nayact isi trapcrop to root-feeditig cultural control for the lesser cornstalk borer and insect pests Acial-fei i ig pests voultId be most affected by pests favored l drought (66 195) Cropping internsity in intrer ropping Shoot aphids leaf beetles sten borers and Brail is low and delayed plraing places the rice crop in the hopplers would be likely ca ntidates to note effcts On the niost stable rainfall period reducing likelihood old rought othlt halnd the corlipalior crop stch~ sitniaie Iiiv be more

In rilniy arca s rinmirsare roi atl to plant other crops benehited than rice from initecropping ( 137) before rice and althNgh it woultl be desirabh to avoid 11echi Weeuling luring the first month after clop infestations by planting rice cuatl lariners lack the labor cstablishent will force pcts such is anyvworris which and time to do so ol a low priority crp prefer grassy cetds ontor tle vorling rice Soil iampsecs will

77age Uplild rice Stils ae light textured and Call be mvnye to the rice crop Cleani citrltirc also lay lore territes plowed duiring the d rv seasor notl only to grin time for to attack I u cetincti inrice ceds i ay he le optinal planting with lie fitst Inilsoon riiains but also to desiccate solition 13) soil insects arid weeds such is ntsedge that foim rhizomes Irtlizr Itneatsing N Iciiliier oin hlwland rice has Htowever becta use most soil insects lie below die plow layer fAtwvored Irigher populains of pliithoppeiv Imlhoppers

26 IRPS No 123 January 1987

and leaffolde rs However because of the need to temper fertilizer use on upland rice o prevent blast fIrtilizer rates will be lower than is comonia r for lowlad rice causing relatixely less hopper and IaIfTfoide r population buildup However inupland rice areas without a history of blast such Is Batangas farmers apply high rates of N and conscqtrently the crop has high infcstation levels of WBPI]and RIF The fhrmrcrs leec insect buildup by splitting the fertili11r in0to two to three applicatiois bit do not applV fertilizer at the vegetati estage because this prodrces a tall crop which illeasily lodge during typhoons Greater crop fertility leads to greater insect survival (better nutrition) larger insects (to cat no re foliage) arid greater fec titd t (to lav nmre eggs) (128 145 171) Not all insect species respondequally hovr Increasing rates (if N l anrid K increased Dialraa s-acchralis and (iaeiurema flea beetle but reduced thrips (66) Zn rcdLuced deadheart density from the lesser cornIstalk borer

Fnldingi Although most Upland rice areas do nIot hawxcaccess to wa ter floodiig the fields is rRrrnriierdcd in areas of Brazil or root bug and white gib control ( 195)

Crop rotatioti Tillage before planting a crop after rc harvest will control stern boreis and other pests renain n the stubble arid will unearth soil pests Planting a non-gratriiaceous crop after rice is recommncrded for termite control (66) Crop rotation is less nttdcd in upland rice culture than in lowlands because atcrop-free period is assured even after harvest of crops following rice

t1ulchilt iryl plastic irli irig designed for weedcoitrol ii upland rice inJapan resulted in ntr highanri liy infestation of (ihilo sulytresvais (93 9-1 95) NIulched Upland rice grew more luxuriantlv and attracted more (Avipositing mothis than u lched rice Also the rice plant and stein borers imatoured 10rC quickly and a partialgeneratiori af vouing ista iNoverwintcred in the stubbleLotili--malitrt varieti Quickly maturing crops reduce the tmbCr of pCst generations that Calt build up Rapid crop establishment olan ca rl v-natu ring vaitvty if)pa 11icLa r will lessen white gib and stem borer damiage White grub larvae intle last iistar are highly destIruCtive so aii early-iiatri ng variety will escape serious root loss Stem borers build 1ip slowlyx in rice to ecormic abundant in a late-Maturing crop anrd caiuse whi tlieads Numibers of somespecies increase exponeitially with each succeeding genera-ti so a xarietyexhat atures I trowh earier thian another will hac less insect damage Early-naturing vaiicties however will not lessen seed pest daiage either ii the soil or oina standing crop

Strwhoi hAIJttam Inrscts tcd irig on thlie aerial portions of rice plats disperse from carly to late plantedfields AS iiiibCirs increase exponentially with eachinsect

gtscnrattm e eiglbrig f rmc~s1(uil upp wh rc generation pests build tIP where neighboring fihrTC e9r1sstagger their plantings II Batangas farmers plant uplandrice within I iro use 120-d s-arieties and plant at the start of the rainy season Ini Misarnis Oriental laricrs stagger their

plantings up to 3 mo apart and use a set of atieties that mature in 4-7 mo Insect pest danage particularly from stem borers is greater in Misarmis Oriental Also seedling maggots damage rice there because rainfall favors frequent planting of maic In Batangas rice escapes seedling maggots bCca usC of distinct drs season and synchron0us early planting with the onset of ains

Therefore combinations of cultural practices ea rlyplanting synchronous planing crop rotation and earlyshymaturing vaneties protect the rice crop against most insect pests

Ilmant resistance Beca tse upland rice has few specialized insect pests and many insect types attack the crop worldwide regional breeding and strong internatiornal cooperation are required

Sown-sced pests arc economically controlled by insectishycide seed treatment and should the relore receive lowxpriori ty inbreeding objectives

Root pests are normally cost ly to control with insecticides but finding resista t sonres mayvbe di fficult Tanaka (229) for exa -ple fi led to find varieties resistant to root aphids Selecting for large root bioniass perhaps should be the strategy for pests sLuch as Whlite grubs that renliove roots Fainter(174) howcvcr cites varietiesof crops tolerant ofor resistant t(i root pests Some sorghun varieties are high in cyanide so rice sorghun might incorporate broadshyspcctruri chemical resistance

Seedling vigor and drought tolerance are high prioritybreeding objectives so tupland rices should have large r root systernsand be able to tolerate ligherlevels of root loss Sonie root loss inolder plants is beneficial if it stimulates new root developnrut and therefore crihances intake of soil nutrients (53) The rice seedling inaggot rtiLotia spp may be

controlled thirough resistant aittics A hi rgc breeding effort on sorghniu has had fair success inideveloping arreties esrstait to Asoccata Rodani (251) A field trial comparing 10 upland ricc varieties showed differences aniong varieties va ring from II to 4(- damaged tilleis (215) Hovver resistarce to 1Iill-0r gtia species may rot cross ox-rti otier

Shiraki (210) reported that upland i-ice stens wcrc harder than lows-land rice stemsperhaps becaIusc (f a higher silicoricontent and thus were more resistant to yellow stern borer hc tortia Iow tillering (f uplaid iice xarneties prodtres a

higher percentage of infested tillers than would occor in higher tillering lowland iice under a siiilar egg density The corollary therefore is to select for high tilleriig upland rices

Stemii bor amcstheh main targes dobrecders inAfrica (571 204 )but nticl oifth e sc reen irig is dotie unde r lo wland co d t osrm s wih A h r~ o l pe i s n acorditions (11 96 218) As with lthrinrnwspecies mayhave to be dealt with independently however initial results show cross resistance between African and Asian species(92)

In Brazil screening Ior insect resistance focttses oil lou I pest problems Varieties tolerant of lasmolwalpus ligi-sellus and resistant to )iatraea saccharalis have been identified (70 153) Current work also emphasies 7iraca limativetntris and Thmialspis (=Iosis) flavouicta (66) These insect pests Ilt onlV citise IIigh yield losses tihey also ire difficult to control chemically ofr by other ilealls

ManV upland rice vaieties aic early maturing and are resistant to pests such as gall midge (I Xl ) Early mIturity is often a highly desirable trait to esuilpe pest buildup but it is not true resista nce bca usc ilthose va reties are plinted late they will be itsdamaged as ain early planted susc ible variety

P~erhaps tile cosiiiopolitall of all upland rice pestsimost are the sced pests (ripenling stagc) which pose a particular problem beta use scuds re ised IsIO for hitnit4ans Ayltoxic substatice introd tcld geticticllyilmay also be toxic to nal iniless it caln he placed old in thfe IdhuffSelectmi fo hairiness or lotng wns kill mIlct sthilrohleiiis dutilig threshing pa rtictfarl maitial threshinfl [hireshers ill complain of itcfimin atid itteascd (fust levels

Attempts itIRRI to fitid tcistatncc to the rice bug plorisa have ilotbeCnl prmitisilg lIIetIattonlids iorc

through the lenilna or palca hillhairs sedstVIor seeds wilh awns that deter bugs promote complaints from threshers Seed bugs liptocorisa and Stcnocoris enter the seed through the opening between tile lemna and palC but varieties with narrow openings are diifficult to mill

Breeding priorities should concentrate ott inotCI gruLtLIps that are most difficult to c nttrol by other meains Steil) horers appear to be the most widespread group that sigiiificantly dalag s rice aind is hard to manage by other Ilnca us

Biological control Very fe stutdic of mitatt~l eneties of rice pests in upland envirollnletits wee in tile wefouiid literature Therefore have not compiled a list of species Ve present a list of natural enemies iOtid in upland ricefields tuic latialln Batangas for some conitno iOldiar pest species ([itble 5)

Muich ofl thc lollowinle itfor ittiol cotlliCs rol Stutdies in the lowlatnds or roi crops otithcr thlin tice

Qu(ailtr o 11ll t11t( Nat trrtf lii hcoeits ight Ile used agaitist ile social insects tilt and termites Sollie predator specice live ill theirf nests n ilnic theirnllt ip-pearanIlCe aMRidbehavior IJtrouMdCing a mote aggressise ant species to displace pest ant codonies is a possibility (169) Patihogens as potential natural encnies have reccived little atteition (I13) Natural ncnuics asa group however hac not been toIl iporta it in Iegulating aintshown be and ternite numbers

The nocturnal behavior aid subteitmien habitat of lole crickets and field crickets protect them from many ntllral enemies There are only limited records of natural cnelics for these groups of pests some pathogens (87) but mainly

IRPS No 123 JattuIny I 271987

parasites - scelionids on eggsand sphecids on ilymphs and adults Sphecid wasps Larra carutmaria (SImitI hI L huzwuensis Rohwer and lsaltiuhta Willi ams speciali in mole crickets pa ralyzing them nd dragging the ninto thcir nests as food lot their voting ()tier sphCcid species Lins aurulenoa (Iabricius) lot maiac (Ashmead ) 1 lhshylssellaus (Smith) and Ml lal)oriosu (Snithf specialie in field clickets

White grUbsO the other hand hae Icti scriuns enuigh to have been extensively studied for cotttroi 1W nati al enemies( 112 140 141) Scoliid parasites C(ajsoMcispp have beetn intrtduced itto tilePhilippites for control of Ielxl ]udisiro la The rcsearchcts w Intttrdced themle during ill era claimed that the white gttionthreak as controlled (142) howevcr it isnormal for pest epidemics to subside and there was no diect evidence to indictte that tfe parasite was responsible A tmore promisitg titctllod might he to identify iso ltcttass prt~f tcartd apply hiactelaln fingal pathogens to tile soil Ihe lalattese beetle in North America has hcci coitrolled hv tite conmnleciall aailable hactcrial prcpa ration itacihlpu pav ofr milksdiscas (71) Spote prtyefott the soil itloted ilrtier reitiain actic fr Veas

Root aphid sardtnctly bugs readilslic tilib to predatots and parlsites once their tetndine ants are controlled The larvac and adtilts of a coccinellid beetle 1Scynms sp prey on llriadlouais in slash and burn upland rice Baits treated with insecticide may be used to control tending ants by killing their ourig (244)

Root-tkcditig blugs false wireoris wireorms and root wccils aic r ofsuch little ifilpoltatice that thenomttll role of liatural Cticniics hias tiot beei asscssed A possibility hoevrl tiglt he tit ItaIIitic niaCIMtodcs lintt1g11en Itese soil-itlhlfbitine mCtitodcs W5uhtfhave to be imiass Produiced

large miltibcr of niatltrll cnemics ircknown ft CltIoriS IC IpblCeii is thtt clttvortnlls ofonlie the crop soott alter land ptel)tritiOti a1tithe bcgit iniig of the raittv

Citsoln hfien tiatllral leniiiy populations aric lo Colltloil WOtuld iii ole mass prodtticing andirclcaslng key spccies

le egg stage is the fpirt (olthe Cedlitgt Ialot life cycle tI hich to f)ctis ftor tittlril elctit cottntrol hplraites o1 pleditors ttsit elleggs highly expoIsYCl I Sgg pailsites or predators stolld probahly e ittass andtbe produced released Littlh i toruialioll is a ailable ott what species to trV o ho to Iniss podLice thiCIi A etiloplhid wasp Icmihlaol sp effctiicly regushy

lates the leatf lcloalapolm2a spioanoiiitr heat andc rice antid should be collsctvcd

leaf beetles whCllose e am] eggs ate laid in the grutindhirO ould be motC difficutlt to control by niatulral enelV Mnll ipuhltiOn But species sUici IS c_gsthispid beetles ith

and larVle oilleascs ate vtilcratble to iparaites and pledators

Natural enemisc of tlirips otl rice are little known

28 IRPS No 123 January 1987

Table S Parasites recorded from upland rice insect pests in Tanauan Batangas Philippines 1977-84

Order Family Species Host stage

I lymttenoptera Braconidae Leaffolders Cnaphalocrocis Marasinia spy

Cardiochilesphilipplinensis Ashmead a Larva

Cotesia angustillasis(Gahan)Cotesia or cypris (Nixon)

Larva Larva

Cotesia or taeniaticornis(Nixon) Larva Cilonusinotakatae Munakata Chelonits SppAlacrocetrts philippinensis (Ashtmead)

Egg to larva Egg to larva Larva

Iclineumonidac Orgilts spp Isclutojppa hireator(Fabricius)

Larva Larva

Itoplectis naraigav(Ashmcad) Larva Tenelhtcha philippiensis (Ashmead) Larva Tenelchha stangli (Ashmead) a

irichomma cnaphalocrosis Uchida Larva Larva

Bethylidac Elasmidac

Goniozms or trianguli r Kieffer lasmtusalhopictus Crawford

Larva Larva

Diptera

Encyrtidac Chalcididac

Tachinidac

Elasmtus spp Copidosomopsisnacohiae (Fidy) Brach nteria evcarinata Gahan Biachyieria lasus (Walker) Ztgohothria ciliata (Wulp)

Larva Egg to larva Pupa Pupa Larva

Argyrophylx nigrotihialis(Blaranov) Larva

l-ymenoptera Braconidac Yellow stern borer Scirpopiaga incertulas

Braco chinensisSzeplegeti Larva CIthelotis spp Chelonus niunakatae Munakata Cotesi flaripes Cameron Stenoracon nicerilli (Bingham) a

Larva Egg to larva Larva Larva

Ichneumonidac Tropobracon schoenobii (Viereck) Amatroinorpiaaccepta metathoracica(Ashmiead)Frihorus sinicus (I lolmgren)

Larva Larva Larva

Ischnojoppaluteator (Fabricius) Larva Isotima nr dammeratiti Rohwer Larva Tenelucha philippinensis (Ashmead) a

Larva Tenehicha stangli (Ashmcad)Trichomma enaphalocrosis Uchida

Larva Lara

Chalcididac Eulophidac Trichogranmmatidae

Xanthopimnpla stemmator (Thunberg) Charops hrachYpterumn Gupta and Malteswary Brachtm eria sppTetrasticttus schoenohii Ferriere Trichorigranm chihunis Ishi

Larva to pupa Larva Pupa Egg Egg

Diptera

Scelionidac Pteromalidae Tachinidae

Tt ichograintnta aponiicuitt (Ashniead) Telenomits rowani (Gahan) a

Trichoinalopsis apattelctena (Crawford) Jeirhaea spp

Egg Fgg Egg pupa Larva

Z4gobothria ciliata (Wulp) Larva

Ilymenoptera lecneumonirlac Gol fringed borer (hilo auricilius

Trichomm enoptalocrosisUchida Larva

Ieioenoptera Braconidae

Aantholimlh steiniator (Thtinberg) a

Pink stet Ioret Sesamia inferens StoInacoil nicrilhi (Bingh am)

Loarva to pupa

larva

Diptera Ilyinenoptera

Armvtnyorms Mythimna separata Spodoptera mauritia acronyetoidesTachinidae Zvgohothriaatropiora( Robinea i -Desvoid y)Eulophidae It-phctrtiscitapadoe (Ashimead)

Larva Larva

(halcididae Braconidac

Brachpin ria spip (oesia spp

Larva larva

IHyntenoptera I)iptera

Rice skipper Ielop ida (halcididae Tachinidae

math ias aidgreen iortedci ciliarNIelan itis eda Brat lhincria sp iii matuqitatt Cameron Irgyrolhirlaxi trotlhiais(marumtv)a1

ismtne Larva Larva

Diptera Ilymenoptera

Tachnidae lelineutionidae

fuown se tilo per Mois frugalisIrgtrophrlax 1igrothiahiv 3 a rat ov) anthopimpa pmnctata (Iabricius)

Larva Larva to pupa

aMost dominant

IRPS No 123 January 1987 29

Normally predatory thrips prey on herbivorous thrips strepsiptera on both Spiders dominate tilenymphaladultNatural enemies of stem bugs --Scolinoplara and Tibraa predators Spittle bugs move to rice from pasture grasses - are in order of importance egg par~isites and predators and biocontrol would be successful only ifcarried out before fungal pathogens and nymphal adult parasites Egg tiley immigrate into ricefields parasites could either be introduced fron other areas or Biological control tactics for hemipterous seed bugs mass produced and released ould follow those outlined for stein bugs

Armyworms are normally lwld in hcck by the activities Coml)arison of environnewtns Natural enemies are of egg and larval parasites When these natural enemies fail perhaps faced with even greaterconstraints than insect pestsusually because of drought armyworns become epidemic to survive ind reproduce in upland environments TheyIt may be worthwhile therefore to release parasites during have adapted ani survived by having wide host ranges the the rainy months following a drought Virus diseases of ability to aestivate or hibernate over unfavorable seasonslarvae would beanotheravnue to exploreforarnyworms and the ability to disperse or by other mechanisms

Parasites have ben traditionallyconsidered for biolegical Comparing the most dominant and therefore the most control attempts against stein borers usually by rekasing adapted species of natural enemies collected in various exotic species( 166) This approach has not met with success environments in the Philippines we can understand how and new avenues should be explored The role of predators different rice environments are arid pathogens is little inderstood and deserves greater Anagqrms olptahiisa mymarid planthopper egg parasiteattention Egg parasitism rates are normally reasonably is cquallyadapted to all Philippine rice environments (Tablehigh but should be supplemented with effective egg 6) It is selective for planthoppers and even parasitizes eggspredators Orthopterans are potential egg predators of the n-iize planthopper Peregrinitsmaidis itt upland areas Metioche vittaticollis (Sta ) Anavipha nifiwnnis (Saus- Alavohis also parasitizes maize planthopper eggs but is sure) and Conocephahs hongipnni (de i-laan) feed on most prevalent on BPH in the irrigated wetlands (onashystem borer eggs etioche and Anaxil)ha specialize in eggs tocerus spp like A optalhilis on BPH and WBPI Iattacks with no hair such as Chilo spp The omnivorous (ono- the eggs of GLH in ell rie environments It is also found cephahsspecializes in eggs cov red with hairs it is a proven parasitizing the eggs ef the white rice licahopper (oimna egg predator of Scirlouphaga spp but has the discouraging spetra Oirosita naias a trichogrammatid egg parasitehabit of eating rice guin (65) attacks BIPH and WBPH in the wetlands and has no other

Grasshoppers have egg (scelionid) and nymphalad ult host A related species 0 aesopi specializes in G1-H in (nemestrinid tachinid and sarcophagid) v irasites Among wetland environments Dryinid strepsipt-an and pipunshypathogens protozoans have been recorded most frequently culid nymphal and adult parasites of planthoppers and Control of locust species by natural enemies could con- leafhoppers also specialize either in hopper species or centrate on their habitual breeding areas environments Among the dryinids Pseuhdgonatoipsflavi-

Leaffolders have rich complexes of natural enemies some feimur prefers the lowlands and BPH while P nuch has no adapted to upland rice (22 27) ranging from egg predators environmental prefirences but is adapted only to WBPH (gryllids coccinellids) to larval parasites (braconids ichncu- The main d ryinid species on GLH aptogonatop)usspp is monids) to larval prcdators (ants and carabids) to larval most dominant in rainfed lowlands and also attacks other pathogeiis (vira and fungal diseases) Efforts to augment leafhoppers within Amnrasca Cicadulinaand Bachaltuathese natural enemies should focus on the egg predators Strepsiptera are more dominant on plarithoppers than (rearing releasing)and viral or fungal pathogens (culturing lealhoppers The Elenchus species in the Philippines attack spraying) all species of planthoppers within the genera Sogawilla

Parasitization of the large conspicuous larvae of Sogatodels NilaparsataHarualia and Opiconsiva Elenshyskippers green horned caterpillars and other polyphagous cusrvasuntaisuiis adapted to the major rice environments Lepidoptera by tachinids and chalcids is normally low ELg but prefers BPH in rainfed arid WBPIH in wetland environshyparasites and predators are perhaps the key to their control ments Ilalictoph(utsmtnroeiand HVpectrus attack G LH

Gall midges are parasitized by pterormalids and platy- and COfimravcctra respectivelygasterids that aestivate within their larval host between Pipunculids are mainly parasitic oii lealhoppers Each seasons Shoot aphids are preyed upon by coccinellids species has a unique environmental preference Tomosshysyrphids and chrysopids varireIa oraetoraprefers Ne7 otetltix nigropictrs and

Planthoppers arid leafhoppers have egg parasites and Pilunwulhtsjavanensisattacks Dehoceplhahls spp which predators as well as nymphialad nlt parasitesand predators are more abundant in upland than irrigated lowland their rich complex of natural enemies includes fungal environments T suhvirescensattacks Nephotettix virescens pathogens The principal egg parasites are myrnarids and and prefers the lowlands Pilpuiulus nttillattls is an trichogramniatids Egg predators are norinally mirid plant upland rice parasite arid attacks only Nephotetix speciesbugs that suck out the yolk Nymphal ad nIt parasites are More than 17 recorded species of leaffolder parasites in dryinids on planthoppers pipunculids on lealhoppers and the Philippines attack species of C(naplwocrocis and

30 IRPS No 123 January 1987

Table 6 Dominant natural enemies of rice pests in upland rainfed lowland and irrigated lowland rice environments of the Philippines

Natural enemy

Host stage Species

Brown planthop-Parasite Egg

MymaridaeAnagrus optabilis(Perkins) Anagrusflaveolus Waterhouse

TrichogrammatidacOligosita naiasGirault

NymphAdult Dryinidac

Pseudogotatopuslaiifemttr Esaki et Hashimoto Elenchidae

Elenchus yasurnatsui Kifune and Hirashima

Environment

Upand Rainfed lowland Irrigated lowland

Ailaparata lugens

x x X x

x

x x

x X X Whitebacked planthoppers Sogatella furcifera and Sogatodes pusanusEgg

MymaridaeAnagrus optabilis(Perkins)

TrichogrammatidacOl~gositanaias Girault

NymphAdult Dryinidae

Pseudogonatopusnudus Perkins Elenchidae

Elenchus yasutatsui Kifune et Hirashima

Parasite Egg

MymaridaeGonatocerus spp

TrichogrammatidaeOligositaaesopi Girault

NymphAdult Pipunculidac

Tomosvaryella subviresccns Loew Tomosvaryella oryzaetora (Koizumi) Pipunclusmutillatus (de Meijere)

DryinidaeIlaplogonatopusspp

Egg Copidosomopsisnacoleiae (Eady)

Larva BraLonidae

Cardiochiles philipphnensisAshnead lchneumonidac

Trichiomnia cnaphalocrosisUchida Temelucha stangli (Ashmead)

Pupa Chalcididae

BrachymeriaexcarinataGahan Brachymerialasus (Walker) Brachymeria nr tachardiaegroup

Parasite Egg

Scelionidae Telenomus rowani (Gahan)

EulophidacTettasticusschoenobhi Ferriere

Larva Braconidae

Stenobraconnicevillei (Bingham) Iclhncurnonidac

Temelucha phiiippinensis(Ashinead)Temelucha stangli (Ashmead)

x x x

x x

X x X

x x Green leafhopper Nephotettix virescens

X x x

x x

X X X X

X

X Rice leaffolders CnaphalocrocisMarasmia

x X x

X

X X

x x x x x X x x x

Yellow stem borer Scirpophaga incertulas

X X x

x

x

x X XAinauromorphaaccepta metathoracica(Ashmead) x

continued on next page

IRPS No 123 January 1987 31

Table 6 cntinued

Natural enemy Environment

Host stage Species Upland Rainfed lowland Irrigated lowland

Larva-pupa Gold fringed borer Chilo attriciiuslchneumonidae

Xanthopimpla stemmator (Thunberg) x aPupa x

Pteromalidae Trichoinalopsis apanteloctena (Crawford) ax

Pink stem borerb Sesamnia inferensParasite

Larva Braconidae

Stenobracon nicevillei (Bingham) x b b Armyworms Mvthinna SpodopteraLarva

Tachinidac Zygobothriaarroptvora (Robineau-Desvoidy) x x

Braconidac Cotesia spp

X Skipper Pelopidas mathias and green horned caterpillar Melanitis leda ismeneF-gg

TrichogramrnatidacTrichogrannaspp x

Scelionidae Undetermined Scelionid

x Larva

Tachinidae Argyroph lax nitrotibialis (Baranov) X x

Pteromalidae Trichomnalopsis apanteloctena (Crawford)

X Predator

CoenagrionidaeAgriocnenis femina femina Brauer x xIschnura senegalensis tambur

Miridae CrtorhinsliidipennisReuter x x

Nabidac Stcnonabis tagalicus (Stal) x

VeliidaeAlicrovelia douglasi atrolineata (Bergroth) x

Coccinellidae Micraspis crocea (Mulsant) X x x

Vespidae Eumenes campaniformis (Fabricius) x

Formicidac Diaecamna spp xOdontoponera transversa (F Smith) x

Carabidae Chaenius spp OphioneaW- aJasciataSchmidt-Goebel

x

Ophionea ishii ishii Habu x x X

Gryllidav Anaxipta longipennis (Saussur-) x xMetioche ittaticollis (Sial) x x x

Tettigoniidae Conocephalus longipennis (de Haan) x x x

aNo rearing made on rainfed lowland bNo rearing made on rainfed lowland or Irrigated lowland

MrasmiaAmong the Laval parasites the braconid Cardio- lowlands and parasitizes oriental maize borer OsIriniachilesphilijpphensis is most adapted to an upland environ- furnacalis (Guenee) larvae The leaffolder pupal parasitesment It also parasitizes Hydelepta indicata a common not only occur equally in all environments but also haveleaffolder of legumes The ichneumonids Trichomma cna- wvide host ranges- parasitizing species of OsIrinia Marucaphalocrosis and Temehcha stangli occur more in the Hydelepta and Honiona - in maize and legumeslowlands T stangli is most adapted to irrigated lowlands The principal egg parasite of yeltow stem borer (YSB)and also parasitizes Chilo and Scirpophaga stem borer Telenomus rowani is widely adapted to all environmentslarvae T cnaphalocrosis is most prevalent in rainfed and has an unusual alternate host -Tabanus eggs (25)

32 IRPS No 123 January 1987

Tetrastictu schoenobii specializes only in YSB and occurs mainly in irrigated lowlands -- further proof that YSB is most adapted to the wetter environments

Larval parasites of YSB are highly environment-specific The braconid Stenohracon nicevillei occurs only in upland rice but also attacks the pink stem borer (PSB) Tieinchwa phiiiplPwnsis an ichneumonid occurs in upland rice and also parasitizes the larvae of (7hiu stqpresaisinatro-morpha accepta melaihoracica like 7 stangi is mnost adapted to the irrigated lowlands

The dark-headed stem borer(l)HSB) (ih pochit3sus was not collected in all rice environments and the gold fringed borer Chilo auricilius nd PSB were collected only in upland rice Xaantiholiplastenrnaloran ichneumonid larval parasite appears to be widely adapted whereas TrihomaslopisapanitI)cteiiaa pteromalid pupal parasite occurs mostly in wetlands and also parasitizes larvae of Chilo suppressais and Pelhuidas mathias

Armyworms Myit hinin and Spodujipera have two main larval parasites - a tachinid Zrgohothriaatropivora in rainfed environments and a braconid Cotesia spp in irrigated lowlands

The butterfly pests Pelopidas and 4elanitishave tricho-grammatid egg parasites found only in irrigated lowland environments

Two larval parasites show environmental preferences --Argrroph av nigrotihialis a tachinid in rainfed habitats and Trichonialopsisapantelkcnela in irrigated ricefields A limgrotibialisalsoattacks the sweet potato hornwormAgrius convolvtdi and T apanteloctena parasitizes DH S1B

Predators as a group show more distinct environmental preferences The aquatic damselflies Coenagrionidae are most abundant in the lowlands Cvriorhinus is also most adapted to the lowlands The nabid Stenonabis tagalicus occurs only in the uplands and isalso prevalent on legumes The coccinellid lady beetles wasps and ants are more prevalent in upland areas AlicrasIpis cit-ea larvae prey ona variety of aphids attacking legumes and maize Etunienes campanonia vespid mud wasp makes nests in trees and is therefore most adapted to the more botanically diverse uplands Soil-dwelling ants cannot tolerate flooding

Upland environments are habitats to arboreal carabid beetles Three species of Chlaeniusbeetles prey on leaffolder larvae and are perhaps more important than parasites (27)

The gryllids and tettigoniids are widely adapted egg predators not only on rice but also on maize and legumes They feed on eggs of most insects that are laid on leaves

The spidercommunity of upland rice environments isrich in species At one site (Batangas Philippines) 31 species have been recorded (23) Of the 176 spider species recorded in Philippine ricefields about one-half (82) occur in upland rice The spiderspecies of upland rice environments overlap more with those of rainfed lowland than with those of irrigated lowland rice (24) The same study showed that of the three environments irrigated rice has (ie greatest spider species diversity foiiowed by rainfed lowland and upland

However of the 10 most prevalent ricefield spiders 9 were abundant in upland rice showing wide adaptation All three spider guilds -- orb-web space-web and huting spiders shywere prevalent in upland ricefields There were however differences in environmental prelerences for some spider species

Of the orb-wcavers Tetragniathamanlihulatawas partishycila rlv abundant 7 japonica was low and Ltucatge dehcorata was absent in upland rice

As in other environments Aiypeiwnaifirtosana was the most dominant space-web spider however its relative numbers were lowerthan for lowland sites Among hunting spiders Ox|opes javanus and Lcosa leatcost fgla were more abundant and Lvcosa lpseudoanmnlata was less abundant in upland compared with lowland rice envishyroniments

Extensive rearing of rice insect pests in upland rainfed lowland and irrigated sites over an 8-yr period in the Philippines provided insight into the effectiveness of parasites as natural enemies (Table 7 8) Most of toe data are from large sample sizes taken over at least one crop which overcomes the pitfalls expressed by Van Driesche (238)

VSB egg parasitization was surprisingly similar across all environments Larval parasitization was highly dynamic among sites and even years within the same site but showed somewhat lowerlevels in uplandsthan in irrigated wetlands This would have been predicted for egg parasites as well since YSB is adapted to the lowlands and so must be its parasites This isevidence that Telenomus rowani is as good at dispersing as its main hosts Trowani clings onto the anal tufts of female YSB and parasitizes eggs as they are laid This phoresy may explain its wide adaptation The more specialized lowland YSB larval parasites therefore appear more effective than their upland counterparts with more alternate hosts

Low collection levels of PSB larvae especially in the wetlands make comparisons between environments difficult Low levels of larval parasifization were recorded in both rainfed environments Except in the Cagayan rainfed wetland site in 1980 low levels ef larval parasitization were evident at irrigated and rainfed sitesalike No differences by environment were evident Higher levels of parasitization occurred in Sarawak during a 1967 outbreak also from tachinids Rothschild (197) indicated that higher parasishytization occurred in upland rice areas

Skipper and green horned caterpillar larval parasitization rates were generally low and showed no environmental effect htighest activity occurred in Cagayan in 1980 (48 parasitization) Although larval parasitization levels of RLF were variable year to year activity appeared greater in the irrigated lowlands than in the uplands Again the highest levels were recorded in Cagayan in 1980 Cagayan has extensive grasslands and fallow around the rice areas and a relatively long rainfall period Rainfall in 1930 was good for crop growth and apparently for increase of larval parasites

IRPS No 123 January 1987 33

Table 7 Parasitization rates of rice planthoppers and green leafhopper in upland rainfed lowland and irrigated lowland environments Philipshypines 1976-840

Environment and site Year Egg parasitization () Nymphaladult parasitization ()

BPH W13P GLH BPII WBPH GLIH

UplandBatangas

Rainfed lowland

1976 1979 1980 1982

15 17 12

2(47)8(39)

4(51)

9(1800)14 (212)

12(304)

8 (101)6 (127)

9(116)

Iloilo

Cagayan

1976 1977 1978 1981 1980 1981 1982

0 (69)

11

44 48 40

-

-

46 55 44

0 (471)

11

52 56 49

0 (200)9 (214)

10 (1259)8(235)6 (900) 3(800)

-7 (462)

11 (270)3 (313)5 (900) 1 (800)

4 (600)10 (725)10 (362)12(870)5 (900) 3 (800)

Pangasinan

Irrigatedlowland

1984 1976 1978 1979 1982

13 (436) 13(379) 14 (457)

6(14)7 (197)

17(198) 11 (486) 18 (275)

3 (81)10 (274)3 (116)

16(871) 19(88)

28(211)9 (764)5 (647)7 (1204) 8(305)

Laguna 1977 11 7 21 1978 31 1979 14 4 4 28 1980 7 8 20 1981 1983

56 7(278)

26 16 (1650)

16 27(1921)

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R I Apostol unpublished 1lo1lo and Pangasinan 136 Cagayan BCanapi unpublished Liuna 175 I C Pantua unpublished 118 234 43 Carino and Shepard unpublished BPH = brown planthopper Nila parvara htgens W131Pi =whitebacked planthopper Sogatelia furcifera GLH = green leafhopper Nephotettrix virescens Figiires in parentheses are sample sizes

Tauher et al (230) reported that internal external and genetic factors influence the seasonal activity of parasitoids

Hopper egg parasites but not nymphaladult parasites were also parlicularly abundant at the Cagayan site possibly the egg parasites have more alternative hopper prey in the weedy areas than nymphaladult paisites

The seasonal dynamics of hopper egg parasites can be seen for upland rice in Batangas(Fig I 2) and for rainfed lowland lice in Pangasinan( 135)and Cagayan( 105) Levels of BPHWBPI-L and GLH egg parasitization were corn-parable in upland and irrigated lowland environments Except for some high rates of paraMsitization early in the season the parasitization rate tended to be steady in all envilonments However hopper nymphaladult parasi-tizution levels were more similar in the rainfed environments and lower than in irrigated rice

Overall parasites do not appear highly effective by themselves against upland or even lowland pests Among natural enemies parasites have attracted the most attention because they can be more readily assesscd

Predators and pathogens lend themselves better to management practices Predators the most important group of natural enemies in riccfields are difficult to quantify On the other hand pathogens appear less im-portant but can be readily cultured and disseminated

The natural enemy commulnity of upland rice is rich in species and differs significantly from that of lowland rice

These beneficial organisms must be conserved by applying insecticides judiciously particularly sprays Seed or soil placement of chemicals minimizes exposure to natural enemies The strategy to derive the greatest benefit from natural enemies isto allow the greatest number of beneficial species to thrive Then perhaps at least one species will be effective against each pest at any tine overcoming the variable abundance of each species within and between years

Programs to introduce exotic species or mass produce indigenous ones are ambitious and expensive because they require trained people on a sustained not ad hoc basis

Chemical control Insecticides offer rapid and efficient control of upland insect pests (190) but-should be used only after other control measures have been considered Insecticides are rarely used on upland rice because of cost (32) Upland rice yields are normally too low to justify the expenditure Also most upland rice is grown as a cash crop Spraying upland rice is more difficult than lowland rice because water is less accessible Government subsidy programs to provide pestishycides to tribal peoples for upland rice production have created problems of toxicity to humans because isolated people have had no experience to be able to handle pesticides safely

However ins-eticide use has beenjustified when a crop is

0Z

Table 8 Parasitization rates of lepidopterous pests in upland rainfed lowland and irrigated lowland environments Philippines 1976-84a

Parasitization (5)

Environment and site Year YSB GFB PSB Earcutting Swarming Skipper Greenborned RLF Brown larva larva caterpillar caterpillar caterpillar caterpillar larva semilooperEgg Larva larva larva larva larva larva

UplandBatangas 1977 - 0 (57) - 0 (63) - 0 (71) - 0 (106) 11(142) 2 (101)1978 43 (4697) 8(451) 5 (38) 3 (40) 6 (61) 2(58) 15 (57) 13 (129) 13 (1286) 4 (30)1979 59(4135) 12(320) 13 (11) 5(28) 9(45) 6(46) 7(23) 8(160) 9(1152) 8(26)

Rainfed lowland Iloilo 1976 - 0 (10) 0(10) 0 (7) 0 (19) 2 13) 15 (180) 5 (76)1978 0 (54) 0 (3) 0 (17) 9 (64) 4 (78) 3 (73) 0 (121) 0 (191)1979 64 (2350) 16 (84) 6 (17) 0(6) 18 (68) 25 (104)

1981 56 (1322) 18(304) 0(5) 12(51) 18(138)Pangasinan 1976 60 (7 7 )b 62(82) 0(111) 5 (39)

1978 58 (53)b 5(51) Cagayan 1979 59(713) 8 (101) 6 (34) 0(7)

1980 0(11) 4 (28) 15 (286)

62 ( 48(177) 61(140)1981 0(5) 12 (2) 13(45) 14 (92)

Irrigatedlowland Laguna 1974 62 8

1974-75 50 41 1975 43 72 1978 96 (623) 21(24) 33 (60)1979 83(2017) 18(110) 8(62) 11 (89) 22(400)1980 78 (1625) 20(75) 0 (16) 4 (72)1983 60

9 (80) 7 (270) 30

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R F Apostol unpublished Iloilo and Pangasnan 136 Cagayan B Canapi unpublished Laguna 175 P CPantua unpublished 118 234 143 Carino and Shepard unpublished YSB = yellow stem borer ScirpophagaincertulasGFB = gold fringed borer Chilo auricilius PSB = pink stem borerSesamia inferens earcutting caterpillar Alythimna separata swarming caterpillar Spcdoptera mauritia acronyctoides skipper Pelopidas mathias reen horned caterpillar Melanitls ledaismene RLF = rice leaffoldersCnaphalocrocismedinalis and Marasmiaspp brown semilooper Mocisfrugalis Figures in parentheses are sample sizes Egg masses

IRPS No 123 January 1987 35

Table 9 Effect of seeding rate and insecticide on plant density and Yield losses from insect pests in upland rice often areyield of Dagge upland rice Batangas Philippines 1976a comparable to those of lowland rainfed or irrigated rice

Seeding rate Insecticide Dosage Plant density Yield (0-Y30o) but usually from different pests The principal (kg aiha) seed treatment (kg aiha) (noIra-row) (tha) groups are soil pests feeding on sown seed and rootsat 14 DE followed by seedling pests These groups have largely gone

50 None - 35 a 20 a unnoticed leading researchers to conclude that stem borers50 Carbofuran F 10 38 ab 23 a and seed bugs are the most important pests (17) Yield loss 10 beCabofuranF100 None - 4551 c 23 a trials that pinpoint major pests need to includc seed

100 Dieldrin WP 10 52 cd 22 a treatments Granules applied in the furrow may not give150 None - 59 d 22 a adequate protection from many seed pests

aAv of 8 fields DE = days after crop emergence In a column means The floristic and pedological diversity of upland rice followed by a common letter are not significantly different at the environments is matched by the diversity of arthropods5level by D)MRT F = flowable Wi = wettable powder both pests and their natural enemies Irrigation has homoshy

genized lowland rice environments Flooded soils tend tothreatened by epidemics of locusts or armyworms New have the same properties and to eliminate soil pests The technology stressing efficiency has minimized insecticide spatial and temporal dominance of cultivated rice has cost benefited specialist pests

Seed treatment is normally inexpensive and can be Upland rice has no specialist arthropod species The economicallyjustified in -nanycases(66) Treating seed with monophagous pests on upland rice prefer the lowlands systemic insecticide (216) but not chlorinated hydrocarbons where rice evolved Rice in the uplands exists only because (247) is most effective in controlling seedling maggots of man and even though lowland rice predates upland riceFoliar sprays have to be repeated to obtain control and specialist species evolved Wheat maize and sorghum do often are washed off by frequent early season rains not have monophagous pests leading us to conclude that Granular insecticides are effective but required dosages are there is no evolutionary advantage for inssects to have too expensive to justify except under heavy infestation narrow host ranges in upland graminaceae(247) When directed against first-instar larvae banding Pests have adapted to the uplands not only by having granules in seed furrows (177) can be effective against white wider plant host ranges but also by having long life cyclesgrubs at low dosages (025 kg aii ha) (134) Seed treatments undergoing dormancy or being dispersive Each pest has a are not promising for white grub control (248) unique life history strategy suited to highly unpredictable

Seed treatment is also effective ainst ants and may be environments In agivenyear someinsect specics an better cheaper than increasing the seeding rate In Batangas exploit upland rice as a food source but no pest is a senious treating 50 kg seed ha was as effective as increasing the one every year on upland rice seeding rate to 100 kgha and using untreated seeds (Table The low yielding potential of upland rice means less 9) In Brazil seed treatments control termites and other soil quantity and lower quality of food for pests to exploit but insects including the lessercornstalk borer(66) Domiciano also less incentive for farmers to undertake control (58) however found seed treatments inconsistent in their The most effective control would be early and synchroshyeffect against high populations of lesser corn stalk borer nous planting of early-maturing upland rice varieties to

Baiting can be an inexpensive way of controlling ants prevent insect pest buildup Breeding for insect resistance seedling maggot flies mole crickets and field crickets Baits should initially focus on stem borers because they are verymade of locally available material and impregnated with difficult to control by other means Biocontrol efforts insecticide can be sparingly distributed taking adantage of should first stress conservation of natural enemies preshythe pests ability to disperse and encounter bait sites dators and pathogens rather than parasites Efficient

The new low-volume hand-held controlled-droplet chemical control includes seed treatment or baiting for seed sprayers may offer an advantage over high-volume knap- seedling and some root pests and using controlled dropletsack sprayes for controlling foliar pests Upland rice sprayers for spot treatments against foliar pests and seed canopies are more open than those in lowland rice and bugs when economic thresholds are reached better droplet penetration should result Controlled droplet applicators would be ideal for seedbugs and defoliators

CONCLUSION

Upland rice is attacked by a wider array of insect pests than is lowland rice mainly beca use of the addition of soil pests Generally population levels of lowland rice pests are lower in upland than lowland rice leading many people to conclude that insect pests are not important on upland rice

36 IRPS No 123 January 1987

REFERENCES CITED

1Abraham C C and K S R Mony 1978 Occurrence ofLeptocorisa acuta Fabr (Coreidae Hemiptera) as a pest of nutmeg trees J Bombay Nat ttistSoc 74553

2 Agyen-Sampong M 1975 HlierogrlphusdiaganensisKrauss (Orthop-tera Acrididae) a new pest of rice in Northern Ghana Ghana J Agric Sci 8249-253

3 Agyen-Samnpong M 1976 Varietal resistance and other inle-ratcd pest control components as factors of rice varietal improvement in West Africa Prcsented at the Second varietal improvement serinar 14-18 Sep 1976 Bouak Ivory Coast West Africa Rice Development Association Monrovia Liberia 18 p

4 Agyen-Sampong M 1977 Outbreak of Ihccrorrdnisorae in rice seedlings near Rokupr Sierra Leone lit Rice Res Newsl 2(6)16-17

5 Agyen-Sampong M 1980 Dirty panicles and rice yield reduction caused by hugs IntRice Res Newsl 5(l)11-12

6 Agyen-Sampong M 1982 The major pest problems of irrigated upland and mangrove swanip rice ecosystems ir the humid tropical Guinea savanna and Salel climate zones Pages 318-328 in Pro-ceedings training course cn concepts techniques and application of irtegrated pest management in rice inWest Africa 10-20 Jan 1982 Fendall Liberia West Africa Rice l)evelopioent Association Monrovia Liberia 505 p

7 Akibo-Betts V I and S A Raymundo 1978 Aphids as rice pests in Sierra Leone lit Rice Res Newsl 3(6)15-16

8 Akinsola F A 1975 Present status of different rice stemborers in parts of Nigeria iicL Entomo Newsl 328

9 Akinsola FA 1979 Ihe biology and ecology of rice stemborers in Nigeria Ph 1) thesis University ol lbadan Nigeria 217 p

10 Akinsola E A 1979 Guidelines of integrated management of insect pests of upland rice in West Africa Presented at the Seminar on integrated management of rice diseases and insect pests at Bobo-Dioulasso Upper Volta 17-22 Sep 1979 West Africa Rice Develop-ment Association Monrovia Liberia

II Akinsola IA 1984 Insect pests of upland rice in Africa Pages 301-305 in An overview of upland rice research Proceedings of the 1982 lIouak Iwsrv Coast upland rice workshop International Rice Research Institute 13O Box 933 Manila Philippines

12 Alam M S V TJohn and Kaung Zan 1985 Insect pests and diseases of rice in Africa Pages 67-82 in Rice improvement in Eastern Central and Southern Africa proceedings of the international rice workshop at Iusaka Zambia 9-19 April 1984

13 Alticri M A A van Schoonhovcn and J D Doll 1977 The ecological role of weeds in insect pest management systems a review illustiated with bean (Phaseohsvulgaris L)cropping systems PANS (Pest Artie News Summ) 23195-205

14 Anmante F 1967 The leaf-cutting ant Atia capiguaraa pest of pastures [in Portuguese] Biol6gico (Sao Paulo) 33113-120 (Rev Appl Entomol A56207)

15Andrade A C and D Puzzi 1951 Preliminary results of experi-merits on the control of Sculptcoris iastantus on sugarcane [in Portuguese] Bioulgico (Sao Paulo) 1744-49 (Rev Appl Entomol A41421422)

16 Appert J 1970 Maliarplia separaelha (rice stem borer) new observations of entomological problems of rice in Madagascar [in French] Agron Trop (Paris) 25329-334

17 Ariadeau Mand Z Harahap 1986 Relevant upland rice breeding objectives Pages 189-197 in Progress in upland rice research Proceedings of the 1985 Jakarta conference International Rice Research Institute 1O Box 933 Manila Philippines

18Asar S and Kisimoto 1975 Seasonal occurrence of the smaller brown planthopper in upland rice [in Japanese] Proc Kanto-Tosan Plant Prot Soc 2228

19 Bakker W 1974 Characterization and ecological aspects of rice yellow mottle virus in Kenya Meded Landbouwhogesh Wage-ningen 152 p

20 Balut F F 1970 White grubs attacking rice Oryza ativa L [in Portuguese] Biol6gico (So Paulo) 36321-322

21 Banerjee S N and L M Pramanik 1967 The lepidopterous stalk borers of rice and their life cycles in the tropics pages 103-124 in The major insect pests ofthe rice plant Proceedings ofa symposium at the International Rice Research Institute September 1964 The Johns Hopkins Press Baltimore Maryland USA

22 Barrion A T and J A Litsinger 1980 Ants a natural enemy of cliaiarac in dryland rice IntRice Res Newsl

5(4)22-23 23 Barrion A T and J A litsinger 1981 The spider fauna of

Philippine rice agroecosystem 1Dryland Philipp Entomnol 5139-166

24 Barrion A T and 1 A Iitsinger 1984 The spider fauna of Philippine rice agroccosystcms II Welland Philipp Eiitomol 611-37

25 Barrion A T1 and J A Litsinger 1984 lihanus (Diptera Tabanidae) eggs an alternative host of rice stei borer (SI3) egg parasite Telenoonis digns (Iliymenoptcra Scelionidae) lit Rice Res Newvsl 9(6)19

26 Barrion A T and J A Litsinger 1985 Identilication of rice leaffolders by wing markings Int Rice Res Newsl 10( 1)24

27 Barrion A T and J A Litsinger 1985 Chctiosspp (Coleoptera Carabidae) a leaffolder ILF)predator lit Rice Res Newsl 10(1)21

28 Bertels A 1970 Insect pests of rice and theircontrol [in Portuguese] Pelotas Brazil Instituto de Expcrimentacao Agropccuararias do Sul Circular 43 24 p

29 Bonzi S M 1982 Ctilo (flisilineus J de Joannis (Lepidoptera Pyralidae) stem borer of irrigated cereals in Upper Volta LioFrench] Agron Trop Riu Rizic Cult Vivrieres Trop 37207-209

30 Bourke T V T L Fenner J N LStihick G Baker EHassan D F OSullivan and C S li 1977 Insect pest survey for the year ending 30th June 1969 Papta New Guinea )epartment of Agrishyculture Stock and Fisheries Port Moresby 57 p

31 Bowden 1 1973 Migration of pests in the tropics Meded Rijksfac Landbouwwet Gent 38785-796

32 Breniere J 1973 The main rice parasites in West Africa and their control Pages 27-42 in Proceedings seminar on plant protection for the rice crop West Africa Rice Development Association Monrovia Liberia

33 Breniere J 1976 The principal insect pests of rice in West Africa and their control West Africa Rice Development Association Monrovia Liberia 52 p

34 Brown ES 1962 The African arinyworn Spoiolptera eXI)Ina (Walker) (Lepidoptera Noctuidae) a review of the literature Cornshymonwealth Institute of Entomology London UK57 p

35 Brown F B 1968 Upland rice in Latin America FAO-International Rice Commission 12th Session of the Working Party on Rice Production and Protection 2-14 Sept Peradeniya Sri lanka Food and Agriculture Organization Rome Italy 6 p

36 Buyckx F J 1 1962 Disease and insect pests of crops inCongo Rwanda and Burundi [iil French] Publications dc LInstilut National pour lestude Agrononiquedu Congo M Weissenbruck S A Brussels Belgium 708 p

37 Cariiio F 0 1981 Role of natural enentics inpopulation suppression and pest management of green rice lealhoppers Ph D thesis University ofthe Philippines at Los Bafitos PhilippinesLaguna 136 -

38 Carpente A J 1983 The development of rice cultivation and extension of vegetable productin in Zanzibar Final Report Vol II1 October 1983 Food and Agricultme Organization Ronte Italy

39 Carroll C P and S1 Risch 1983 Tropical annual cropping systems ant ecology Environ Manage 751-57

4(0 Cates R G 198 1 Host plant predictability and the feeding patterns of monophagous oligophagous and polyphagous insect herbivores Oecologia (Berl) 48319-326

IRPS No 123 January 1987 37

41 Cendcafia S M and F B Caora 1967 Insect pests of rice in the 62 Ene 1 C 1981 I)iiorldhol)ierusihilis Slater (licteroplera Philippines Pages 591-616 in The najor insect pests ofthe rice plant lygacidue Illissinac) a new rice pest front Nigeria lit Rice Res Proceedings of a symposium at the International Rice Research Newsl N3)15 Institute September 1964 The Johns Hopkins Press Baltimore 63 Fugenio T S 1965 rhe effect oi e-vCgetation on rice production Maryland USA 729 p and the possibility of overcoming it by leaf pruning MS thesis

42 Centro Ihternacional de Agricklittira Tropical 1984 Upland rice in University ol the Philippines at Los Baijos ILaguna Ihilippins 69 p Iatin Anerica Pages 93-119 hi An os ervies of uplaind rice research 04 Farrow R A 1975 The Alrican nmigratory locust ii is main Proceedings of the 1982 Boinakamp Ivory Coast upland rice workshop outbreak area of the middle Niger qlinlitatise studies of solitary International Rice Research Institute P0 Blox 933 Manila populations in relation to environmental factors Locusla No II Philippines 198 p

43 Chandra G 1979 Taxonomy ad hn riononlics ofthe imect parasites of 65 Fennal R G 1943 Food crop investigation Windward and rice Icalloppers and planthoppers in the Pihilippines and their Ieeward Islands First report Report onl work dunring the periodiiiportairce iii natural biol)gical cointrol Philipp Fntoltol 4119-139 April 1942-April 1943 Imperial College of Tropical Agriculture

44 Chang T T 1976 The originii colut ion enlIivation dissemination Trinidad 16 p (Rev of Appl Fintornol A3222-25) and diersificition of Asian and African rices -uphytica 25425-44 1 66 Ferreira F 1983 Integrated pest ritaageiret of upland rice [ill

45 Chatierjee P B 1979 Rice leaffolder attacks in India lit Rice Res Portuguese English summary] Pages 323-341 in Upland riceculttre Newsl 4(3)21 factors affecting production E Ferreira T Yamada and F-

46 Cheng Cbien-pan 19(4 Upland rice in Taiwan International Rice Malmolta eds Institutoda Potassae Foslato Institnto Internacional Cnmmission Wiorking Party oit Rice Production ard Protection I()th da Potassa Piracicaha Sao Paulo lraid Meeting 3-10 March 1964 Manila Philippines Food and Agricul- 67 Ferreira E arid F dl Silva Martins 1979 Occurrence of ture Organiaion Rome Italy 5 p Sogatohs orvziola (Muir) inupland rice in Goias Ilrail lit Rice

47 Chcrrett 1 M and 1) 1Peregrine 1976 A review of the status of Rcs Ncwssl 4(4)18 leaf-cutting ants and their control Ann Appl Biol 84124-128 68 Ferreira E and 1F da Silva Martiris 1984 Insect pestof rice ill

48 Chipeta F M 1974 Pests of rice in Malawi Pages 90-95 in Brazil and their control Jill Portugucsej FMIlRAtPA-CNPAF Proceedings Sith eastern Africa cereals research conference 10-15 Goiania Brmiil 67 p March 1974 DRB Mialda ed Ministry of Agriculture lilongwe 69 Ferreira FI F ca Silha Miartins S S Nero and 1 It P de Malawi Carvallio 1982 M tltiplc regression analysis todeterimine the effect of

49 Chu Yau-i 1979 The d atiaging period of the rice seedling fly insect populatiois ott upland rice yield [inIPortuguese Pesqui (Atlterigotiavxigu Stcin) (NItiscide Diptera) oil cori) at Fast Java Agropecu Bras 17671-675 Indonesia Plant Prot Bill 21403413 70 Ferreira F 1F da Silva Martins and F 1PZimmermann 1979

50 Conklin H C 1957 IHaninoo agriculture a report on aln integral Rice varietal resistance to tle lesser corn stalk borer [in Port uguese] systent of shi fti rig cult ivationi in the Pltilippincs Food and Agriculture Pesqui Agropccu Biras 14317-321 Organization Rome Italy 209 p 71 Fleming W F 1968 Biological control of lie hapanese beetle U S

51 Cramer H II 1967 Plant proection and world crop production Dep Agric RLs Sers [ech Bull 1383 78 p Pilaniensclhutz-Nachr 201-524 72 Food and Agriculture Organiation of the United Nations 1982

52 l)ani R C and Majunidar 1978 Prelininary observations on tie Country report surmimary Pages 27-37 in Report on the first incidence of root aphids on differeit rice cultnrcs Sci Cult 4488-89 FAOIUN)IPhail and regional training coiuirse in iniproved ctltural

53 Davidson R I 1979 lffects of root feeding on foliage yield Pages practices for upland rice 5-23 Oct 1981 Northern Region Agrculturd117-120 in T K Ci osb and R B Pottinger eds Proceedings of the Developririt Center Chiang Mai Food and Agricuilture Organiiashy2nd Australian conlerence on grassland invertebrate ecology Govt tion Rome Italy 37 n Printer Wellington New Zealand 73 Franco do Ariaral S 195 1 Residual action of insecticides in turfy

54 l)ean G 1W 1976 Rice insect pests inlaos Int Rice Res Newsl soil--the results of a field experiment [in Portuguese] Biol6gico (Sao 1(2)15 Paulo) 17209-214 (Rev Appl Entomol A41423)

55 I)ean G J W 1978 Insect pests of rice in Laos PANS 24(3)280-289 74 Freeman ) 1970 Report onithe Iban london School of -coroiics 56 )ingle H 1984 Behavior genes and lifc histories conplex adapta- Monographs on Social Anthropology No 41 Athlane Press

tions to uncertain environments Pages 169-194 in A new ecology London UK 317 p novel approaches to interactive systems P W Price C N Slobod- 75 Fuchs T W1 A Harding and 1W Smithlr 1979 Induction and chikoff and W S Gaud cdsl ohn Wiley New York USA termination oldiapause in the sugarcane borer Ann Flitonlol Soc

57 Dobelmann-JP and M Falais 1968 Upland rice cultivation ill tie Am 72271-274 North-western part of Madagascar in 1966-1967 [in French English 76 (hbriel B1) 1975 A review tf the major insect pests of some tpland summary] Trop Agron 2313-21 crops in tlc Philippines Pages 86-111 in Ad Htoc panel of experts on

58 Doniiciano N L 1979 Chemical control of tie lesser corn stalk pest disease arnd weed problems in some rainfed crops Bangkok (lolsonno~paloIgnowslhts) (Zeller 1848) (I-epid(ptera Ihycitid ae) Thailand Food aid Agriculture Organization Roire Italy

on rice in Ponta Gressa Parara 1977-1978 [in Portuguese] Pages 77 Garg D K and N K Shal 1983 White grub outbreak on rainfed 291-303 in Anais 1 Reuniao de tecnicos cmiirizicultura do estado de dryland rice in Uttar Pradeslh Int Rice Res Ncwsl 81(4)16-17 Sao Paulo-Campinas Cati Brazil 78 Garg I) K and IP Tandon 1982 Major insect pests of rice on hilly

59 Dunsmore 1 R 1970 Investigations on tlie varieties pests and tracts of Uttar Pradesh India lilt Rice Res Newsl 7(11)11-12diseases of upland rice (hill padi) in Sarawak Malaysia Int Rice 79 Garrity 1) P 1983 Upland rice research an international bibliograpy Comm Newsl 1929-35 1975-1982 Iternational Rice Research hnstitnte PO Box 933

60 Elias 1R1965 The new decline of rice seedlings is not caused by Manila Philippines 121 p parasites [in Portuguese] Biol6gico (Sao Paulo) 313-6 80 Ghauri M S K 1982 A new subspecies of Iinorh)rone bull

61 Empresa Brasiliera de Pesquisa Agropecuaria 1984 Upland rice itt cornlus Slater (Hemiptera Iygaeidae) front Papua New Guinea on Brazil Pages 121-134 it Av overview of upland rice research rice and carpet grass Ihll lEntoniol Res 72133-137 Proceedings ofthe 1982 Blouak Ivory Coast Upland rice workshop 81 Gonzales F 0 PArregoces R I ihernandef and OT Parada International Rice Research Institute PO Box 933 Manila 1983 Insect and imite pests of rice arid their control in Latin America Philippines [in Spanish] la Federacion Nacional de Arrocerts (Fedearroz)

Bogota Colombia 60 n

38 IRPS No 123 January 1987

82 Grist D H and R J A W Lever 1969 Pests of rice Longmans London UK520 p

83 Hale P R 1979 Two subterranean pests of upland rice in Papua New Guirnca IntRice Res Newsl 4(2)16-17

84 Hale P R and J T Hale 1975 Insect pests of rice in the New Guinea Islands Rice Entomol Newsl 34

85 Harris W V 1969 Termites as pests of crops and trees Common-wealth Institute of Entomology London UK 41 p

86 Hattori 1 1968 A method to distinguish between grass leaf roller (Ctaplha rocis medinalisGuenee) and the rice leaf roller (Susurtia exigua Butler) [in Japanese] Plant Prot (Japan) 23167-170

87 Henry I E and 1A Onsager 1984 Experimental control of the mormon cricket Iahrosshnph by NJsena locustae(Aficrospora Microsporidae) a protozoan parasite of grasshoppers (Ort Acri-didae) Entomophaga 27197-201

88 Hidaka 1 P Vungsilabulr and SKadkao 1974 Studies on ecology and control of the rice gall midge in Thailand Tech Bull 6 TropicalAgricultural Research Center Tokyo Japan 113 p

89 Hincklcy A 1) 1963 Ecology and control of rice planthoppers in Fiji Bull Entomrol Res 53467 481

90 Hirao J 1982 Ecology and chtInical control of the rice leaffolder Jpn Pestic Inf 4114-17

91 Ho DT 1984 Sten borer incifience in rice ecosystems in KenyaEast Africa Pages 307-320 in A overview of upland rice research Proceedings of the 1982 Bouac Ivory Coast upland rice workshop International Rice Research Institute PO Box 933 Manila Philippines

92 Ho )T J 1Njokah and J GKibuka 1983 Studies on rice sten borers in Kenya with emphasis on ifaliarpiaipiraellRag Insect Sci Appl 465-73

93 Ichihara I T Ito A NIatsuo and M Sawada 1971 Chemical control of the rice striped stem borer in upland rice grown in plastic mulch [iii Japanese] Proc Kanto-Tosan Plant Prot Soc 1882

94 IclhiharaIand A Matsuo 1969 Damage caused by rice striped stett borer to upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1686-87

95 Ichhara I atid A MIatsun11970 Conside ration oft lie extraordinary prevalence of rice striped stem borer in upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1778-79

96 Institut de Recherches Agrolnliqes Iropicale et des Cultures Vivrieres 1978 Insect resistance Pages 12-13 in Upland rice 1978 atnual report Bouak Ivory Coast 27 p

97 International Institute for Tropical Agriculture 1983 Aphids itt rice roots I ITA Res Briefs 4(1-3)4-5

98 International Institite fi Tropical Agriculture 1983 Hopperburn in rice caused by brown plathopper IITA Res Briefs 4(l-3)6-7

99 International Rice Research Institute 1975 Annual report for 1974 PO Box 933 Manila Philippines p 216

100 International Rice Restarch Institute 1976 Annual repiort for 1975 PO Box 933 Manila Philippines p 228

101 International Rice Research Institute 1979 Cropping systems program Component technology development and evaluation Page 404 i Annual report for 1978 11O Box 933 Manila Philippines

102 International Rice Research Institute Department of Entomology 1981) Insecticid evaluation 1979 Tables 73 74 PO Box 933Manila Philipes

103 International Rice Research Institute Department of Entonmology 1981 Insecticide evalomtion 1980 Table 81 POBox 933 Manila Philippines

104 International Rice Research Institute 1981 Report of a meeting of Asian rice-based cropping systems entomologists Eleventh cropping systems working group meeting 18-22 May 1981 PO Box 933 Manila Philippines p 132 133 144

105 International Rice Research Institute 1984 Cropping systems program Pages 368-373 in Annual report for 1983 PO Box 933 Manila Philippines

106 International Rice Research Institutc 1984 Upland rice in Asia Pages 45-68 in An overview of upland rice research Proceedings of the 1982 Bouakamp Ivory Coast upland rice workiiop loternational Rice Research Institute 11O Box 933 Manila Philippines

107 Irshad M 1977 Grasshoppers in rice in Pakistan lit Rice Res Newsl 2(3)8-9

108 Ito R and TOno 1964 Upland lice its culture and varieties in Japan lIternational Rice Cormmission loth Meeting of the Working Party on Rice Production and Protection 3-10 March 1964 Manila Philippines Food and Agriculture Organization Rome Italy 12 p

109 Jeffrey M I and V A Dyck 1983 Effect of moonlight on lightshytrap catches of brown planthopper lit Rice Res Newsl 8(6) 13

110 Jennings P R and T Alicia Pineda 1983 Management ofthe rice plnthoppers Sogatodsorivi(coIl in Latin America Presented at the meeting on stable plant resistance FAO and lIndocsian I)inectorate of Food Crops Protection Bali lndonesia 29 Novcniber-2 l)ccemshyber 9 p

IllJimenez Gomez J A 1979 Rice insect pests i lla ios ()riCntales lil Spanish] Pages 99-103 in Rice ciltivation Instituto (olotibiatto Agropecuario Compndio No 29

112 Jongeleen F JJ Soesanto and A Mockidjo 1979 Studies oti pathogens of two species of Scarabacidac froinIndonesia Z Angew Enlomnol 87412-417

113 Jouvenaz 1) PC S lofgren and W A Baiks 1981 Biological control of imported fire antsa review of cirrent knowledge Bull Entomol Soc Am 27203-208

114 Kainoh Y K Shiniiu S MarUani Y Taiaki 1980 lost finding behavior of tie rice bu lplotori+JchinvittjI)allas (liciniptera Corcidac) with special reference to diet pattern ol aggregation and feeding on the rice plant Appl Entomol Zool 15225-233

115 Kakkar K 0 P Sharma and P I Sharma 1977 Note on Ihteronyrchus liodresRedt (Coleopicra I)ynastidae) as asporadic pest of paddy and maie inHimachal Pradesh Corr Agric 156-57

116 Kalode M131 79 Major insect pest problems and their control in upland rice (Unpublishcd mineogriph CWI oJNaaiat lni Chandaratat (198 1)original not seen)

117 Kalshoven L i 98 I The pests of crops in I ldoniesia (translated by P A vatn der Laan) 1 T Ichtiar B3aru-vati Iloeve lakarta Indonesia 701 p

118 Kamal N Q 1981 Suppression of whitebacked planto ner Sogatella fircaifra (Horvath) aLId rice lealfolder (nhal -cis licdb lis (Guenee) populations by natural cnclmie hlI) tiesis Grcgorio Araneta University Foundation Victoncta Park Metto Manila Philippines 179 Ip

119 Karim A N M Reaul 1975 A species ofliealy bug Trioni i70s ceres new oii rice in the Philippines Rice Entoniol Newsl 247

120 Katanyukul Wand C Chandaratat 1981 Insect pestsof uplaid rice and their control Report firstFAOUNlFi Tliailand training course on improved cultural practices for tplad rice Chiang Mai Thailand 5-23 Oct 1981 Food and Agriculture Organiation Rome Italy 22 p

121 Kern F 1955 lhletwioa(I)uninemts) bhhit Biturm(Coleoptera) a new pest of sugarcane inVenezuela [inSpanish with Etnglish sunmary] Agron Trop (Maracay) 527-29 (Rev Appl Eitoniol A4572)

122 Khan K M and S Ghai 1974 White grubs ald their control in India Pesticides (1Bombay) 819-25

123 Kiritani K 1975 Integrated pest management of rice pests Pages 66-70 itt Proceedings platning workshop ott cooperative hIeld research in pest managementP SMotooka ed East-West Center Honololu Hawaii USA

124 Kisimoto R 1967 Ecology of insect vectors forecasting and chemical control Pages 243-255 in The virus diseases of tle rice plant Proceedings of a symposium atthe International Rice Research Institute April 1967 The Johns Hopkins Press altimore Maryshyland USA

125 Kisimoto R and V A Dyck 1976 Climate and rice insects Pages 367-391 in Climate and rice International Rice Research Institute PO Box 933 Manila Philippines

126 Kureha S S Kobayashi SAsari and T Ishihara 1977 Life history and control of the mole cricket Grrllotalpa africana Palisot de Beauvois [in Japanese English translation 7520 1RRI library] Proc Kanto-Tosan Plant Prot Soc 21192-197

127 Kushwaha K S 1981 The black beetle as a pest of rice in Haryana India Int Rice Res Newsl 6(4)14

128 Lawani S M 1982 A review of tile effects of various agronomic practices on cereal stem borers Trop Pest Manage 28266-276

129 Leuck D B 1966 Biology of the lesser cornstalk borer in South Georgia J Econ Entomol 59 797-801

130 lever R J A W 1955 Insect pests of the rice plant in Malaya Int Rice Comm Newsl 1612-21

131 Li Ching-Sing and Shin-Foon Chiu 1951 The study ofthe rice gall midge Iachydiplosis orvza Wood Mason [in Chinese with English summary] 1Taiwan Agric Res 21-13

132 Iitsinger J A 1984 Assesament of the need-based application for rice Proceedings TechnologyTransfer Workshop 15-16 March 1984 International Rice Research Institute 1PO Box 933 Manila Philippines 6 p

133 Litsinger J A 1984 Environmental diversity and insect pest abundance with reference to the Pacific Pages 285-307 in Proceedings sub-regional training course on methods of controlling diseases insects and other pests of plants in the South Pacific 4-20 Oct 1982 Vaini Kingdom of Tonga Nukualofa

134 Litsinger J A RF Apostol and M BObusan 1983 White grub Leucopholis irrorata (Coleoptera Scarabaeidae) pest status popula-tion dynamics and chemical conllol in arice-maize cropping pattern in the Philippines J Econ Entoinol 761133-1138

135 Litsinger J A C de la Cruz F Raymundo A T Barrion M D Lumaban J P Bandong M S Venugopal F Paragna and E Balete 1982 Insect pests and insecticide reponse of dry-seeded bunded rice Pages 179-193 in Report of workshop on cropping systems research in Asia International Rice Research Institute Los Bafos Philippine

136 Litsinger J A M 1) Luinaban J P Bandong A T Barrion W Katanyukul and N Panda 1978 Insect pest management in rainfcd lowland rice agroecosystms of tropical Asia International Rice Research Conference 17-21 April 1978 International Rice Research Institute Los Bafios Philippines

137 Litsinger J A and K Moody 1976 Integrated pest management in multiple cropping systems Pages 293-317 in R I Papendick P A Sanchez and G B Triplett eds Multiple cropping American Society of Agronomy Madison Wisconsin USA

138 Loevinsohn M E 1984 The ecology and control of rice pests in relation to the intensity and synchrony of cultivation Ph D thesis University of London 36) p

139 Loevinsohn M E J A Litsinger N Panda M A Austria R F Apostol C Andrion and J Bandong 1982 A basis for predicting rice insect pest abundance in cropping systems Paper presented at the 1982 International rice research conference on rice and cropping systems 19-23 April 1982 International Rice Research Institute Los Bafios Iaguna Philippines 25 p

140 Lopez A W 1930 Repot of the Entomology Department Pages 145-172 in Annual report of the Philippine Sugar Association 1929-30 Manila

141 Lopez AW 1931 Entomology Department Pages 227-266 in Annual report of the director or research 1930-31 Philippine Sugar Association Manila

142 Lopez A W 1932 Entomology Deparment Pages 252-279 in Annual report of the Philippine Sugar Association 1931-32 Manila

143 Lungibill P and G G Ainslee 1917 The lessercornstalk borer US Dep Agric Agric Res Serv Tech Bull 539 27 p

IRPS No 123 January 1987 39

144 McGuire J U Jr and BS Crandall 1967 Survey of insect pests and plant diseases ofselected food crops of Mexico Central America and Panama U SDepartment of Agriculture International Agriculshytural Development Service Agricultuial Researrh Service October 1967 11 p

145 McNeill Sand T R ESouthwood 1978 The role of nitrogen in the development of insectplant relations Pages 77-9 in lliochemical aspects of plant and animal coevolution J B Ilarbornu ed Academic Press London UK

146 Maebara H 1955 Spatial distribution of lastinstiarlanaeof purplish stem borer (Sesamia iqferens Walker) hibernating in an upland rice field Bull Kagoshima Univ Fac Agric 438-41

147 Mallick E H and N G Hajra 1977 Response of rice plant to defoliation treatments at different developmental stages Indian Agric 2151-56

148 Mansingh A 1971 Physiological classification of dormancies in insects Canl Entomol 103983-10)9

149 Martell C G 1974 Catalogue of plant feeding insects of Mexico [in Spanish] Fitofilo 27(69)1-176

150 Martinez 0 J 1982 Principal rice pests in Ilanos orientalis [in Spanish] P~ages 80-94 in Curso deact nalization de conocimientos en arroz Villaniencis Instituto (olonmbiano Agropecuario

151 Martins J F da Silva F Ferrcira A S Prabhu F 1 P Zimmermann 1980 Use of pesticides for the control of the main subterranean upland rice pests esqui Agropecu Bras 1553-u

152 Martins I F da Silva F Fe eira rod BdamSilveira Pinheiro I Q2 Damage simulation caused by defoliators of upland rice [in Portuguese with English summary] Pesqui Agropecu Bras 171113-1119

153 Martins 1F da Silva N(iyuen van Tan and 1Ida Silveira Pinheiro 1981 Stem borer resistance in upland rice associated with plant morphological characters [in Portuuce Iesqui Agropecu Bras 16187-192

154 Masaki S 1980 Sumnmer diapause Ann Rev Entomiol 251-25 155 Masuzawa T 11 Suwa and F Nakasoji 1983 l)ifkleences of

oviposition preferencc and survival rate of two skipper butterflies Parnara guttata and PIchlpidai iathiais (Iepidoptera HIesperiidac) on rice plant and cogon grass New Entomol 3247-56

156 Maurya D M and C P Vaish 1984 Upland rice in India Pages 69-92 in An overview oftpland rice research Proceedings ofthe 1982 Bouakamp Ivory Coast upland rice workshop International Rice Research Institute IO Box 933 Mnila Philippines

157 Menon M G R R N Katiyai S Kuar and S S Misra 1970 Termites attacking the paddy crop Sci Cult 36294-295

158 Miyahara Y and 1 Nishikubo 1967 Behavior of young nymphs of the red rice root aphid Iholahsipiun ri liabhdontinalisSasaki in relation to movement from the leaf to the roots of upland rice [in Japanese] Pioc Assoc Plant Prot Kyushu 1363-66

159 Mo ljoa Tjien 1952 White grub of upland rice (Holotrichiahelleri Brsk) [in Indonesian with English summary] Contrib Agric Res Stn Bogor No 13134 p

160 Mochida 0 and T Suryana 1979 The occurrence of the brown planthopper and ragged stunt on lowland and upland rice ill Central Java and logyakarta in tile late of January 1979 Entomnolgy Survey Report No 15 Central Research Institute of Agriculture (CRIA) Sukamandi Indonesia I February 1979 9 p

161Monet C 1979 Varietal resistance to upland rice pests [inI French] Pages 35-39 in Annual report 1977 Institut des Savanes Department des Cultures Vivrieres Bouak6 Ivory Coast

162 Monte 0 1942 A caterpillar ofrice-fields[in Portuguese] Biol6gico (Sao Paulo) 8161-163 (Rev Appl Entomol A31414)

163 Munroe I) D 1979 Pest management strategy for hill rice in four resettlement areas in Sarawak Malaysia Pages 89-102 i1 Guidelines for integrated control of rice insect pests Food and Agriculture Organization Plant Production and Protection Paper 14 Rome Italy 115 p

40 IRPS No 123 January 1987

164 Nakasuji F M Ishii and Jen-Zon Ito 1984 Rice skippers in Taiwan and their life histories ZZrC (Taipei) Tech Bull 82 10 p

165 Navas I) 1976 Fall armyworum in rice Pages 99-106 il Ecological animal control by habitat management Proc Tall Timbers Con-ference 28 Feb-I Mar 1974 Fall Timbers Res Stn Misc Publ 6 Tallahassee Horida USA

166 Nickel 1 1 1961 Biological control of rice stem borer A feasibility study Tcch Bull 2 International Rice Rcarch Institute PO Box 933 Manila Philippines IIl p

167 Nigan P M and R A Vernia 1985 Insect pests of uplaild rice in Uttar Pradesh Int Rice Res Newsl 10(6)22

168 Nilakhe S S 1985 -cological observations on spittle bugs with emphasis on their occirence in rice Pcsqtui Agropecu Bras 20407-414

169 Nixon G EJ 195 1 lie association of ats with aphids and coccids Commonwealth Iusti lcte of Entonology Iondon UK 36 p

170 Nurdin F an Ishak Manti 1985 Study oit seedling fly of upland rice Interlational Upland Rice Moitoring Tour Report 26 lebruary-2 MarLI Sukaranti West Sumatra

171 Oka 1 N 1979 Ciltural conttol of the brown planthopper Pages 357-369 in Brown planthopper threat to rice production in Asia International Rice Research Institutc 1p0 Box 933 Manila Philippines

172 Ohni M and A Vilnaii 1975 Rice insect pests of Italy biology of Sipha grcria (Kaltenbach) and its control [in Italian] Riso (Milan) 2459-71

173 Otanes F Q atnd 1 I Sison 194 1 Pests of rice Philipp J Agric 12211-259

174 Painter R I 1951 Insect icsistance in crop plants University Press of Kansits Lawrence Kansas USA 520 p

175 Pantua P C and 1A litsinger 1981 Arthropod abundance in continuous vs biannual rice cropping systeis Int Rice Res Ncwsl 6(2)20-21

176 Pathak M ) and V A l)yck 1975 Sttudics ott insect pests of upland rice Pages 186-191 in Major research in upland rice International Rice Research Institute PO Box 933 Ilanila Philippines

177 Patil B R B M IHasabe and 1) S Ajri 198 I Chemical control of white gritb lIditriihinhorraia Fabr intfesting groundntut paddy atnd sorghum Indian 1Entomol 43232-236

178 Pawar A 1) 1974 (ollection and identification of insect pests of rice rlerminal repirt lInternational Rice Research Institute PO Box 933 Manila Philippines 72 p (tnimco)

179 IerrajU A aid A S Reedy 1965 Observations ott diapausc itt tie steti borer ohrice Rice News Teller (lai-Mar)9-13

180 Pont A C 1972 A review of the Oriental species of theri ona Rondani (I)iptera Muscidac) of econtomic importance Pages 27-102 in Control of sorglium shoot fly M G lomtani and W R Young eds Oxford atd IH1 Ptthl Co New I)clhi India 324 p

181 Prasad S C 1PChand aid M 1PSingh 1983 Incidence of rice gall midge in the upland plateau region of Hihar Oryza 20235-236

182 Rai P S 198 0 Nisaga simpex damage to rice in the hill tracts of South India Int Rice kles News[ 5(I)12

183 Rathorc Y S 1969 Gonocplhahli dcprssunt Fabr as itpredator of (tilo zonwIlhis (Swinhoc) and other moths in field cage at Patnagar Uttar Pradesh Indian 1Entotnol 3127

184 Reed W 1983 Crop losses causcd by insect pists in the developing world Proceedings 10th Internatial Congress of Plant Protection lrighton UK 174-80

185 Reedy M V 1981 Ateactogasterinhens(Walker)a new root-grub pest of tipland paddy and inaize Curr Sci 501076

186 Reveno M A 1968 Catalogue of the insect pests of economic crops in Colombia [in Spaiish] Asociacion Latino Americana de Entomo-logia (AIAE) 129-33

187 Reyes G M 1957 Rice dwarf disease in the Philippines Food and Agriculture Organiiation Plant Prot Bull 617-19

188 Rice S E A A Grigarick and M 0 Way 1982 Effect of leaf and panicle feeding byarmyworm (Lepidoptera Noctuidae) larvae on rice grain yield Environ Entomol 7593-595

189 Risch S J 1) Andow and M A Altieri 1983 Agioccosystem diversity arnd pest control data tentative conclusions and nizw research dirctions Environ Entomol 12625-629

190 Rivera C T 1956 Three organic insecticides in thecontrol of insects affecting upland rice Philipp Agric 39465-472

191 Roffrey 1 1972 Locusta outbreaks in the Philippines Acrida 1177-188

192 Rojas R 1) 1941 The effect of pruning at varying ages upon the yield of Ramairio Phipp Agric 29851-860

193 Roonwal M i 1979 Ternite life and termite control in tropical South Asia Scientilic Pnbl odhpr India 139 p

194 Rosetto C1 F da Silva Martins N C Schmidt and L EAzzini 1978 Damage caused by pasturespittle bugs 4Dvoi)lavopictaandDJ schach) on rice lBragantia Nota 6 (Matcli)25-27

195 Rosztto C1- S S Neto I- Amantc I) Iink 1Garcia-Vieira D M de SoLza N V Bansa t and A M de Olivcira 1971 Rice pests of Brazil [in Portuguese] Pages 149-238 inrFA()-International Rice Commission 2nd meeting of the Rice Committee for the Americas 6-11 l)ecember 1971 Pelotas Rio Grande do Sul Brazil

196 Rothschild G H L 1969 Obseratiois on the armyworm Spodoptera matritia acronrctoiiles Gn (Lep Noctuidae) Bull Entomol Res 59143-160

197 Rothschild G 1- L 1974 Parasites of the armworm Spolopiera mauritiaacroncitoihes in Malaysia Fntornopliaga 19293-299

198 Sachan i N S K Gangwar and 1N Katujar 1980 Field crickets damaging cages in North Eastern IHill Region Indian J Entomol 42526-527

199 Sands 1) P A 1977 The biology and ecology of Lepocorisa (Hemiptera Alydidac) in Papua New Guinea Res Bull 18 Departshyment of Primary Industry Port Moresby Papua New Guinea 104 p

200 Sarawak 1968 titomology division Pages 97-106 in Report of the research branch of the Department of Altriculture 1966 Kuching (Rev AIppl Eintomol A56603)

201 Satpathy JM M S Das and K Naik 1977 Effect of multiple and mixed cropping on the incidence of scine important pestsI Entomol Res 178-85

202 Sauer IHF G 1939 Notes Ott Elasmpnlpualtsligniosellths Zeller (Lep Pyr) a serious pest ofcereals in Sao Paolo [in Portuguese] Arq Inst Biol 10199-206 (Rev Appl Entotnol A28378)

203 Satinders 1L A 1 SKing and C L Vargas S 1983 Crop pests of Central Anerica with t bibliography [in Spanish] Centro Agroshynotnico Tropical de Investigacion y Ensenanza Turrialba Costa Rica (CATIE) Tec Bol 9 90 p

204 Sauphanor B 1985 Some factors of upland rice tolerttce to stem borers in West Africa Inseu Sci Appl 6429434

205 Sen P atnd S Chakravorty 1976 Stentflics in Haritighata Nadia West Bengal India [it Rice Res Newsl 1(12) 15

206 Senapai B and 1M Satpathy 1983 The seedling fly Atherigona orrzae Malloch is a pest of rice in Orissa (India) J Entomol Res 777-79

207 Shhjahan M S Alatn M Ah nad and BN Islam 1983 A new scarabaeid beetle pest of dryland rice in Bangladesh Int Rice Res Ncwsl 8(I)16-17

208 Sharma G L S C Modgol and R C Gautam 1979 Effect of depth of submergence ott incidence of bacterial blight and white grub infestation in transplanted rice Int Rice Res Newsl 4(6)12-13

209 Sharma S 1985 Scarabaeid beetle outbreak itt upland rice Int Rice Res Newsl 1(1(l)24

210 Shiraki T 1917 The paddy borer Schoenobhits incertulas Walker [achaku Agric Exp Stn Formosa 256 p

211 Silva A de Brito and 11P Magalhaes 1981 Insect pests of rice [in Portuguese] Circular Tecnica No 22 EMBRAPA Centro de Pesquisa t ecuaria de Tropico Unido (CPAJU) Belem Brazil 14 p

212 Singh S A 1979 Emergence of rice leaf butterfly as a rice pest in Manipur India int Rice Res Newsl 4(4)16-17

213 SinghS A 1983 Changing trends of insect pestsof rice in Manipur Int Rice Res Newsl 8(f)12

214 Siwi S S and P H van Doesburg 1984 Lenovorisa Latreille in Indonesia (Heteroptera Coreidae Alydinae) Zool Meded (Leiden) 58117-129

215 Soejitno J 1977 Reaction of some upland rice varieties toseedling fly eltherikonaexigua Stein [in Indonesian] Siri Hana I Penyakit 630-35 Lembga I osat Fenclilian Pertanian Bogor Indonesia

216 Socnardi 1967 Insect pests of rice in Indonesia Pages675-683 in The major inse pests of thu rice plant Proceedings of a symposium at the Internatital Rice Rec Wi itt S pternber 1964 The Johns Hopkins Press Baltimore Maryland USA

217 Soto P E A T Petcz and I W Buddenhagen 1976 Survey of insect pests and diseases of rice in different ecological zones in Nigeria Rice Entomol Ncwsl 435-46

218 Soto 11E and Z Ziddiqi 1976 Screening for resistance to African rice insect pests Varietal improsetnent sCmi n r Bouak Ivory Coast 13-17 Sept 1976 West Africa Rice l)evelopment Association Monrovia Iiberia

219 Squire F A 1972 Entomological problems in Bolivia PANS (PestArtic News Sumr) 18249-268

220 Stapely 1 II 1978 The rice leaf roller in the Solomon Islands Int Rice Res Newsl 3(l)13

221 Suenaga H 1937 On aphids attacking the roots of rice Oyo-I)oubts Zasshi 9150-152 [in Japanese] (Rev Appl Entomol A25745)

222 Suenaga H 1952 On the seasonal abundance of three subterranean aphids on upland rice Bull Kyushu Agric Exp Sti 1249-262

223 Sutynso Yas and Dandi Soekaria 1964 Beberapa hana yang ttenperang tana man padi tanah kering [in Indonesian] Rapat kerja Padi Tanah Kering 6 sd 8 Juily 1964 Kaliurang 9 p

224 Tallamy I) W and R F Denno 1983 Alternative life history patterns in risky environments in example from lace bugs Pages 129-147 in Insect life history patterns R F l)cnio and If Dingle eds Spring-Verlag 225 p

225 Tamura 1 1939 Observations on some insect pests of field crops [in Japanese] 1 Plant Prot 26414-420 (Rev Appl EntomolA27-556)

226 Tamura and Suzuki 1964 Influences of the early-transplanting practice in rice culture upon development of the rice stem borer Chilo siql)ressalisWalker Bull Ilokuriku Natil Agric Exp Stn 761-94

227 Tan J P 1924 The rice root aphid Drowhi ihirsoia Philipp Agric 13277-288

228 Tanaka 1 1957 Taxonomy and distribution of some subterranean aphids injurious to upland rice in Japan with description of a new species [in Japanese English summary] Botyu-Kagaku 22168-176

229 Tanaka T 1961 The rice root aphids their ecology and control Spec [full College of Agriculture Utsunomiya University Japan F3 p

230 Tauber M IC A Tauber R Nechols and 1Obrycki 1983 Seasonal activity of parasitoids control by external internal and genetic factors Pages 87-108 itt I)iapause and life cycle strategies in insects V K Brown and I Ilodek eds )r W Junk The Hague Netherlands

231 Taylor W F 1972 Effects of artificial defoliation (simulating pest damage) on varieties of upland rice Exp Agric 879-83

232 feotia T P S and S Natud 1966 Biotoomics of the rice skipper Parnaraniathiis Fabricius (Lepidoptera Hesperiidac) Indian J Entomol 28181-186

233 Teran F 0 1971 Rice pests of Santa Cruz [in Spanish Ministerio de Agricultura Sita Cruz Bolivia Bull 45 8 p

234 Than Itun 1976 Population dynamics of the yellow rice borer Tryporza itcertulas (Walker) and its damage to the rice plant MS thesis University of the Philippines at Ios Bafios Laguna Philippines 92 p

235 Uichanco L B 1911 Pests Pages 55-74 it Rice Philipp Agric 11-200

IRPS No 123 January 1987 41

236 van der Laan P A 195 1 Life history of the rice seedling fly Contrib Gen Agric Res Stn Boor Indonesia 118 15 p

237 van der Laan 11A 1959 Correlation between rainfall in the dry season and the occurrence of the white rice borer (Scirpolhaga innotala WIk) in lava Entomol Fxp Appl 212-20

238 Van Driesche R G 1983 Meaning of percent parasitismin studies of insect parasitism Fnviron Entomol 121 I1- 122

239 Venugopal M S and A litsingcr 1984 Fffect ofcarbofuran on rice growth Prot Ecol 7313-317

240 Vietnam Directorate of Rural Affairs 1962 Annual work progress report on crop improvement program of rice sugarcane vegetable and field crops (lor the period from July 1961 to June 1962) (11) 299 p (Rev Appl Entomol A51526-527)

241 Wan M T K 1972 Observations on rice leaf- and planthoppers in Sarawak (Malaysian Borneo) Malay Agric 148308-335

242 West Africa Rice I)esclopnient Association 1978 Annual research report Vol 2 Statistics Monrovia Liberia

243 West Africa Rice Development Association 1979 Annual research report p47 Monrovia Liberia

244 Williams ) F and CS lofgren 1981 Fli I illy l-468 anew bait toxicant for control of the red imported lire at Fla Entomol 64472477

245 Williams I) 1 1970 The mealybugs OFlomoptera Coccoidea Iscudococcidac) of sugar-cane rice and sorghum Bull Entomol Res 0109-188

246 Williams D) 1 1982 Pulinaria icervi (Signoret) and its allies ott sugar cane and other grasses Btll Entoinol Res 72111-117

247 Wirjosuiardjo S 1976 Study ofchenmical cont rot of insect pests ott upland rice Imiu Pertanian (Jogjakarta) 1323-329

248 Wirjosuhardjo S and N Wirarta 1976 Chemical control of the white grub tIolorithiatlltriin upland rice in M uneng Irobolinggo Indonesia) [in Indonesian] Imut Pertanian (Jogjakarta) 1330-332

249 Yano K T Miyake and V F Eastop 1983 The biology and economic importance of rice aphids (Henuiptera Aphid idae) a review Bull Entomol Res 73539-566

250 Yelp D and P Rhodes 19(4 lhe success of drilled dryland rice iii the Fiji Islands International Rice Commission Working Party on Rice Production and Protection Manila Philippines 3-10 March1964 Food and Agriculture Organization Rome Italy 13 p

251 Young W R 1981 Fifty-five years of research on the sorghum shootfly Insect Sci Appi 23-9

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No 101The economics of hybrid rice production in (hina No 113 Boron toxicity in rice No 102 Rice ratooning No 114 Energy analysis rice production systems and rice research No 103 Growth and development of the (Jeep water rice plant No 115 Production risk and optimal fertilizer rates an application of the

No 104 Faridpur acomputer-assisted instruction nmodel forrainfed lowland random coefficient model No 116 Consumer demand for rice grain quality in Thailand Indonesia andrice

No 105 A readingand listening comprehcnsion test inEnglish for non native the Philippines speakers applying for tiraining at IRIRI No 117 Morphological changes in rice panicle development a review of

No 106 Rice grassy stunt virus 2 a new strain of rice grassy stunt in tire literature

Philippines No 118 IRRI-Korca collaborative project for the development of cold-No 107 Physical losses and quality deterioration in rice plstprodnCtion tolerant lines through anthei culture

systemis No 119 Problen soils as potenial areas for adverse soils-tolerant rice No 108 (olptblication ofIRRI materials a survey of translators and varieties in South and Ssuthcasi Asia

publishers No 120 Changes isinall-farm iLtireliing techii)hgy in Thailand and [he No 109 Classification of Philippine rainfall patterns Philippines No 110 Contributions of modern rice varieties to rutrition in Asia No 121 Landforms and modern rice varieties No Ill Changes in rice breeding in 10 Asian countries 1965-84 No 122 Yield stability and modern rice technology No 112 Design parameters affecting the performance of the IRRI-designed

axial-flow pump

Page 14: UPLANDRICE. INSECTl--1i M T -E!R.EOLOU t RTANCE ,AND …delphacid.s3.amazonaws.com › 6958.pdf · The significance of abiotic factors in upland rice insect ecology will be apparent

14 IRPS No 123 January 1987

upland savannas whereas Chilo zacconio prefers the lowlands (II) Aaliarpha is most abundant in the mangrove swamp habitats (91) coastal regions (16) or rain forest zones (II) of Africa and appears to be the homologue ofScirlpophaga incertulas The yellow stem borr can also be prevalent in upland rice (120 167) The sugarcane borer Eldanasaccharinais so far only a potential pest of uplandrice (91) despite Grist and evers (82) warning

Grasshotl)ers andI locitas Short-horned grasshoppers and locusts -- Locvta Patanrga Schistocerca - layeggs in the soil inhabit grasslands and develop into swarms to seriously damage upland rice from time to time (173) Largenumbers can destroy a field leaving only the stubble In low cmuoroer s giassnoppeirs an cut panicles (173) Farmers may not plant if locust swarms are imminent Prolonged dryweather followed by favorable rain favors development of large swarms (173) Usually breeding in di) grasslands theydisperse toattack crops including upland rice(191) Eggsaresusceptible to desiccation d uring drough t and toinundation during seasonal rains With favorable soil moisture over consecutive seasons natural enemies ca nnot prevent a largepopulation increase leading to migratory swarms (64)

Most grasshoppers on rice are adapted to uplands (107)but lowland rice grasshoppers iervgl(rlhusspp and Ora spp occur occasionally on upland rice (2 82) They lay eggson rice foliage Nymphs are scmiaquatic

The gryllid Lsctrtus coucinnus is a pest of lowland and deep water rice in Bangladesh and Thailand In the Philippines it damages upland rice grown in isolated pockets near lakes The gryllid nymphs and ad ults feed on the central portions of folrage leasing only the midrib and leaf nargins

Leal ler Pvralid moths whose larvae fold leaves to make a feeding shelter occur worldwide (217) but arc appareritly most common in Asia Leaffolde rsattacking rice in Asia are a complex of species (26) (Ctalhahcrcisinedinalis the best-known rice Ia flolde r in Asia may prefer grassy weeds over rice (86) It is a late colonizer of uplandrice U1arasmja eviua a lowland species (220) nreflers rice to grass specres (86) and colonizes upland rice during il early growth stages llarasnria patnalis is the most corn-rnonly encountered species in upland rice in the Philippines Perennial grasses are their alternate hosts and they can survive year-round in upland rice environments The leaf-folders 11arasmia ruralisand Alarasmia spp are least aburndant Dormancy is unknown in leaffolders Cnaplha-ocrocis medhnalis however is known as a migrant (90)Leaffolders remove photosynthetic tissue attack during the flag leaf stage is particularly injurious Popurlations are normally held in check by natural enemies but leaffolders become parlicularly abundant in conditions of high plantfertility and shade

Butterflies Skippers PelopidasInathiasand Borhofanta are more prevalent in African lowland than in uplandhabitats (6) but extensive studies in Japan and China

showed that Pelopidas mathias prefers an upland habitat (155) Parnaragutata seasonally migrates between lowland and upland areas and is equally abundant in both environshyments (164)

The green horned caterpillar Melanitis leda ismene is more abundant in lowlands (212) but our experience in the Philippines suggests that Pelopidas niathiasand 11elanifis leda ismene are often more abundant in uplands than in lowlands Bamboo iaunbuiva spp annually sustains P mathiaspopulations in the Philippine uplands Adults feed on nectar from flowers and migrate to upland rice during the late vegetative stage

iPi erlled h i-1 1d dill L tIC i 7-0 l(become numerous probably because of high egg predationEgg and larval parasitization are often low Butterflies can readily disperse long distances to seek renlote uplandricefields However they have low biotic potential and even it predation rates are low the rarely become abundant Upland environments with their diverse microhabitats can provide more favorable sites than lowland rice plains

tutterfly pests of rice have broad host ranges and become dormant Juring urnfavorable times of the year

IolhagrotslaidopwraThe larvae of many polyshyphagous moths are reported to defoliate upland rice tussock moths lsaispelnnatula Iawlia sUifita luproctisirgtwltaE mnorand P vathorrhoeaslug caterpillar

litotiahicolorhairy caterpillars Nisaga sinp vandA mata spp and wooly bears l)iacrisiaohliqtia and Crvatonotts gangis Many of these polyphgous larvac migrate and are transients in rice including HelhWhis arm iera and 1t zea highly pol phagous species (82)

Gall nidges The Asian rice gall midge Orseoliaorrzae (88 131) and its African counterpart 0 ortzivora(6 11 12 32217) prelerlowland to upland rice Gall midges in rainfed areas are prevalent only during tle rainy season and dispersepoorly Their alternate hosts (wild rices LeersiaIschaetnuan and Pasahmt)are mostlyaquatic grasses and are less likelyto be near upland rice areas In an area of mixed uplandfloating and rainfed lowland rice cultures in northern Thailand upland rice which is ilanted a month earlier was attacked firstbut subsequcent populations were much lower than in rainfed lowla id or floating rice Low numbers of gallmidges passed the dry season orr perennial weeds such as Paspahim (88)

Smot (hid(s Reports of aphids attacking upland rice plants are few (249) Without maize the maize leaf aphidRho alosihhunm maidis attacks upland rice in the Philipshypines The plum peach aphid IIrsteroneuraselariae was reported in Sierra Leerne toinfest leavesand tnriperied grain(7) Siphaglrctriaecaused transverse linrea r necrotic striae iir rice leaves in Italy (172) I-lea v rains held populations in check Rice is not a preferred host for shoot aphids

Leafhoper andI )anthol)pers Cicadellids and delshyphacidsdispersecasilyand readilycolonie upland ricefrom nearby lowland areas A study in Sarawak (241) showed

IRPS No 123 January 1987 15

similar species in upland and lowland rice Nephoiettix The principal vectors of rice virus diseases generally have species dominated The most prevalent species accounting narrow host ranges and alternate hosts in the grass family for 90-95 of the sweep net samples wre N virescens N are mostly annuals dying in the dr3 season Some virus umropictus Nilaparvia hgens Soatellafiircifera Cfana vectors such as BPH can disperse lontg distances but few do

sectraand Recilia dorsalisLowlands had higher popula- Th restriction of upland rice and weed hosts to the rainy tions Katanyukul and Chandaratat (120) monitored hopper season the small area planted to upland rice and the hopper numbers in upland and lowland fields of northern Thaiiand vectors having narrow host ranges all work against virus and found similar results A comparative study of plant- diseases in upland rice hoppers in Fiji showed more brown planthoppe BPH) and In Latin America Sogctdhs o(rzicola and S cuhanus whitebacked planthopper (WBPH) in wetland rice than in transmit hoja blanca virus disease but reports on upland lowland rice and hopperburn was prevalent in lowland rice are rare except in the favored upland areas of Colombia areas with standing water (89) Peru and Venezuela (35 42) S rricolaocurs in Brazil

Hopperburn in upland ricefields in Asia is rare (84) In on upland rice but hojaI blanca has not been reported (67) Batangas Philippines isolated fields of upland rice become S orzicola has a wide host ramgc (10) similar to WBPI- in hopperburned infrequently from WBPH A report from Fiji Asia WBPH however is not known to transmit any virus cites hopperburn from BPH and WBPH (250) In these two disease situations upland rice was grown on highly fertic soil and In Africa pale yellow mosaic virus is endemic to swampy N fertilizer was applied Hoppers respond to better nutrition lowland areas where rice ratoons year-round (19) and is Normally upland rice is grown on poor soil and receives no mechanically vectored mainly by hispid beetles A minor fertilizer vector a flea beetle Chaeiocnema spp is an upland rice

In Asia upland rice ras are normally not far from pest and upland ricetields near endemic areas occasionally lowland ricefields-- a source of hoppers Otherwise hopper contract the virus species with greater polyphagy than BPH or Nelhotettiv Spittle bugs Cercopid nymphs producc a protective virescens would be dominant frothy covering that looks like saliva In Latin America

In Africa leathoppers and planthoppers do not become spittle bugs damage upland rice In parts of Brazil where abundant on upland rice (6 II) probably because lowland upland ricefields are surrounded by pastures adults rice is not widely rlanted nearby The only reported migrating from grasses can damage rice severely and cause hopperburn as from Nilapanrata maeander in breeder complete hopperburn in a young rice crop within a week of plots receiving high N rates in Nigeria (98) Hoppers colonization (194 195) Dispersal to rice occurs even if however are a greter problem in Latin America where pastures are verdant and spittle bugs prefer pasture grasses large grassland areas breed polyphagous species (Gra- to rice (168) Nymphs are rarely abundant enough to phocephala spp Hortensia spp lEvitianus ohscurunervis damage rice Regular rainfall favors their development Balclutha spp and Draeculacephala spp) vhich then (233) disperse to cause hopperburn on seedling rice (68) Rosetto et al (195) reported Graphocep7hala spp and Hortensia spp Seed pests to be equally abundant on upland and lowland rice Alydid and pentatomid bugs are seed pests of upland rice

The leafliopper and planthopper epidemics in Asia that worldwide In Asia Leptocorisa sp prevail Their habits caused severe losses from hopperburn and vijus dicases make them particularly suitable for upland rice 1)thev feed over the past several decades occurred principally on on many grasses (and therefore can survive the earl y wet lowland rice Except in a few isolated cases (160) upland season before rice sets its grain) 2)adults are long-ied and rice has been spared from tungro grassy stunt and ragged mobile allowing them to find isolated plantings 0 rice stunt virus diseases vectorcd by green lcafhopper (GLH) (besides seed bugs only birds and rats appear to have shyand BPI] (123) Virus diseases have occurred in upland rice capability) and 3) adults aestivate in forested areas or where lowland rice was nearby Rice dwarf disease however sugarcane fields during the dry sca son when neither rice nor was first recorded on upland rice in the Philippines (187) grassy weeds are present (I199) Rice seed bugs concentrate Virus diseases can be perpetuated only on livir plants or on small-scale upland riceficlds because they can actively insect vectors and no living rice - plans tooned or search them out The prevalence of groves of trees characshyvolunteer - grows year-round in upland arcas excert in a teristic of upland environments ensures nearby aostivation few with 12 mo of rainfall sites keeping the seed bugs close by

The smaller brown planthopper Laodcelphax striatellu Leptocorisae are uncommon in lowland rice plains occurs in upland rice in Japan (18) It spreads black- because 1) their populations are diluted in a sea of rice streaked dwarf and stripe viruses between winter wheat and planted more or less at the same time (they ca n only feed on barley to lowland rice Early planting of lowland rice rice during the milk to hard dough stages) 2) they lack nurseries spreads these viruses but upland ice has remained aestivation sites and 3) weedy fallow areas are limited uninfected The lowands of Asia are dominated by L oratorius

16 IRPS No 123 January 1987

which lays its eggs hign on tile foliage (199) L acuta and L solonlonlesis oviposit at ground level the former on litter and the latter loose on the ground These egg laying habis explain theirtenvironmental prefcrencecs(214) In Japan the postdiapause adults of L chinensis migrate from moun-tainous areas to upland ricefields during floweiing (1 14)

Pentatornid bugs prevail in Latin America (8182 195) and in Africa pentatomid and coreid bugs arc equally cited (6 82) l)amage is often characterstic of the species (5)rcoris ventralis and Mnida spp are the common

pentatomid seed bugs of Philippine upland rice but they prefer lEchinochva spp grass seed to rice Vearaviritda occurs worldwide and hoften more abundant in the uplands(54) We saw few reports of seed bugs on upland rice but see(d bugs apparently opelrate cqually between by land and upland rice Good dispersal is necessary ior species that call feed for only 2-3 xon a crop and must first find those crops at ripening stages

Other pct species feed n flowers In Mato Grosso Brazil (194) and Santa Cruz Bolivia the panicle weevil Aeo-hariuia amllitarsis attacks rice during flowering (68 233)The larva e rice sten horers adutlts feed at the bases of spikelets caIusing empty grains Also inLatin America thecydnid -lkiituttatralusisa seed bug as anad tilt and nymph (194)

A number of beetles ecd on poilen - coccinellids lialrotia Aulac lhora and Monohpl-a (55) - but becatise riCe is self-pollinating and fcrtiliation occurs before pikelets open they pose no threat to yield

IST ECOLOGY

We now look at characteiistics of the life cycles of upland rice insect pests to lea rn hew each plays an adaptive role in upland rice ecology

Life history strategies Orzasativadoes not tolerate drodgh well so upland rice is highly seasonal normally grown in the wettest months of the year Usually rice is present for less than half of the year The rice-fiee fallow poses serious problems of survival to upland rice insects which have evolved att least four mechanisms to ovrcolne tle cvcl ic-l lack of a hostI) polyphagy 2) longevity 3) dormancy and 4) vagility (dispersal)

Poldt)agy lolyphagy is defined as having hosts of at least two botanical families oligophagy of more than one gents and ionophagy of only one ventus (40) - inour case Orvza Most wild rices however are illi tiC and theieiirc are normally far removed from upland rice hahitat othewise more would be alternate hosts for uplanJ iice insects Thte highly mornophagous rice pests shySciriol)haita hiceritlas Nilal)arvaiahiens Iand Nealo--tettiyvircsctws in Asia and Aalirpha secparatella and 1ioj)sis lngicornis inAfrica - occur on upland rice but

are more abundant in lowland culture They can specialize in rice because they are highly vagile and can attack all growth stages By specializing in rice monophagous rice pests can outcompete related but polyphagous speciesIntensive andI extensive lowland rice culture has favored monophagous species Riqwla alhinella (Cramer) the onlymonophagous species in Latin America is rarely reported on upland ice (I1)

Because of its limited temporal and spatial existence upland rice has not favored the evolution of specialized species Clhang(44)shows evidence that upland rice has only recently been cultured by man from the lowlands thus pests have had little time to fully adapt to the crop All but the lowland-adapted insect pest species have alternate planthosts to rice Root pests with limited mobility are highly polyphagous Pests that specialize in one growth stage eg secdling maggots seed bugs are oligophagous or polyphagous

Most alternat hosts of rice pests are annual grasseswhich also are seasonal Upla_rice is also a host but not the preferred one of the most highly polypliagous spcciessoil pests grass hoppers armvwarrs and Vearaviirielda

The d ry season poses igreat obstacle to ani upland riceinsects survival and only sonic perennial plant hosts are suitable food during the off-seasoin Plolyphagy allows upland rice insects to suLrx-ixe during thC rainy season when plants- both annual and perennial -areactivelygrowing and therefore more nutritious

Loiugevitr The ability of aiinsect to live 2-3 mo without undergoing dormancy helps species sich as root-feedingwhite grbs (as larvae) and secd-feeding bugs (as adtilt) to survive unfavorable periods White grubs arc not highly fecund and their life strategy is to improve survivorship of limited progeny Their subterranean habitat hides them from manv natural enemies lBtut root tissue is not highly nutritious so they ltilst eat great quantities and need a largeinsect biomass to digest it Other soil-inhabiting rootshyfeeding insects have short life cycles but are more fecund

Seed pests need greater longcvi vas adults to locate ahost at early grain dcclopment arnd give sedentary offspring time to develop Few groups of upland rice insects live longin active development Polyphagy and longevity only increase their ability to survive during the rainy seasonOther mechanisms are needed to survive the dry season

orantc In the tropics the nonrice season is the dry season while intemperate areas tile off-+season is winter Many upland rice insects undergo dormancy inlsummer (aestivation) or winter (hibernation) Dormancy is simplyinactivity during winters with prolonged temperatures below 10 C insects find shelter arnd cease movement Activity is resu med with warmer t empcera tu res A deep state of irnactivity termed diapauise occurs in sole species that prepare physiologically (havxing low metabolism and convcrting gonad s to fatty fuel reserves) for the unfavorable season (148) They enter diapaUse inanticipation of unshy

IRPS No 123 January 1987 17

favorable weather and resume activity after favorable Armyworms skippers and locusts actively travel hundreds weather begins usually in response to photoperiod or of kilometers in air fronts such as intertropical convergencethrough a delayed response to a stimulus such as heavy zones to descend onto upland rice fields far from their rainfall (237) Not all individuals of a population have the breeding grounds (31) Several generations can build ip on same threshold for environmental responses for entering or alternate hosts in tile grass family before rice is plantedterminating diapause -- this is seen as a mechanism to These large insects are active fliers Smaller insects such as prevent total mortality if a false cue appears such as a thrips aphids and planthoppers passively migrate long sudden warm spell in winter or rain in the middle of the dry distances by the wind season (154) Otherspecies ean readily travel tens of kilometers - stem

Probably more upland rice insects undL go dormancy borers leaffolders leafhoppes cutworms spittle bugs leaf than we realize Candidates are Atherigona spp root beetles seed bugs ants and termites aphids root weevils crickets leaf beetles thrips and seed The least dispersive species are white grabs mealybugSbugs Insects known to entervarious states of aestivation are seedling maggots crickets weevils and gall midgesleptocorisaspp adults (199) Maliarphascparatellalarvae Each species has evolved a unique set of attributes to (9) Scirpophaga innotata larvae (117) Chilo partelhtsv enable it to survive and adapt to the changing uplandlarvae (21) ScirpopThagainccetulas (179) white grub larvae environments Several evolutionary avenues lead to fitness (134) Patangasuccincia adults (6 82) and Schistocerca (Table 2) but there is no reason to believe one set of gregariaeggs (82) attributes is better than another (224)

Insects known to enter hibernation are LodeIphaDstriatellus nymphs (124) Chtio Tvlpressalis larivae (125) DroughtSesaia inferens larvae (146) but not S calavisis(2) Well-drained rainfed upland soils are subject to drought ifIDiaraea saccharalis larvae (75) Scirpophaga- tcerlttas rains do not fall within 2-3 wk Prolonged drought followedlarae(5 lacchrli s lae (75)saSlOhagasshoperseggs (82)and felo pidas athias pOpae (232) by favorable rains stimulates armyworm and locust outshybreaks but mealybtigs root aphids and thrips become

Lcpmotorisa oratorisand L acuta adults aggregate on numerous onlyafter dry spells for two reasons First heavy trees sugarcane or other shady moist sites L paluwanmsis rain normally kills soft-bodied foliage-feeding thrips and confined to Sulu and Palawan Philippines lives on the mealybugs and soil-inhabiting root aphids and root mealyshygrass Brachiariamutica (Forsk) Stapf in the absence of bugs Drought eliminates this source of mortality Second rice The adults can live for 5 mo When aestivating their the rice plant responds to drought by breaking down rate of metabolism drops and they do not feed They will proteins into soluble N compounds which enter the phloem however puncture plant tissues of their aestivation host in and are taken up by these sap-feeding insects (245) Greater search of water This behavior has led to errors of the host nutrition therefore leads to greater survivorship and range of rice bugs (I) Aestivation quickly terminates with fecundity rain probably meaning aestivation is not deep Small rice area

Some stem borers aestivate as mature larvae in rice Except in the highly mechanized large landholdings of stubble or in straw stacked as livestock feed However Latin America upland ricefieldstypically are patchy within moriality increases proportionally with the duration of the a highly diverse flora The small fields result in higher ratios dry season even if the stubble remains undisturbed If the of perimeter to area than with the typically larger lowland stubble is plowed for a crop following rice few stem borers ricefields Upland ricefields themselves may be intercroppedsurvive Rainfall terminates dormancy but it normally with cereals legumes or root crops Mixed intercroppingtakes several weeks for moths to develop and emerge with a wide variety of species also occ in tribal slash-andshy

Last-instar white grub larvae tunnel 1-2 m deep in the soil burn agriculture (50 74) to construct pupal chambers in which they aestivate The Upland ricefields tend to be small because of the labor larvae are very sensitive to dryness so pupal chambers are needed to clear land in slash-and-burn areas and to till land sealed to coierve body moisture Aestivation terminates to minimize weeds Rice requires more tillage than does with the first heavy rains but as in stem borer 3-4 wk pass maize because rice competes less well with weeds Uplandbefore adults develop rice is more a subsistence than a cash crop and small areas

Insects may respond differently in each region to cold (I ha) can feed a family temperature or drought Biotypes or loal populations may Small fields favor some insect pests such as some ant evolve (56) Local populations of Scirpophagaincertulasor species that can forage from more permanent field borders Patangasuccincta may have become adapted to prevailing Trees provide food and mating sites for white grubs (134) as conditions well as aestivation sites for LetTocorisa spp (199) Field

iagiliti The ability to disperse combined with high border grasses plus tIhose within a ricefield provide alternate fecundity orshort life cycles enable some upland rice pests to food for all but the highly monophagous upland rice better exploit the temporarily favorable upland habitat colonizers and a habitat for natural enemies

0 z

Table 2 Life history patterns of upland rice insects or insect groups

Host range Longevity Dormancy Vagility FertilityMonophagy Oligophagy Polyphagy Low Medium High Yes No Low Dispersive Migratory Low Medium High(Oryza spp) (grasses sedges) (angiosperms) (lt3 wk) (gt2 mo) (lt1 kin) (lt50 eggs) (gt500 eggs)

Ants xTermites x x x

Mole crickets x x x xx x xField crickets x xx

White grubs x x x xx xRoot aphids x x xxMealybugs x

x x x xxSeedling maggots x

x x x x xLeaf beetles

x xx xArmyworms x x xx x

Thrips x x xx xChilo spp x x xx xMaliarpha x x x x

xLocusts x x Xx x xGall midges x x xxiVephotettL iirescens x x xx x x xLeprocorisaspp x

x xLeaftolders x x x

Sogatellafurcifera x

xx x x x x x x

00

IRPS No 123 January 1Q87 19

All this appeals to favor insect pest buildup but the notwithstanding the release of stored nutrients in response contrary is true Loevinsohn (138) showed that insect pest to drought that temporarily benefits philoem feeders populations respond exponentially with the proportion of land devoted to rice up to about 75 of the area then the M POR ANCE I I P1A 1) RWI P1St S

rate of response declines In most upland rico areas tie rice crop occupies less than 50i of the area- therefore the Visits to upland ricefields normally reveal 1kw insect pests potential for population buildup is low This relationship Loevinsohn (138) and Locvinsohn ct al (139) compared may not hold for seed pests that can locate small ricefields insect abundance using annual light trap atchcs in upland

Another outcome of the highly diverse flora and phical rainfed lowland and irrigated lowland sites in the Philipshyenvironment of upland rice isthateach upland ricearca has pines Although only one upland site was studied data a unique composition of insect pests showed equal or fewer insects at the upland site than at the

For example in Tanauan Batangas Philippines the rainfcd lowland sites and fewer than at the irrigated sites main upland rice pests are rice leaffolder(RLF) armvworm (Table 3) M Lltiple regression analysis showed that the key Aythntnaseparata WBPH ants Soli psis gentitataand factors explaining insect abundance among a set of cropping white grub Leucopholis irrorata Sixty kilometers away in intensity variables were the number of rice crops grown per Real Quezon the main pests are seedling maggot Athe- year followed by area devoted to rice Table 3 shows that rigona orae flea beetle (Thacitcmnra hasalis thrips the upland rice site had only I rice crop perym rand that rice Stenchaetothrips hiurnmis and rice bug Leptocorisa was planted in only 20Vi of the area Fertiliter and oratorius In a third Philippine upland rice site in Claveria insecticide use had little bea ring on the trend The armters at Misamis Oriental Mindanao the main pests are ants the upland site applied 60 kg N ha higher than at most Solenopsis gemtinata seedling maggot Alieriona orvzae irrigated sites white grubs Holotrichia in)iUaiuoan t and Lucopholiv Insect occurrence at the upland site was omparable to irrorata stem borers Sesamia ifiArelr (hio auriciliusC that at the Cagayan rainfed lowland site parlicularly in 1982stpessallis Scirpophaga incertulas S innoiata and when drought prevented many farmers fron planting(only

4c ona chrrsurapella root aphids Tetraneura niriah- 40(of the rice area eventually was planted that year) M ore dunrinalisand Rhupaosiphunm riahbdminalis lcaffolders favorable weather occurred in 1980 and 1981 which Marasniapataltis and Chaphalocrocis Medinalis army- supported greater rice areas and consequently more insects worm MIthinna separata and rice bug Leltucorisa The most intensive site for insect pests was South Cotabato oratoriusThe great diversity of insect problems is typical Mindanao where farmers plant 23-25 rice crops per year fer upland rice and makes control efforts more difficult with irrigation The Zaragoa Nutva Ecija site also had particularly regarding breeding for insect resistance high pest abundance and at this site the fields were highly

asynchronous Asynchrony was the third most important Low yield potential Iactor in explaining rice insect abundance Most upland rice is grown on low-nutrient and mineral- Rice cropping intensity in upland areas is bound to toxic soils which when combined with eiratic rainfall remain low Only one rainid upland crop is possible per make upland rice a highly riskycrop for the farmerto invest year (Farmers in Claveria Misamis Oriental Mindanao in costly soil amendments and land management Often Philippines tried asecond upland ricecrop i 1984a ycarof upland rice isgrown far from markets making inputs even favorable rains but no harvest was possible b-cause of more expensive Farmers growing a subsistence crop hosi- drought blast and birds) Rice area isaso bound to remain tate to invest in costly inputs to raise yields low because of the high labor and power rcquirelmenlts to

Even having varieties that could double existing yield prepare the land (because of weeds)and because upland rice potential under present management levels would probably wvill not compete with lowland rice as a cash crop Some not prompt many farmers to use purchased insecticide upland rice sites with prolonged rainy seasons howeverare unless tile risk of crop failure were reduced such as it was planted asynchronously such as the Clacra site with irrigation systems in lowland rice The seasonal abundance of Upland rice insect pests

The Tanauan Batangas site is atypical for upland rice attracted toa light trap wasgrapied for crop year 1980-81 in The soils and rainfall pattern are favorable and because of Tanauan FBatangas (Fig I 2) W13P11 was he most its nearness to Manila farmers have cash resources ftm common rice hopper and its population peaked at the sales of vegetables to purchase tertilier They applied an midgrowth stage as it does in lowland rice D-Vac suction average of 60 kg Nha after particle initiation and obtained samples showed an earlier peak than that from a light trap 25-35 tha yields The high fertility isone reason why RLF indicating emigration W13PI usually emigrate after the armyworms and WBPH developed into large numbers vegetative stage Populations in 1980 reached levels that Farmers however used no insecticide on upland rice caused patches of slight yellowing in some fields WBPH although they did on vegetables probably cause hopperburn in Batangas They immigrate

The poor growing conditions fOr rice generally mean year-round as shown by low levels in the light trap BPH poor growth and low fecundity for insect pests fIeding on it however were scarce in the light trap and not collected with

Table 3 Abundance of rice insect pests and factors in rice cropping intensity of Philippine upland rainfed lowland and irrigated lowland locations where light traps were operated daily for atleast I yr -I

0Rainfed Rainfed lowland Irrigated lowlandupland

Pangasinan Cagayan lloilo Santa Maria Nueva Ecija South Cotabato Tanauan Caaringayan Caaringayan Bangag Bangag Bangag Buray

LagunaCale

Batangas Manaoag 1980-81 Santa Motiica 1982-83 Cabanatuan Zaragoza Avancena NaniamanSolana 1981-82 1982-83 Oton

1980-81 1979-80 Oton 1981-82 1981-82 Koronadal Koronadal1980-81 1979-80 1979-80 1983 1983

Insect (kerosene light trapNephotcttixcatch nolyr)virescens 650 1240 500 1800 330 210 3400 1600 160N nigropictus 210 - 360 18400 5500 2900- 140 90 20 - shy 1700 -Reciliadorsalis 710 - 3200 1700180 890 630 140 170 2400 800Nilaparratalugens 130 690 - - 4200 4600300 520 750 180 60 3300 1600 740 310Sogatellafiircifera 1700 31000 13000 8200310 210 3300 610 270 980 1600 6300 -Cyrrorhinus - 12000 6700600 540 3200 290 160 330 120 410Scirpophaga spp 50 410 620 - - 21000 12000150 180 220 270 30 1000Cnaphalocrocismedinalis 70 70 540 610 270 11000 6100200 140 100 6 60 250Chilo suppressalis 0 60 60 1900 300 200- - 30 20 0 6 3 160 -C auricilius - 0 00 - - 20 3 60 20 3 0Sesania inferens 0 - 80 60 3 80 10 51 -Cropping intensity _ -

Rice crops (noyr) 10 10 12 10 10 10 13Rice area (7c) 17 19 20 20 25 2320 85 85 60 70 40 85 85 90Fertilizer (kg Nha) 60 20 85 80 70 7020 0 0 0 30 30Insecticide (no of 40 60 40 300 04 04 0 03 300 13 13 30 50 40 33 33applicationscrop)

__________

IRPS No 123 January 1987 21

Light trapped Insects Drytnd __ 2(notrap-nght) Suctioned Insects (nom )

50 -)o 9 jO40- SOW ffurciWO Light trp

D-Vc suction

20 to 4

20

Egg porasitizahion (M) 75 60

300 SOOampReOftwciffr

45 30shy

15

-insects (notrop-night) 4 Nileporvata Iuen5

Egg parastization()

Light trapped insects (no trop-nght) Suchoned insects (no m 2 )

6 8 Cytorh~nuls hvidipennii 0

4 05 2

50 0 Insects (noltrop-night)40-ui Ipanh pe g

Moohase 1 Seasonal abundance measured by light trap

Rainfall(etod) and D-Vac suction sampler of whitebacked

to ~ hopper Nilaparvata lugens and lea ffolder60)-t Cnaphalocrocis mnedinalis andplthpe g20 L I II I I I parasites and mtrid predator Ortorhjntus livi-Apr May Jun Jul Aug Sep Oct Nov Dec JaIn Feb Mar dipennis set against cropping pattern daily1 9 80o 19 8 1rainfall and moon phase Tanauan Batangas

1980-81

the D- Vac Cyrtorhinus build up on the crop coterminous are collected in light traps during full moon (109)with WBPH buildup Cyrtorhinus disperses during the Because upland rice shares most of the lowland rice pestsrainy season RLF not readily collected in light traps was and those pest species are generally less impotant to uplandprevalent only during the rice crop rice than lowland rice there is a general belief that insect

Nephotettix virescens was more abundant than Nnigro- pests are less important onpicLus in the light trap collections and on the crop GLH upland than lowland rice Arcview of limited data on upland rice shows an averag of population peaked at the reproductive stage as did zigzag 10-21 yield loss from insect pests on a par with that ofleafhopper (ZLH) Crtorhinus tracked the populations of lowland uee determined by the same methodology (132) planthoppers and leafhoppers Both GLH species were (Table 4) McGuire and Crandall (144) made a similarcollected year-round in the light trap indicating immigra- estimate for Central America Cramer (51) howevertion from nearby irrigated rice areas Possibly because of the estimated 35 yieldlow populations loss in South America based on ainsect number did not correlate with similar pest complex - stem borers and seed bugs shymoon phase although reports indicate that more hoppers attacking irrigated rice in North America This estimate did

22 IRPS No 123 January 1987

Light trapped insects D~~nd (no trap-night )

St plht Suctined insects (nom 2

)

D-Voc suction 1520

1o

0 Lih trap 5 ~105 o5

00 Eggparositizotion()5040 NephOtgtha virneens

30 20

I010 A-Insects (notrop -night)

4 Nephatelh nigropccent i

2 o I vV^J - -- ^ VA____ f _

Light trapped insects (no trap-night) Suctioned insects (no rnE

10 -1208 ReCdO doRsol

6 15

4 10 052 iwxALAiamp 0

Light trapped insects (notrap-night) 2Suc oned nsects (nom )

8 CyrtorhmnusI6o nn]10 i0075

4 05

2 0 0

025

2 Seasonal abundance measured by light trap Roinfall (mmd) Moon phase and D-Vac suction sampler of green leafhoppersNephotettix virescens and N ngropicrus and I zigzag leafhopper Recilia dorsalis their egg 60

parasites and micid predator Cyrtorhinus livi-dipennis set against cropping pattern dailyrainfall and moon phase Tanauan Batangas1980-81

40

5 Lf Apr May Jun

1980 Jul Aug Sp Oct Nov I

-aMar

1981

not account for soil pests and was not based on field data in the absence of pests Carbofuran granules are phytotonicAkinsola (I1) cited mainly data from irrigated rice areas in to rice (239) The prevailing hypothesis is that carbofuranAfrica but estimated yield losses of upland rice between 15 prevents soil bacteria from consuming fertilizer The phytoshyand 30 from insects However losses in Thailand over a tonic effect is through greater availability of fertilizer to the4-yr period ranged from I to 13 averaging 5(106) plant Denitrifying bacteria are known to be more active inYield loss data however should be interpreted with aerobic than anaerobic soils therefore it may not be merecaution (184) The Philippine data were all derived from coincidence that the highest recorded yield losses (22-69)trials on farmers fields under farmers normal agronomic occurred in those trials with 12-30 kg basal Nha On themanagement practices except for Bukidnon and Capiz other hand contro with carbofuran granules may not be aswhere researchers used fertilizer Batangas farmers are good as a seed treatment against seed pests and perhapsatypical because ofhigh levels of fertilizer used The trials in yields would have been higher if a systemic seed treatmentBrazil were carried out at an experiment station but insecticide had been used conditions were similar to those of local farmers The 1979 and 1980 trials in Batangas had two full-

Carbofuran granules were used in every trial as a broad- protection treatments one using carbofuran granules andspectrum insecticide applied basally with the seed An the other using bendiocarb WP as a seed treatment Plotsobjection to using insecticides to measure yield losses isthat protected with bendiocarb yielded higher than those proshyinsecticidescandirectlyorindirectlystimulateplantgrowth tected with carbofuran granules The other trials in the

Table 4 Yield losses to insect pests and yield responses to insecticide in upland rice in the Philippines Brazil and Upper Volta

Yield (tha) Yield loss (5) Location Year Cultivar Protected

with Protected

without Check Total Vegetative Reproductive Ripening

Basal N (kgha) Reference

carbofuran carbofuran Protected No with carbofuran

carbofuran

Philippines Pili Camarines Sur Tanauan Batangas

Pangantucan Bukidnon

Dumarao t-apiz

BrazilGoias Minas Ge~ais

Upper Volta6

1975 1974 1974 1975 1975 1976 1977 1978

1979

1980

1979 1980 1979

1977 1978 1979

Bursiging puti Kinanda Dagge Kinanda Dagge Dagge Dagge Dagge UPL Ri5 Dagge UPL Ri5 Dagge UPL RiS UPL Ri5 UPL Ri5 UPL RiS

IAC47 IAC47 IAC47

14 33 11 36 35 31 30 27 37 26 40 30 40 42 46 35

19 21 09

--

-34 24 24 27 --

29 43 31 42 -

47 25

---

10 28 05 27 27 30 26 29 33 28 33 29 43 23 36 11

15 15 06

29 15 55 25 23

3 13 -7 11

-7 18

3 -7 45 22 69

24 28 35

--

21 -11 -20

4 -

-3

23 7

-2 -23 56

-

-

-__0

6 --

-

-23 14

5 0 8 2

22

-

--

0

-

--18 11

3 -10

4 10

7

--7 -

14 -

-1 6

10 -5

8 9 7

-

--

0 0 0 0 0 0 0 0 0 0 0

30 30 30

12 12 12

99 99

100 100 101 101 102 102 102 102 103 103 104 104 104

66 66 66

Farako-Ba

Mean

1977 1978

IRAT10 IRATIO

38 66

--

40 43

-5 35

21

-

10

--

--

--

- 242 243

Z

tJ

-w

24 IRPS No 123 January 1987

Philippines which had two treatments offering protection In the 9 Philippine trials where yield losses were measuredwith and without carbofuran granules relied on further by growth stage 10 of the yield loss (21) occurred at thecontrol with foliar spray and not on seed dressings The vegetative stage and 517 occurred at each of the reproshycarbofuran-protected plots had an 18yield los compared ductive and ripening stages Ibis result is similar to thewith only 10 with foliar sprayS This yield difference is pattern of yield loss in lowland rice measured by successiveprobably due to differences in protection against sown-seed treatments where the crop is unprotected at each growthpests (foliar spray do not control sown-seed pests) rather stage but proticted during the other growth stagesthan phytotonic effects became basal fertilizer wa used in However the insect complexes r sponsible for vegetativeonly 2 of the 10 comparisons stage yield losses in upland rice are entirely diftfrent than in

The recorded yield losses were not related to yield lowland rice The key vegetative stage pests in lowland ricepotential (Fig 3) One might expect higheryielding crops to arc aquatic whereas those in upland rice are sown--seedhave higher percentage of yield losses but this ws not tile (sown in soil) root and seedling pests This dift rence maycase One reason is that insect damage exacerbates plant explain why agricul torists have overlooked tile ilmportanccinjury from drought stress low yielding fields from d ought of upland rice insect pests Many reports of upland rice pestsstress would show disproportionate differences between in Asia ha Ve focused on major lowland species (46 108)insect-protected and -unprotected treatments Shoot and It isalso surprising that tei vegctative stage of upland rice root pests would be more responsible for losses from would record the highest yield losses becatuseseeral studiesdrought stress than sown-scd pests A second reason have shown that upland rice can readily recover from foliageconcerns seed pests which would have a greater impact on reinoval during that stage (117 152 165 188) Upland ricelow tillering low yielding varieties such as )aggc Low poscsses a hitgh obilitV to compensate for ca rly loss oftillering varieties nnot fill in the space created by removal foliage and will even be stinulated by foliage removal toof seeds oweer 1) R i 5 is high yielding because it tillersP1 produce a higher viel than plants ithout any foliageactively which can close the canopy to provide greater weed removed (63 192 231) [ie reason for this apparentcontrol discrepancy hovce is that yield loss at the vegetative

stage does not octUr frol leat arca loss but fron pestsYield loss () removing sown seed and feeding on devcloping tillers and roots -- damage that cannot be readily conpLtnsatCd for

70 - Measuring yield losses in upland rice cnvironrnents is further complicated by a report from Brazil that greater yield losses occurred in response to better insect control60 - =-09 as Insecticide seed treatments resuled in denser plant stands

(n=21) which in turn created a selectively more favorable environshyment for blast (69) This resuIt was parlticurla rlxexacerbated50 when cirbofuran granules were used in seed furrows Aside

ffrom a greater plant stand ca rbofuran iay have auigmented the N lecl and stimulated blast If so ca rbofuran should be40 - replaced with another insecticide or the seeding or fertilizerI rates decreased

Other methods to determine the impact of upland rice30 shy pests wvcrc to interview farners or to develop single speciesbull bull correlations of population levels and yield loss Feneira (66)2 NIM)st (75(1) Golas farmers reporteddid both in Brazil

20- insect losses in ufpland rice and 251 said those lossesS avcragcd 37(1 [his result is consistent with tie yield losses 0 0 measured by field trials in the same statc(Table4) For every

I0 4 of seedlings removC d by leaf-cutting ants a I 7yield loss is predicted Similarly Ferreira (66) found that ai

0 - average of 53 thrips per panicle before panicle initiation meant 3 times more unfillh d grains

-IO0 I 1 1 I I-101 0 2 3 4 5 6 7

Yield (tho) C)N [ROt IM FI IODS 3 Correlation between upland rice yield and yield lols caused by Upland rice insect pests may be controlled by culturalinsects measured in 21 trials in the Philippines Ivory Coast andBrazil genetic biological and chemical methods

IRPS No 123 January 1987 25

Cultural control dry season plowing will have little effect on them Plowing Crop husbandry to reduce upland insect pest populations when the soil is wet from the first monsoon rains will expose can be classified into practices effective in a single field and soil insects that tunnel close to the soil surface to predatory those effective only at the community level Cultural birds chickens dogs and even man (field crickets nole controls should come first in a pest management program crickets and white grubs a re delicacies in man tra ditional They have broad stable effects as pests have little pos- diets) sibility of overcoming them through selection of biotyps The greatest degree i1 inIscct control fron tillage comes (138) On the other hand some cultu ral controls work only from plowing soon after rice ha rvest when most soil insects with high labor or power inputs and connunity-wilt lie close to the soil surface anrd stern borers are in the rice methods require coordination of iany farniters stuble Whitegrubsare mature and tillage exposesthem to

Pantin time The luxury of being able to shift the time of predators Tirnelv control will prtec the crop following pla ntring upland rice is open onl hr la rniers in areas with a rice and lowr the populationIfor succeedinig years Plowingprolonged rainifall pa ttern lI copping intensity or underthe ice stulble helps it decompose killing sten borer rnechanied land preparatiot Gcnerally planlting as soon larvae inside as the rFiinv season begins will lower populatiois of 1110st [requent tillage destroys ait nests bit not termites insect pests Insect populationsarelo aftcra dlvseoll or Tillage and clean culture howes er rcliove food for winter fallow delayed plaiting lets them build tip on termites which then mtay attack a young rice crop Scarab alIternate hosts that gi vigorotsly with the first seasonial beetles also prefer to miposit in eceitly tilled fields Noshyratiris This first flush iof weds cros lItxuttriartlv with iceCult is dibbling faors the survival oftillace Clre Such rineralied N released during the dr season aid made stils i aels tnd insects thtt pttpate or lay eggs- in the soil

available to the plants with the rains It iliso igrtmitcally Zero tilla hio eerl conserves predators ofthe GIH (37) advanttgeous for the rice ciop to tap this trlLItlaI fertility nt it mcttoi lo lessen the impact o0 soil insects and grow vigorotsly to compete with wrus and to tolerate brotdcasting seed or planting in ftltrrows is prefc ired over pest infestation dihlirig in hills Crickets are large enotlgh to eat seedlings in

Larly plantngs terd to escape seedling maggolts Upland a hill atrd tillering normnally ca iriot fill in tlhe missing space rice in Bitangas is planted in Ma and ine itli the Iirst Oftcn dibbll rd is not well covered and iseasily found by rains Fairners are highly mtotiated to plant early tocscape foraging insets ioIdents and birds drlitght and typlotns ricar harct Arn Atrgtst planting Plan uit sii Incrcatsini sced density protects tile cropwould be evcely attacked by It riim orivzo At tti againrst seed and seelling pests especially ants particularly other Philippiie tipland sites Rea l Qteot antd (laelit if the crop will germinate and ermerge quickly M isainis Oriental seedlig mnaggots arnrige ric boe t se hr trftinll itI India rice intcrclopped witi cotton or farnes have a longer growing season and plnit 2 rin or pigeon pci had lower 6IIt and WBIlII poptrlations than mote after the ortset of the rainy season liaaiangas ftrmners rice alone (211) NfailC arid Urplald ir_ 2re coilltori in the plant ca rly becia use ofa shorter raiinfall period arnd the desi re Philippines he chatngC in pest status folii iniercropping is to lliarvet before the typhoor sea son peaks ltighly locatioll-specific (133) cmianarid the net elfc be either

Early plantings cornbind with early-iatiuring varictics nil bcnelficial ortdetriential (189) For example BIatangias provide higher crp tolerartce for white grubs Aestiatior farmers intercrip Itro in their ipland ricefields A liornshyterminates with the first rains but 5-6 moare ineeded for the nrii iotd-Ahis i (I iiiacius) which norlially feeds resting larvae to develop into beetles emerge and lay eggs on taro ill feed or miarbv rice plants Intercropping and for larvae to reach the danriigii g ird-instar stage probably would not afft0 seed pests A crluanion crop (134 In lataigas this normually iseiough tiite for harvest planted ith rice and haing gleater tolerarnce for root

It Brazil howecver delayed planting is a suggested dariage or being toxic nayact isi trapcrop to root-feeditig cultural control for the lesser cornstalk borer and insect pests Acial-fei i ig pests voultId be most affected by pests favored l drought (66 195) Cropping internsity in intrer ropping Shoot aphids leaf beetles sten borers and Brail is low and delayed plraing places the rice crop in the hopplers would be likely ca ntidates to note effcts On the niost stable rainfall period reducing likelihood old rought othlt halnd the corlipalior crop stch~ sitniaie Iiiv be more

In rilniy arca s rinmirsare roi atl to plant other crops benehited than rice from initecropping ( 137) before rice and althNgh it woultl be desirabh to avoid 11echi Weeuling luring the first month after clop infestations by planting rice cuatl lariners lack the labor cstablishent will force pcts such is anyvworris which and time to do so ol a low priority crp prefer grassy cetds ontor tle vorling rice Soil iampsecs will

77age Uplild rice Stils ae light textured and Call be mvnye to the rice crop Cleani citrltirc also lay lore territes plowed duiring the d rv seasor notl only to grin time for to attack I u cetincti inrice ceds i ay he le optinal planting with lie fitst Inilsoon riiains but also to desiccate solition 13) soil insects arid weeds such is ntsedge that foim rhizomes Irtlizr Itneatsing N Iciiliier oin hlwland rice has Htowever becta use most soil insects lie below die plow layer fAtwvored Irigher populains of pliithoppeiv Imlhoppers

26 IRPS No 123 January 1987

and leaffolde rs However because of the need to temper fertilizer use on upland rice o prevent blast fIrtilizer rates will be lower than is comonia r for lowlad rice causing relatixely less hopper and IaIfTfoide r population buildup However inupland rice areas without a history of blast such Is Batangas farmers apply high rates of N and conscqtrently the crop has high infcstation levels of WBPI]and RIF The fhrmrcrs leec insect buildup by splitting the fertili11r in0to two to three applicatiois bit do not applV fertilizer at the vegetati estage because this prodrces a tall crop which illeasily lodge during typhoons Greater crop fertility leads to greater insect survival (better nutrition) larger insects (to cat no re foliage) arid greater fec titd t (to lav nmre eggs) (128 145 171) Not all insect species respondequally hovr Increasing rates (if N l anrid K increased Dialraa s-acchralis and (iaeiurema flea beetle but reduced thrips (66) Zn rcdLuced deadheart density from the lesser cornIstalk borer

Fnldingi Although most Upland rice areas do nIot hawxcaccess to wa ter floodiig the fields is rRrrnriierdcd in areas of Brazil or root bug and white gib control ( 195)

Crop rotatioti Tillage before planting a crop after rc harvest will control stern boreis and other pests renain n the stubble arid will unearth soil pests Planting a non-gratriiaceous crop after rice is recommncrded for termite control (66) Crop rotation is less nttdcd in upland rice culture than in lowlands because atcrop-free period is assured even after harvest of crops following rice

t1ulchilt iryl plastic irli irig designed for weedcoitrol ii upland rice inJapan resulted in ntr highanri liy infestation of (ihilo sulytresvais (93 9-1 95) NIulched Upland rice grew more luxuriantlv and attracted more (Avipositing mothis than u lched rice Also the rice plant and stein borers imatoured 10rC quickly and a partialgeneratiori af vouing ista iNoverwintcred in the stubbleLotili--malitrt varieti Quickly maturing crops reduce the tmbCr of pCst generations that Calt build up Rapid crop establishment olan ca rl v-natu ring vaitvty if)pa 11icLa r will lessen white gib and stem borer damiage White grub larvae intle last iistar are highly destIruCtive so aii early-iiatri ng variety will escape serious root loss Stem borers build 1ip slowlyx in rice to ecormic abundant in a late-Maturing crop anrd caiuse whi tlieads Numibers of somespecies increase exponeitially with each succeeding genera-ti so a xarietyexhat atures I trowh earier thian another will hac less insect damage Early-naturing vaiicties however will not lessen seed pest daiage either ii the soil or oina standing crop

Strwhoi hAIJttam Inrscts tcd irig on thlie aerial portions of rice plats disperse from carly to late plantedfields AS iiiibCirs increase exponentially with eachinsect

gtscnrattm e eiglbrig f rmc~s1(uil upp wh rc generation pests build tIP where neighboring fihrTC e9r1sstagger their plantings II Batangas farmers plant uplandrice within I iro use 120-d s-arieties and plant at the start of the rainy season Ini Misarnis Oriental laricrs stagger their

plantings up to 3 mo apart and use a set of atieties that mature in 4-7 mo Insect pest danage particularly from stem borers is greater in Misarmis Oriental Also seedling maggots damage rice there because rainfall favors frequent planting of maic In Batangas rice escapes seedling maggots bCca usC of distinct drs season and synchron0us early planting with the onset of ains

Therefore combinations of cultural practices ea rlyplanting synchronous planing crop rotation and earlyshymaturing vaneties protect the rice crop against most insect pests

Ilmant resistance Beca tse upland rice has few specialized insect pests and many insect types attack the crop worldwide regional breeding and strong internatiornal cooperation are required

Sown-sced pests arc economically controlled by insectishycide seed treatment and should the relore receive lowxpriori ty inbreeding objectives

Root pests are normally cost ly to control with insecticides but finding resista t sonres mayvbe di fficult Tanaka (229) for exa -ple fi led to find varieties resistant to root aphids Selecting for large root bioniass perhaps should be the strategy for pests sLuch as Whlite grubs that renliove roots Fainter(174) howcvcr cites varietiesof crops tolerant ofor resistant t(i root pests Some sorghun varieties are high in cyanide so rice sorghun might incorporate broadshyspcctruri chemical resistance

Seedling vigor and drought tolerance are high prioritybreeding objectives so tupland rices should have large r root systernsand be able to tolerate ligherlevels of root loss Sonie root loss inolder plants is beneficial if it stimulates new root developnrut and therefore crihances intake of soil nutrients (53) The rice seedling inaggot rtiLotia spp may be

controlled thirough resistant aittics A hi rgc breeding effort on sorghniu has had fair success inideveloping arreties esrstait to Asoccata Rodani (251) A field trial comparing 10 upland ricc varieties showed differences aniong varieties va ring from II to 4(- damaged tilleis (215) Hovver resistarce to 1Iill-0r gtia species may rot cross ox-rti otier

Shiraki (210) reported that upland i-ice stens wcrc harder than lows-land rice stemsperhaps becaIusc (f a higher silicoricontent and thus were more resistant to yellow stern borer hc tortia Iow tillering (f uplaid iice xarneties prodtres a

higher percentage of infested tillers than would occor in higher tillering lowland iice under a siiilar egg density The corollary therefore is to select for high tilleriig upland rices

Stemii bor amcstheh main targes dobrecders inAfrica (571 204 )but nticl oifth e sc reen irig is dotie unde r lo wland co d t osrm s wih A h r~ o l pe i s n acorditions (11 96 218) As with lthrinrnwspecies mayhave to be dealt with independently however initial results show cross resistance between African and Asian species(92)

In Brazil screening Ior insect resistance focttses oil lou I pest problems Varieties tolerant of lasmolwalpus ligi-sellus and resistant to )iatraea saccharalis have been identified (70 153) Current work also emphasies 7iraca limativetntris and Thmialspis (=Iosis) flavouicta (66) These insect pests Ilt onlV citise IIigh yield losses tihey also ire difficult to control chemically ofr by other ilealls

ManV upland rice vaieties aic early maturing and are resistant to pests such as gall midge (I Xl ) Early mIturity is often a highly desirable trait to esuilpe pest buildup but it is not true resista nce bca usc ilthose va reties are plinted late they will be itsdamaged as ain early planted susc ible variety

P~erhaps tile cosiiiopolitall of all upland rice pestsimost are the sced pests (ripenling stagc) which pose a particular problem beta use scuds re ised IsIO for hitnit4ans Ayltoxic substatice introd tcld geticticllyilmay also be toxic to nal iniless it caln he placed old in thfe IdhuffSelectmi fo hairiness or lotng wns kill mIlct sthilrohleiiis dutilig threshing pa rtictfarl maitial threshinfl [hireshers ill complain of itcfimin atid itteascd (fust levels

Attempts itIRRI to fitid tcistatncc to the rice bug plorisa have ilotbeCnl prmitisilg lIIetIattonlids iorc

through the lenilna or palca hillhairs sedstVIor seeds wilh awns that deter bugs promote complaints from threshers Seed bugs liptocorisa and Stcnocoris enter the seed through the opening between tile lemna and palC but varieties with narrow openings are diifficult to mill

Breeding priorities should concentrate ott inotCI gruLtLIps that are most difficult to c nttrol by other meains Steil) horers appear to be the most widespread group that sigiiificantly dalag s rice aind is hard to manage by other Ilnca us

Biological control Very fe stutdic of mitatt~l eneties of rice pests in upland envirollnletits wee in tile wefouiid literature Therefore have not compiled a list of species Ve present a list of natural enemies iOtid in upland ricefields tuic latialln Batangas for some conitno iOldiar pest species ([itble 5)

Muich ofl thc lollowinle itfor ittiol cotlliCs rol Stutdies in the lowlatnds or roi crops otithcr thlin tice

Qu(ailtr o 11ll t11t( Nat trrtf lii hcoeits ight Ile used agaitist ile social insects tilt and termites Sollie predator specice live ill theirf nests n ilnic theirnllt ip-pearanIlCe aMRidbehavior IJtrouMdCing a mote aggressise ant species to displace pest ant codonies is a possibility (169) Patihogens as potential natural encnies have reccived little atteition (I13) Natural ncnuics asa group however hac not been toIl iporta it in Iegulating aintshown be and ternite numbers

The nocturnal behavior aid subteitmien habitat of lole crickets and field crickets protect them from many ntllral enemies There are only limited records of natural cnelics for these groups of pests some pathogens (87) but mainly

IRPS No 123 JattuIny I 271987

parasites - scelionids on eggsand sphecids on ilymphs and adults Sphecid wasps Larra carutmaria (SImitI hI L huzwuensis Rohwer and lsaltiuhta Willi ams speciali in mole crickets pa ralyzing them nd dragging the ninto thcir nests as food lot their voting ()tier sphCcid species Lins aurulenoa (Iabricius) lot maiac (Ashmead ) 1 lhshylssellaus (Smith) and Ml lal)oriosu (Snithf specialie in field clickets

White grUbsO the other hand hae Icti scriuns enuigh to have been extensively studied for cotttroi 1W nati al enemies( 112 140 141) Scoliid parasites C(ajsoMcispp have beetn intrtduced itto tilePhilippites for control of Ielxl ]udisiro la The rcsearchcts w Intttrdced themle during ill era claimed that the white gttionthreak as controlled (142) howevcr it isnormal for pest epidemics to subside and there was no diect evidence to indictte that tfe parasite was responsible A tmore promisitg titctllod might he to identify iso ltcttass prt~f tcartd apply hiactelaln fingal pathogens to tile soil Ihe lalattese beetle in North America has hcci coitrolled hv tite conmnleciall aailable hactcrial prcpa ration itacihlpu pav ofr milksdiscas (71) Spote prtyefott the soil itloted ilrtier reitiain actic fr Veas

Root aphid sardtnctly bugs readilslic tilib to predatots and parlsites once their tetndine ants are controlled The larvac and adtilts of a coccinellid beetle 1Scynms sp prey on llriadlouais in slash and burn upland rice Baits treated with insecticide may be used to control tending ants by killing their ourig (244)

Root-tkcditig blugs false wireoris wireorms and root wccils aic r ofsuch little ifilpoltatice that thenomttll role of liatural Cticniics hias tiot beei asscssed A possibility hoevrl tiglt he tit ItaIIitic niaCIMtodcs lintt1g11en Itese soil-itlhlfbitine mCtitodcs W5uhtfhave to be imiass Produiced

large miltibcr of niatltrll cnemics ircknown ft CltIoriS IC IpblCeii is thtt clttvortnlls ofonlie the crop soott alter land ptel)tritiOti a1tithe bcgit iniig of the raittv

Citsoln hfien tiatllral leniiiy populations aric lo Colltloil WOtuld iii ole mass prodtticing andirclcaslng key spccies

le egg stage is the fpirt (olthe Cedlitgt Ialot life cycle tI hich to f)ctis ftor tittlril elctit cottntrol hplraites o1 pleditors ttsit elleggs highly expoIsYCl I Sgg pailsites or predators stolld probahly e ittass andtbe produced released Littlh i toruialioll is a ailable ott what species to trV o ho to Iniss podLice thiCIi A etiloplhid wasp Icmihlaol sp effctiicly regushy

lates the leatf lcloalapolm2a spioanoiiitr heat andc rice antid should be collsctvcd

leaf beetles whCllose e am] eggs ate laid in the grutindhirO ould be motC difficutlt to control by niatulral enelV Mnll ipuhltiOn But species sUici IS c_gsthispid beetles ith

and larVle oilleascs ate vtilcratble to iparaites and pledators

Natural enemisc of tlirips otl rice are little known

28 IRPS No 123 January 1987

Table S Parasites recorded from upland rice insect pests in Tanauan Batangas Philippines 1977-84

Order Family Species Host stage

I lymttenoptera Braconidae Leaffolders Cnaphalocrocis Marasinia spy

Cardiochilesphilipplinensis Ashmead a Larva

Cotesia angustillasis(Gahan)Cotesia or cypris (Nixon)

Larva Larva

Cotesia or taeniaticornis(Nixon) Larva Cilonusinotakatae Munakata Chelonits SppAlacrocetrts philippinensis (Ashtmead)

Egg to larva Egg to larva Larva

Iclineumonidac Orgilts spp Isclutojppa hireator(Fabricius)

Larva Larva

Itoplectis naraigav(Ashmcad) Larva Tenelhtcha philippiensis (Ashmead) Larva Tenelchha stangli (Ashmead) a

irichomma cnaphalocrosis Uchida Larva Larva

Bethylidac Elasmidac

Goniozms or trianguli r Kieffer lasmtusalhopictus Crawford

Larva Larva

Diptera

Encyrtidac Chalcididac

Tachinidac

Elasmtus spp Copidosomopsisnacohiae (Fidy) Brach nteria evcarinata Gahan Biachyieria lasus (Walker) Ztgohothria ciliata (Wulp)

Larva Egg to larva Pupa Pupa Larva

Argyrophylx nigrotihialis(Blaranov) Larva

l-ymenoptera Braconidac Yellow stern borer Scirpopiaga incertulas

Braco chinensisSzeplegeti Larva CIthelotis spp Chelonus niunakatae Munakata Cotesi flaripes Cameron Stenoracon nicerilli (Bingham) a

Larva Egg to larva Larva Larva

Ichneumonidac Tropobracon schoenobii (Viereck) Amatroinorpiaaccepta metathoracica(Ashmiead)Frihorus sinicus (I lolmgren)

Larva Larva Larva

Ischnojoppaluteator (Fabricius) Larva Isotima nr dammeratiti Rohwer Larva Tenelucha philippinensis (Ashmead) a

Larva Tenehicha stangli (Ashmcad)Trichomma enaphalocrosis Uchida

Larva Lara

Chalcididac Eulophidac Trichogranmmatidae

Xanthopimnpla stemmator (Thunberg) Charops hrachYpterumn Gupta and Malteswary Brachtm eria sppTetrasticttus schoenohii Ferriere Trichorigranm chihunis Ishi

Larva to pupa Larva Pupa Egg Egg

Diptera

Scelionidac Pteromalidae Tachinidae

Tt ichograintnta aponiicuitt (Ashniead) Telenomits rowani (Gahan) a

Trichoinalopsis apattelctena (Crawford) Jeirhaea spp

Egg Fgg Egg pupa Larva

Z4gobothria ciliata (Wulp) Larva

Ilymenoptera lecneumonirlac Gol fringed borer (hilo auricilius

Trichomm enoptalocrosisUchida Larva

Ieioenoptera Braconidae

Aantholimlh steiniator (Thtinberg) a

Pink stet Ioret Sesamia inferens StoInacoil nicrilhi (Bingh am)

Loarva to pupa

larva

Diptera Ilyinenoptera

Armvtnyorms Mythimna separata Spodoptera mauritia acronyetoidesTachinidae Zvgohothriaatropiora( Robinea i -Desvoid y)Eulophidae It-phctrtiscitapadoe (Ashimead)

Larva Larva

(halcididae Braconidac

Brachpin ria spip (oesia spp

Larva larva

IHyntenoptera I)iptera

Rice skipper Ielop ida (halcididae Tachinidae

math ias aidgreen iortedci ciliarNIelan itis eda Brat lhincria sp iii matuqitatt Cameron Irgyrolhirlaxi trotlhiais(marumtv)a1

ismtne Larva Larva

Diptera Ilymenoptera

Tachnidae lelineutionidae

fuown se tilo per Mois frugalisIrgtrophrlax 1igrothiahiv 3 a rat ov) anthopimpa pmnctata (Iabricius)

Larva Larva to pupa

aMost dominant

IRPS No 123 January 1987 29

Normally predatory thrips prey on herbivorous thrips strepsiptera on both Spiders dominate tilenymphaladultNatural enemies of stem bugs --Scolinoplara and Tibraa predators Spittle bugs move to rice from pasture grasses - are in order of importance egg par~isites and predators and biocontrol would be successful only ifcarried out before fungal pathogens and nymphal adult parasites Egg tiley immigrate into ricefields parasites could either be introduced fron other areas or Biological control tactics for hemipterous seed bugs mass produced and released ould follow those outlined for stein bugs

Armyworms are normally lwld in hcck by the activities Coml)arison of environnewtns Natural enemies are of egg and larval parasites When these natural enemies fail perhaps faced with even greaterconstraints than insect pestsusually because of drought armyworns become epidemic to survive ind reproduce in upland environments TheyIt may be worthwhile therefore to release parasites during have adapted ani survived by having wide host ranges the the rainy months following a drought Virus diseases of ability to aestivate or hibernate over unfavorable seasonslarvae would beanotheravnue to exploreforarnyworms and the ability to disperse or by other mechanisms

Parasites have ben traditionallyconsidered for biolegical Comparing the most dominant and therefore the most control attempts against stein borers usually by rekasing adapted species of natural enemies collected in various exotic species( 166) This approach has not met with success environments in the Philippines we can understand how and new avenues should be explored The role of predators different rice environments are arid pathogens is little inderstood and deserves greater Anagqrms olptahiisa mymarid planthopper egg parasiteattention Egg parasitism rates are normally reasonably is cquallyadapted to all Philippine rice environments (Tablehigh but should be supplemented with effective egg 6) It is selective for planthoppers and even parasitizes eggspredators Orthopterans are potential egg predators of the n-iize planthopper Peregrinitsmaidis itt upland areas Metioche vittaticollis (Sta ) Anavipha nifiwnnis (Saus- Alavohis also parasitizes maize planthopper eggs but is sure) and Conocephahs hongipnni (de i-laan) feed on most prevalent on BPH in the irrigated wetlands (onashystem borer eggs etioche and Anaxil)ha specialize in eggs tocerus spp like A optalhilis on BPH and WBPI Iattacks with no hair such as Chilo spp The omnivorous (ono- the eggs of GLH in ell rie environments It is also found cephahsspecializes in eggs cov red with hairs it is a proven parasitizing the eggs ef the white rice licahopper (oimna egg predator of Scirlouphaga spp but has the discouraging spetra Oirosita naias a trichogrammatid egg parasitehabit of eating rice guin (65) attacks BIPH and WBPH in the wetlands and has no other

Grasshoppers have egg (scelionid) and nymphalad ult host A related species 0 aesopi specializes in G1-H in (nemestrinid tachinid and sarcophagid) v irasites Among wetland environments Dryinid strepsipt-an and pipunshypathogens protozoans have been recorded most frequently culid nymphal and adult parasites of planthoppers and Control of locust species by natural enemies could con- leafhoppers also specialize either in hopper species or centrate on their habitual breeding areas environments Among the dryinids Pseuhdgonatoipsflavi-

Leaffolders have rich complexes of natural enemies some feimur prefers the lowlands and BPH while P nuch has no adapted to upland rice (22 27) ranging from egg predators environmental prefirences but is adapted only to WBPH (gryllids coccinellids) to larval parasites (braconids ichncu- The main d ryinid species on GLH aptogonatop)usspp is monids) to larval prcdators (ants and carabids) to larval most dominant in rainfed lowlands and also attacks other pathogeiis (vira and fungal diseases) Efforts to augment leafhoppers within Amnrasca Cicadulinaand Bachaltuathese natural enemies should focus on the egg predators Strepsiptera are more dominant on plarithoppers than (rearing releasing)and viral or fungal pathogens (culturing lealhoppers The Elenchus species in the Philippines attack spraying) all species of planthoppers within the genera Sogawilla

Parasitization of the large conspicuous larvae of Sogatodels NilaparsataHarualia and Opiconsiva Elenshyskippers green horned caterpillars and other polyphagous cusrvasuntaisuiis adapted to the major rice environments Lepidoptera by tachinids and chalcids is normally low ELg but prefers BPH in rainfed arid WBPIH in wetland environshyparasites and predators are perhaps the key to their control ments Ilalictoph(utsmtnroeiand HVpectrus attack G LH

Gall midges are parasitized by pterormalids and platy- and COfimravcctra respectivelygasterids that aestivate within their larval host between Pipunculids are mainly parasitic oii lealhoppers Each seasons Shoot aphids are preyed upon by coccinellids species has a unique environmental preference Tomosshysyrphids and chrysopids varireIa oraetoraprefers Ne7 otetltix nigropictrs and

Planthoppers arid leafhoppers have egg parasites and Pilunwulhtsjavanensisattacks Dehoceplhahls spp which predators as well as nymphialad nlt parasitesand predators are more abundant in upland than irrigated lowland their rich complex of natural enemies includes fungal environments T suhvirescensattacks Nephotettix virescens pathogens The principal egg parasites are myrnarids and and prefers the lowlands Pilpuiulus nttillattls is an trichogramniatids Egg predators are norinally mirid plant upland rice parasite arid attacks only Nephotetix speciesbugs that suck out the yolk Nymphal ad nIt parasites are More than 17 recorded species of leaffolder parasites in dryinids on planthoppers pipunculids on lealhoppers and the Philippines attack species of C(naplwocrocis and

30 IRPS No 123 January 1987

Table 6 Dominant natural enemies of rice pests in upland rainfed lowland and irrigated lowland rice environments of the Philippines

Natural enemy

Host stage Species

Brown planthop-Parasite Egg

MymaridaeAnagrus optabilis(Perkins) Anagrusflaveolus Waterhouse

TrichogrammatidacOligosita naiasGirault

NymphAdult Dryinidac

Pseudogotatopuslaiifemttr Esaki et Hashimoto Elenchidae

Elenchus yasurnatsui Kifune and Hirashima

Environment

Upand Rainfed lowland Irrigated lowland

Ailaparata lugens

x x X x

x

x x

x X X Whitebacked planthoppers Sogatella furcifera and Sogatodes pusanusEgg

MymaridaeAnagrus optabilis(Perkins)

TrichogrammatidacOl~gositanaias Girault

NymphAdult Dryinidae

Pseudogonatopusnudus Perkins Elenchidae

Elenchus yasutatsui Kifune et Hirashima

Parasite Egg

MymaridaeGonatocerus spp

TrichogrammatidaeOligositaaesopi Girault

NymphAdult Pipunculidac

Tomosvaryella subviresccns Loew Tomosvaryella oryzaetora (Koizumi) Pipunclusmutillatus (de Meijere)

DryinidaeIlaplogonatopusspp

Egg Copidosomopsisnacoleiae (Eady)

Larva BraLonidae

Cardiochiles philipphnensisAshnead lchneumonidac

Trichiomnia cnaphalocrosisUchida Temelucha stangli (Ashmead)

Pupa Chalcididae

BrachymeriaexcarinataGahan Brachymerialasus (Walker) Brachymeria nr tachardiaegroup

Parasite Egg

Scelionidae Telenomus rowani (Gahan)

EulophidacTettasticusschoenobhi Ferriere

Larva Braconidae

Stenobraconnicevillei (Bingham) Iclhncurnonidac

Temelucha phiiippinensis(Ashinead)Temelucha stangli (Ashmead)

x x x

x x

X x X

x x Green leafhopper Nephotettix virescens

X x x

x x

X X X X

X

X Rice leaffolders CnaphalocrocisMarasmia

x X x

X

X X

x x x x x X x x x

Yellow stem borer Scirpophaga incertulas

X X x

x

x

x X XAinauromorphaaccepta metathoracica(Ashmead) x

continued on next page

IRPS No 123 January 1987 31

Table 6 cntinued

Natural enemy Environment

Host stage Species Upland Rainfed lowland Irrigated lowland

Larva-pupa Gold fringed borer Chilo attriciiuslchneumonidae

Xanthopimpla stemmator (Thunberg) x aPupa x

Pteromalidae Trichoinalopsis apanteloctena (Crawford) ax

Pink stem borerb Sesamnia inferensParasite

Larva Braconidae

Stenobracon nicevillei (Bingham) x b b Armyworms Mvthinna SpodopteraLarva

Tachinidac Zygobothriaarroptvora (Robineau-Desvoidy) x x

Braconidac Cotesia spp

X Skipper Pelopidas mathias and green horned caterpillar Melanitis leda ismeneF-gg

TrichogramrnatidacTrichogrannaspp x

Scelionidae Undetermined Scelionid

x Larva

Tachinidae Argyroph lax nitrotibialis (Baranov) X x

Pteromalidae Trichomnalopsis apanteloctena (Crawford)

X Predator

CoenagrionidaeAgriocnenis femina femina Brauer x xIschnura senegalensis tambur

Miridae CrtorhinsliidipennisReuter x x

Nabidac Stcnonabis tagalicus (Stal) x

VeliidaeAlicrovelia douglasi atrolineata (Bergroth) x

Coccinellidae Micraspis crocea (Mulsant) X x x

Vespidae Eumenes campaniformis (Fabricius) x

Formicidac Diaecamna spp xOdontoponera transversa (F Smith) x

Carabidae Chaenius spp OphioneaW- aJasciataSchmidt-Goebel

x

Ophionea ishii ishii Habu x x X

Gryllidav Anaxipta longipennis (Saussur-) x xMetioche ittaticollis (Sial) x x x

Tettigoniidae Conocephalus longipennis (de Haan) x x x

aNo rearing made on rainfed lowland bNo rearing made on rainfed lowland or Irrigated lowland

MrasmiaAmong the Laval parasites the braconid Cardio- lowlands and parasitizes oriental maize borer OsIriniachilesphilijpphensis is most adapted to an upland environ- furnacalis (Guenee) larvae The leaffolder pupal parasitesment It also parasitizes Hydelepta indicata a common not only occur equally in all environments but also haveleaffolder of legumes The ichneumonids Trichomma cna- wvide host ranges- parasitizing species of OsIrinia Marucaphalocrosis and Temehcha stangli occur more in the Hydelepta and Honiona - in maize and legumeslowlands T stangli is most adapted to irrigated lowlands The principal egg parasite of yeltow stem borer (YSB)and also parasitizes Chilo and Scirpophaga stem borer Telenomus rowani is widely adapted to all environmentslarvae T cnaphalocrosis is most prevalent in rainfed and has an unusual alternate host -Tabanus eggs (25)

32 IRPS No 123 January 1987

Tetrastictu schoenobii specializes only in YSB and occurs mainly in irrigated lowlands -- further proof that YSB is most adapted to the wetter environments

Larval parasites of YSB are highly environment-specific The braconid Stenohracon nicevillei occurs only in upland rice but also attacks the pink stem borer (PSB) Tieinchwa phiiiplPwnsis an ichneumonid occurs in upland rice and also parasitizes the larvae of (7hiu stqpresaisinatro-morpha accepta melaihoracica like 7 stangi is mnost adapted to the irrigated lowlands

The dark-headed stem borer(l)HSB) (ih pochit3sus was not collected in all rice environments and the gold fringed borer Chilo auricilius nd PSB were collected only in upland rice Xaantiholiplastenrnaloran ichneumonid larval parasite appears to be widely adapted whereas TrihomaslopisapanitI)cteiiaa pteromalid pupal parasite occurs mostly in wetlands and also parasitizes larvae of Chilo suppressais and Pelhuidas mathias

Armyworms Myit hinin and Spodujipera have two main larval parasites - a tachinid Zrgohothriaatropivora in rainfed environments and a braconid Cotesia spp in irrigated lowlands

The butterfly pests Pelopidas and 4elanitishave tricho-grammatid egg parasites found only in irrigated lowland environments

Two larval parasites show environmental preferences --Argrroph av nigrotihialis a tachinid in rainfed habitats and Trichonialopsisapantelkcnela in irrigated ricefields A limgrotibialisalsoattacks the sweet potato hornwormAgrius convolvtdi and T apanteloctena parasitizes DH S1B

Predators as a group show more distinct environmental preferences The aquatic damselflies Coenagrionidae are most abundant in the lowlands Cvriorhinus is also most adapted to the lowlands The nabid Stenonabis tagalicus occurs only in the uplands and isalso prevalent on legumes The coccinellid lady beetles wasps and ants are more prevalent in upland areas AlicrasIpis cit-ea larvae prey ona variety of aphids attacking legumes and maize Etunienes campanonia vespid mud wasp makes nests in trees and is therefore most adapted to the more botanically diverse uplands Soil-dwelling ants cannot tolerate flooding

Upland environments are habitats to arboreal carabid beetles Three species of Chlaeniusbeetles prey on leaffolder larvae and are perhaps more important than parasites (27)

The gryllids and tettigoniids are widely adapted egg predators not only on rice but also on maize and legumes They feed on eggs of most insects that are laid on leaves

The spidercommunity of upland rice environments isrich in species At one site (Batangas Philippines) 31 species have been recorded (23) Of the 176 spider species recorded in Philippine ricefields about one-half (82) occur in upland rice The spiderspecies of upland rice environments overlap more with those of rainfed lowland than with those of irrigated lowland rice (24) The same study showed that of the three environments irrigated rice has (ie greatest spider species diversity foiiowed by rainfed lowland and upland

However of the 10 most prevalent ricefield spiders 9 were abundant in upland rice showing wide adaptation All three spider guilds -- orb-web space-web and huting spiders shywere prevalent in upland ricefields There were however differences in environmental prelerences for some spider species

Of the orb-wcavers Tetragniathamanlihulatawas partishycila rlv abundant 7 japonica was low and Ltucatge dehcorata was absent in upland rice

As in other environments Aiypeiwnaifirtosana was the most dominant space-web spider however its relative numbers were lowerthan for lowland sites Among hunting spiders Ox|opes javanus and Lcosa leatcost fgla were more abundant and Lvcosa lpseudoanmnlata was less abundant in upland compared with lowland rice envishyroniments

Extensive rearing of rice insect pests in upland rainfed lowland and irrigated sites over an 8-yr period in the Philippines provided insight into the effectiveness of parasites as natural enemies (Table 7 8) Most of toe data are from large sample sizes taken over at least one crop which overcomes the pitfalls expressed by Van Driesche (238)

VSB egg parasitization was surprisingly similar across all environments Larval parasitization was highly dynamic among sites and even years within the same site but showed somewhat lowerlevels in uplandsthan in irrigated wetlands This would have been predicted for egg parasites as well since YSB is adapted to the lowlands and so must be its parasites This isevidence that Telenomus rowani is as good at dispersing as its main hosts Trowani clings onto the anal tufts of female YSB and parasitizes eggs as they are laid This phoresy may explain its wide adaptation The more specialized lowland YSB larval parasites therefore appear more effective than their upland counterparts with more alternate hosts

Low collection levels of PSB larvae especially in the wetlands make comparisons between environments difficult Low levels of larval parasifization were recorded in both rainfed environments Except in the Cagayan rainfed wetland site in 1980 low levels ef larval parasitization were evident at irrigated and rainfed sitesalike No differences by environment were evident Higher levels of parasitization occurred in Sarawak during a 1967 outbreak also from tachinids Rothschild (197) indicated that higher parasishytization occurred in upland rice areas

Skipper and green horned caterpillar larval parasitization rates were generally low and showed no environmental effect htighest activity occurred in Cagayan in 1980 (48 parasitization) Although larval parasitization levels of RLF were variable year to year activity appeared greater in the irrigated lowlands than in the uplands Again the highest levels were recorded in Cagayan in 1980 Cagayan has extensive grasslands and fallow around the rice areas and a relatively long rainfall period Rainfall in 1930 was good for crop growth and apparently for increase of larval parasites

IRPS No 123 January 1987 33

Table 7 Parasitization rates of rice planthoppers and green leafhopper in upland rainfed lowland and irrigated lowland environments Philipshypines 1976-840

Environment and site Year Egg parasitization () Nymphaladult parasitization ()

BPH W13P GLH BPII WBPH GLIH

UplandBatangas

Rainfed lowland

1976 1979 1980 1982

15 17 12

2(47)8(39)

4(51)

9(1800)14 (212)

12(304)

8 (101)6 (127)

9(116)

Iloilo

Cagayan

1976 1977 1978 1981 1980 1981 1982

0 (69)

11

44 48 40

-

-

46 55 44

0 (471)

11

52 56 49

0 (200)9 (214)

10 (1259)8(235)6 (900) 3(800)

-7 (462)

11 (270)3 (313)5 (900) 1 (800)

4 (600)10 (725)10 (362)12(870)5 (900) 3 (800)

Pangasinan

Irrigatedlowland

1984 1976 1978 1979 1982

13 (436) 13(379) 14 (457)

6(14)7 (197)

17(198) 11 (486) 18 (275)

3 (81)10 (274)3 (116)

16(871) 19(88)

28(211)9 (764)5 (647)7 (1204) 8(305)

Laguna 1977 11 7 21 1978 31 1979 14 4 4 28 1980 7 8 20 1981 1983

56 7(278)

26 16 (1650)

16 27(1921)

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R I Apostol unpublished 1lo1lo and Pangasinan 136 Cagayan BCanapi unpublished Liuna 175 I C Pantua unpublished 118 234 43 Carino and Shepard unpublished BPH = brown planthopper Nila parvara htgens W131Pi =whitebacked planthopper Sogatelia furcifera GLH = green leafhopper Nephotettrix virescens Figiires in parentheses are sample sizes

Tauher et al (230) reported that internal external and genetic factors influence the seasonal activity of parasitoids

Hopper egg parasites but not nymphaladult parasites were also parlicularly abundant at the Cagayan site possibly the egg parasites have more alternative hopper prey in the weedy areas than nymphaladult paisites

The seasonal dynamics of hopper egg parasites can be seen for upland rice in Batangas(Fig I 2) and for rainfed lowland lice in Pangasinan( 135)and Cagayan( 105) Levels of BPHWBPI-L and GLH egg parasitization were corn-parable in upland and irrigated lowland environments Except for some high rates of paraMsitization early in the season the parasitization rate tended to be steady in all envilonments However hopper nymphaladult parasi-tizution levels were more similar in the rainfed environments and lower than in irrigated rice

Overall parasites do not appear highly effective by themselves against upland or even lowland pests Among natural enemies parasites have attracted the most attention because they can be more readily assesscd

Predators and pathogens lend themselves better to management practices Predators the most important group of natural enemies in riccfields are difficult to quantify On the other hand pathogens appear less im-portant but can be readily cultured and disseminated

The natural enemy commulnity of upland rice is rich in species and differs significantly from that of lowland rice

These beneficial organisms must be conserved by applying insecticides judiciously particularly sprays Seed or soil placement of chemicals minimizes exposure to natural enemies The strategy to derive the greatest benefit from natural enemies isto allow the greatest number of beneficial species to thrive Then perhaps at least one species will be effective against each pest at any tine overcoming the variable abundance of each species within and between years

Programs to introduce exotic species or mass produce indigenous ones are ambitious and expensive because they require trained people on a sustained not ad hoc basis

Chemical control Insecticides offer rapid and efficient control of upland insect pests (190) but-should be used only after other control measures have been considered Insecticides are rarely used on upland rice because of cost (32) Upland rice yields are normally too low to justify the expenditure Also most upland rice is grown as a cash crop Spraying upland rice is more difficult than lowland rice because water is less accessible Government subsidy programs to provide pestishycides to tribal peoples for upland rice production have created problems of toxicity to humans because isolated people have had no experience to be able to handle pesticides safely

However ins-eticide use has beenjustified when a crop is

0Z

Table 8 Parasitization rates of lepidopterous pests in upland rainfed lowland and irrigated lowland environments Philippines 1976-84a

Parasitization (5)

Environment and site Year YSB GFB PSB Earcutting Swarming Skipper Greenborned RLF Brown larva larva caterpillar caterpillar caterpillar caterpillar larva semilooperEgg Larva larva larva larva larva larva

UplandBatangas 1977 - 0 (57) - 0 (63) - 0 (71) - 0 (106) 11(142) 2 (101)1978 43 (4697) 8(451) 5 (38) 3 (40) 6 (61) 2(58) 15 (57) 13 (129) 13 (1286) 4 (30)1979 59(4135) 12(320) 13 (11) 5(28) 9(45) 6(46) 7(23) 8(160) 9(1152) 8(26)

Rainfed lowland Iloilo 1976 - 0 (10) 0(10) 0 (7) 0 (19) 2 13) 15 (180) 5 (76)1978 0 (54) 0 (3) 0 (17) 9 (64) 4 (78) 3 (73) 0 (121) 0 (191)1979 64 (2350) 16 (84) 6 (17) 0(6) 18 (68) 25 (104)

1981 56 (1322) 18(304) 0(5) 12(51) 18(138)Pangasinan 1976 60 (7 7 )b 62(82) 0(111) 5 (39)

1978 58 (53)b 5(51) Cagayan 1979 59(713) 8 (101) 6 (34) 0(7)

1980 0(11) 4 (28) 15 (286)

62 ( 48(177) 61(140)1981 0(5) 12 (2) 13(45) 14 (92)

Irrigatedlowland Laguna 1974 62 8

1974-75 50 41 1975 43 72 1978 96 (623) 21(24) 33 (60)1979 83(2017) 18(110) 8(62) 11 (89) 22(400)1980 78 (1625) 20(75) 0 (16) 4 (72)1983 60

9 (80) 7 (270) 30

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R F Apostol unpublished Iloilo and Pangasnan 136 Cagayan B Canapi unpublished Laguna 175 P CPantua unpublished 118 234 143 Carino and Shepard unpublished YSB = yellow stem borer ScirpophagaincertulasGFB = gold fringed borer Chilo auricilius PSB = pink stem borerSesamia inferens earcutting caterpillar Alythimna separata swarming caterpillar Spcdoptera mauritia acronyctoides skipper Pelopidas mathias reen horned caterpillar Melanitls ledaismene RLF = rice leaffoldersCnaphalocrocismedinalis and Marasmiaspp brown semilooper Mocisfrugalis Figures in parentheses are sample sizes Egg masses

IRPS No 123 January 1987 35

Table 9 Effect of seeding rate and insecticide on plant density and Yield losses from insect pests in upland rice often areyield of Dagge upland rice Batangas Philippines 1976a comparable to those of lowland rainfed or irrigated rice

Seeding rate Insecticide Dosage Plant density Yield (0-Y30o) but usually from different pests The principal (kg aiha) seed treatment (kg aiha) (noIra-row) (tha) groups are soil pests feeding on sown seed and rootsat 14 DE followed by seedling pests These groups have largely gone

50 None - 35 a 20 a unnoticed leading researchers to conclude that stem borers50 Carbofuran F 10 38 ab 23 a and seed bugs are the most important pests (17) Yield loss 10 beCabofuranF100 None - 4551 c 23 a trials that pinpoint major pests need to includc seed

100 Dieldrin WP 10 52 cd 22 a treatments Granules applied in the furrow may not give150 None - 59 d 22 a adequate protection from many seed pests

aAv of 8 fields DE = days after crop emergence In a column means The floristic and pedological diversity of upland rice followed by a common letter are not significantly different at the environments is matched by the diversity of arthropods5level by D)MRT F = flowable Wi = wettable powder both pests and their natural enemies Irrigation has homoshy

genized lowland rice environments Flooded soils tend tothreatened by epidemics of locusts or armyworms New have the same properties and to eliminate soil pests The technology stressing efficiency has minimized insecticide spatial and temporal dominance of cultivated rice has cost benefited specialist pests

Seed treatment is normally inexpensive and can be Upland rice has no specialist arthropod species The economicallyjustified in -nanycases(66) Treating seed with monophagous pests on upland rice prefer the lowlands systemic insecticide (216) but not chlorinated hydrocarbons where rice evolved Rice in the uplands exists only because (247) is most effective in controlling seedling maggots of man and even though lowland rice predates upland riceFoliar sprays have to be repeated to obtain control and specialist species evolved Wheat maize and sorghum do often are washed off by frequent early season rains not have monophagous pests leading us to conclude that Granular insecticides are effective but required dosages are there is no evolutionary advantage for inssects to have too expensive to justify except under heavy infestation narrow host ranges in upland graminaceae(247) When directed against first-instar larvae banding Pests have adapted to the uplands not only by having granules in seed furrows (177) can be effective against white wider plant host ranges but also by having long life cyclesgrubs at low dosages (025 kg aii ha) (134) Seed treatments undergoing dormancy or being dispersive Each pest has a are not promising for white grub control (248) unique life history strategy suited to highly unpredictable

Seed treatment is also effective ainst ants and may be environments In agivenyear someinsect specics an better cheaper than increasing the seeding rate In Batangas exploit upland rice as a food source but no pest is a senious treating 50 kg seed ha was as effective as increasing the one every year on upland rice seeding rate to 100 kgha and using untreated seeds (Table The low yielding potential of upland rice means less 9) In Brazil seed treatments control termites and other soil quantity and lower quality of food for pests to exploit but insects including the lessercornstalk borer(66) Domiciano also less incentive for farmers to undertake control (58) however found seed treatments inconsistent in their The most effective control would be early and synchroshyeffect against high populations of lesser corn stalk borer nous planting of early-maturing upland rice varieties to

Baiting can be an inexpensive way of controlling ants prevent insect pest buildup Breeding for insect resistance seedling maggot flies mole crickets and field crickets Baits should initially focus on stem borers because they are verymade of locally available material and impregnated with difficult to control by other means Biocontrol efforts insecticide can be sparingly distributed taking adantage of should first stress conservation of natural enemies preshythe pests ability to disperse and encounter bait sites dators and pathogens rather than parasites Efficient

The new low-volume hand-held controlled-droplet chemical control includes seed treatment or baiting for seed sprayers may offer an advantage over high-volume knap- seedling and some root pests and using controlled dropletsack sprayes for controlling foliar pests Upland rice sprayers for spot treatments against foliar pests and seed canopies are more open than those in lowland rice and bugs when economic thresholds are reached better droplet penetration should result Controlled droplet applicators would be ideal for seedbugs and defoliators

CONCLUSION

Upland rice is attacked by a wider array of insect pests than is lowland rice mainly beca use of the addition of soil pests Generally population levels of lowland rice pests are lower in upland than lowland rice leading many people to conclude that insect pests are not important on upland rice

36 IRPS No 123 January 1987

REFERENCES CITED

1Abraham C C and K S R Mony 1978 Occurrence ofLeptocorisa acuta Fabr (Coreidae Hemiptera) as a pest of nutmeg trees J Bombay Nat ttistSoc 74553

2 Agyen-Sampong M 1975 HlierogrlphusdiaganensisKrauss (Orthop-tera Acrididae) a new pest of rice in Northern Ghana Ghana J Agric Sci 8249-253

3 Agyen-Samnpong M 1976 Varietal resistance and other inle-ratcd pest control components as factors of rice varietal improvement in West Africa Prcsented at the Second varietal improvement serinar 14-18 Sep 1976 Bouak Ivory Coast West Africa Rice Development Association Monrovia Liberia 18 p

4 Agyen-Sampong M 1977 Outbreak of Ihccrorrdnisorae in rice seedlings near Rokupr Sierra Leone lit Rice Res Newsl 2(6)16-17

5 Agyen-Sampong M 1980 Dirty panicles and rice yield reduction caused by hugs IntRice Res Newsl 5(l)11-12

6 Agyen-Sampong M 1982 The major pest problems of irrigated upland and mangrove swanip rice ecosystems ir the humid tropical Guinea savanna and Salel climate zones Pages 318-328 in Pro-ceedings training course cn concepts techniques and application of irtegrated pest management in rice inWest Africa 10-20 Jan 1982 Fendall Liberia West Africa Rice l)evelopioent Association Monrovia Liberia 505 p

7 Akibo-Betts V I and S A Raymundo 1978 Aphids as rice pests in Sierra Leone lit Rice Res Newsl 3(6)15-16

8 Akinsola F A 1975 Present status of different rice stemborers in parts of Nigeria iicL Entomo Newsl 328

9 Akinsola FA 1979 Ihe biology and ecology of rice stemborers in Nigeria Ph 1) thesis University ol lbadan Nigeria 217 p

10 Akinsola E A 1979 Guidelines of integrated management of insect pests of upland rice in West Africa Presented at the Seminar on integrated management of rice diseases and insect pests at Bobo-Dioulasso Upper Volta 17-22 Sep 1979 West Africa Rice Develop-ment Association Monrovia Liberia

II Akinsola IA 1984 Insect pests of upland rice in Africa Pages 301-305 in An overview of upland rice research Proceedings of the 1982 lIouak Iwsrv Coast upland rice workshop International Rice Research Institute 13O Box 933 Manila Philippines

12 Alam M S V TJohn and Kaung Zan 1985 Insect pests and diseases of rice in Africa Pages 67-82 in Rice improvement in Eastern Central and Southern Africa proceedings of the international rice workshop at Iusaka Zambia 9-19 April 1984

13 Alticri M A A van Schoonhovcn and J D Doll 1977 The ecological role of weeds in insect pest management systems a review illustiated with bean (Phaseohsvulgaris L)cropping systems PANS (Pest Artie News Summ) 23195-205

14 Anmante F 1967 The leaf-cutting ant Atia capiguaraa pest of pastures [in Portuguese] Biol6gico (Sao Paulo) 33113-120 (Rev Appl Entomol A56207)

15Andrade A C and D Puzzi 1951 Preliminary results of experi-merits on the control of Sculptcoris iastantus on sugarcane [in Portuguese] Bioulgico (Sao Paulo) 1744-49 (Rev Appl Entomol A41421422)

16 Appert J 1970 Maliarplia separaelha (rice stem borer) new observations of entomological problems of rice in Madagascar [in French] Agron Trop (Paris) 25329-334

17 Ariadeau Mand Z Harahap 1986 Relevant upland rice breeding objectives Pages 189-197 in Progress in upland rice research Proceedings of the 1985 Jakarta conference International Rice Research Institute 1O Box 933 Manila Philippines

18Asar S and Kisimoto 1975 Seasonal occurrence of the smaller brown planthopper in upland rice [in Japanese] Proc Kanto-Tosan Plant Prot Soc 2228

19 Bakker W 1974 Characterization and ecological aspects of rice yellow mottle virus in Kenya Meded Landbouwhogesh Wage-ningen 152 p

20 Balut F F 1970 White grubs attacking rice Oryza ativa L [in Portuguese] Biol6gico (So Paulo) 36321-322

21 Banerjee S N and L M Pramanik 1967 The lepidopterous stalk borers of rice and their life cycles in the tropics pages 103-124 in The major insect pests ofthe rice plant Proceedings ofa symposium at the International Rice Research Institute September 1964 The Johns Hopkins Press Baltimore Maryland USA

22 Barrion A T and J A Litsinger 1980 Ants a natural enemy of cliaiarac in dryland rice IntRice Res Newsl

5(4)22-23 23 Barrion A T and J A litsinger 1981 The spider fauna of

Philippine rice agroecosystem 1Dryland Philipp Entomnol 5139-166

24 Barrion A T and 1 A Iitsinger 1984 The spider fauna of Philippine rice agroccosystcms II Welland Philipp Eiitomol 611-37

25 Barrion A T1 and J A Litsinger 1984 lihanus (Diptera Tabanidae) eggs an alternative host of rice stei borer (SI3) egg parasite Telenoonis digns (Iliymenoptcra Scelionidae) lit Rice Res Newvsl 9(6)19

26 Barrion A T and J A Litsinger 1985 Identilication of rice leaffolders by wing markings Int Rice Res Newsl 10( 1)24

27 Barrion A T and J A Litsinger 1985 Chctiosspp (Coleoptera Carabidae) a leaffolder ILF)predator lit Rice Res Newsl 10(1)21

28 Bertels A 1970 Insect pests of rice and theircontrol [in Portuguese] Pelotas Brazil Instituto de Expcrimentacao Agropccuararias do Sul Circular 43 24 p

29 Bonzi S M 1982 Ctilo (flisilineus J de Joannis (Lepidoptera Pyralidae) stem borer of irrigated cereals in Upper Volta LioFrench] Agron Trop Riu Rizic Cult Vivrieres Trop 37207-209

30 Bourke T V T L Fenner J N LStihick G Baker EHassan D F OSullivan and C S li 1977 Insect pest survey for the year ending 30th June 1969 Papta New Guinea )epartment of Agrishyculture Stock and Fisheries Port Moresby 57 p

31 Bowden 1 1973 Migration of pests in the tropics Meded Rijksfac Landbouwwet Gent 38785-796

32 Breniere J 1973 The main rice parasites in West Africa and their control Pages 27-42 in Proceedings seminar on plant protection for the rice crop West Africa Rice Development Association Monrovia Liberia

33 Breniere J 1976 The principal insect pests of rice in West Africa and their control West Africa Rice Development Association Monrovia Liberia 52 p

34 Brown ES 1962 The African arinyworn Spoiolptera eXI)Ina (Walker) (Lepidoptera Noctuidae) a review of the literature Cornshymonwealth Institute of Entomology London UK57 p

35 Brown F B 1968 Upland rice in Latin America FAO-International Rice Commission 12th Session of the Working Party on Rice Production and Protection 2-14 Sept Peradeniya Sri lanka Food and Agriculture Organization Rome Italy 6 p

36 Buyckx F J 1 1962 Disease and insect pests of crops inCongo Rwanda and Burundi [iil French] Publications dc LInstilut National pour lestude Agrononiquedu Congo M Weissenbruck S A Brussels Belgium 708 p

37 Cariiio F 0 1981 Role of natural enentics inpopulation suppression and pest management of green rice lealhoppers Ph D thesis University ofthe Philippines at Los Bafitos PhilippinesLaguna 136 -

38 Carpente A J 1983 The development of rice cultivation and extension of vegetable productin in Zanzibar Final Report Vol II1 October 1983 Food and Agricultme Organization Ronte Italy

39 Carroll C P and S1 Risch 1983 Tropical annual cropping systems ant ecology Environ Manage 751-57

4(0 Cates R G 198 1 Host plant predictability and the feeding patterns of monophagous oligophagous and polyphagous insect herbivores Oecologia (Berl) 48319-326

IRPS No 123 January 1987 37

41 Cendcafia S M and F B Caora 1967 Insect pests of rice in the 62 Ene 1 C 1981 I)iiorldhol)ierusihilis Slater (licteroplera Philippines Pages 591-616 in The najor insect pests ofthe rice plant lygacidue Illissinac) a new rice pest front Nigeria lit Rice Res Proceedings of a symposium at the International Rice Research Newsl N3)15 Institute September 1964 The Johns Hopkins Press Baltimore 63 Fugenio T S 1965 rhe effect oi e-vCgetation on rice production Maryland USA 729 p and the possibility of overcoming it by leaf pruning MS thesis

42 Centro Ihternacional de Agricklittira Tropical 1984 Upland rice in University ol the Philippines at Los Baijos ILaguna Ihilippins 69 p Iatin Anerica Pages 93-119 hi An os ervies of uplaind rice research 04 Farrow R A 1975 The Alrican nmigratory locust ii is main Proceedings of the 1982 Boinakamp Ivory Coast upland rice workshop outbreak area of the middle Niger qlinlitatise studies of solitary International Rice Research Institute P0 Blox 933 Manila populations in relation to environmental factors Locusla No II Philippines 198 p

43 Chandra G 1979 Taxonomy ad hn riononlics ofthe imect parasites of 65 Fennal R G 1943 Food crop investigation Windward and rice Icalloppers and planthoppers in the Pihilippines and their Ieeward Islands First report Report onl work dunring the periodiiiportairce iii natural biol)gical cointrol Philipp Fntoltol 4119-139 April 1942-April 1943 Imperial College of Tropical Agriculture

44 Chang T T 1976 The originii colut ion enlIivation dissemination Trinidad 16 p (Rev of Appl Fintornol A3222-25) and diersificition of Asian and African rices -uphytica 25425-44 1 66 Ferreira F 1983 Integrated pest ritaageiret of upland rice [ill

45 Chatierjee P B 1979 Rice leaffolder attacks in India lit Rice Res Portuguese English summary] Pages 323-341 in Upland riceculttre Newsl 4(3)21 factors affecting production E Ferreira T Yamada and F-

46 Cheng Cbien-pan 19(4 Upland rice in Taiwan International Rice Malmolta eds Institutoda Potassae Foslato Institnto Internacional Cnmmission Wiorking Party oit Rice Production ard Protection I()th da Potassa Piracicaha Sao Paulo lraid Meeting 3-10 March 1964 Manila Philippines Food and Agricul- 67 Ferreira E arid F dl Silva Martins 1979 Occurrence of ture Organiaion Rome Italy 5 p Sogatohs orvziola (Muir) inupland rice in Goias Ilrail lit Rice

47 Chcrrett 1 M and 1) 1Peregrine 1976 A review of the status of Rcs Ncwssl 4(4)18 leaf-cutting ants and their control Ann Appl Biol 84124-128 68 Ferreira E and 1F da Silva Martiris 1984 Insect pestof rice ill

48 Chipeta F M 1974 Pests of rice in Malawi Pages 90-95 in Brazil and their control Jill Portugucsej FMIlRAtPA-CNPAF Proceedings Sith eastern Africa cereals research conference 10-15 Goiania Brmiil 67 p March 1974 DRB Mialda ed Ministry of Agriculture lilongwe 69 Ferreira FI F ca Silha Miartins S S Nero and 1 It P de Malawi Carvallio 1982 M tltiplc regression analysis todeterimine the effect of

49 Chu Yau-i 1979 The d atiaging period of the rice seedling fly insect populatiois ott upland rice yield [inIPortuguese Pesqui (Atlterigotiavxigu Stcin) (NItiscide Diptera) oil cori) at Fast Java Agropecu Bras 17671-675 Indonesia Plant Prot Bill 21403413 70 Ferreira F 1F da Silva Martins and F 1PZimmermann 1979

50 Conklin H C 1957 IHaninoo agriculture a report on aln integral Rice varietal resistance to tle lesser corn stalk borer [in Port uguese] systent of shi fti rig cult ivationi in the Pltilippincs Food and Agriculture Pesqui Agropccu Biras 14317-321 Organization Rome Italy 209 p 71 Fleming W F 1968 Biological control of lie hapanese beetle U S

51 Cramer H II 1967 Plant proection and world crop production Dep Agric RLs Sers [ech Bull 1383 78 p Pilaniensclhutz-Nachr 201-524 72 Food and Agriculture Organiation of the United Nations 1982

52 l)ani R C and Majunidar 1978 Prelininary observations on tie Country report surmimary Pages 27-37 in Report on the first incidence of root aphids on differeit rice cultnrcs Sci Cult 4488-89 FAOIUN)IPhail and regional training coiuirse in iniproved ctltural

53 Davidson R I 1979 lffects of root feeding on foliage yield Pages practices for upland rice 5-23 Oct 1981 Northern Region Agrculturd117-120 in T K Ci osb and R B Pottinger eds Proceedings of the Developririt Center Chiang Mai Food and Agricuilture Organiiashy2nd Australian conlerence on grassland invertebrate ecology Govt tion Rome Italy 37 n Printer Wellington New Zealand 73 Franco do Ariaral S 195 1 Residual action of insecticides in turfy

54 l)ean G 1W 1976 Rice insect pests inlaos Int Rice Res Newsl soil--the results of a field experiment [in Portuguese] Biol6gico (Sao 1(2)15 Paulo) 17209-214 (Rev Appl Entomol A41423)

55 I)ean G J W 1978 Insect pests of rice in Laos PANS 24(3)280-289 74 Freeman ) 1970 Report onithe Iban london School of -coroiics 56 )ingle H 1984 Behavior genes and lifc histories conplex adapta- Monographs on Social Anthropology No 41 Athlane Press

tions to uncertain environments Pages 169-194 in A new ecology London UK 317 p novel approaches to interactive systems P W Price C N Slobod- 75 Fuchs T W1 A Harding and 1W Smithlr 1979 Induction and chikoff and W S Gaud cdsl ohn Wiley New York USA termination oldiapause in the sugarcane borer Ann Flitonlol Soc

57 Dobelmann-JP and M Falais 1968 Upland rice cultivation ill tie Am 72271-274 North-western part of Madagascar in 1966-1967 [in French English 76 (hbriel B1) 1975 A review tf the major insect pests of some tpland summary] Trop Agron 2313-21 crops in tlc Philippines Pages 86-111 in Ad Htoc panel of experts on

58 Doniiciano N L 1979 Chemical control of tie lesser corn stalk pest disease arnd weed problems in some rainfed crops Bangkok (lolsonno~paloIgnowslhts) (Zeller 1848) (I-epid(ptera Ihycitid ae) Thailand Food aid Agriculture Organization Roire Italy

on rice in Ponta Gressa Parara 1977-1978 [in Portuguese] Pages 77 Garg D K and N K Shal 1983 White grub outbreak on rainfed 291-303 in Anais 1 Reuniao de tecnicos cmiirizicultura do estado de dryland rice in Uttar Pradeslh Int Rice Res Ncwsl 81(4)16-17 Sao Paulo-Campinas Cati Brazil 78 Garg I) K and IP Tandon 1982 Major insect pests of rice on hilly

59 Dunsmore 1 R 1970 Investigations on tlie varieties pests and tracts of Uttar Pradesh India lilt Rice Res Newsl 7(11)11-12diseases of upland rice (hill padi) in Sarawak Malaysia Int Rice 79 Garrity 1) P 1983 Upland rice research an international bibliograpy Comm Newsl 1929-35 1975-1982 Iternational Rice Research hnstitnte PO Box 933

60 Elias 1R1965 The new decline of rice seedlings is not caused by Manila Philippines 121 p parasites [in Portuguese] Biol6gico (Sao Paulo) 313-6 80 Ghauri M S K 1982 A new subspecies of Iinorh)rone bull

61 Empresa Brasiliera de Pesquisa Agropecuaria 1984 Upland rice itt cornlus Slater (Hemiptera Iygaeidae) front Papua New Guinea on Brazil Pages 121-134 it Av overview of upland rice research rice and carpet grass Ihll lEntoniol Res 72133-137 Proceedings ofthe 1982 Blouak Ivory Coast Upland rice workshop 81 Gonzales F 0 PArregoces R I ihernandef and OT Parada International Rice Research Institute PO Box 933 Manila 1983 Insect and imite pests of rice arid their control in Latin America Philippines [in Spanish] la Federacion Nacional de Arrocerts (Fedearroz)

Bogota Colombia 60 n

38 IRPS No 123 January 1987

82 Grist D H and R J A W Lever 1969 Pests of rice Longmans London UK520 p

83 Hale P R 1979 Two subterranean pests of upland rice in Papua New Guirnca IntRice Res Newsl 4(2)16-17

84 Hale P R and J T Hale 1975 Insect pests of rice in the New Guinea Islands Rice Entomol Newsl 34

85 Harris W V 1969 Termites as pests of crops and trees Common-wealth Institute of Entomology London UK 41 p

86 Hattori 1 1968 A method to distinguish between grass leaf roller (Ctaplha rocis medinalisGuenee) and the rice leaf roller (Susurtia exigua Butler) [in Japanese] Plant Prot (Japan) 23167-170

87 Henry I E and 1A Onsager 1984 Experimental control of the mormon cricket Iahrosshnph by NJsena locustae(Aficrospora Microsporidae) a protozoan parasite of grasshoppers (Ort Acri-didae) Entomophaga 27197-201

88 Hidaka 1 P Vungsilabulr and SKadkao 1974 Studies on ecology and control of the rice gall midge in Thailand Tech Bull 6 TropicalAgricultural Research Center Tokyo Japan 113 p

89 Hincklcy A 1) 1963 Ecology and control of rice planthoppers in Fiji Bull Entomrol Res 53467 481

90 Hirao J 1982 Ecology and chtInical control of the rice leaffolder Jpn Pestic Inf 4114-17

91 Ho DT 1984 Sten borer incifience in rice ecosystems in KenyaEast Africa Pages 307-320 in A overview of upland rice research Proceedings of the 1982 Bouac Ivory Coast upland rice workshop International Rice Research Institute PO Box 933 Manila Philippines

92 Ho )T J 1Njokah and J GKibuka 1983 Studies on rice sten borers in Kenya with emphasis on ifaliarpiaipiraellRag Insect Sci Appl 465-73

93 Ichihara I T Ito A NIatsuo and M Sawada 1971 Chemical control of the rice striped stem borer in upland rice grown in plastic mulch [iii Japanese] Proc Kanto-Tosan Plant Prot Soc 1882

94 IclhiharaIand A Matsuo 1969 Damage caused by rice striped stett borer to upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1686-87

95 Ichhara I atid A MIatsun11970 Conside ration oft lie extraordinary prevalence of rice striped stem borer in upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1778-79

96 Institut de Recherches Agrolnliqes Iropicale et des Cultures Vivrieres 1978 Insect resistance Pages 12-13 in Upland rice 1978 atnual report Bouak Ivory Coast 27 p

97 International Institute for Tropical Agriculture 1983 Aphids itt rice roots I ITA Res Briefs 4(1-3)4-5

98 International Institite fi Tropical Agriculture 1983 Hopperburn in rice caused by brown plathopper IITA Res Briefs 4(l-3)6-7

99 International Rice Research Institute 1975 Annual report for 1974 PO Box 933 Manila Philippines p 216

100 International Rice Restarch Institute 1976 Annual repiort for 1975 PO Box 933 Manila Philippines p 228

101 International Rice Research Institute 1979 Cropping systems program Component technology development and evaluation Page 404 i Annual report for 1978 11O Box 933 Manila Philippines

102 International Rice Research Institute Department of Entomology 1981) Insecticid evaluation 1979 Tables 73 74 PO Box 933Manila Philipes

103 International Rice Research Institute Department of Entonmology 1981 Insecticide evalomtion 1980 Table 81 POBox 933 Manila Philippines

104 International Rice Research Institute 1981 Report of a meeting of Asian rice-based cropping systems entomologists Eleventh cropping systems working group meeting 18-22 May 1981 PO Box 933 Manila Philippines p 132 133 144

105 International Rice Research Institute 1984 Cropping systems program Pages 368-373 in Annual report for 1983 PO Box 933 Manila Philippines

106 International Rice Research Institutc 1984 Upland rice in Asia Pages 45-68 in An overview of upland rice research Proceedings of the 1982 Bouakamp Ivory Coast upland rice workiiop loternational Rice Research Institute 11O Box 933 Manila Philippines

107 Irshad M 1977 Grasshoppers in rice in Pakistan lit Rice Res Newsl 2(3)8-9

108 Ito R and TOno 1964 Upland lice its culture and varieties in Japan lIternational Rice Cormmission loth Meeting of the Working Party on Rice Production and Protection 3-10 March 1964 Manila Philippines Food and Agriculture Organization Rome Italy 12 p

109 Jeffrey M I and V A Dyck 1983 Effect of moonlight on lightshytrap catches of brown planthopper lit Rice Res Newsl 8(6) 13

110 Jennings P R and T Alicia Pineda 1983 Management ofthe rice plnthoppers Sogatodsorivi(coIl in Latin America Presented at the meeting on stable plant resistance FAO and lIndocsian I)inectorate of Food Crops Protection Bali lndonesia 29 Novcniber-2 l)ccemshyber 9 p

IllJimenez Gomez J A 1979 Rice insect pests i lla ios ()riCntales lil Spanish] Pages 99-103 in Rice ciltivation Instituto (olotibiatto Agropecuario Compndio No 29

112 Jongeleen F JJ Soesanto and A Mockidjo 1979 Studies oti pathogens of two species of Scarabacidac froinIndonesia Z Angew Enlomnol 87412-417

113 Jouvenaz 1) PC S lofgren and W A Baiks 1981 Biological control of imported fire antsa review of cirrent knowledge Bull Entomol Soc Am 27203-208

114 Kainoh Y K Shiniiu S MarUani Y Taiaki 1980 lost finding behavior of tie rice bu lplotori+JchinvittjI)allas (liciniptera Corcidac) with special reference to diet pattern ol aggregation and feeding on the rice plant Appl Entomol Zool 15225-233

115 Kakkar K 0 P Sharma and P I Sharma 1977 Note on Ihteronyrchus liodresRedt (Coleopicra I)ynastidae) as asporadic pest of paddy and maie inHimachal Pradesh Corr Agric 156-57

116 Kalode M131 79 Major insect pest problems and their control in upland rice (Unpublishcd mineogriph CWI oJNaaiat lni Chandaratat (198 1)original not seen)

117 Kalshoven L i 98 I The pests of crops in I ldoniesia (translated by P A vatn der Laan) 1 T Ichtiar B3aru-vati Iloeve lakarta Indonesia 701 p

118 Kamal N Q 1981 Suppression of whitebacked planto ner Sogatella fircaifra (Horvath) aLId rice lealfolder (nhal -cis licdb lis (Guenee) populations by natural cnclmie hlI) tiesis Grcgorio Araneta University Foundation Victoncta Park Metto Manila Philippines 179 Ip

119 Karim A N M Reaul 1975 A species ofliealy bug Trioni i70s ceres new oii rice in the Philippines Rice Entoniol Newsl 247

120 Katanyukul Wand C Chandaratat 1981 Insect pestsof uplaid rice and their control Report firstFAOUNlFi Tliailand training course on improved cultural practices for tplad rice Chiang Mai Thailand 5-23 Oct 1981 Food and Agriculture Organiation Rome Italy 22 p

121 Kern F 1955 lhletwioa(I)uninemts) bhhit Biturm(Coleoptera) a new pest of sugarcane inVenezuela [inSpanish with Etnglish sunmary] Agron Trop (Maracay) 527-29 (Rev Appl Eitoniol A4572)

122 Khan K M and S Ghai 1974 White grubs ald their control in India Pesticides (1Bombay) 819-25

123 Kiritani K 1975 Integrated pest management of rice pests Pages 66-70 itt Proceedings platning workshop ott cooperative hIeld research in pest managementP SMotooka ed East-West Center Honololu Hawaii USA

124 Kisimoto R 1967 Ecology of insect vectors forecasting and chemical control Pages 243-255 in The virus diseases of tle rice plant Proceedings of a symposium atthe International Rice Research Institute April 1967 The Johns Hopkins Press altimore Maryshyland USA

125 Kisimoto R and V A Dyck 1976 Climate and rice insects Pages 367-391 in Climate and rice International Rice Research Institute PO Box 933 Manila Philippines

126 Kureha S S Kobayashi SAsari and T Ishihara 1977 Life history and control of the mole cricket Grrllotalpa africana Palisot de Beauvois [in Japanese English translation 7520 1RRI library] Proc Kanto-Tosan Plant Prot Soc 21192-197

127 Kushwaha K S 1981 The black beetle as a pest of rice in Haryana India Int Rice Res Newsl 6(4)14

128 Lawani S M 1982 A review of tile effects of various agronomic practices on cereal stem borers Trop Pest Manage 28266-276

129 Leuck D B 1966 Biology of the lesser cornstalk borer in South Georgia J Econ Entomol 59 797-801

130 lever R J A W 1955 Insect pests of the rice plant in Malaya Int Rice Comm Newsl 1612-21

131 Li Ching-Sing and Shin-Foon Chiu 1951 The study ofthe rice gall midge Iachydiplosis orvza Wood Mason [in Chinese with English summary] 1Taiwan Agric Res 21-13

132 Iitsinger J A 1984 Assesament of the need-based application for rice Proceedings TechnologyTransfer Workshop 15-16 March 1984 International Rice Research Institute 1PO Box 933 Manila Philippines 6 p

133 Litsinger J A 1984 Environmental diversity and insect pest abundance with reference to the Pacific Pages 285-307 in Proceedings sub-regional training course on methods of controlling diseases insects and other pests of plants in the South Pacific 4-20 Oct 1982 Vaini Kingdom of Tonga Nukualofa

134 Litsinger J A RF Apostol and M BObusan 1983 White grub Leucopholis irrorata (Coleoptera Scarabaeidae) pest status popula-tion dynamics and chemical conllol in arice-maize cropping pattern in the Philippines J Econ Entoinol 761133-1138

135 Litsinger J A C de la Cruz F Raymundo A T Barrion M D Lumaban J P Bandong M S Venugopal F Paragna and E Balete 1982 Insect pests and insecticide reponse of dry-seeded bunded rice Pages 179-193 in Report of workshop on cropping systems research in Asia International Rice Research Institute Los Bafos Philippine

136 Litsinger J A M 1) Luinaban J P Bandong A T Barrion W Katanyukul and N Panda 1978 Insect pest management in rainfcd lowland rice agroecosystms of tropical Asia International Rice Research Conference 17-21 April 1978 International Rice Research Institute Los Bafios Philippines

137 Litsinger J A and K Moody 1976 Integrated pest management in multiple cropping systems Pages 293-317 in R I Papendick P A Sanchez and G B Triplett eds Multiple cropping American Society of Agronomy Madison Wisconsin USA

138 Loevinsohn M E 1984 The ecology and control of rice pests in relation to the intensity and synchrony of cultivation Ph D thesis University of London 36) p

139 Loevinsohn M E J A Litsinger N Panda M A Austria R F Apostol C Andrion and J Bandong 1982 A basis for predicting rice insect pest abundance in cropping systems Paper presented at the 1982 International rice research conference on rice and cropping systems 19-23 April 1982 International Rice Research Institute Los Bafios Iaguna Philippines 25 p

140 Lopez A W 1930 Repot of the Entomology Department Pages 145-172 in Annual report of the Philippine Sugar Association 1929-30 Manila

141 Lopez AW 1931 Entomology Department Pages 227-266 in Annual report of the director or research 1930-31 Philippine Sugar Association Manila

142 Lopez A W 1932 Entomology Deparment Pages 252-279 in Annual report of the Philippine Sugar Association 1931-32 Manila

143 Lungibill P and G G Ainslee 1917 The lessercornstalk borer US Dep Agric Agric Res Serv Tech Bull 539 27 p

IRPS No 123 January 1987 39

144 McGuire J U Jr and BS Crandall 1967 Survey of insect pests and plant diseases ofselected food crops of Mexico Central America and Panama U SDepartment of Agriculture International Agriculshytural Development Service Agricultuial Researrh Service October 1967 11 p

145 McNeill Sand T R ESouthwood 1978 The role of nitrogen in the development of insectplant relations Pages 77-9 in lliochemical aspects of plant and animal coevolution J B Ilarbornu ed Academic Press London UK

146 Maebara H 1955 Spatial distribution of lastinstiarlanaeof purplish stem borer (Sesamia iqferens Walker) hibernating in an upland rice field Bull Kagoshima Univ Fac Agric 438-41

147 Mallick E H and N G Hajra 1977 Response of rice plant to defoliation treatments at different developmental stages Indian Agric 2151-56

148 Mansingh A 1971 Physiological classification of dormancies in insects Canl Entomol 103983-10)9

149 Martell C G 1974 Catalogue of plant feeding insects of Mexico [in Spanish] Fitofilo 27(69)1-176

150 Martinez 0 J 1982 Principal rice pests in Ilanos orientalis [in Spanish] P~ages 80-94 in Curso deact nalization de conocimientos en arroz Villaniencis Instituto (olonmbiano Agropecuario

151 Martins J F da Silva F Ferrcira A S Prabhu F 1 P Zimmermann 1980 Use of pesticides for the control of the main subterranean upland rice pests esqui Agropecu Bras 1553-u

152 Martins I F da Silva F Fe eira rod BdamSilveira Pinheiro I Q2 Damage simulation caused by defoliators of upland rice [in Portuguese with English summary] Pesqui Agropecu Bras 171113-1119

153 Martins 1F da Silva N(iyuen van Tan and 1Ida Silveira Pinheiro 1981 Stem borer resistance in upland rice associated with plant morphological characters [in Portuuce Iesqui Agropecu Bras 16187-192

154 Masaki S 1980 Sumnmer diapause Ann Rev Entomiol 251-25 155 Masuzawa T 11 Suwa and F Nakasoji 1983 l)ifkleences of

oviposition preferencc and survival rate of two skipper butterflies Parnara guttata and PIchlpidai iathiais (Iepidoptera HIesperiidac) on rice plant and cogon grass New Entomol 3247-56

156 Maurya D M and C P Vaish 1984 Upland rice in India Pages 69-92 in An overview oftpland rice research Proceedings ofthe 1982 Bouakamp Ivory Coast upland rice workshop International Rice Research Institute IO Box 933 Mnila Philippines

157 Menon M G R R N Katiyai S Kuar and S S Misra 1970 Termites attacking the paddy crop Sci Cult 36294-295

158 Miyahara Y and 1 Nishikubo 1967 Behavior of young nymphs of the red rice root aphid Iholahsipiun ri liabhdontinalisSasaki in relation to movement from the leaf to the roots of upland rice [in Japanese] Pioc Assoc Plant Prot Kyushu 1363-66

159 Mo ljoa Tjien 1952 White grub of upland rice (Holotrichiahelleri Brsk) [in Indonesian with English summary] Contrib Agric Res Stn Bogor No 13134 p

160 Mochida 0 and T Suryana 1979 The occurrence of the brown planthopper and ragged stunt on lowland and upland rice ill Central Java and logyakarta in tile late of January 1979 Entomnolgy Survey Report No 15 Central Research Institute of Agriculture (CRIA) Sukamandi Indonesia I February 1979 9 p

161Monet C 1979 Varietal resistance to upland rice pests [inI French] Pages 35-39 in Annual report 1977 Institut des Savanes Department des Cultures Vivrieres Bouak6 Ivory Coast

162 Monte 0 1942 A caterpillar ofrice-fields[in Portuguese] Biol6gico (Sao Paulo) 8161-163 (Rev Appl Entomol A31414)

163 Munroe I) D 1979 Pest management strategy for hill rice in four resettlement areas in Sarawak Malaysia Pages 89-102 i1 Guidelines for integrated control of rice insect pests Food and Agriculture Organization Plant Production and Protection Paper 14 Rome Italy 115 p

40 IRPS No 123 January 1987

164 Nakasuji F M Ishii and Jen-Zon Ito 1984 Rice skippers in Taiwan and their life histories ZZrC (Taipei) Tech Bull 82 10 p

165 Navas I) 1976 Fall armyworum in rice Pages 99-106 il Ecological animal control by habitat management Proc Tall Timbers Con-ference 28 Feb-I Mar 1974 Fall Timbers Res Stn Misc Publ 6 Tallahassee Horida USA

166 Nickel 1 1 1961 Biological control of rice stem borer A feasibility study Tcch Bull 2 International Rice Rcarch Institute PO Box 933 Manila Philippines IIl p

167 Nigan P M and R A Vernia 1985 Insect pests of uplaild rice in Uttar Pradesh Int Rice Res Newsl 10(6)22

168 Nilakhe S S 1985 -cological observations on spittle bugs with emphasis on their occirence in rice Pcsqtui Agropecu Bras 20407-414

169 Nixon G EJ 195 1 lie association of ats with aphids and coccids Commonwealth Iusti lcte of Entonology Iondon UK 36 p

170 Nurdin F an Ishak Manti 1985 Study oit seedling fly of upland rice Interlational Upland Rice Moitoring Tour Report 26 lebruary-2 MarLI Sukaranti West Sumatra

171 Oka 1 N 1979 Ciltural conttol of the brown planthopper Pages 357-369 in Brown planthopper threat to rice production in Asia International Rice Research Institutc 1p0 Box 933 Manila Philippines

172 Ohni M and A Vilnaii 1975 Rice insect pests of Italy biology of Sipha grcria (Kaltenbach) and its control [in Italian] Riso (Milan) 2459-71

173 Otanes F Q atnd 1 I Sison 194 1 Pests of rice Philipp J Agric 12211-259

174 Painter R I 1951 Insect icsistance in crop plants University Press of Kansits Lawrence Kansas USA 520 p

175 Pantua P C and 1A litsinger 1981 Arthropod abundance in continuous vs biannual rice cropping systeis Int Rice Res Ncwsl 6(2)20-21

176 Pathak M ) and V A l)yck 1975 Sttudics ott insect pests of upland rice Pages 186-191 in Major research in upland rice International Rice Research Institute PO Box 933 Ilanila Philippines

177 Patil B R B M IHasabe and 1) S Ajri 198 I Chemical control of white gritb lIditriihinhorraia Fabr intfesting groundntut paddy atnd sorghum Indian 1Entomol 43232-236

178 Pawar A 1) 1974 (ollection and identification of insect pests of rice rlerminal repirt lInternational Rice Research Institute PO Box 933 Manila Philippines 72 p (tnimco)

179 IerrajU A aid A S Reedy 1965 Observations ott diapausc itt tie steti borer ohrice Rice News Teller (lai-Mar)9-13

180 Pont A C 1972 A review of the Oriental species of theri ona Rondani (I)iptera Muscidac) of econtomic importance Pages 27-102 in Control of sorglium shoot fly M G lomtani and W R Young eds Oxford atd IH1 Ptthl Co New I)clhi India 324 p

181 Prasad S C 1PChand aid M 1PSingh 1983 Incidence of rice gall midge in the upland plateau region of Hihar Oryza 20235-236

182 Rai P S 198 0 Nisaga simpex damage to rice in the hill tracts of South India Int Rice kles News[ 5(I)12

183 Rathorc Y S 1969 Gonocplhahli dcprssunt Fabr as itpredator of (tilo zonwIlhis (Swinhoc) and other moths in field cage at Patnagar Uttar Pradesh Indian 1Entotnol 3127

184 Reed W 1983 Crop losses causcd by insect pists in the developing world Proceedings 10th Internatial Congress of Plant Protection lrighton UK 174-80

185 Reedy M V 1981 Ateactogasterinhens(Walker)a new root-grub pest of tipland paddy and inaize Curr Sci 501076

186 Reveno M A 1968 Catalogue of the insect pests of economic crops in Colombia [in Spaiish] Asociacion Latino Americana de Entomo-logia (AIAE) 129-33

187 Reyes G M 1957 Rice dwarf disease in the Philippines Food and Agriculture Organiiation Plant Prot Bull 617-19

188 Rice S E A A Grigarick and M 0 Way 1982 Effect of leaf and panicle feeding byarmyworm (Lepidoptera Noctuidae) larvae on rice grain yield Environ Entomol 7593-595

189 Risch S J 1) Andow and M A Altieri 1983 Agioccosystem diversity arnd pest control data tentative conclusions and nizw research dirctions Environ Entomol 12625-629

190 Rivera C T 1956 Three organic insecticides in thecontrol of insects affecting upland rice Philipp Agric 39465-472

191 Roffrey 1 1972 Locusta outbreaks in the Philippines Acrida 1177-188

192 Rojas R 1) 1941 The effect of pruning at varying ages upon the yield of Ramairio Phipp Agric 29851-860

193 Roonwal M i 1979 Ternite life and termite control in tropical South Asia Scientilic Pnbl odhpr India 139 p

194 Rosetto C1 F da Silva Martins N C Schmidt and L EAzzini 1978 Damage caused by pasturespittle bugs 4Dvoi)lavopictaandDJ schach) on rice lBragantia Nota 6 (Matcli)25-27

195 Rosztto C1- S S Neto I- Amantc I) Iink 1Garcia-Vieira D M de SoLza N V Bansa t and A M de Olivcira 1971 Rice pests of Brazil [in Portuguese] Pages 149-238 inrFA()-International Rice Commission 2nd meeting of the Rice Committee for the Americas 6-11 l)ecember 1971 Pelotas Rio Grande do Sul Brazil

196 Rothschild G H L 1969 Obseratiois on the armyworm Spodoptera matritia acronrctoiiles Gn (Lep Noctuidae) Bull Entomol Res 59143-160

197 Rothschild G 1- L 1974 Parasites of the armworm Spolopiera mauritiaacroncitoihes in Malaysia Fntornopliaga 19293-299

198 Sachan i N S K Gangwar and 1N Katujar 1980 Field crickets damaging cages in North Eastern IHill Region Indian J Entomol 42526-527

199 Sands 1) P A 1977 The biology and ecology of Lepocorisa (Hemiptera Alydidac) in Papua New Guinea Res Bull 18 Departshyment of Primary Industry Port Moresby Papua New Guinea 104 p

200 Sarawak 1968 titomology division Pages 97-106 in Report of the research branch of the Department of Altriculture 1966 Kuching (Rev AIppl Eintomol A56603)

201 Satpathy JM M S Das and K Naik 1977 Effect of multiple and mixed cropping on the incidence of scine important pestsI Entomol Res 178-85

202 Sauer IHF G 1939 Notes Ott Elasmpnlpualtsligniosellths Zeller (Lep Pyr) a serious pest ofcereals in Sao Paolo [in Portuguese] Arq Inst Biol 10199-206 (Rev Appl Entotnol A28378)

203 Satinders 1L A 1 SKing and C L Vargas S 1983 Crop pests of Central Anerica with t bibliography [in Spanish] Centro Agroshynotnico Tropical de Investigacion y Ensenanza Turrialba Costa Rica (CATIE) Tec Bol 9 90 p

204 Sauphanor B 1985 Some factors of upland rice tolerttce to stem borers in West Africa Inseu Sci Appl 6429434

205 Sen P atnd S Chakravorty 1976 Stentflics in Haritighata Nadia West Bengal India [it Rice Res Newsl 1(12) 15

206 Senapai B and 1M Satpathy 1983 The seedling fly Atherigona orrzae Malloch is a pest of rice in Orissa (India) J Entomol Res 777-79

207 Shhjahan M S Alatn M Ah nad and BN Islam 1983 A new scarabaeid beetle pest of dryland rice in Bangladesh Int Rice Res Ncwsl 8(I)16-17

208 Sharma G L S C Modgol and R C Gautam 1979 Effect of depth of submergence ott incidence of bacterial blight and white grub infestation in transplanted rice Int Rice Res Newsl 4(6)12-13

209 Sharma S 1985 Scarabaeid beetle outbreak itt upland rice Int Rice Res Newsl 1(1(l)24

210 Shiraki T 1917 The paddy borer Schoenobhits incertulas Walker [achaku Agric Exp Stn Formosa 256 p

211 Silva A de Brito and 11P Magalhaes 1981 Insect pests of rice [in Portuguese] Circular Tecnica No 22 EMBRAPA Centro de Pesquisa t ecuaria de Tropico Unido (CPAJU) Belem Brazil 14 p

212 Singh S A 1979 Emergence of rice leaf butterfly as a rice pest in Manipur India int Rice Res Newsl 4(4)16-17

213 SinghS A 1983 Changing trends of insect pestsof rice in Manipur Int Rice Res Newsl 8(f)12

214 Siwi S S and P H van Doesburg 1984 Lenovorisa Latreille in Indonesia (Heteroptera Coreidae Alydinae) Zool Meded (Leiden) 58117-129

215 Soejitno J 1977 Reaction of some upland rice varieties toseedling fly eltherikonaexigua Stein [in Indonesian] Siri Hana I Penyakit 630-35 Lembga I osat Fenclilian Pertanian Bogor Indonesia

216 Socnardi 1967 Insect pests of rice in Indonesia Pages675-683 in The major inse pests of thu rice plant Proceedings of a symposium at the Internatital Rice Rec Wi itt S pternber 1964 The Johns Hopkins Press Baltimore Maryland USA

217 Soto P E A T Petcz and I W Buddenhagen 1976 Survey of insect pests and diseases of rice in different ecological zones in Nigeria Rice Entomol Ncwsl 435-46

218 Soto 11E and Z Ziddiqi 1976 Screening for resistance to African rice insect pests Varietal improsetnent sCmi n r Bouak Ivory Coast 13-17 Sept 1976 West Africa Rice l)evelopment Association Monrovia Iiberia

219 Squire F A 1972 Entomological problems in Bolivia PANS (PestArtic News Sumr) 18249-268

220 Stapely 1 II 1978 The rice leaf roller in the Solomon Islands Int Rice Res Newsl 3(l)13

221 Suenaga H 1937 On aphids attacking the roots of rice Oyo-I)oubts Zasshi 9150-152 [in Japanese] (Rev Appl Entomol A25745)

222 Suenaga H 1952 On the seasonal abundance of three subterranean aphids on upland rice Bull Kyushu Agric Exp Sti 1249-262

223 Sutynso Yas and Dandi Soekaria 1964 Beberapa hana yang ttenperang tana man padi tanah kering [in Indonesian] Rapat kerja Padi Tanah Kering 6 sd 8 Juily 1964 Kaliurang 9 p

224 Tallamy I) W and R F Denno 1983 Alternative life history patterns in risky environments in example from lace bugs Pages 129-147 in Insect life history patterns R F l)cnio and If Dingle eds Spring-Verlag 225 p

225 Tamura 1 1939 Observations on some insect pests of field crops [in Japanese] 1 Plant Prot 26414-420 (Rev Appl EntomolA27-556)

226 Tamura and Suzuki 1964 Influences of the early-transplanting practice in rice culture upon development of the rice stem borer Chilo siql)ressalisWalker Bull Ilokuriku Natil Agric Exp Stn 761-94

227 Tan J P 1924 The rice root aphid Drowhi ihirsoia Philipp Agric 13277-288

228 Tanaka 1 1957 Taxonomy and distribution of some subterranean aphids injurious to upland rice in Japan with description of a new species [in Japanese English summary] Botyu-Kagaku 22168-176

229 Tanaka T 1961 The rice root aphids their ecology and control Spec [full College of Agriculture Utsunomiya University Japan F3 p

230 Tauber M IC A Tauber R Nechols and 1Obrycki 1983 Seasonal activity of parasitoids control by external internal and genetic factors Pages 87-108 itt I)iapause and life cycle strategies in insects V K Brown and I Ilodek eds )r W Junk The Hague Netherlands

231 Taylor W F 1972 Effects of artificial defoliation (simulating pest damage) on varieties of upland rice Exp Agric 879-83

232 feotia T P S and S Natud 1966 Biotoomics of the rice skipper Parnaraniathiis Fabricius (Lepidoptera Hesperiidac) Indian J Entomol 28181-186

233 Teran F 0 1971 Rice pests of Santa Cruz [in Spanish Ministerio de Agricultura Sita Cruz Bolivia Bull 45 8 p

234 Than Itun 1976 Population dynamics of the yellow rice borer Tryporza itcertulas (Walker) and its damage to the rice plant MS thesis University of the Philippines at Ios Bafios Laguna Philippines 92 p

235 Uichanco L B 1911 Pests Pages 55-74 it Rice Philipp Agric 11-200

IRPS No 123 January 1987 41

236 van der Laan P A 195 1 Life history of the rice seedling fly Contrib Gen Agric Res Stn Boor Indonesia 118 15 p

237 van der Laan 11A 1959 Correlation between rainfall in the dry season and the occurrence of the white rice borer (Scirpolhaga innotala WIk) in lava Entomol Fxp Appl 212-20

238 Van Driesche R G 1983 Meaning of percent parasitismin studies of insect parasitism Fnviron Entomol 121 I1- 122

239 Venugopal M S and A litsingcr 1984 Fffect ofcarbofuran on rice growth Prot Ecol 7313-317

240 Vietnam Directorate of Rural Affairs 1962 Annual work progress report on crop improvement program of rice sugarcane vegetable and field crops (lor the period from July 1961 to June 1962) (11) 299 p (Rev Appl Entomol A51526-527)

241 Wan M T K 1972 Observations on rice leaf- and planthoppers in Sarawak (Malaysian Borneo) Malay Agric 148308-335

242 West Africa Rice I)esclopnient Association 1978 Annual research report Vol 2 Statistics Monrovia Liberia

243 West Africa Rice Development Association 1979 Annual research report p47 Monrovia Liberia

244 Williams ) F and CS lofgren 1981 Fli I illy l-468 anew bait toxicant for control of the red imported lire at Fla Entomol 64472477

245 Williams I) 1 1970 The mealybugs OFlomoptera Coccoidea Iscudococcidac) of sugar-cane rice and sorghum Bull Entomol Res 0109-188

246 Williams D) 1 1982 Pulinaria icervi (Signoret) and its allies ott sugar cane and other grasses Btll Entoinol Res 72111-117

247 Wirjosuiardjo S 1976 Study ofchenmical cont rot of insect pests ott upland rice Imiu Pertanian (Jogjakarta) 1323-329

248 Wirjosuhardjo S and N Wirarta 1976 Chemical control of the white grub tIolorithiatlltriin upland rice in M uneng Irobolinggo Indonesia) [in Indonesian] Imut Pertanian (Jogjakarta) 1330-332

249 Yano K T Miyake and V F Eastop 1983 The biology and economic importance of rice aphids (Henuiptera Aphid idae) a review Bull Entomol Res 73539-566

250 Yelp D and P Rhodes 19(4 lhe success of drilled dryland rice iii the Fiji Islands International Rice Commission Working Party on Rice Production and Protection Manila Philippines 3-10 March1964 Food and Agriculture Organization Rome Italy 13 p

251 Young W R 1981 Fifty-five years of research on the sorghum shootfly Insect Sci Appi 23-9

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No 101The economics of hybrid rice production in (hina No 113 Boron toxicity in rice No 102 Rice ratooning No 114 Energy analysis rice production systems and rice research No 103 Growth and development of the (Jeep water rice plant No 115 Production risk and optimal fertilizer rates an application of the

No 104 Faridpur acomputer-assisted instruction nmodel forrainfed lowland random coefficient model No 116 Consumer demand for rice grain quality in Thailand Indonesia andrice

No 105 A readingand listening comprehcnsion test inEnglish for non native the Philippines speakers applying for tiraining at IRIRI No 117 Morphological changes in rice panicle development a review of

No 106 Rice grassy stunt virus 2 a new strain of rice grassy stunt in tire literature

Philippines No 118 IRRI-Korca collaborative project for the development of cold-No 107 Physical losses and quality deterioration in rice plstprodnCtion tolerant lines through anthei culture

systemis No 119 Problen soils as potenial areas for adverse soils-tolerant rice No 108 (olptblication ofIRRI materials a survey of translators and varieties in South and Ssuthcasi Asia

publishers No 120 Changes isinall-farm iLtireliing techii)hgy in Thailand and [he No 109 Classification of Philippine rainfall patterns Philippines No 110 Contributions of modern rice varieties to rutrition in Asia No 121 Landforms and modern rice varieties No Ill Changes in rice breeding in 10 Asian countries 1965-84 No 122 Yield stability and modern rice technology No 112 Design parameters affecting the performance of the IRRI-designed

axial-flow pump

Page 15: UPLANDRICE. INSECTl--1i M T -E!R.EOLOU t RTANCE ,AND …delphacid.s3.amazonaws.com › 6958.pdf · The significance of abiotic factors in upland rice insect ecology will be apparent

IRPS No 123 January 1987 15

similar species in upland and lowland rice Nephoiettix The principal vectors of rice virus diseases generally have species dominated The most prevalent species accounting narrow host ranges and alternate hosts in the grass family for 90-95 of the sweep net samples wre N virescens N are mostly annuals dying in the dr3 season Some virus umropictus Nilaparvia hgens Soatellafiircifera Cfana vectors such as BPH can disperse lontg distances but few do

sectraand Recilia dorsalisLowlands had higher popula- Th restriction of upland rice and weed hosts to the rainy tions Katanyukul and Chandaratat (120) monitored hopper season the small area planted to upland rice and the hopper numbers in upland and lowland fields of northern Thaiiand vectors having narrow host ranges all work against virus and found similar results A comparative study of plant- diseases in upland rice hoppers in Fiji showed more brown planthoppe BPH) and In Latin America Sogctdhs o(rzicola and S cuhanus whitebacked planthopper (WBPH) in wetland rice than in transmit hoja blanca virus disease but reports on upland lowland rice and hopperburn was prevalent in lowland rice are rare except in the favored upland areas of Colombia areas with standing water (89) Peru and Venezuela (35 42) S rricolaocurs in Brazil

Hopperburn in upland ricefields in Asia is rare (84) In on upland rice but hojaI blanca has not been reported (67) Batangas Philippines isolated fields of upland rice become S orzicola has a wide host ramgc (10) similar to WBPI- in hopperburned infrequently from WBPH A report from Fiji Asia WBPH however is not known to transmit any virus cites hopperburn from BPH and WBPH (250) In these two disease situations upland rice was grown on highly fertic soil and In Africa pale yellow mosaic virus is endemic to swampy N fertilizer was applied Hoppers respond to better nutrition lowland areas where rice ratoons year-round (19) and is Normally upland rice is grown on poor soil and receives no mechanically vectored mainly by hispid beetles A minor fertilizer vector a flea beetle Chaeiocnema spp is an upland rice

In Asia upland rice ras are normally not far from pest and upland ricetields near endemic areas occasionally lowland ricefields-- a source of hoppers Otherwise hopper contract the virus species with greater polyphagy than BPH or Nelhotettiv Spittle bugs Cercopid nymphs producc a protective virescens would be dominant frothy covering that looks like saliva In Latin America

In Africa leathoppers and planthoppers do not become spittle bugs damage upland rice In parts of Brazil where abundant on upland rice (6 II) probably because lowland upland ricefields are surrounded by pastures adults rice is not widely rlanted nearby The only reported migrating from grasses can damage rice severely and cause hopperburn as from Nilapanrata maeander in breeder complete hopperburn in a young rice crop within a week of plots receiving high N rates in Nigeria (98) Hoppers colonization (194 195) Dispersal to rice occurs even if however are a greter problem in Latin America where pastures are verdant and spittle bugs prefer pasture grasses large grassland areas breed polyphagous species (Gra- to rice (168) Nymphs are rarely abundant enough to phocephala spp Hortensia spp lEvitianus ohscurunervis damage rice Regular rainfall favors their development Balclutha spp and Draeculacephala spp) vhich then (233) disperse to cause hopperburn on seedling rice (68) Rosetto et al (195) reported Graphocep7hala spp and Hortensia spp Seed pests to be equally abundant on upland and lowland rice Alydid and pentatomid bugs are seed pests of upland rice

The leafliopper and planthopper epidemics in Asia that worldwide In Asia Leptocorisa sp prevail Their habits caused severe losses from hopperburn and vijus dicases make them particularly suitable for upland rice 1)thev feed over the past several decades occurred principally on on many grasses (and therefore can survive the earl y wet lowland rice Except in a few isolated cases (160) upland season before rice sets its grain) 2)adults are long-ied and rice has been spared from tungro grassy stunt and ragged mobile allowing them to find isolated plantings 0 rice stunt virus diseases vectorcd by green lcafhopper (GLH) (besides seed bugs only birds and rats appear to have shyand BPI] (123) Virus diseases have occurred in upland rice capability) and 3) adults aestivate in forested areas or where lowland rice was nearby Rice dwarf disease however sugarcane fields during the dry sca son when neither rice nor was first recorded on upland rice in the Philippines (187) grassy weeds are present (I199) Rice seed bugs concentrate Virus diseases can be perpetuated only on livir plants or on small-scale upland riceficlds because they can actively insect vectors and no living rice - plans tooned or search them out The prevalence of groves of trees characshyvolunteer - grows year-round in upland arcas excert in a teristic of upland environments ensures nearby aostivation few with 12 mo of rainfall sites keeping the seed bugs close by

The smaller brown planthopper Laodcelphax striatellu Leptocorisae are uncommon in lowland rice plains occurs in upland rice in Japan (18) It spreads black- because 1) their populations are diluted in a sea of rice streaked dwarf and stripe viruses between winter wheat and planted more or less at the same time (they ca n only feed on barley to lowland rice Early planting of lowland rice rice during the milk to hard dough stages) 2) they lack nurseries spreads these viruses but upland ice has remained aestivation sites and 3) weedy fallow areas are limited uninfected The lowands of Asia are dominated by L oratorius

16 IRPS No 123 January 1987

which lays its eggs hign on tile foliage (199) L acuta and L solonlonlesis oviposit at ground level the former on litter and the latter loose on the ground These egg laying habis explain theirtenvironmental prefcrencecs(214) In Japan the postdiapause adults of L chinensis migrate from moun-tainous areas to upland ricefields during floweiing (1 14)

Pentatornid bugs prevail in Latin America (8182 195) and in Africa pentatomid and coreid bugs arc equally cited (6 82) l)amage is often characterstic of the species (5)rcoris ventralis and Mnida spp are the common

pentatomid seed bugs of Philippine upland rice but they prefer lEchinochva spp grass seed to rice Vearaviritda occurs worldwide and hoften more abundant in the uplands(54) We saw few reports of seed bugs on upland rice but see(d bugs apparently opelrate cqually between by land and upland rice Good dispersal is necessary ior species that call feed for only 2-3 xon a crop and must first find those crops at ripening stages

Other pct species feed n flowers In Mato Grosso Brazil (194) and Santa Cruz Bolivia the panicle weevil Aeo-hariuia amllitarsis attacks rice during flowering (68 233)The larva e rice sten horers adutlts feed at the bases of spikelets caIusing empty grains Also inLatin America thecydnid -lkiituttatralusisa seed bug as anad tilt and nymph (194)

A number of beetles ecd on poilen - coccinellids lialrotia Aulac lhora and Monohpl-a (55) - but becatise riCe is self-pollinating and fcrtiliation occurs before pikelets open they pose no threat to yield

IST ECOLOGY

We now look at characteiistics of the life cycles of upland rice insect pests to lea rn hew each plays an adaptive role in upland rice ecology

Life history strategies Orzasativadoes not tolerate drodgh well so upland rice is highly seasonal normally grown in the wettest months of the year Usually rice is present for less than half of the year The rice-fiee fallow poses serious problems of survival to upland rice insects which have evolved att least four mechanisms to ovrcolne tle cvcl ic-l lack of a hostI) polyphagy 2) longevity 3) dormancy and 4) vagility (dispersal)

Poldt)agy lolyphagy is defined as having hosts of at least two botanical families oligophagy of more than one gents and ionophagy of only one ventus (40) - inour case Orvza Most wild rices however are illi tiC and theieiirc are normally far removed from upland rice hahitat othewise more would be alternate hosts for uplanJ iice insects Thte highly mornophagous rice pests shySciriol)haita hiceritlas Nilal)arvaiahiens Iand Nealo--tettiyvircsctws in Asia and Aalirpha secparatella and 1ioj)sis lngicornis inAfrica - occur on upland rice but

are more abundant in lowland culture They can specialize in rice because they are highly vagile and can attack all growth stages By specializing in rice monophagous rice pests can outcompete related but polyphagous speciesIntensive andI extensive lowland rice culture has favored monophagous species Riqwla alhinella (Cramer) the onlymonophagous species in Latin America is rarely reported on upland ice (I1)

Because of its limited temporal and spatial existence upland rice has not favored the evolution of specialized species Clhang(44)shows evidence that upland rice has only recently been cultured by man from the lowlands thus pests have had little time to fully adapt to the crop All but the lowland-adapted insect pest species have alternate planthosts to rice Root pests with limited mobility are highly polyphagous Pests that specialize in one growth stage eg secdling maggots seed bugs are oligophagous or polyphagous

Most alternat hosts of rice pests are annual grasseswhich also are seasonal Upla_rice is also a host but not the preferred one of the most highly polypliagous spcciessoil pests grass hoppers armvwarrs and Vearaviirielda

The d ry season poses igreat obstacle to ani upland riceinsects survival and only sonic perennial plant hosts are suitable food during the off-seasoin Plolyphagy allows upland rice insects to suLrx-ixe during thC rainy season when plants- both annual and perennial -areactivelygrowing and therefore more nutritious

Loiugevitr The ability of aiinsect to live 2-3 mo without undergoing dormancy helps species sich as root-feedingwhite grbs (as larvae) and secd-feeding bugs (as adtilt) to survive unfavorable periods White grubs arc not highly fecund and their life strategy is to improve survivorship of limited progeny Their subterranean habitat hides them from manv natural enemies lBtut root tissue is not highly nutritious so they ltilst eat great quantities and need a largeinsect biomass to digest it Other soil-inhabiting rootshyfeeding insects have short life cycles but are more fecund

Seed pests need greater longcvi vas adults to locate ahost at early grain dcclopment arnd give sedentary offspring time to develop Few groups of upland rice insects live longin active development Polyphagy and longevity only increase their ability to survive during the rainy seasonOther mechanisms are needed to survive the dry season

orantc In the tropics the nonrice season is the dry season while intemperate areas tile off-+season is winter Many upland rice insects undergo dormancy inlsummer (aestivation) or winter (hibernation) Dormancy is simplyinactivity during winters with prolonged temperatures below 10 C insects find shelter arnd cease movement Activity is resu med with warmer t empcera tu res A deep state of irnactivity termed diapauise occurs in sole species that prepare physiologically (havxing low metabolism and convcrting gonad s to fatty fuel reserves) for the unfavorable season (148) They enter diapaUse inanticipation of unshy

IRPS No 123 January 1987 17

favorable weather and resume activity after favorable Armyworms skippers and locusts actively travel hundreds weather begins usually in response to photoperiod or of kilometers in air fronts such as intertropical convergencethrough a delayed response to a stimulus such as heavy zones to descend onto upland rice fields far from their rainfall (237) Not all individuals of a population have the breeding grounds (31) Several generations can build ip on same threshold for environmental responses for entering or alternate hosts in tile grass family before rice is plantedterminating diapause -- this is seen as a mechanism to These large insects are active fliers Smaller insects such as prevent total mortality if a false cue appears such as a thrips aphids and planthoppers passively migrate long sudden warm spell in winter or rain in the middle of the dry distances by the wind season (154) Otherspecies ean readily travel tens of kilometers - stem

Probably more upland rice insects undL go dormancy borers leaffolders leafhoppes cutworms spittle bugs leaf than we realize Candidates are Atherigona spp root beetles seed bugs ants and termites aphids root weevils crickets leaf beetles thrips and seed The least dispersive species are white grabs mealybugSbugs Insects known to entervarious states of aestivation are seedling maggots crickets weevils and gall midgesleptocorisaspp adults (199) Maliarphascparatellalarvae Each species has evolved a unique set of attributes to (9) Scirpophaga innotata larvae (117) Chilo partelhtsv enable it to survive and adapt to the changing uplandlarvae (21) ScirpopThagainccetulas (179) white grub larvae environments Several evolutionary avenues lead to fitness (134) Patangasuccincia adults (6 82) and Schistocerca (Table 2) but there is no reason to believe one set of gregariaeggs (82) attributes is better than another (224)

Insects known to enter hibernation are LodeIphaDstriatellus nymphs (124) Chtio Tvlpressalis larivae (125) DroughtSesaia inferens larvae (146) but not S calavisis(2) Well-drained rainfed upland soils are subject to drought ifIDiaraea saccharalis larvae (75) Scirpophaga- tcerlttas rains do not fall within 2-3 wk Prolonged drought followedlarae(5 lacchrli s lae (75)saSlOhagasshoperseggs (82)and felo pidas athias pOpae (232) by favorable rains stimulates armyworm and locust outshybreaks but mealybtigs root aphids and thrips become

Lcpmotorisa oratorisand L acuta adults aggregate on numerous onlyafter dry spells for two reasons First heavy trees sugarcane or other shady moist sites L paluwanmsis rain normally kills soft-bodied foliage-feeding thrips and confined to Sulu and Palawan Philippines lives on the mealybugs and soil-inhabiting root aphids and root mealyshygrass Brachiariamutica (Forsk) Stapf in the absence of bugs Drought eliminates this source of mortality Second rice The adults can live for 5 mo When aestivating their the rice plant responds to drought by breaking down rate of metabolism drops and they do not feed They will proteins into soluble N compounds which enter the phloem however puncture plant tissues of their aestivation host in and are taken up by these sap-feeding insects (245) Greater search of water This behavior has led to errors of the host nutrition therefore leads to greater survivorship and range of rice bugs (I) Aestivation quickly terminates with fecundity rain probably meaning aestivation is not deep Small rice area

Some stem borers aestivate as mature larvae in rice Except in the highly mechanized large landholdings of stubble or in straw stacked as livestock feed However Latin America upland ricefieldstypically are patchy within moriality increases proportionally with the duration of the a highly diverse flora The small fields result in higher ratios dry season even if the stubble remains undisturbed If the of perimeter to area than with the typically larger lowland stubble is plowed for a crop following rice few stem borers ricefields Upland ricefields themselves may be intercroppedsurvive Rainfall terminates dormancy but it normally with cereals legumes or root crops Mixed intercroppingtakes several weeks for moths to develop and emerge with a wide variety of species also occ in tribal slash-andshy

Last-instar white grub larvae tunnel 1-2 m deep in the soil burn agriculture (50 74) to construct pupal chambers in which they aestivate The Upland ricefields tend to be small because of the labor larvae are very sensitive to dryness so pupal chambers are needed to clear land in slash-and-burn areas and to till land sealed to coierve body moisture Aestivation terminates to minimize weeds Rice requires more tillage than does with the first heavy rains but as in stem borer 3-4 wk pass maize because rice competes less well with weeds Uplandbefore adults develop rice is more a subsistence than a cash crop and small areas

Insects may respond differently in each region to cold (I ha) can feed a family temperature or drought Biotypes or loal populations may Small fields favor some insect pests such as some ant evolve (56) Local populations of Scirpophagaincertulasor species that can forage from more permanent field borders Patangasuccincta may have become adapted to prevailing Trees provide food and mating sites for white grubs (134) as conditions well as aestivation sites for LetTocorisa spp (199) Field

iagiliti The ability to disperse combined with high border grasses plus tIhose within a ricefield provide alternate fecundity orshort life cycles enable some upland rice pests to food for all but the highly monophagous upland rice better exploit the temporarily favorable upland habitat colonizers and a habitat for natural enemies

0 z

Table 2 Life history patterns of upland rice insects or insect groups

Host range Longevity Dormancy Vagility FertilityMonophagy Oligophagy Polyphagy Low Medium High Yes No Low Dispersive Migratory Low Medium High(Oryza spp) (grasses sedges) (angiosperms) (lt3 wk) (gt2 mo) (lt1 kin) (lt50 eggs) (gt500 eggs)

Ants xTermites x x x

Mole crickets x x x xx x xField crickets x xx

White grubs x x x xx xRoot aphids x x xxMealybugs x

x x x xxSeedling maggots x

x x x x xLeaf beetles

x xx xArmyworms x x xx x

Thrips x x xx xChilo spp x x xx xMaliarpha x x x x

xLocusts x x Xx x xGall midges x x xxiVephotettL iirescens x x xx x x xLeprocorisaspp x

x xLeaftolders x x x

Sogatellafurcifera x

xx x x x x x x

00

IRPS No 123 January 1Q87 19

All this appeals to favor insect pest buildup but the notwithstanding the release of stored nutrients in response contrary is true Loevinsohn (138) showed that insect pest to drought that temporarily benefits philoem feeders populations respond exponentially with the proportion of land devoted to rice up to about 75 of the area then the M POR ANCE I I P1A 1) RWI P1St S

rate of response declines In most upland rico areas tie rice crop occupies less than 50i of the area- therefore the Visits to upland ricefields normally reveal 1kw insect pests potential for population buildup is low This relationship Loevinsohn (138) and Locvinsohn ct al (139) compared may not hold for seed pests that can locate small ricefields insect abundance using annual light trap atchcs in upland

Another outcome of the highly diverse flora and phical rainfed lowland and irrigated lowland sites in the Philipshyenvironment of upland rice isthateach upland ricearca has pines Although only one upland site was studied data a unique composition of insect pests showed equal or fewer insects at the upland site than at the

For example in Tanauan Batangas Philippines the rainfcd lowland sites and fewer than at the irrigated sites main upland rice pests are rice leaffolder(RLF) armvworm (Table 3) M Lltiple regression analysis showed that the key Aythntnaseparata WBPH ants Soli psis gentitataand factors explaining insect abundance among a set of cropping white grub Leucopholis irrorata Sixty kilometers away in intensity variables were the number of rice crops grown per Real Quezon the main pests are seedling maggot Athe- year followed by area devoted to rice Table 3 shows that rigona orae flea beetle (Thacitcmnra hasalis thrips the upland rice site had only I rice crop perym rand that rice Stenchaetothrips hiurnmis and rice bug Leptocorisa was planted in only 20Vi of the area Fertiliter and oratorius In a third Philippine upland rice site in Claveria insecticide use had little bea ring on the trend The armters at Misamis Oriental Mindanao the main pests are ants the upland site applied 60 kg N ha higher than at most Solenopsis gemtinata seedling maggot Alieriona orvzae irrigated sites white grubs Holotrichia in)iUaiuoan t and Lucopholiv Insect occurrence at the upland site was omparable to irrorata stem borers Sesamia ifiArelr (hio auriciliusC that at the Cagayan rainfed lowland site parlicularly in 1982stpessallis Scirpophaga incertulas S innoiata and when drought prevented many farmers fron planting(only

4c ona chrrsurapella root aphids Tetraneura niriah- 40(of the rice area eventually was planted that year) M ore dunrinalisand Rhupaosiphunm riahbdminalis lcaffolders favorable weather occurred in 1980 and 1981 which Marasniapataltis and Chaphalocrocis Medinalis army- supported greater rice areas and consequently more insects worm MIthinna separata and rice bug Leltucorisa The most intensive site for insect pests was South Cotabato oratoriusThe great diversity of insect problems is typical Mindanao where farmers plant 23-25 rice crops per year fer upland rice and makes control efforts more difficult with irrigation The Zaragoa Nutva Ecija site also had particularly regarding breeding for insect resistance high pest abundance and at this site the fields were highly

asynchronous Asynchrony was the third most important Low yield potential Iactor in explaining rice insect abundance Most upland rice is grown on low-nutrient and mineral- Rice cropping intensity in upland areas is bound to toxic soils which when combined with eiratic rainfall remain low Only one rainid upland crop is possible per make upland rice a highly riskycrop for the farmerto invest year (Farmers in Claveria Misamis Oriental Mindanao in costly soil amendments and land management Often Philippines tried asecond upland ricecrop i 1984a ycarof upland rice isgrown far from markets making inputs even favorable rains but no harvest was possible b-cause of more expensive Farmers growing a subsistence crop hosi- drought blast and birds) Rice area isaso bound to remain tate to invest in costly inputs to raise yields low because of the high labor and power rcquirelmenlts to

Even having varieties that could double existing yield prepare the land (because of weeds)and because upland rice potential under present management levels would probably wvill not compete with lowland rice as a cash crop Some not prompt many farmers to use purchased insecticide upland rice sites with prolonged rainy seasons howeverare unless tile risk of crop failure were reduced such as it was planted asynchronously such as the Clacra site with irrigation systems in lowland rice The seasonal abundance of Upland rice insect pests

The Tanauan Batangas site is atypical for upland rice attracted toa light trap wasgrapied for crop year 1980-81 in The soils and rainfall pattern are favorable and because of Tanauan FBatangas (Fig I 2) W13P11 was he most its nearness to Manila farmers have cash resources ftm common rice hopper and its population peaked at the sales of vegetables to purchase tertilier They applied an midgrowth stage as it does in lowland rice D-Vac suction average of 60 kg Nha after particle initiation and obtained samples showed an earlier peak than that from a light trap 25-35 tha yields The high fertility isone reason why RLF indicating emigration W13PI usually emigrate after the armyworms and WBPH developed into large numbers vegetative stage Populations in 1980 reached levels that Farmers however used no insecticide on upland rice caused patches of slight yellowing in some fields WBPH although they did on vegetables probably cause hopperburn in Batangas They immigrate

The poor growing conditions fOr rice generally mean year-round as shown by low levels in the light trap BPH poor growth and low fecundity for insect pests fIeding on it however were scarce in the light trap and not collected with

Table 3 Abundance of rice insect pests and factors in rice cropping intensity of Philippine upland rainfed lowland and irrigated lowland locations where light traps were operated daily for atleast I yr -I

0Rainfed Rainfed lowland Irrigated lowlandupland

Pangasinan Cagayan lloilo Santa Maria Nueva Ecija South Cotabato Tanauan Caaringayan Caaringayan Bangag Bangag Bangag Buray

LagunaCale

Batangas Manaoag 1980-81 Santa Motiica 1982-83 Cabanatuan Zaragoza Avancena NaniamanSolana 1981-82 1982-83 Oton

1980-81 1979-80 Oton 1981-82 1981-82 Koronadal Koronadal1980-81 1979-80 1979-80 1983 1983

Insect (kerosene light trapNephotcttixcatch nolyr)virescens 650 1240 500 1800 330 210 3400 1600 160N nigropictus 210 - 360 18400 5500 2900- 140 90 20 - shy 1700 -Reciliadorsalis 710 - 3200 1700180 890 630 140 170 2400 800Nilaparratalugens 130 690 - - 4200 4600300 520 750 180 60 3300 1600 740 310Sogatellafiircifera 1700 31000 13000 8200310 210 3300 610 270 980 1600 6300 -Cyrrorhinus - 12000 6700600 540 3200 290 160 330 120 410Scirpophaga spp 50 410 620 - - 21000 12000150 180 220 270 30 1000Cnaphalocrocismedinalis 70 70 540 610 270 11000 6100200 140 100 6 60 250Chilo suppressalis 0 60 60 1900 300 200- - 30 20 0 6 3 160 -C auricilius - 0 00 - - 20 3 60 20 3 0Sesania inferens 0 - 80 60 3 80 10 51 -Cropping intensity _ -

Rice crops (noyr) 10 10 12 10 10 10 13Rice area (7c) 17 19 20 20 25 2320 85 85 60 70 40 85 85 90Fertilizer (kg Nha) 60 20 85 80 70 7020 0 0 0 30 30Insecticide (no of 40 60 40 300 04 04 0 03 300 13 13 30 50 40 33 33applicationscrop)

__________

IRPS No 123 January 1987 21

Light trapped Insects Drytnd __ 2(notrap-nght) Suctioned Insects (nom )

50 -)o 9 jO40- SOW ffurciWO Light trp

D-Vc suction

20 to 4

20

Egg porasitizahion (M) 75 60

300 SOOampReOftwciffr

45 30shy

15

-insects (notrop-night) 4 Nileporvata Iuen5

Egg parastization()

Light trapped insects (no trop-nght) Suchoned insects (no m 2 )

6 8 Cytorh~nuls hvidipennii 0

4 05 2

50 0 Insects (noltrop-night)40-ui Ipanh pe g

Moohase 1 Seasonal abundance measured by light trap

Rainfall(etod) and D-Vac suction sampler of whitebacked

to ~ hopper Nilaparvata lugens and lea ffolder60)-t Cnaphalocrocis mnedinalis andplthpe g20 L I II I I I parasites and mtrid predator Ortorhjntus livi-Apr May Jun Jul Aug Sep Oct Nov Dec JaIn Feb Mar dipennis set against cropping pattern daily1 9 80o 19 8 1rainfall and moon phase Tanauan Batangas

1980-81

the D- Vac Cyrtorhinus build up on the crop coterminous are collected in light traps during full moon (109)with WBPH buildup Cyrtorhinus disperses during the Because upland rice shares most of the lowland rice pestsrainy season RLF not readily collected in light traps was and those pest species are generally less impotant to uplandprevalent only during the rice crop rice than lowland rice there is a general belief that insect

Nephotettix virescens was more abundant than Nnigro- pests are less important onpicLus in the light trap collections and on the crop GLH upland than lowland rice Arcview of limited data on upland rice shows an averag of population peaked at the reproductive stage as did zigzag 10-21 yield loss from insect pests on a par with that ofleafhopper (ZLH) Crtorhinus tracked the populations of lowland uee determined by the same methodology (132) planthoppers and leafhoppers Both GLH species were (Table 4) McGuire and Crandall (144) made a similarcollected year-round in the light trap indicating immigra- estimate for Central America Cramer (51) howevertion from nearby irrigated rice areas Possibly because of the estimated 35 yieldlow populations loss in South America based on ainsect number did not correlate with similar pest complex - stem borers and seed bugs shymoon phase although reports indicate that more hoppers attacking irrigated rice in North America This estimate did

22 IRPS No 123 January 1987

Light trapped insects D~~nd (no trap-night )

St plht Suctined insects (nom 2

)

D-Voc suction 1520

1o

0 Lih trap 5 ~105 o5

00 Eggparositizotion()5040 NephOtgtha virneens

30 20

I010 A-Insects (notrop -night)

4 Nephatelh nigropccent i

2 o I vV^J - -- ^ VA____ f _

Light trapped insects (no trap-night) Suctioned insects (no rnE

10 -1208 ReCdO doRsol

6 15

4 10 052 iwxALAiamp 0

Light trapped insects (notrap-night) 2Suc oned nsects (nom )

8 CyrtorhmnusI6o nn]10 i0075

4 05

2 0 0

025

2 Seasonal abundance measured by light trap Roinfall (mmd) Moon phase and D-Vac suction sampler of green leafhoppersNephotettix virescens and N ngropicrus and I zigzag leafhopper Recilia dorsalis their egg 60

parasites and micid predator Cyrtorhinus livi-dipennis set against cropping pattern dailyrainfall and moon phase Tanauan Batangas1980-81

40

5 Lf Apr May Jun

1980 Jul Aug Sp Oct Nov I

-aMar

1981

not account for soil pests and was not based on field data in the absence of pests Carbofuran granules are phytotonicAkinsola (I1) cited mainly data from irrigated rice areas in to rice (239) The prevailing hypothesis is that carbofuranAfrica but estimated yield losses of upland rice between 15 prevents soil bacteria from consuming fertilizer The phytoshyand 30 from insects However losses in Thailand over a tonic effect is through greater availability of fertilizer to the4-yr period ranged from I to 13 averaging 5(106) plant Denitrifying bacteria are known to be more active inYield loss data however should be interpreted with aerobic than anaerobic soils therefore it may not be merecaution (184) The Philippine data were all derived from coincidence that the highest recorded yield losses (22-69)trials on farmers fields under farmers normal agronomic occurred in those trials with 12-30 kg basal Nha On themanagement practices except for Bukidnon and Capiz other hand contro with carbofuran granules may not be aswhere researchers used fertilizer Batangas farmers are good as a seed treatment against seed pests and perhapsatypical because ofhigh levels of fertilizer used The trials in yields would have been higher if a systemic seed treatmentBrazil were carried out at an experiment station but insecticide had been used conditions were similar to those of local farmers The 1979 and 1980 trials in Batangas had two full-

Carbofuran granules were used in every trial as a broad- protection treatments one using carbofuran granules andspectrum insecticide applied basally with the seed An the other using bendiocarb WP as a seed treatment Plotsobjection to using insecticides to measure yield losses isthat protected with bendiocarb yielded higher than those proshyinsecticidescandirectlyorindirectlystimulateplantgrowth tected with carbofuran granules The other trials in the

Table 4 Yield losses to insect pests and yield responses to insecticide in upland rice in the Philippines Brazil and Upper Volta

Yield (tha) Yield loss (5) Location Year Cultivar Protected

with Protected

without Check Total Vegetative Reproductive Ripening

Basal N (kgha) Reference

carbofuran carbofuran Protected No with carbofuran

carbofuran

Philippines Pili Camarines Sur Tanauan Batangas

Pangantucan Bukidnon

Dumarao t-apiz

BrazilGoias Minas Ge~ais

Upper Volta6

1975 1974 1974 1975 1975 1976 1977 1978

1979

1980

1979 1980 1979

1977 1978 1979

Bursiging puti Kinanda Dagge Kinanda Dagge Dagge Dagge Dagge UPL Ri5 Dagge UPL Ri5 Dagge UPL RiS UPL Ri5 UPL Ri5 UPL RiS

IAC47 IAC47 IAC47

14 33 11 36 35 31 30 27 37 26 40 30 40 42 46 35

19 21 09

--

-34 24 24 27 --

29 43 31 42 -

47 25

---

10 28 05 27 27 30 26 29 33 28 33 29 43 23 36 11

15 15 06

29 15 55 25 23

3 13 -7 11

-7 18

3 -7 45 22 69

24 28 35

--

21 -11 -20

4 -

-3

23 7

-2 -23 56

-

-

-__0

6 --

-

-23 14

5 0 8 2

22

-

--

0

-

--18 11

3 -10

4 10

7

--7 -

14 -

-1 6

10 -5

8 9 7

-

--

0 0 0 0 0 0 0 0 0 0 0

30 30 30

12 12 12

99 99

100 100 101 101 102 102 102 102 103 103 104 104 104

66 66 66

Farako-Ba

Mean

1977 1978

IRAT10 IRATIO

38 66

--

40 43

-5 35

21

-

10

--

--

--

- 242 243

Z

tJ

-w

24 IRPS No 123 January 1987

Philippines which had two treatments offering protection In the 9 Philippine trials where yield losses were measuredwith and without carbofuran granules relied on further by growth stage 10 of the yield loss (21) occurred at thecontrol with foliar spray and not on seed dressings The vegetative stage and 517 occurred at each of the reproshycarbofuran-protected plots had an 18yield los compared ductive and ripening stages Ibis result is similar to thewith only 10 with foliar sprayS This yield difference is pattern of yield loss in lowland rice measured by successiveprobably due to differences in protection against sown-seed treatments where the crop is unprotected at each growthpests (foliar spray do not control sown-seed pests) rather stage but proticted during the other growth stagesthan phytotonic effects became basal fertilizer wa used in However the insect complexes r sponsible for vegetativeonly 2 of the 10 comparisons stage yield losses in upland rice are entirely diftfrent than in

The recorded yield losses were not related to yield lowland rice The key vegetative stage pests in lowland ricepotential (Fig 3) One might expect higheryielding crops to arc aquatic whereas those in upland rice are sown--seedhave higher percentage of yield losses but this ws not tile (sown in soil) root and seedling pests This dift rence maycase One reason is that insect damage exacerbates plant explain why agricul torists have overlooked tile ilmportanccinjury from drought stress low yielding fields from d ought of upland rice insect pests Many reports of upland rice pestsstress would show disproportionate differences between in Asia ha Ve focused on major lowland species (46 108)insect-protected and -unprotected treatments Shoot and It isalso surprising that tei vegctative stage of upland rice root pests would be more responsible for losses from would record the highest yield losses becatuseseeral studiesdrought stress than sown-scd pests A second reason have shown that upland rice can readily recover from foliageconcerns seed pests which would have a greater impact on reinoval during that stage (117 152 165 188) Upland ricelow tillering low yielding varieties such as )aggc Low poscsses a hitgh obilitV to compensate for ca rly loss oftillering varieties nnot fill in the space created by removal foliage and will even be stinulated by foliage removal toof seeds oweer 1) R i 5 is high yielding because it tillersP1 produce a higher viel than plants ithout any foliageactively which can close the canopy to provide greater weed removed (63 192 231) [ie reason for this apparentcontrol discrepancy hovce is that yield loss at the vegetative

stage does not octUr frol leat arca loss but fron pestsYield loss () removing sown seed and feeding on devcloping tillers and roots -- damage that cannot be readily conpLtnsatCd for

70 - Measuring yield losses in upland rice cnvironrnents is further complicated by a report from Brazil that greater yield losses occurred in response to better insect control60 - =-09 as Insecticide seed treatments resuled in denser plant stands

(n=21) which in turn created a selectively more favorable environshyment for blast (69) This resuIt was parlticurla rlxexacerbated50 when cirbofuran granules were used in seed furrows Aside

ffrom a greater plant stand ca rbofuran iay have auigmented the N lecl and stimulated blast If so ca rbofuran should be40 - replaced with another insecticide or the seeding or fertilizerI rates decreased

Other methods to determine the impact of upland rice30 shy pests wvcrc to interview farners or to develop single speciesbull bull correlations of population levels and yield loss Feneira (66)2 NIM)st (75(1) Golas farmers reporteddid both in Brazil

20- insect losses in ufpland rice and 251 said those lossesS avcragcd 37(1 [his result is consistent with tie yield losses 0 0 measured by field trials in the same statc(Table4) For every

I0 4 of seedlings removC d by leaf-cutting ants a I 7yield loss is predicted Similarly Ferreira (66) found that ai

0 - average of 53 thrips per panicle before panicle initiation meant 3 times more unfillh d grains

-IO0 I 1 1 I I-101 0 2 3 4 5 6 7

Yield (tho) C)N [ROt IM FI IODS 3 Correlation between upland rice yield and yield lols caused by Upland rice insect pests may be controlled by culturalinsects measured in 21 trials in the Philippines Ivory Coast andBrazil genetic biological and chemical methods

IRPS No 123 January 1987 25

Cultural control dry season plowing will have little effect on them Plowing Crop husbandry to reduce upland insect pest populations when the soil is wet from the first monsoon rains will expose can be classified into practices effective in a single field and soil insects that tunnel close to the soil surface to predatory those effective only at the community level Cultural birds chickens dogs and even man (field crickets nole controls should come first in a pest management program crickets and white grubs a re delicacies in man tra ditional They have broad stable effects as pests have little pos- diets) sibility of overcoming them through selection of biotyps The greatest degree i1 inIscct control fron tillage comes (138) On the other hand some cultu ral controls work only from plowing soon after rice ha rvest when most soil insects with high labor or power inputs and connunity-wilt lie close to the soil surface anrd stern borers are in the rice methods require coordination of iany farniters stuble Whitegrubsare mature and tillage exposesthem to

Pantin time The luxury of being able to shift the time of predators Tirnelv control will prtec the crop following pla ntring upland rice is open onl hr la rniers in areas with a rice and lowr the populationIfor succeedinig years Plowingprolonged rainifall pa ttern lI copping intensity or underthe ice stulble helps it decompose killing sten borer rnechanied land preparatiot Gcnerally planlting as soon larvae inside as the rFiinv season begins will lower populatiois of 1110st [requent tillage destroys ait nests bit not termites insect pests Insect populationsarelo aftcra dlvseoll or Tillage and clean culture howes er rcliove food for winter fallow delayed plaiting lets them build tip on termites which then mtay attack a young rice crop Scarab alIternate hosts that gi vigorotsly with the first seasonial beetles also prefer to miposit in eceitly tilled fields Noshyratiris This first flush iof weds cros lItxuttriartlv with iceCult is dibbling faors the survival oftillace Clre Such rineralied N released during the dr season aid made stils i aels tnd insects thtt pttpate or lay eggs- in the soil

available to the plants with the rains It iliso igrtmitcally Zero tilla hio eerl conserves predators ofthe GIH (37) advanttgeous for the rice ciop to tap this trlLItlaI fertility nt it mcttoi lo lessen the impact o0 soil insects and grow vigorotsly to compete with wrus and to tolerate brotdcasting seed or planting in ftltrrows is prefc ired over pest infestation dihlirig in hills Crickets are large enotlgh to eat seedlings in

Larly plantngs terd to escape seedling maggolts Upland a hill atrd tillering normnally ca iriot fill in tlhe missing space rice in Bitangas is planted in Ma and ine itli the Iirst Oftcn dibbll rd is not well covered and iseasily found by rains Fairners are highly mtotiated to plant early tocscape foraging insets ioIdents and birds drlitght and typlotns ricar harct Arn Atrgtst planting Plan uit sii Incrcatsini sced density protects tile cropwould be evcely attacked by It riim orivzo At tti againrst seed and seelling pests especially ants particularly other Philippiie tipland sites Rea l Qteot antd (laelit if the crop will germinate and ermerge quickly M isainis Oriental seedlig mnaggots arnrige ric boe t se hr trftinll itI India rice intcrclopped witi cotton or farnes have a longer growing season and plnit 2 rin or pigeon pci had lower 6IIt and WBIlII poptrlations than mote after the ortset of the rainy season liaaiangas ftrmners rice alone (211) NfailC arid Urplald ir_ 2re coilltori in the plant ca rly becia use ofa shorter raiinfall period arnd the desi re Philippines he chatngC in pest status folii iniercropping is to lliarvet before the typhoor sea son peaks ltighly locatioll-specific (133) cmianarid the net elfc be either

Early plantings cornbind with early-iatiuring varictics nil bcnelficial ortdetriential (189) For example BIatangias provide higher crp tolerartce for white grubs Aestiatior farmers intercrip Itro in their ipland ricefields A liornshyterminates with the first rains but 5-6 moare ineeded for the nrii iotd-Ahis i (I iiiacius) which norlially feeds resting larvae to develop into beetles emerge and lay eggs on taro ill feed or miarbv rice plants Intercropping and for larvae to reach the danriigii g ird-instar stage probably would not afft0 seed pests A crluanion crop (134 In lataigas this normually iseiough tiite for harvest planted ith rice and haing gleater tolerarnce for root

It Brazil howecver delayed planting is a suggested dariage or being toxic nayact isi trapcrop to root-feeditig cultural control for the lesser cornstalk borer and insect pests Acial-fei i ig pests voultId be most affected by pests favored l drought (66 195) Cropping internsity in intrer ropping Shoot aphids leaf beetles sten borers and Brail is low and delayed plraing places the rice crop in the hopplers would be likely ca ntidates to note effcts On the niost stable rainfall period reducing likelihood old rought othlt halnd the corlipalior crop stch~ sitniaie Iiiv be more

In rilniy arca s rinmirsare roi atl to plant other crops benehited than rice from initecropping ( 137) before rice and althNgh it woultl be desirabh to avoid 11echi Weeuling luring the first month after clop infestations by planting rice cuatl lariners lack the labor cstablishent will force pcts such is anyvworris which and time to do so ol a low priority crp prefer grassy cetds ontor tle vorling rice Soil iampsecs will

77age Uplild rice Stils ae light textured and Call be mvnye to the rice crop Cleani citrltirc also lay lore territes plowed duiring the d rv seasor notl only to grin time for to attack I u cetincti inrice ceds i ay he le optinal planting with lie fitst Inilsoon riiains but also to desiccate solition 13) soil insects arid weeds such is ntsedge that foim rhizomes Irtlizr Itneatsing N Iciiliier oin hlwland rice has Htowever becta use most soil insects lie below die plow layer fAtwvored Irigher populains of pliithoppeiv Imlhoppers

26 IRPS No 123 January 1987

and leaffolde rs However because of the need to temper fertilizer use on upland rice o prevent blast fIrtilizer rates will be lower than is comonia r for lowlad rice causing relatixely less hopper and IaIfTfoide r population buildup However inupland rice areas without a history of blast such Is Batangas farmers apply high rates of N and conscqtrently the crop has high infcstation levels of WBPI]and RIF The fhrmrcrs leec insect buildup by splitting the fertili11r in0to two to three applicatiois bit do not applV fertilizer at the vegetati estage because this prodrces a tall crop which illeasily lodge during typhoons Greater crop fertility leads to greater insect survival (better nutrition) larger insects (to cat no re foliage) arid greater fec titd t (to lav nmre eggs) (128 145 171) Not all insect species respondequally hovr Increasing rates (if N l anrid K increased Dialraa s-acchralis and (iaeiurema flea beetle but reduced thrips (66) Zn rcdLuced deadheart density from the lesser cornIstalk borer

Fnldingi Although most Upland rice areas do nIot hawxcaccess to wa ter floodiig the fields is rRrrnriierdcd in areas of Brazil or root bug and white gib control ( 195)

Crop rotatioti Tillage before planting a crop after rc harvest will control stern boreis and other pests renain n the stubble arid will unearth soil pests Planting a non-gratriiaceous crop after rice is recommncrded for termite control (66) Crop rotation is less nttdcd in upland rice culture than in lowlands because atcrop-free period is assured even after harvest of crops following rice

t1ulchilt iryl plastic irli irig designed for weedcoitrol ii upland rice inJapan resulted in ntr highanri liy infestation of (ihilo sulytresvais (93 9-1 95) NIulched Upland rice grew more luxuriantlv and attracted more (Avipositing mothis than u lched rice Also the rice plant and stein borers imatoured 10rC quickly and a partialgeneratiori af vouing ista iNoverwintcred in the stubbleLotili--malitrt varieti Quickly maturing crops reduce the tmbCr of pCst generations that Calt build up Rapid crop establishment olan ca rl v-natu ring vaitvty if)pa 11icLa r will lessen white gib and stem borer damiage White grub larvae intle last iistar are highly destIruCtive so aii early-iiatri ng variety will escape serious root loss Stem borers build 1ip slowlyx in rice to ecormic abundant in a late-Maturing crop anrd caiuse whi tlieads Numibers of somespecies increase exponeitially with each succeeding genera-ti so a xarietyexhat atures I trowh earier thian another will hac less insect damage Early-naturing vaiicties however will not lessen seed pest daiage either ii the soil or oina standing crop

Strwhoi hAIJttam Inrscts tcd irig on thlie aerial portions of rice plats disperse from carly to late plantedfields AS iiiibCirs increase exponentially with eachinsect

gtscnrattm e eiglbrig f rmc~s1(uil upp wh rc generation pests build tIP where neighboring fihrTC e9r1sstagger their plantings II Batangas farmers plant uplandrice within I iro use 120-d s-arieties and plant at the start of the rainy season Ini Misarnis Oriental laricrs stagger their

plantings up to 3 mo apart and use a set of atieties that mature in 4-7 mo Insect pest danage particularly from stem borers is greater in Misarmis Oriental Also seedling maggots damage rice there because rainfall favors frequent planting of maic In Batangas rice escapes seedling maggots bCca usC of distinct drs season and synchron0us early planting with the onset of ains

Therefore combinations of cultural practices ea rlyplanting synchronous planing crop rotation and earlyshymaturing vaneties protect the rice crop against most insect pests

Ilmant resistance Beca tse upland rice has few specialized insect pests and many insect types attack the crop worldwide regional breeding and strong internatiornal cooperation are required

Sown-sced pests arc economically controlled by insectishycide seed treatment and should the relore receive lowxpriori ty inbreeding objectives

Root pests are normally cost ly to control with insecticides but finding resista t sonres mayvbe di fficult Tanaka (229) for exa -ple fi led to find varieties resistant to root aphids Selecting for large root bioniass perhaps should be the strategy for pests sLuch as Whlite grubs that renliove roots Fainter(174) howcvcr cites varietiesof crops tolerant ofor resistant t(i root pests Some sorghun varieties are high in cyanide so rice sorghun might incorporate broadshyspcctruri chemical resistance

Seedling vigor and drought tolerance are high prioritybreeding objectives so tupland rices should have large r root systernsand be able to tolerate ligherlevels of root loss Sonie root loss inolder plants is beneficial if it stimulates new root developnrut and therefore crihances intake of soil nutrients (53) The rice seedling inaggot rtiLotia spp may be

controlled thirough resistant aittics A hi rgc breeding effort on sorghniu has had fair success inideveloping arreties esrstait to Asoccata Rodani (251) A field trial comparing 10 upland ricc varieties showed differences aniong varieties va ring from II to 4(- damaged tilleis (215) Hovver resistarce to 1Iill-0r gtia species may rot cross ox-rti otier

Shiraki (210) reported that upland i-ice stens wcrc harder than lows-land rice stemsperhaps becaIusc (f a higher silicoricontent and thus were more resistant to yellow stern borer hc tortia Iow tillering (f uplaid iice xarneties prodtres a

higher percentage of infested tillers than would occor in higher tillering lowland iice under a siiilar egg density The corollary therefore is to select for high tilleriig upland rices

Stemii bor amcstheh main targes dobrecders inAfrica (571 204 )but nticl oifth e sc reen irig is dotie unde r lo wland co d t osrm s wih A h r~ o l pe i s n acorditions (11 96 218) As with lthrinrnwspecies mayhave to be dealt with independently however initial results show cross resistance between African and Asian species(92)

In Brazil screening Ior insect resistance focttses oil lou I pest problems Varieties tolerant of lasmolwalpus ligi-sellus and resistant to )iatraea saccharalis have been identified (70 153) Current work also emphasies 7iraca limativetntris and Thmialspis (=Iosis) flavouicta (66) These insect pests Ilt onlV citise IIigh yield losses tihey also ire difficult to control chemically ofr by other ilealls

ManV upland rice vaieties aic early maturing and are resistant to pests such as gall midge (I Xl ) Early mIturity is often a highly desirable trait to esuilpe pest buildup but it is not true resista nce bca usc ilthose va reties are plinted late they will be itsdamaged as ain early planted susc ible variety

P~erhaps tile cosiiiopolitall of all upland rice pestsimost are the sced pests (ripenling stagc) which pose a particular problem beta use scuds re ised IsIO for hitnit4ans Ayltoxic substatice introd tcld geticticllyilmay also be toxic to nal iniless it caln he placed old in thfe IdhuffSelectmi fo hairiness or lotng wns kill mIlct sthilrohleiiis dutilig threshing pa rtictfarl maitial threshinfl [hireshers ill complain of itcfimin atid itteascd (fust levels

Attempts itIRRI to fitid tcistatncc to the rice bug plorisa have ilotbeCnl prmitisilg lIIetIattonlids iorc

through the lenilna or palca hillhairs sedstVIor seeds wilh awns that deter bugs promote complaints from threshers Seed bugs liptocorisa and Stcnocoris enter the seed through the opening between tile lemna and palC but varieties with narrow openings are diifficult to mill

Breeding priorities should concentrate ott inotCI gruLtLIps that are most difficult to c nttrol by other meains Steil) horers appear to be the most widespread group that sigiiificantly dalag s rice aind is hard to manage by other Ilnca us

Biological control Very fe stutdic of mitatt~l eneties of rice pests in upland envirollnletits wee in tile wefouiid literature Therefore have not compiled a list of species Ve present a list of natural enemies iOtid in upland ricefields tuic latialln Batangas for some conitno iOldiar pest species ([itble 5)

Muich ofl thc lollowinle itfor ittiol cotlliCs rol Stutdies in the lowlatnds or roi crops otithcr thlin tice

Qu(ailtr o 11ll t11t( Nat trrtf lii hcoeits ight Ile used agaitist ile social insects tilt and termites Sollie predator specice live ill theirf nests n ilnic theirnllt ip-pearanIlCe aMRidbehavior IJtrouMdCing a mote aggressise ant species to displace pest ant codonies is a possibility (169) Patihogens as potential natural encnies have reccived little atteition (I13) Natural ncnuics asa group however hac not been toIl iporta it in Iegulating aintshown be and ternite numbers

The nocturnal behavior aid subteitmien habitat of lole crickets and field crickets protect them from many ntllral enemies There are only limited records of natural cnelics for these groups of pests some pathogens (87) but mainly

IRPS No 123 JattuIny I 271987

parasites - scelionids on eggsand sphecids on ilymphs and adults Sphecid wasps Larra carutmaria (SImitI hI L huzwuensis Rohwer and lsaltiuhta Willi ams speciali in mole crickets pa ralyzing them nd dragging the ninto thcir nests as food lot their voting ()tier sphCcid species Lins aurulenoa (Iabricius) lot maiac (Ashmead ) 1 lhshylssellaus (Smith) and Ml lal)oriosu (Snithf specialie in field clickets

White grUbsO the other hand hae Icti scriuns enuigh to have been extensively studied for cotttroi 1W nati al enemies( 112 140 141) Scoliid parasites C(ajsoMcispp have beetn intrtduced itto tilePhilippites for control of Ielxl ]udisiro la The rcsearchcts w Intttrdced themle during ill era claimed that the white gttionthreak as controlled (142) howevcr it isnormal for pest epidemics to subside and there was no diect evidence to indictte that tfe parasite was responsible A tmore promisitg titctllod might he to identify iso ltcttass prt~f tcartd apply hiactelaln fingal pathogens to tile soil Ihe lalattese beetle in North America has hcci coitrolled hv tite conmnleciall aailable hactcrial prcpa ration itacihlpu pav ofr milksdiscas (71) Spote prtyefott the soil itloted ilrtier reitiain actic fr Veas

Root aphid sardtnctly bugs readilslic tilib to predatots and parlsites once their tetndine ants are controlled The larvac and adtilts of a coccinellid beetle 1Scynms sp prey on llriadlouais in slash and burn upland rice Baits treated with insecticide may be used to control tending ants by killing their ourig (244)

Root-tkcditig blugs false wireoris wireorms and root wccils aic r ofsuch little ifilpoltatice that thenomttll role of liatural Cticniics hias tiot beei asscssed A possibility hoevrl tiglt he tit ItaIIitic niaCIMtodcs lintt1g11en Itese soil-itlhlfbitine mCtitodcs W5uhtfhave to be imiass Produiced

large miltibcr of niatltrll cnemics ircknown ft CltIoriS IC IpblCeii is thtt clttvortnlls ofonlie the crop soott alter land ptel)tritiOti a1tithe bcgit iniig of the raittv

Citsoln hfien tiatllral leniiiy populations aric lo Colltloil WOtuld iii ole mass prodtticing andirclcaslng key spccies

le egg stage is the fpirt (olthe Cedlitgt Ialot life cycle tI hich to f)ctis ftor tittlril elctit cottntrol hplraites o1 pleditors ttsit elleggs highly expoIsYCl I Sgg pailsites or predators stolld probahly e ittass andtbe produced released Littlh i toruialioll is a ailable ott what species to trV o ho to Iniss podLice thiCIi A etiloplhid wasp Icmihlaol sp effctiicly regushy

lates the leatf lcloalapolm2a spioanoiiitr heat andc rice antid should be collsctvcd

leaf beetles whCllose e am] eggs ate laid in the grutindhirO ould be motC difficutlt to control by niatulral enelV Mnll ipuhltiOn But species sUici IS c_gsthispid beetles ith

and larVle oilleascs ate vtilcratble to iparaites and pledators

Natural enemisc of tlirips otl rice are little known

28 IRPS No 123 January 1987

Table S Parasites recorded from upland rice insect pests in Tanauan Batangas Philippines 1977-84

Order Family Species Host stage

I lymttenoptera Braconidae Leaffolders Cnaphalocrocis Marasinia spy

Cardiochilesphilipplinensis Ashmead a Larva

Cotesia angustillasis(Gahan)Cotesia or cypris (Nixon)

Larva Larva

Cotesia or taeniaticornis(Nixon) Larva Cilonusinotakatae Munakata Chelonits SppAlacrocetrts philippinensis (Ashtmead)

Egg to larva Egg to larva Larva

Iclineumonidac Orgilts spp Isclutojppa hireator(Fabricius)

Larva Larva

Itoplectis naraigav(Ashmcad) Larva Tenelhtcha philippiensis (Ashmead) Larva Tenelchha stangli (Ashmead) a

irichomma cnaphalocrosis Uchida Larva Larva

Bethylidac Elasmidac

Goniozms or trianguli r Kieffer lasmtusalhopictus Crawford

Larva Larva

Diptera

Encyrtidac Chalcididac

Tachinidac

Elasmtus spp Copidosomopsisnacohiae (Fidy) Brach nteria evcarinata Gahan Biachyieria lasus (Walker) Ztgohothria ciliata (Wulp)

Larva Egg to larva Pupa Pupa Larva

Argyrophylx nigrotihialis(Blaranov) Larva

l-ymenoptera Braconidac Yellow stern borer Scirpopiaga incertulas

Braco chinensisSzeplegeti Larva CIthelotis spp Chelonus niunakatae Munakata Cotesi flaripes Cameron Stenoracon nicerilli (Bingham) a

Larva Egg to larva Larva Larva

Ichneumonidac Tropobracon schoenobii (Viereck) Amatroinorpiaaccepta metathoracica(Ashmiead)Frihorus sinicus (I lolmgren)

Larva Larva Larva

Ischnojoppaluteator (Fabricius) Larva Isotima nr dammeratiti Rohwer Larva Tenelucha philippinensis (Ashmead) a

Larva Tenehicha stangli (Ashmcad)Trichomma enaphalocrosis Uchida

Larva Lara

Chalcididac Eulophidac Trichogranmmatidae

Xanthopimnpla stemmator (Thunberg) Charops hrachYpterumn Gupta and Malteswary Brachtm eria sppTetrasticttus schoenohii Ferriere Trichorigranm chihunis Ishi

Larva to pupa Larva Pupa Egg Egg

Diptera

Scelionidac Pteromalidae Tachinidae

Tt ichograintnta aponiicuitt (Ashniead) Telenomits rowani (Gahan) a

Trichoinalopsis apattelctena (Crawford) Jeirhaea spp

Egg Fgg Egg pupa Larva

Z4gobothria ciliata (Wulp) Larva

Ilymenoptera lecneumonirlac Gol fringed borer (hilo auricilius

Trichomm enoptalocrosisUchida Larva

Ieioenoptera Braconidae

Aantholimlh steiniator (Thtinberg) a

Pink stet Ioret Sesamia inferens StoInacoil nicrilhi (Bingh am)

Loarva to pupa

larva

Diptera Ilyinenoptera

Armvtnyorms Mythimna separata Spodoptera mauritia acronyetoidesTachinidae Zvgohothriaatropiora( Robinea i -Desvoid y)Eulophidae It-phctrtiscitapadoe (Ashimead)

Larva Larva

(halcididae Braconidac

Brachpin ria spip (oesia spp

Larva larva

IHyntenoptera I)iptera

Rice skipper Ielop ida (halcididae Tachinidae

math ias aidgreen iortedci ciliarNIelan itis eda Brat lhincria sp iii matuqitatt Cameron Irgyrolhirlaxi trotlhiais(marumtv)a1

ismtne Larva Larva

Diptera Ilymenoptera

Tachnidae lelineutionidae

fuown se tilo per Mois frugalisIrgtrophrlax 1igrothiahiv 3 a rat ov) anthopimpa pmnctata (Iabricius)

Larva Larva to pupa

aMost dominant

IRPS No 123 January 1987 29

Normally predatory thrips prey on herbivorous thrips strepsiptera on both Spiders dominate tilenymphaladultNatural enemies of stem bugs --Scolinoplara and Tibraa predators Spittle bugs move to rice from pasture grasses - are in order of importance egg par~isites and predators and biocontrol would be successful only ifcarried out before fungal pathogens and nymphal adult parasites Egg tiley immigrate into ricefields parasites could either be introduced fron other areas or Biological control tactics for hemipterous seed bugs mass produced and released ould follow those outlined for stein bugs

Armyworms are normally lwld in hcck by the activities Coml)arison of environnewtns Natural enemies are of egg and larval parasites When these natural enemies fail perhaps faced with even greaterconstraints than insect pestsusually because of drought armyworns become epidemic to survive ind reproduce in upland environments TheyIt may be worthwhile therefore to release parasites during have adapted ani survived by having wide host ranges the the rainy months following a drought Virus diseases of ability to aestivate or hibernate over unfavorable seasonslarvae would beanotheravnue to exploreforarnyworms and the ability to disperse or by other mechanisms

Parasites have ben traditionallyconsidered for biolegical Comparing the most dominant and therefore the most control attempts against stein borers usually by rekasing adapted species of natural enemies collected in various exotic species( 166) This approach has not met with success environments in the Philippines we can understand how and new avenues should be explored The role of predators different rice environments are arid pathogens is little inderstood and deserves greater Anagqrms olptahiisa mymarid planthopper egg parasiteattention Egg parasitism rates are normally reasonably is cquallyadapted to all Philippine rice environments (Tablehigh but should be supplemented with effective egg 6) It is selective for planthoppers and even parasitizes eggspredators Orthopterans are potential egg predators of the n-iize planthopper Peregrinitsmaidis itt upland areas Metioche vittaticollis (Sta ) Anavipha nifiwnnis (Saus- Alavohis also parasitizes maize planthopper eggs but is sure) and Conocephahs hongipnni (de i-laan) feed on most prevalent on BPH in the irrigated wetlands (onashystem borer eggs etioche and Anaxil)ha specialize in eggs tocerus spp like A optalhilis on BPH and WBPI Iattacks with no hair such as Chilo spp The omnivorous (ono- the eggs of GLH in ell rie environments It is also found cephahsspecializes in eggs cov red with hairs it is a proven parasitizing the eggs ef the white rice licahopper (oimna egg predator of Scirlouphaga spp but has the discouraging spetra Oirosita naias a trichogrammatid egg parasitehabit of eating rice guin (65) attacks BIPH and WBPH in the wetlands and has no other

Grasshoppers have egg (scelionid) and nymphalad ult host A related species 0 aesopi specializes in G1-H in (nemestrinid tachinid and sarcophagid) v irasites Among wetland environments Dryinid strepsipt-an and pipunshypathogens protozoans have been recorded most frequently culid nymphal and adult parasites of planthoppers and Control of locust species by natural enemies could con- leafhoppers also specialize either in hopper species or centrate on their habitual breeding areas environments Among the dryinids Pseuhdgonatoipsflavi-

Leaffolders have rich complexes of natural enemies some feimur prefers the lowlands and BPH while P nuch has no adapted to upland rice (22 27) ranging from egg predators environmental prefirences but is adapted only to WBPH (gryllids coccinellids) to larval parasites (braconids ichncu- The main d ryinid species on GLH aptogonatop)usspp is monids) to larval prcdators (ants and carabids) to larval most dominant in rainfed lowlands and also attacks other pathogeiis (vira and fungal diseases) Efforts to augment leafhoppers within Amnrasca Cicadulinaand Bachaltuathese natural enemies should focus on the egg predators Strepsiptera are more dominant on plarithoppers than (rearing releasing)and viral or fungal pathogens (culturing lealhoppers The Elenchus species in the Philippines attack spraying) all species of planthoppers within the genera Sogawilla

Parasitization of the large conspicuous larvae of Sogatodels NilaparsataHarualia and Opiconsiva Elenshyskippers green horned caterpillars and other polyphagous cusrvasuntaisuiis adapted to the major rice environments Lepidoptera by tachinids and chalcids is normally low ELg but prefers BPH in rainfed arid WBPIH in wetland environshyparasites and predators are perhaps the key to their control ments Ilalictoph(utsmtnroeiand HVpectrus attack G LH

Gall midges are parasitized by pterormalids and platy- and COfimravcctra respectivelygasterids that aestivate within their larval host between Pipunculids are mainly parasitic oii lealhoppers Each seasons Shoot aphids are preyed upon by coccinellids species has a unique environmental preference Tomosshysyrphids and chrysopids varireIa oraetoraprefers Ne7 otetltix nigropictrs and

Planthoppers arid leafhoppers have egg parasites and Pilunwulhtsjavanensisattacks Dehoceplhahls spp which predators as well as nymphialad nlt parasitesand predators are more abundant in upland than irrigated lowland their rich complex of natural enemies includes fungal environments T suhvirescensattacks Nephotettix virescens pathogens The principal egg parasites are myrnarids and and prefers the lowlands Pilpuiulus nttillattls is an trichogramniatids Egg predators are norinally mirid plant upland rice parasite arid attacks only Nephotetix speciesbugs that suck out the yolk Nymphal ad nIt parasites are More than 17 recorded species of leaffolder parasites in dryinids on planthoppers pipunculids on lealhoppers and the Philippines attack species of C(naplwocrocis and

30 IRPS No 123 January 1987

Table 6 Dominant natural enemies of rice pests in upland rainfed lowland and irrigated lowland rice environments of the Philippines

Natural enemy

Host stage Species

Brown planthop-Parasite Egg

MymaridaeAnagrus optabilis(Perkins) Anagrusflaveolus Waterhouse

TrichogrammatidacOligosita naiasGirault

NymphAdult Dryinidac

Pseudogotatopuslaiifemttr Esaki et Hashimoto Elenchidae

Elenchus yasurnatsui Kifune and Hirashima

Environment

Upand Rainfed lowland Irrigated lowland

Ailaparata lugens

x x X x

x

x x

x X X Whitebacked planthoppers Sogatella furcifera and Sogatodes pusanusEgg

MymaridaeAnagrus optabilis(Perkins)

TrichogrammatidacOl~gositanaias Girault

NymphAdult Dryinidae

Pseudogonatopusnudus Perkins Elenchidae

Elenchus yasutatsui Kifune et Hirashima

Parasite Egg

MymaridaeGonatocerus spp

TrichogrammatidaeOligositaaesopi Girault

NymphAdult Pipunculidac

Tomosvaryella subviresccns Loew Tomosvaryella oryzaetora (Koizumi) Pipunclusmutillatus (de Meijere)

DryinidaeIlaplogonatopusspp

Egg Copidosomopsisnacoleiae (Eady)

Larva BraLonidae

Cardiochiles philipphnensisAshnead lchneumonidac

Trichiomnia cnaphalocrosisUchida Temelucha stangli (Ashmead)

Pupa Chalcididae

BrachymeriaexcarinataGahan Brachymerialasus (Walker) Brachymeria nr tachardiaegroup

Parasite Egg

Scelionidae Telenomus rowani (Gahan)

EulophidacTettasticusschoenobhi Ferriere

Larva Braconidae

Stenobraconnicevillei (Bingham) Iclhncurnonidac

Temelucha phiiippinensis(Ashinead)Temelucha stangli (Ashmead)

x x x

x x

X x X

x x Green leafhopper Nephotettix virescens

X x x

x x

X X X X

X

X Rice leaffolders CnaphalocrocisMarasmia

x X x

X

X X

x x x x x X x x x

Yellow stem borer Scirpophaga incertulas

X X x

x

x

x X XAinauromorphaaccepta metathoracica(Ashmead) x

continued on next page

IRPS No 123 January 1987 31

Table 6 cntinued

Natural enemy Environment

Host stage Species Upland Rainfed lowland Irrigated lowland

Larva-pupa Gold fringed borer Chilo attriciiuslchneumonidae

Xanthopimpla stemmator (Thunberg) x aPupa x

Pteromalidae Trichoinalopsis apanteloctena (Crawford) ax

Pink stem borerb Sesamnia inferensParasite

Larva Braconidae

Stenobracon nicevillei (Bingham) x b b Armyworms Mvthinna SpodopteraLarva

Tachinidac Zygobothriaarroptvora (Robineau-Desvoidy) x x

Braconidac Cotesia spp

X Skipper Pelopidas mathias and green horned caterpillar Melanitis leda ismeneF-gg

TrichogramrnatidacTrichogrannaspp x

Scelionidae Undetermined Scelionid

x Larva

Tachinidae Argyroph lax nitrotibialis (Baranov) X x

Pteromalidae Trichomnalopsis apanteloctena (Crawford)

X Predator

CoenagrionidaeAgriocnenis femina femina Brauer x xIschnura senegalensis tambur

Miridae CrtorhinsliidipennisReuter x x

Nabidac Stcnonabis tagalicus (Stal) x

VeliidaeAlicrovelia douglasi atrolineata (Bergroth) x

Coccinellidae Micraspis crocea (Mulsant) X x x

Vespidae Eumenes campaniformis (Fabricius) x

Formicidac Diaecamna spp xOdontoponera transversa (F Smith) x

Carabidae Chaenius spp OphioneaW- aJasciataSchmidt-Goebel

x

Ophionea ishii ishii Habu x x X

Gryllidav Anaxipta longipennis (Saussur-) x xMetioche ittaticollis (Sial) x x x

Tettigoniidae Conocephalus longipennis (de Haan) x x x

aNo rearing made on rainfed lowland bNo rearing made on rainfed lowland or Irrigated lowland

MrasmiaAmong the Laval parasites the braconid Cardio- lowlands and parasitizes oriental maize borer OsIriniachilesphilijpphensis is most adapted to an upland environ- furnacalis (Guenee) larvae The leaffolder pupal parasitesment It also parasitizes Hydelepta indicata a common not only occur equally in all environments but also haveleaffolder of legumes The ichneumonids Trichomma cna- wvide host ranges- parasitizing species of OsIrinia Marucaphalocrosis and Temehcha stangli occur more in the Hydelepta and Honiona - in maize and legumeslowlands T stangli is most adapted to irrigated lowlands The principal egg parasite of yeltow stem borer (YSB)and also parasitizes Chilo and Scirpophaga stem borer Telenomus rowani is widely adapted to all environmentslarvae T cnaphalocrosis is most prevalent in rainfed and has an unusual alternate host -Tabanus eggs (25)

32 IRPS No 123 January 1987

Tetrastictu schoenobii specializes only in YSB and occurs mainly in irrigated lowlands -- further proof that YSB is most adapted to the wetter environments

Larval parasites of YSB are highly environment-specific The braconid Stenohracon nicevillei occurs only in upland rice but also attacks the pink stem borer (PSB) Tieinchwa phiiiplPwnsis an ichneumonid occurs in upland rice and also parasitizes the larvae of (7hiu stqpresaisinatro-morpha accepta melaihoracica like 7 stangi is mnost adapted to the irrigated lowlands

The dark-headed stem borer(l)HSB) (ih pochit3sus was not collected in all rice environments and the gold fringed borer Chilo auricilius nd PSB were collected only in upland rice Xaantiholiplastenrnaloran ichneumonid larval parasite appears to be widely adapted whereas TrihomaslopisapanitI)cteiiaa pteromalid pupal parasite occurs mostly in wetlands and also parasitizes larvae of Chilo suppressais and Pelhuidas mathias

Armyworms Myit hinin and Spodujipera have two main larval parasites - a tachinid Zrgohothriaatropivora in rainfed environments and a braconid Cotesia spp in irrigated lowlands

The butterfly pests Pelopidas and 4elanitishave tricho-grammatid egg parasites found only in irrigated lowland environments

Two larval parasites show environmental preferences --Argrroph av nigrotihialis a tachinid in rainfed habitats and Trichonialopsisapantelkcnela in irrigated ricefields A limgrotibialisalsoattacks the sweet potato hornwormAgrius convolvtdi and T apanteloctena parasitizes DH S1B

Predators as a group show more distinct environmental preferences The aquatic damselflies Coenagrionidae are most abundant in the lowlands Cvriorhinus is also most adapted to the lowlands The nabid Stenonabis tagalicus occurs only in the uplands and isalso prevalent on legumes The coccinellid lady beetles wasps and ants are more prevalent in upland areas AlicrasIpis cit-ea larvae prey ona variety of aphids attacking legumes and maize Etunienes campanonia vespid mud wasp makes nests in trees and is therefore most adapted to the more botanically diverse uplands Soil-dwelling ants cannot tolerate flooding

Upland environments are habitats to arboreal carabid beetles Three species of Chlaeniusbeetles prey on leaffolder larvae and are perhaps more important than parasites (27)

The gryllids and tettigoniids are widely adapted egg predators not only on rice but also on maize and legumes They feed on eggs of most insects that are laid on leaves

The spidercommunity of upland rice environments isrich in species At one site (Batangas Philippines) 31 species have been recorded (23) Of the 176 spider species recorded in Philippine ricefields about one-half (82) occur in upland rice The spiderspecies of upland rice environments overlap more with those of rainfed lowland than with those of irrigated lowland rice (24) The same study showed that of the three environments irrigated rice has (ie greatest spider species diversity foiiowed by rainfed lowland and upland

However of the 10 most prevalent ricefield spiders 9 were abundant in upland rice showing wide adaptation All three spider guilds -- orb-web space-web and huting spiders shywere prevalent in upland ricefields There were however differences in environmental prelerences for some spider species

Of the orb-wcavers Tetragniathamanlihulatawas partishycila rlv abundant 7 japonica was low and Ltucatge dehcorata was absent in upland rice

As in other environments Aiypeiwnaifirtosana was the most dominant space-web spider however its relative numbers were lowerthan for lowland sites Among hunting spiders Ox|opes javanus and Lcosa leatcost fgla were more abundant and Lvcosa lpseudoanmnlata was less abundant in upland compared with lowland rice envishyroniments

Extensive rearing of rice insect pests in upland rainfed lowland and irrigated sites over an 8-yr period in the Philippines provided insight into the effectiveness of parasites as natural enemies (Table 7 8) Most of toe data are from large sample sizes taken over at least one crop which overcomes the pitfalls expressed by Van Driesche (238)

VSB egg parasitization was surprisingly similar across all environments Larval parasitization was highly dynamic among sites and even years within the same site but showed somewhat lowerlevels in uplandsthan in irrigated wetlands This would have been predicted for egg parasites as well since YSB is adapted to the lowlands and so must be its parasites This isevidence that Telenomus rowani is as good at dispersing as its main hosts Trowani clings onto the anal tufts of female YSB and parasitizes eggs as they are laid This phoresy may explain its wide adaptation The more specialized lowland YSB larval parasites therefore appear more effective than their upland counterparts with more alternate hosts

Low collection levels of PSB larvae especially in the wetlands make comparisons between environments difficult Low levels of larval parasifization were recorded in both rainfed environments Except in the Cagayan rainfed wetland site in 1980 low levels ef larval parasitization were evident at irrigated and rainfed sitesalike No differences by environment were evident Higher levels of parasitization occurred in Sarawak during a 1967 outbreak also from tachinids Rothschild (197) indicated that higher parasishytization occurred in upland rice areas

Skipper and green horned caterpillar larval parasitization rates were generally low and showed no environmental effect htighest activity occurred in Cagayan in 1980 (48 parasitization) Although larval parasitization levels of RLF were variable year to year activity appeared greater in the irrigated lowlands than in the uplands Again the highest levels were recorded in Cagayan in 1980 Cagayan has extensive grasslands and fallow around the rice areas and a relatively long rainfall period Rainfall in 1930 was good for crop growth and apparently for increase of larval parasites

IRPS No 123 January 1987 33

Table 7 Parasitization rates of rice planthoppers and green leafhopper in upland rainfed lowland and irrigated lowland environments Philipshypines 1976-840

Environment and site Year Egg parasitization () Nymphaladult parasitization ()

BPH W13P GLH BPII WBPH GLIH

UplandBatangas

Rainfed lowland

1976 1979 1980 1982

15 17 12

2(47)8(39)

4(51)

9(1800)14 (212)

12(304)

8 (101)6 (127)

9(116)

Iloilo

Cagayan

1976 1977 1978 1981 1980 1981 1982

0 (69)

11

44 48 40

-

-

46 55 44

0 (471)

11

52 56 49

0 (200)9 (214)

10 (1259)8(235)6 (900) 3(800)

-7 (462)

11 (270)3 (313)5 (900) 1 (800)

4 (600)10 (725)10 (362)12(870)5 (900) 3 (800)

Pangasinan

Irrigatedlowland

1984 1976 1978 1979 1982

13 (436) 13(379) 14 (457)

6(14)7 (197)

17(198) 11 (486) 18 (275)

3 (81)10 (274)3 (116)

16(871) 19(88)

28(211)9 (764)5 (647)7 (1204) 8(305)

Laguna 1977 11 7 21 1978 31 1979 14 4 4 28 1980 7 8 20 1981 1983

56 7(278)

26 16 (1650)

16 27(1921)

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R I Apostol unpublished 1lo1lo and Pangasinan 136 Cagayan BCanapi unpublished Liuna 175 I C Pantua unpublished 118 234 43 Carino and Shepard unpublished BPH = brown planthopper Nila parvara htgens W131Pi =whitebacked planthopper Sogatelia furcifera GLH = green leafhopper Nephotettrix virescens Figiires in parentheses are sample sizes

Tauher et al (230) reported that internal external and genetic factors influence the seasonal activity of parasitoids

Hopper egg parasites but not nymphaladult parasites were also parlicularly abundant at the Cagayan site possibly the egg parasites have more alternative hopper prey in the weedy areas than nymphaladult paisites

The seasonal dynamics of hopper egg parasites can be seen for upland rice in Batangas(Fig I 2) and for rainfed lowland lice in Pangasinan( 135)and Cagayan( 105) Levels of BPHWBPI-L and GLH egg parasitization were corn-parable in upland and irrigated lowland environments Except for some high rates of paraMsitization early in the season the parasitization rate tended to be steady in all envilonments However hopper nymphaladult parasi-tizution levels were more similar in the rainfed environments and lower than in irrigated rice

Overall parasites do not appear highly effective by themselves against upland or even lowland pests Among natural enemies parasites have attracted the most attention because they can be more readily assesscd

Predators and pathogens lend themselves better to management practices Predators the most important group of natural enemies in riccfields are difficult to quantify On the other hand pathogens appear less im-portant but can be readily cultured and disseminated

The natural enemy commulnity of upland rice is rich in species and differs significantly from that of lowland rice

These beneficial organisms must be conserved by applying insecticides judiciously particularly sprays Seed or soil placement of chemicals minimizes exposure to natural enemies The strategy to derive the greatest benefit from natural enemies isto allow the greatest number of beneficial species to thrive Then perhaps at least one species will be effective against each pest at any tine overcoming the variable abundance of each species within and between years

Programs to introduce exotic species or mass produce indigenous ones are ambitious and expensive because they require trained people on a sustained not ad hoc basis

Chemical control Insecticides offer rapid and efficient control of upland insect pests (190) but-should be used only after other control measures have been considered Insecticides are rarely used on upland rice because of cost (32) Upland rice yields are normally too low to justify the expenditure Also most upland rice is grown as a cash crop Spraying upland rice is more difficult than lowland rice because water is less accessible Government subsidy programs to provide pestishycides to tribal peoples for upland rice production have created problems of toxicity to humans because isolated people have had no experience to be able to handle pesticides safely

However ins-eticide use has beenjustified when a crop is

0Z

Table 8 Parasitization rates of lepidopterous pests in upland rainfed lowland and irrigated lowland environments Philippines 1976-84a

Parasitization (5)

Environment and site Year YSB GFB PSB Earcutting Swarming Skipper Greenborned RLF Brown larva larva caterpillar caterpillar caterpillar caterpillar larva semilooperEgg Larva larva larva larva larva larva

UplandBatangas 1977 - 0 (57) - 0 (63) - 0 (71) - 0 (106) 11(142) 2 (101)1978 43 (4697) 8(451) 5 (38) 3 (40) 6 (61) 2(58) 15 (57) 13 (129) 13 (1286) 4 (30)1979 59(4135) 12(320) 13 (11) 5(28) 9(45) 6(46) 7(23) 8(160) 9(1152) 8(26)

Rainfed lowland Iloilo 1976 - 0 (10) 0(10) 0 (7) 0 (19) 2 13) 15 (180) 5 (76)1978 0 (54) 0 (3) 0 (17) 9 (64) 4 (78) 3 (73) 0 (121) 0 (191)1979 64 (2350) 16 (84) 6 (17) 0(6) 18 (68) 25 (104)

1981 56 (1322) 18(304) 0(5) 12(51) 18(138)Pangasinan 1976 60 (7 7 )b 62(82) 0(111) 5 (39)

1978 58 (53)b 5(51) Cagayan 1979 59(713) 8 (101) 6 (34) 0(7)

1980 0(11) 4 (28) 15 (286)

62 ( 48(177) 61(140)1981 0(5) 12 (2) 13(45) 14 (92)

Irrigatedlowland Laguna 1974 62 8

1974-75 50 41 1975 43 72 1978 96 (623) 21(24) 33 (60)1979 83(2017) 18(110) 8(62) 11 (89) 22(400)1980 78 (1625) 20(75) 0 (16) 4 (72)1983 60

9 (80) 7 (270) 30

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R F Apostol unpublished Iloilo and Pangasnan 136 Cagayan B Canapi unpublished Laguna 175 P CPantua unpublished 118 234 143 Carino and Shepard unpublished YSB = yellow stem borer ScirpophagaincertulasGFB = gold fringed borer Chilo auricilius PSB = pink stem borerSesamia inferens earcutting caterpillar Alythimna separata swarming caterpillar Spcdoptera mauritia acronyctoides skipper Pelopidas mathias reen horned caterpillar Melanitls ledaismene RLF = rice leaffoldersCnaphalocrocismedinalis and Marasmiaspp brown semilooper Mocisfrugalis Figures in parentheses are sample sizes Egg masses

IRPS No 123 January 1987 35

Table 9 Effect of seeding rate and insecticide on plant density and Yield losses from insect pests in upland rice often areyield of Dagge upland rice Batangas Philippines 1976a comparable to those of lowland rainfed or irrigated rice

Seeding rate Insecticide Dosage Plant density Yield (0-Y30o) but usually from different pests The principal (kg aiha) seed treatment (kg aiha) (noIra-row) (tha) groups are soil pests feeding on sown seed and rootsat 14 DE followed by seedling pests These groups have largely gone

50 None - 35 a 20 a unnoticed leading researchers to conclude that stem borers50 Carbofuran F 10 38 ab 23 a and seed bugs are the most important pests (17) Yield loss 10 beCabofuranF100 None - 4551 c 23 a trials that pinpoint major pests need to includc seed

100 Dieldrin WP 10 52 cd 22 a treatments Granules applied in the furrow may not give150 None - 59 d 22 a adequate protection from many seed pests

aAv of 8 fields DE = days after crop emergence In a column means The floristic and pedological diversity of upland rice followed by a common letter are not significantly different at the environments is matched by the diversity of arthropods5level by D)MRT F = flowable Wi = wettable powder both pests and their natural enemies Irrigation has homoshy

genized lowland rice environments Flooded soils tend tothreatened by epidemics of locusts or armyworms New have the same properties and to eliminate soil pests The technology stressing efficiency has minimized insecticide spatial and temporal dominance of cultivated rice has cost benefited specialist pests

Seed treatment is normally inexpensive and can be Upland rice has no specialist arthropod species The economicallyjustified in -nanycases(66) Treating seed with monophagous pests on upland rice prefer the lowlands systemic insecticide (216) but not chlorinated hydrocarbons where rice evolved Rice in the uplands exists only because (247) is most effective in controlling seedling maggots of man and even though lowland rice predates upland riceFoliar sprays have to be repeated to obtain control and specialist species evolved Wheat maize and sorghum do often are washed off by frequent early season rains not have monophagous pests leading us to conclude that Granular insecticides are effective but required dosages are there is no evolutionary advantage for inssects to have too expensive to justify except under heavy infestation narrow host ranges in upland graminaceae(247) When directed against first-instar larvae banding Pests have adapted to the uplands not only by having granules in seed furrows (177) can be effective against white wider plant host ranges but also by having long life cyclesgrubs at low dosages (025 kg aii ha) (134) Seed treatments undergoing dormancy or being dispersive Each pest has a are not promising for white grub control (248) unique life history strategy suited to highly unpredictable

Seed treatment is also effective ainst ants and may be environments In agivenyear someinsect specics an better cheaper than increasing the seeding rate In Batangas exploit upland rice as a food source but no pest is a senious treating 50 kg seed ha was as effective as increasing the one every year on upland rice seeding rate to 100 kgha and using untreated seeds (Table The low yielding potential of upland rice means less 9) In Brazil seed treatments control termites and other soil quantity and lower quality of food for pests to exploit but insects including the lessercornstalk borer(66) Domiciano also less incentive for farmers to undertake control (58) however found seed treatments inconsistent in their The most effective control would be early and synchroshyeffect against high populations of lesser corn stalk borer nous planting of early-maturing upland rice varieties to

Baiting can be an inexpensive way of controlling ants prevent insect pest buildup Breeding for insect resistance seedling maggot flies mole crickets and field crickets Baits should initially focus on stem borers because they are verymade of locally available material and impregnated with difficult to control by other means Biocontrol efforts insecticide can be sparingly distributed taking adantage of should first stress conservation of natural enemies preshythe pests ability to disperse and encounter bait sites dators and pathogens rather than parasites Efficient

The new low-volume hand-held controlled-droplet chemical control includes seed treatment or baiting for seed sprayers may offer an advantage over high-volume knap- seedling and some root pests and using controlled dropletsack sprayes for controlling foliar pests Upland rice sprayers for spot treatments against foliar pests and seed canopies are more open than those in lowland rice and bugs when economic thresholds are reached better droplet penetration should result Controlled droplet applicators would be ideal for seedbugs and defoliators

CONCLUSION

Upland rice is attacked by a wider array of insect pests than is lowland rice mainly beca use of the addition of soil pests Generally population levels of lowland rice pests are lower in upland than lowland rice leading many people to conclude that insect pests are not important on upland rice

36 IRPS No 123 January 1987

REFERENCES CITED

1Abraham C C and K S R Mony 1978 Occurrence ofLeptocorisa acuta Fabr (Coreidae Hemiptera) as a pest of nutmeg trees J Bombay Nat ttistSoc 74553

2 Agyen-Sampong M 1975 HlierogrlphusdiaganensisKrauss (Orthop-tera Acrididae) a new pest of rice in Northern Ghana Ghana J Agric Sci 8249-253

3 Agyen-Samnpong M 1976 Varietal resistance and other inle-ratcd pest control components as factors of rice varietal improvement in West Africa Prcsented at the Second varietal improvement serinar 14-18 Sep 1976 Bouak Ivory Coast West Africa Rice Development Association Monrovia Liberia 18 p

4 Agyen-Sampong M 1977 Outbreak of Ihccrorrdnisorae in rice seedlings near Rokupr Sierra Leone lit Rice Res Newsl 2(6)16-17

5 Agyen-Sampong M 1980 Dirty panicles and rice yield reduction caused by hugs IntRice Res Newsl 5(l)11-12

6 Agyen-Sampong M 1982 The major pest problems of irrigated upland and mangrove swanip rice ecosystems ir the humid tropical Guinea savanna and Salel climate zones Pages 318-328 in Pro-ceedings training course cn concepts techniques and application of irtegrated pest management in rice inWest Africa 10-20 Jan 1982 Fendall Liberia West Africa Rice l)evelopioent Association Monrovia Liberia 505 p

7 Akibo-Betts V I and S A Raymundo 1978 Aphids as rice pests in Sierra Leone lit Rice Res Newsl 3(6)15-16

8 Akinsola F A 1975 Present status of different rice stemborers in parts of Nigeria iicL Entomo Newsl 328

9 Akinsola FA 1979 Ihe biology and ecology of rice stemborers in Nigeria Ph 1) thesis University ol lbadan Nigeria 217 p

10 Akinsola E A 1979 Guidelines of integrated management of insect pests of upland rice in West Africa Presented at the Seminar on integrated management of rice diseases and insect pests at Bobo-Dioulasso Upper Volta 17-22 Sep 1979 West Africa Rice Develop-ment Association Monrovia Liberia

II Akinsola IA 1984 Insect pests of upland rice in Africa Pages 301-305 in An overview of upland rice research Proceedings of the 1982 lIouak Iwsrv Coast upland rice workshop International Rice Research Institute 13O Box 933 Manila Philippines

12 Alam M S V TJohn and Kaung Zan 1985 Insect pests and diseases of rice in Africa Pages 67-82 in Rice improvement in Eastern Central and Southern Africa proceedings of the international rice workshop at Iusaka Zambia 9-19 April 1984

13 Alticri M A A van Schoonhovcn and J D Doll 1977 The ecological role of weeds in insect pest management systems a review illustiated with bean (Phaseohsvulgaris L)cropping systems PANS (Pest Artie News Summ) 23195-205

14 Anmante F 1967 The leaf-cutting ant Atia capiguaraa pest of pastures [in Portuguese] Biol6gico (Sao Paulo) 33113-120 (Rev Appl Entomol A56207)

15Andrade A C and D Puzzi 1951 Preliminary results of experi-merits on the control of Sculptcoris iastantus on sugarcane [in Portuguese] Bioulgico (Sao Paulo) 1744-49 (Rev Appl Entomol A41421422)

16 Appert J 1970 Maliarplia separaelha (rice stem borer) new observations of entomological problems of rice in Madagascar [in French] Agron Trop (Paris) 25329-334

17 Ariadeau Mand Z Harahap 1986 Relevant upland rice breeding objectives Pages 189-197 in Progress in upland rice research Proceedings of the 1985 Jakarta conference International Rice Research Institute 1O Box 933 Manila Philippines

18Asar S and Kisimoto 1975 Seasonal occurrence of the smaller brown planthopper in upland rice [in Japanese] Proc Kanto-Tosan Plant Prot Soc 2228

19 Bakker W 1974 Characterization and ecological aspects of rice yellow mottle virus in Kenya Meded Landbouwhogesh Wage-ningen 152 p

20 Balut F F 1970 White grubs attacking rice Oryza ativa L [in Portuguese] Biol6gico (So Paulo) 36321-322

21 Banerjee S N and L M Pramanik 1967 The lepidopterous stalk borers of rice and their life cycles in the tropics pages 103-124 in The major insect pests ofthe rice plant Proceedings ofa symposium at the International Rice Research Institute September 1964 The Johns Hopkins Press Baltimore Maryland USA

22 Barrion A T and J A Litsinger 1980 Ants a natural enemy of cliaiarac in dryland rice IntRice Res Newsl

5(4)22-23 23 Barrion A T and J A litsinger 1981 The spider fauna of

Philippine rice agroecosystem 1Dryland Philipp Entomnol 5139-166

24 Barrion A T and 1 A Iitsinger 1984 The spider fauna of Philippine rice agroccosystcms II Welland Philipp Eiitomol 611-37

25 Barrion A T1 and J A Litsinger 1984 lihanus (Diptera Tabanidae) eggs an alternative host of rice stei borer (SI3) egg parasite Telenoonis digns (Iliymenoptcra Scelionidae) lit Rice Res Newvsl 9(6)19

26 Barrion A T and J A Litsinger 1985 Identilication of rice leaffolders by wing markings Int Rice Res Newsl 10( 1)24

27 Barrion A T and J A Litsinger 1985 Chctiosspp (Coleoptera Carabidae) a leaffolder ILF)predator lit Rice Res Newsl 10(1)21

28 Bertels A 1970 Insect pests of rice and theircontrol [in Portuguese] Pelotas Brazil Instituto de Expcrimentacao Agropccuararias do Sul Circular 43 24 p

29 Bonzi S M 1982 Ctilo (flisilineus J de Joannis (Lepidoptera Pyralidae) stem borer of irrigated cereals in Upper Volta LioFrench] Agron Trop Riu Rizic Cult Vivrieres Trop 37207-209

30 Bourke T V T L Fenner J N LStihick G Baker EHassan D F OSullivan and C S li 1977 Insect pest survey for the year ending 30th June 1969 Papta New Guinea )epartment of Agrishyculture Stock and Fisheries Port Moresby 57 p

31 Bowden 1 1973 Migration of pests in the tropics Meded Rijksfac Landbouwwet Gent 38785-796

32 Breniere J 1973 The main rice parasites in West Africa and their control Pages 27-42 in Proceedings seminar on plant protection for the rice crop West Africa Rice Development Association Monrovia Liberia

33 Breniere J 1976 The principal insect pests of rice in West Africa and their control West Africa Rice Development Association Monrovia Liberia 52 p

34 Brown ES 1962 The African arinyworn Spoiolptera eXI)Ina (Walker) (Lepidoptera Noctuidae) a review of the literature Cornshymonwealth Institute of Entomology London UK57 p

35 Brown F B 1968 Upland rice in Latin America FAO-International Rice Commission 12th Session of the Working Party on Rice Production and Protection 2-14 Sept Peradeniya Sri lanka Food and Agriculture Organization Rome Italy 6 p

36 Buyckx F J 1 1962 Disease and insect pests of crops inCongo Rwanda and Burundi [iil French] Publications dc LInstilut National pour lestude Agrononiquedu Congo M Weissenbruck S A Brussels Belgium 708 p

37 Cariiio F 0 1981 Role of natural enentics inpopulation suppression and pest management of green rice lealhoppers Ph D thesis University ofthe Philippines at Los Bafitos PhilippinesLaguna 136 -

38 Carpente A J 1983 The development of rice cultivation and extension of vegetable productin in Zanzibar Final Report Vol II1 October 1983 Food and Agricultme Organization Ronte Italy

39 Carroll C P and S1 Risch 1983 Tropical annual cropping systems ant ecology Environ Manage 751-57

4(0 Cates R G 198 1 Host plant predictability and the feeding patterns of monophagous oligophagous and polyphagous insect herbivores Oecologia (Berl) 48319-326

IRPS No 123 January 1987 37

41 Cendcafia S M and F B Caora 1967 Insect pests of rice in the 62 Ene 1 C 1981 I)iiorldhol)ierusihilis Slater (licteroplera Philippines Pages 591-616 in The najor insect pests ofthe rice plant lygacidue Illissinac) a new rice pest front Nigeria lit Rice Res Proceedings of a symposium at the International Rice Research Newsl N3)15 Institute September 1964 The Johns Hopkins Press Baltimore 63 Fugenio T S 1965 rhe effect oi e-vCgetation on rice production Maryland USA 729 p and the possibility of overcoming it by leaf pruning MS thesis

42 Centro Ihternacional de Agricklittira Tropical 1984 Upland rice in University ol the Philippines at Los Baijos ILaguna Ihilippins 69 p Iatin Anerica Pages 93-119 hi An os ervies of uplaind rice research 04 Farrow R A 1975 The Alrican nmigratory locust ii is main Proceedings of the 1982 Boinakamp Ivory Coast upland rice workshop outbreak area of the middle Niger qlinlitatise studies of solitary International Rice Research Institute P0 Blox 933 Manila populations in relation to environmental factors Locusla No II Philippines 198 p

43 Chandra G 1979 Taxonomy ad hn riononlics ofthe imect parasites of 65 Fennal R G 1943 Food crop investigation Windward and rice Icalloppers and planthoppers in the Pihilippines and their Ieeward Islands First report Report onl work dunring the periodiiiportairce iii natural biol)gical cointrol Philipp Fntoltol 4119-139 April 1942-April 1943 Imperial College of Tropical Agriculture

44 Chang T T 1976 The originii colut ion enlIivation dissemination Trinidad 16 p (Rev of Appl Fintornol A3222-25) and diersificition of Asian and African rices -uphytica 25425-44 1 66 Ferreira F 1983 Integrated pest ritaageiret of upland rice [ill

45 Chatierjee P B 1979 Rice leaffolder attacks in India lit Rice Res Portuguese English summary] Pages 323-341 in Upland riceculttre Newsl 4(3)21 factors affecting production E Ferreira T Yamada and F-

46 Cheng Cbien-pan 19(4 Upland rice in Taiwan International Rice Malmolta eds Institutoda Potassae Foslato Institnto Internacional Cnmmission Wiorking Party oit Rice Production ard Protection I()th da Potassa Piracicaha Sao Paulo lraid Meeting 3-10 March 1964 Manila Philippines Food and Agricul- 67 Ferreira E arid F dl Silva Martins 1979 Occurrence of ture Organiaion Rome Italy 5 p Sogatohs orvziola (Muir) inupland rice in Goias Ilrail lit Rice

47 Chcrrett 1 M and 1) 1Peregrine 1976 A review of the status of Rcs Ncwssl 4(4)18 leaf-cutting ants and their control Ann Appl Biol 84124-128 68 Ferreira E and 1F da Silva Martiris 1984 Insect pestof rice ill

48 Chipeta F M 1974 Pests of rice in Malawi Pages 90-95 in Brazil and their control Jill Portugucsej FMIlRAtPA-CNPAF Proceedings Sith eastern Africa cereals research conference 10-15 Goiania Brmiil 67 p March 1974 DRB Mialda ed Ministry of Agriculture lilongwe 69 Ferreira FI F ca Silha Miartins S S Nero and 1 It P de Malawi Carvallio 1982 M tltiplc regression analysis todeterimine the effect of

49 Chu Yau-i 1979 The d atiaging period of the rice seedling fly insect populatiois ott upland rice yield [inIPortuguese Pesqui (Atlterigotiavxigu Stcin) (NItiscide Diptera) oil cori) at Fast Java Agropecu Bras 17671-675 Indonesia Plant Prot Bill 21403413 70 Ferreira F 1F da Silva Martins and F 1PZimmermann 1979

50 Conklin H C 1957 IHaninoo agriculture a report on aln integral Rice varietal resistance to tle lesser corn stalk borer [in Port uguese] systent of shi fti rig cult ivationi in the Pltilippincs Food and Agriculture Pesqui Agropccu Biras 14317-321 Organization Rome Italy 209 p 71 Fleming W F 1968 Biological control of lie hapanese beetle U S

51 Cramer H II 1967 Plant proection and world crop production Dep Agric RLs Sers [ech Bull 1383 78 p Pilaniensclhutz-Nachr 201-524 72 Food and Agriculture Organiation of the United Nations 1982

52 l)ani R C and Majunidar 1978 Prelininary observations on tie Country report surmimary Pages 27-37 in Report on the first incidence of root aphids on differeit rice cultnrcs Sci Cult 4488-89 FAOIUN)IPhail and regional training coiuirse in iniproved ctltural

53 Davidson R I 1979 lffects of root feeding on foliage yield Pages practices for upland rice 5-23 Oct 1981 Northern Region Agrculturd117-120 in T K Ci osb and R B Pottinger eds Proceedings of the Developririt Center Chiang Mai Food and Agricuilture Organiiashy2nd Australian conlerence on grassland invertebrate ecology Govt tion Rome Italy 37 n Printer Wellington New Zealand 73 Franco do Ariaral S 195 1 Residual action of insecticides in turfy

54 l)ean G 1W 1976 Rice insect pests inlaos Int Rice Res Newsl soil--the results of a field experiment [in Portuguese] Biol6gico (Sao 1(2)15 Paulo) 17209-214 (Rev Appl Entomol A41423)

55 I)ean G J W 1978 Insect pests of rice in Laos PANS 24(3)280-289 74 Freeman ) 1970 Report onithe Iban london School of -coroiics 56 )ingle H 1984 Behavior genes and lifc histories conplex adapta- Monographs on Social Anthropology No 41 Athlane Press

tions to uncertain environments Pages 169-194 in A new ecology London UK 317 p novel approaches to interactive systems P W Price C N Slobod- 75 Fuchs T W1 A Harding and 1W Smithlr 1979 Induction and chikoff and W S Gaud cdsl ohn Wiley New York USA termination oldiapause in the sugarcane borer Ann Flitonlol Soc

57 Dobelmann-JP and M Falais 1968 Upland rice cultivation ill tie Am 72271-274 North-western part of Madagascar in 1966-1967 [in French English 76 (hbriel B1) 1975 A review tf the major insect pests of some tpland summary] Trop Agron 2313-21 crops in tlc Philippines Pages 86-111 in Ad Htoc panel of experts on

58 Doniiciano N L 1979 Chemical control of tie lesser corn stalk pest disease arnd weed problems in some rainfed crops Bangkok (lolsonno~paloIgnowslhts) (Zeller 1848) (I-epid(ptera Ihycitid ae) Thailand Food aid Agriculture Organization Roire Italy

on rice in Ponta Gressa Parara 1977-1978 [in Portuguese] Pages 77 Garg D K and N K Shal 1983 White grub outbreak on rainfed 291-303 in Anais 1 Reuniao de tecnicos cmiirizicultura do estado de dryland rice in Uttar Pradeslh Int Rice Res Ncwsl 81(4)16-17 Sao Paulo-Campinas Cati Brazil 78 Garg I) K and IP Tandon 1982 Major insect pests of rice on hilly

59 Dunsmore 1 R 1970 Investigations on tlie varieties pests and tracts of Uttar Pradesh India lilt Rice Res Newsl 7(11)11-12diseases of upland rice (hill padi) in Sarawak Malaysia Int Rice 79 Garrity 1) P 1983 Upland rice research an international bibliograpy Comm Newsl 1929-35 1975-1982 Iternational Rice Research hnstitnte PO Box 933

60 Elias 1R1965 The new decline of rice seedlings is not caused by Manila Philippines 121 p parasites [in Portuguese] Biol6gico (Sao Paulo) 313-6 80 Ghauri M S K 1982 A new subspecies of Iinorh)rone bull

61 Empresa Brasiliera de Pesquisa Agropecuaria 1984 Upland rice itt cornlus Slater (Hemiptera Iygaeidae) front Papua New Guinea on Brazil Pages 121-134 it Av overview of upland rice research rice and carpet grass Ihll lEntoniol Res 72133-137 Proceedings ofthe 1982 Blouak Ivory Coast Upland rice workshop 81 Gonzales F 0 PArregoces R I ihernandef and OT Parada International Rice Research Institute PO Box 933 Manila 1983 Insect and imite pests of rice arid their control in Latin America Philippines [in Spanish] la Federacion Nacional de Arrocerts (Fedearroz)

Bogota Colombia 60 n

38 IRPS No 123 January 1987

82 Grist D H and R J A W Lever 1969 Pests of rice Longmans London UK520 p

83 Hale P R 1979 Two subterranean pests of upland rice in Papua New Guirnca IntRice Res Newsl 4(2)16-17

84 Hale P R and J T Hale 1975 Insect pests of rice in the New Guinea Islands Rice Entomol Newsl 34

85 Harris W V 1969 Termites as pests of crops and trees Common-wealth Institute of Entomology London UK 41 p

86 Hattori 1 1968 A method to distinguish between grass leaf roller (Ctaplha rocis medinalisGuenee) and the rice leaf roller (Susurtia exigua Butler) [in Japanese] Plant Prot (Japan) 23167-170

87 Henry I E and 1A Onsager 1984 Experimental control of the mormon cricket Iahrosshnph by NJsena locustae(Aficrospora Microsporidae) a protozoan parasite of grasshoppers (Ort Acri-didae) Entomophaga 27197-201

88 Hidaka 1 P Vungsilabulr and SKadkao 1974 Studies on ecology and control of the rice gall midge in Thailand Tech Bull 6 TropicalAgricultural Research Center Tokyo Japan 113 p

89 Hincklcy A 1) 1963 Ecology and control of rice planthoppers in Fiji Bull Entomrol Res 53467 481

90 Hirao J 1982 Ecology and chtInical control of the rice leaffolder Jpn Pestic Inf 4114-17

91 Ho DT 1984 Sten borer incifience in rice ecosystems in KenyaEast Africa Pages 307-320 in A overview of upland rice research Proceedings of the 1982 Bouac Ivory Coast upland rice workshop International Rice Research Institute PO Box 933 Manila Philippines

92 Ho )T J 1Njokah and J GKibuka 1983 Studies on rice sten borers in Kenya with emphasis on ifaliarpiaipiraellRag Insect Sci Appl 465-73

93 Ichihara I T Ito A NIatsuo and M Sawada 1971 Chemical control of the rice striped stem borer in upland rice grown in plastic mulch [iii Japanese] Proc Kanto-Tosan Plant Prot Soc 1882

94 IclhiharaIand A Matsuo 1969 Damage caused by rice striped stett borer to upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1686-87

95 Ichhara I atid A MIatsun11970 Conside ration oft lie extraordinary prevalence of rice striped stem borer in upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1778-79

96 Institut de Recherches Agrolnliqes Iropicale et des Cultures Vivrieres 1978 Insect resistance Pages 12-13 in Upland rice 1978 atnual report Bouak Ivory Coast 27 p

97 International Institute for Tropical Agriculture 1983 Aphids itt rice roots I ITA Res Briefs 4(1-3)4-5

98 International Institite fi Tropical Agriculture 1983 Hopperburn in rice caused by brown plathopper IITA Res Briefs 4(l-3)6-7

99 International Rice Research Institute 1975 Annual report for 1974 PO Box 933 Manila Philippines p 216

100 International Rice Restarch Institute 1976 Annual repiort for 1975 PO Box 933 Manila Philippines p 228

101 International Rice Research Institute 1979 Cropping systems program Component technology development and evaluation Page 404 i Annual report for 1978 11O Box 933 Manila Philippines

102 International Rice Research Institute Department of Entomology 1981) Insecticid evaluation 1979 Tables 73 74 PO Box 933Manila Philipes

103 International Rice Research Institute Department of Entonmology 1981 Insecticide evalomtion 1980 Table 81 POBox 933 Manila Philippines

104 International Rice Research Institute 1981 Report of a meeting of Asian rice-based cropping systems entomologists Eleventh cropping systems working group meeting 18-22 May 1981 PO Box 933 Manila Philippines p 132 133 144

105 International Rice Research Institute 1984 Cropping systems program Pages 368-373 in Annual report for 1983 PO Box 933 Manila Philippines

106 International Rice Research Institutc 1984 Upland rice in Asia Pages 45-68 in An overview of upland rice research Proceedings of the 1982 Bouakamp Ivory Coast upland rice workiiop loternational Rice Research Institute 11O Box 933 Manila Philippines

107 Irshad M 1977 Grasshoppers in rice in Pakistan lit Rice Res Newsl 2(3)8-9

108 Ito R and TOno 1964 Upland lice its culture and varieties in Japan lIternational Rice Cormmission loth Meeting of the Working Party on Rice Production and Protection 3-10 March 1964 Manila Philippines Food and Agriculture Organization Rome Italy 12 p

109 Jeffrey M I and V A Dyck 1983 Effect of moonlight on lightshytrap catches of brown planthopper lit Rice Res Newsl 8(6) 13

110 Jennings P R and T Alicia Pineda 1983 Management ofthe rice plnthoppers Sogatodsorivi(coIl in Latin America Presented at the meeting on stable plant resistance FAO and lIndocsian I)inectorate of Food Crops Protection Bali lndonesia 29 Novcniber-2 l)ccemshyber 9 p

IllJimenez Gomez J A 1979 Rice insect pests i lla ios ()riCntales lil Spanish] Pages 99-103 in Rice ciltivation Instituto (olotibiatto Agropecuario Compndio No 29

112 Jongeleen F JJ Soesanto and A Mockidjo 1979 Studies oti pathogens of two species of Scarabacidac froinIndonesia Z Angew Enlomnol 87412-417

113 Jouvenaz 1) PC S lofgren and W A Baiks 1981 Biological control of imported fire antsa review of cirrent knowledge Bull Entomol Soc Am 27203-208

114 Kainoh Y K Shiniiu S MarUani Y Taiaki 1980 lost finding behavior of tie rice bu lplotori+JchinvittjI)allas (liciniptera Corcidac) with special reference to diet pattern ol aggregation and feeding on the rice plant Appl Entomol Zool 15225-233

115 Kakkar K 0 P Sharma and P I Sharma 1977 Note on Ihteronyrchus liodresRedt (Coleopicra I)ynastidae) as asporadic pest of paddy and maie inHimachal Pradesh Corr Agric 156-57

116 Kalode M131 79 Major insect pest problems and their control in upland rice (Unpublishcd mineogriph CWI oJNaaiat lni Chandaratat (198 1)original not seen)

117 Kalshoven L i 98 I The pests of crops in I ldoniesia (translated by P A vatn der Laan) 1 T Ichtiar B3aru-vati Iloeve lakarta Indonesia 701 p

118 Kamal N Q 1981 Suppression of whitebacked planto ner Sogatella fircaifra (Horvath) aLId rice lealfolder (nhal -cis licdb lis (Guenee) populations by natural cnclmie hlI) tiesis Grcgorio Araneta University Foundation Victoncta Park Metto Manila Philippines 179 Ip

119 Karim A N M Reaul 1975 A species ofliealy bug Trioni i70s ceres new oii rice in the Philippines Rice Entoniol Newsl 247

120 Katanyukul Wand C Chandaratat 1981 Insect pestsof uplaid rice and their control Report firstFAOUNlFi Tliailand training course on improved cultural practices for tplad rice Chiang Mai Thailand 5-23 Oct 1981 Food and Agriculture Organiation Rome Italy 22 p

121 Kern F 1955 lhletwioa(I)uninemts) bhhit Biturm(Coleoptera) a new pest of sugarcane inVenezuela [inSpanish with Etnglish sunmary] Agron Trop (Maracay) 527-29 (Rev Appl Eitoniol A4572)

122 Khan K M and S Ghai 1974 White grubs ald their control in India Pesticides (1Bombay) 819-25

123 Kiritani K 1975 Integrated pest management of rice pests Pages 66-70 itt Proceedings platning workshop ott cooperative hIeld research in pest managementP SMotooka ed East-West Center Honololu Hawaii USA

124 Kisimoto R 1967 Ecology of insect vectors forecasting and chemical control Pages 243-255 in The virus diseases of tle rice plant Proceedings of a symposium atthe International Rice Research Institute April 1967 The Johns Hopkins Press altimore Maryshyland USA

125 Kisimoto R and V A Dyck 1976 Climate and rice insects Pages 367-391 in Climate and rice International Rice Research Institute PO Box 933 Manila Philippines

126 Kureha S S Kobayashi SAsari and T Ishihara 1977 Life history and control of the mole cricket Grrllotalpa africana Palisot de Beauvois [in Japanese English translation 7520 1RRI library] Proc Kanto-Tosan Plant Prot Soc 21192-197

127 Kushwaha K S 1981 The black beetle as a pest of rice in Haryana India Int Rice Res Newsl 6(4)14

128 Lawani S M 1982 A review of tile effects of various agronomic practices on cereal stem borers Trop Pest Manage 28266-276

129 Leuck D B 1966 Biology of the lesser cornstalk borer in South Georgia J Econ Entomol 59 797-801

130 lever R J A W 1955 Insect pests of the rice plant in Malaya Int Rice Comm Newsl 1612-21

131 Li Ching-Sing and Shin-Foon Chiu 1951 The study ofthe rice gall midge Iachydiplosis orvza Wood Mason [in Chinese with English summary] 1Taiwan Agric Res 21-13

132 Iitsinger J A 1984 Assesament of the need-based application for rice Proceedings TechnologyTransfer Workshop 15-16 March 1984 International Rice Research Institute 1PO Box 933 Manila Philippines 6 p

133 Litsinger J A 1984 Environmental diversity and insect pest abundance with reference to the Pacific Pages 285-307 in Proceedings sub-regional training course on methods of controlling diseases insects and other pests of plants in the South Pacific 4-20 Oct 1982 Vaini Kingdom of Tonga Nukualofa

134 Litsinger J A RF Apostol and M BObusan 1983 White grub Leucopholis irrorata (Coleoptera Scarabaeidae) pest status popula-tion dynamics and chemical conllol in arice-maize cropping pattern in the Philippines J Econ Entoinol 761133-1138

135 Litsinger J A C de la Cruz F Raymundo A T Barrion M D Lumaban J P Bandong M S Venugopal F Paragna and E Balete 1982 Insect pests and insecticide reponse of dry-seeded bunded rice Pages 179-193 in Report of workshop on cropping systems research in Asia International Rice Research Institute Los Bafos Philippine

136 Litsinger J A M 1) Luinaban J P Bandong A T Barrion W Katanyukul and N Panda 1978 Insect pest management in rainfcd lowland rice agroecosystms of tropical Asia International Rice Research Conference 17-21 April 1978 International Rice Research Institute Los Bafios Philippines

137 Litsinger J A and K Moody 1976 Integrated pest management in multiple cropping systems Pages 293-317 in R I Papendick P A Sanchez and G B Triplett eds Multiple cropping American Society of Agronomy Madison Wisconsin USA

138 Loevinsohn M E 1984 The ecology and control of rice pests in relation to the intensity and synchrony of cultivation Ph D thesis University of London 36) p

139 Loevinsohn M E J A Litsinger N Panda M A Austria R F Apostol C Andrion and J Bandong 1982 A basis for predicting rice insect pest abundance in cropping systems Paper presented at the 1982 International rice research conference on rice and cropping systems 19-23 April 1982 International Rice Research Institute Los Bafios Iaguna Philippines 25 p

140 Lopez A W 1930 Repot of the Entomology Department Pages 145-172 in Annual report of the Philippine Sugar Association 1929-30 Manila

141 Lopez AW 1931 Entomology Department Pages 227-266 in Annual report of the director or research 1930-31 Philippine Sugar Association Manila

142 Lopez A W 1932 Entomology Deparment Pages 252-279 in Annual report of the Philippine Sugar Association 1931-32 Manila

143 Lungibill P and G G Ainslee 1917 The lessercornstalk borer US Dep Agric Agric Res Serv Tech Bull 539 27 p

IRPS No 123 January 1987 39

144 McGuire J U Jr and BS Crandall 1967 Survey of insect pests and plant diseases ofselected food crops of Mexico Central America and Panama U SDepartment of Agriculture International Agriculshytural Development Service Agricultuial Researrh Service October 1967 11 p

145 McNeill Sand T R ESouthwood 1978 The role of nitrogen in the development of insectplant relations Pages 77-9 in lliochemical aspects of plant and animal coevolution J B Ilarbornu ed Academic Press London UK

146 Maebara H 1955 Spatial distribution of lastinstiarlanaeof purplish stem borer (Sesamia iqferens Walker) hibernating in an upland rice field Bull Kagoshima Univ Fac Agric 438-41

147 Mallick E H and N G Hajra 1977 Response of rice plant to defoliation treatments at different developmental stages Indian Agric 2151-56

148 Mansingh A 1971 Physiological classification of dormancies in insects Canl Entomol 103983-10)9

149 Martell C G 1974 Catalogue of plant feeding insects of Mexico [in Spanish] Fitofilo 27(69)1-176

150 Martinez 0 J 1982 Principal rice pests in Ilanos orientalis [in Spanish] P~ages 80-94 in Curso deact nalization de conocimientos en arroz Villaniencis Instituto (olonmbiano Agropecuario

151 Martins J F da Silva F Ferrcira A S Prabhu F 1 P Zimmermann 1980 Use of pesticides for the control of the main subterranean upland rice pests esqui Agropecu Bras 1553-u

152 Martins I F da Silva F Fe eira rod BdamSilveira Pinheiro I Q2 Damage simulation caused by defoliators of upland rice [in Portuguese with English summary] Pesqui Agropecu Bras 171113-1119

153 Martins 1F da Silva N(iyuen van Tan and 1Ida Silveira Pinheiro 1981 Stem borer resistance in upland rice associated with plant morphological characters [in Portuuce Iesqui Agropecu Bras 16187-192

154 Masaki S 1980 Sumnmer diapause Ann Rev Entomiol 251-25 155 Masuzawa T 11 Suwa and F Nakasoji 1983 l)ifkleences of

oviposition preferencc and survival rate of two skipper butterflies Parnara guttata and PIchlpidai iathiais (Iepidoptera HIesperiidac) on rice plant and cogon grass New Entomol 3247-56

156 Maurya D M and C P Vaish 1984 Upland rice in India Pages 69-92 in An overview oftpland rice research Proceedings ofthe 1982 Bouakamp Ivory Coast upland rice workshop International Rice Research Institute IO Box 933 Mnila Philippines

157 Menon M G R R N Katiyai S Kuar and S S Misra 1970 Termites attacking the paddy crop Sci Cult 36294-295

158 Miyahara Y and 1 Nishikubo 1967 Behavior of young nymphs of the red rice root aphid Iholahsipiun ri liabhdontinalisSasaki in relation to movement from the leaf to the roots of upland rice [in Japanese] Pioc Assoc Plant Prot Kyushu 1363-66

159 Mo ljoa Tjien 1952 White grub of upland rice (Holotrichiahelleri Brsk) [in Indonesian with English summary] Contrib Agric Res Stn Bogor No 13134 p

160 Mochida 0 and T Suryana 1979 The occurrence of the brown planthopper and ragged stunt on lowland and upland rice ill Central Java and logyakarta in tile late of January 1979 Entomnolgy Survey Report No 15 Central Research Institute of Agriculture (CRIA) Sukamandi Indonesia I February 1979 9 p

161Monet C 1979 Varietal resistance to upland rice pests [inI French] Pages 35-39 in Annual report 1977 Institut des Savanes Department des Cultures Vivrieres Bouak6 Ivory Coast

162 Monte 0 1942 A caterpillar ofrice-fields[in Portuguese] Biol6gico (Sao Paulo) 8161-163 (Rev Appl Entomol A31414)

163 Munroe I) D 1979 Pest management strategy for hill rice in four resettlement areas in Sarawak Malaysia Pages 89-102 i1 Guidelines for integrated control of rice insect pests Food and Agriculture Organization Plant Production and Protection Paper 14 Rome Italy 115 p

40 IRPS No 123 January 1987

164 Nakasuji F M Ishii and Jen-Zon Ito 1984 Rice skippers in Taiwan and their life histories ZZrC (Taipei) Tech Bull 82 10 p

165 Navas I) 1976 Fall armyworum in rice Pages 99-106 il Ecological animal control by habitat management Proc Tall Timbers Con-ference 28 Feb-I Mar 1974 Fall Timbers Res Stn Misc Publ 6 Tallahassee Horida USA

166 Nickel 1 1 1961 Biological control of rice stem borer A feasibility study Tcch Bull 2 International Rice Rcarch Institute PO Box 933 Manila Philippines IIl p

167 Nigan P M and R A Vernia 1985 Insect pests of uplaild rice in Uttar Pradesh Int Rice Res Newsl 10(6)22

168 Nilakhe S S 1985 -cological observations on spittle bugs with emphasis on their occirence in rice Pcsqtui Agropecu Bras 20407-414

169 Nixon G EJ 195 1 lie association of ats with aphids and coccids Commonwealth Iusti lcte of Entonology Iondon UK 36 p

170 Nurdin F an Ishak Manti 1985 Study oit seedling fly of upland rice Interlational Upland Rice Moitoring Tour Report 26 lebruary-2 MarLI Sukaranti West Sumatra

171 Oka 1 N 1979 Ciltural conttol of the brown planthopper Pages 357-369 in Brown planthopper threat to rice production in Asia International Rice Research Institutc 1p0 Box 933 Manila Philippines

172 Ohni M and A Vilnaii 1975 Rice insect pests of Italy biology of Sipha grcria (Kaltenbach) and its control [in Italian] Riso (Milan) 2459-71

173 Otanes F Q atnd 1 I Sison 194 1 Pests of rice Philipp J Agric 12211-259

174 Painter R I 1951 Insect icsistance in crop plants University Press of Kansits Lawrence Kansas USA 520 p

175 Pantua P C and 1A litsinger 1981 Arthropod abundance in continuous vs biannual rice cropping systeis Int Rice Res Ncwsl 6(2)20-21

176 Pathak M ) and V A l)yck 1975 Sttudics ott insect pests of upland rice Pages 186-191 in Major research in upland rice International Rice Research Institute PO Box 933 Ilanila Philippines

177 Patil B R B M IHasabe and 1) S Ajri 198 I Chemical control of white gritb lIditriihinhorraia Fabr intfesting groundntut paddy atnd sorghum Indian 1Entomol 43232-236

178 Pawar A 1) 1974 (ollection and identification of insect pests of rice rlerminal repirt lInternational Rice Research Institute PO Box 933 Manila Philippines 72 p (tnimco)

179 IerrajU A aid A S Reedy 1965 Observations ott diapausc itt tie steti borer ohrice Rice News Teller (lai-Mar)9-13

180 Pont A C 1972 A review of the Oriental species of theri ona Rondani (I)iptera Muscidac) of econtomic importance Pages 27-102 in Control of sorglium shoot fly M G lomtani and W R Young eds Oxford atd IH1 Ptthl Co New I)clhi India 324 p

181 Prasad S C 1PChand aid M 1PSingh 1983 Incidence of rice gall midge in the upland plateau region of Hihar Oryza 20235-236

182 Rai P S 198 0 Nisaga simpex damage to rice in the hill tracts of South India Int Rice kles News[ 5(I)12

183 Rathorc Y S 1969 Gonocplhahli dcprssunt Fabr as itpredator of (tilo zonwIlhis (Swinhoc) and other moths in field cage at Patnagar Uttar Pradesh Indian 1Entotnol 3127

184 Reed W 1983 Crop losses causcd by insect pists in the developing world Proceedings 10th Internatial Congress of Plant Protection lrighton UK 174-80

185 Reedy M V 1981 Ateactogasterinhens(Walker)a new root-grub pest of tipland paddy and inaize Curr Sci 501076

186 Reveno M A 1968 Catalogue of the insect pests of economic crops in Colombia [in Spaiish] Asociacion Latino Americana de Entomo-logia (AIAE) 129-33

187 Reyes G M 1957 Rice dwarf disease in the Philippines Food and Agriculture Organiiation Plant Prot Bull 617-19

188 Rice S E A A Grigarick and M 0 Way 1982 Effect of leaf and panicle feeding byarmyworm (Lepidoptera Noctuidae) larvae on rice grain yield Environ Entomol 7593-595

189 Risch S J 1) Andow and M A Altieri 1983 Agioccosystem diversity arnd pest control data tentative conclusions and nizw research dirctions Environ Entomol 12625-629

190 Rivera C T 1956 Three organic insecticides in thecontrol of insects affecting upland rice Philipp Agric 39465-472

191 Roffrey 1 1972 Locusta outbreaks in the Philippines Acrida 1177-188

192 Rojas R 1) 1941 The effect of pruning at varying ages upon the yield of Ramairio Phipp Agric 29851-860

193 Roonwal M i 1979 Ternite life and termite control in tropical South Asia Scientilic Pnbl odhpr India 139 p

194 Rosetto C1 F da Silva Martins N C Schmidt and L EAzzini 1978 Damage caused by pasturespittle bugs 4Dvoi)lavopictaandDJ schach) on rice lBragantia Nota 6 (Matcli)25-27

195 Rosztto C1- S S Neto I- Amantc I) Iink 1Garcia-Vieira D M de SoLza N V Bansa t and A M de Olivcira 1971 Rice pests of Brazil [in Portuguese] Pages 149-238 inrFA()-International Rice Commission 2nd meeting of the Rice Committee for the Americas 6-11 l)ecember 1971 Pelotas Rio Grande do Sul Brazil

196 Rothschild G H L 1969 Obseratiois on the armyworm Spodoptera matritia acronrctoiiles Gn (Lep Noctuidae) Bull Entomol Res 59143-160

197 Rothschild G 1- L 1974 Parasites of the armworm Spolopiera mauritiaacroncitoihes in Malaysia Fntornopliaga 19293-299

198 Sachan i N S K Gangwar and 1N Katujar 1980 Field crickets damaging cages in North Eastern IHill Region Indian J Entomol 42526-527

199 Sands 1) P A 1977 The biology and ecology of Lepocorisa (Hemiptera Alydidac) in Papua New Guinea Res Bull 18 Departshyment of Primary Industry Port Moresby Papua New Guinea 104 p

200 Sarawak 1968 titomology division Pages 97-106 in Report of the research branch of the Department of Altriculture 1966 Kuching (Rev AIppl Eintomol A56603)

201 Satpathy JM M S Das and K Naik 1977 Effect of multiple and mixed cropping on the incidence of scine important pestsI Entomol Res 178-85

202 Sauer IHF G 1939 Notes Ott Elasmpnlpualtsligniosellths Zeller (Lep Pyr) a serious pest ofcereals in Sao Paolo [in Portuguese] Arq Inst Biol 10199-206 (Rev Appl Entotnol A28378)

203 Satinders 1L A 1 SKing and C L Vargas S 1983 Crop pests of Central Anerica with t bibliography [in Spanish] Centro Agroshynotnico Tropical de Investigacion y Ensenanza Turrialba Costa Rica (CATIE) Tec Bol 9 90 p

204 Sauphanor B 1985 Some factors of upland rice tolerttce to stem borers in West Africa Inseu Sci Appl 6429434

205 Sen P atnd S Chakravorty 1976 Stentflics in Haritighata Nadia West Bengal India [it Rice Res Newsl 1(12) 15

206 Senapai B and 1M Satpathy 1983 The seedling fly Atherigona orrzae Malloch is a pest of rice in Orissa (India) J Entomol Res 777-79

207 Shhjahan M S Alatn M Ah nad and BN Islam 1983 A new scarabaeid beetle pest of dryland rice in Bangladesh Int Rice Res Ncwsl 8(I)16-17

208 Sharma G L S C Modgol and R C Gautam 1979 Effect of depth of submergence ott incidence of bacterial blight and white grub infestation in transplanted rice Int Rice Res Newsl 4(6)12-13

209 Sharma S 1985 Scarabaeid beetle outbreak itt upland rice Int Rice Res Newsl 1(1(l)24

210 Shiraki T 1917 The paddy borer Schoenobhits incertulas Walker [achaku Agric Exp Stn Formosa 256 p

211 Silva A de Brito and 11P Magalhaes 1981 Insect pests of rice [in Portuguese] Circular Tecnica No 22 EMBRAPA Centro de Pesquisa t ecuaria de Tropico Unido (CPAJU) Belem Brazil 14 p

212 Singh S A 1979 Emergence of rice leaf butterfly as a rice pest in Manipur India int Rice Res Newsl 4(4)16-17

213 SinghS A 1983 Changing trends of insect pestsof rice in Manipur Int Rice Res Newsl 8(f)12

214 Siwi S S and P H van Doesburg 1984 Lenovorisa Latreille in Indonesia (Heteroptera Coreidae Alydinae) Zool Meded (Leiden) 58117-129

215 Soejitno J 1977 Reaction of some upland rice varieties toseedling fly eltherikonaexigua Stein [in Indonesian] Siri Hana I Penyakit 630-35 Lembga I osat Fenclilian Pertanian Bogor Indonesia

216 Socnardi 1967 Insect pests of rice in Indonesia Pages675-683 in The major inse pests of thu rice plant Proceedings of a symposium at the Internatital Rice Rec Wi itt S pternber 1964 The Johns Hopkins Press Baltimore Maryland USA

217 Soto P E A T Petcz and I W Buddenhagen 1976 Survey of insect pests and diseases of rice in different ecological zones in Nigeria Rice Entomol Ncwsl 435-46

218 Soto 11E and Z Ziddiqi 1976 Screening for resistance to African rice insect pests Varietal improsetnent sCmi n r Bouak Ivory Coast 13-17 Sept 1976 West Africa Rice l)evelopment Association Monrovia Iiberia

219 Squire F A 1972 Entomological problems in Bolivia PANS (PestArtic News Sumr) 18249-268

220 Stapely 1 II 1978 The rice leaf roller in the Solomon Islands Int Rice Res Newsl 3(l)13

221 Suenaga H 1937 On aphids attacking the roots of rice Oyo-I)oubts Zasshi 9150-152 [in Japanese] (Rev Appl Entomol A25745)

222 Suenaga H 1952 On the seasonal abundance of three subterranean aphids on upland rice Bull Kyushu Agric Exp Sti 1249-262

223 Sutynso Yas and Dandi Soekaria 1964 Beberapa hana yang ttenperang tana man padi tanah kering [in Indonesian] Rapat kerja Padi Tanah Kering 6 sd 8 Juily 1964 Kaliurang 9 p

224 Tallamy I) W and R F Denno 1983 Alternative life history patterns in risky environments in example from lace bugs Pages 129-147 in Insect life history patterns R F l)cnio and If Dingle eds Spring-Verlag 225 p

225 Tamura 1 1939 Observations on some insect pests of field crops [in Japanese] 1 Plant Prot 26414-420 (Rev Appl EntomolA27-556)

226 Tamura and Suzuki 1964 Influences of the early-transplanting practice in rice culture upon development of the rice stem borer Chilo siql)ressalisWalker Bull Ilokuriku Natil Agric Exp Stn 761-94

227 Tan J P 1924 The rice root aphid Drowhi ihirsoia Philipp Agric 13277-288

228 Tanaka 1 1957 Taxonomy and distribution of some subterranean aphids injurious to upland rice in Japan with description of a new species [in Japanese English summary] Botyu-Kagaku 22168-176

229 Tanaka T 1961 The rice root aphids their ecology and control Spec [full College of Agriculture Utsunomiya University Japan F3 p

230 Tauber M IC A Tauber R Nechols and 1Obrycki 1983 Seasonal activity of parasitoids control by external internal and genetic factors Pages 87-108 itt I)iapause and life cycle strategies in insects V K Brown and I Ilodek eds )r W Junk The Hague Netherlands

231 Taylor W F 1972 Effects of artificial defoliation (simulating pest damage) on varieties of upland rice Exp Agric 879-83

232 feotia T P S and S Natud 1966 Biotoomics of the rice skipper Parnaraniathiis Fabricius (Lepidoptera Hesperiidac) Indian J Entomol 28181-186

233 Teran F 0 1971 Rice pests of Santa Cruz [in Spanish Ministerio de Agricultura Sita Cruz Bolivia Bull 45 8 p

234 Than Itun 1976 Population dynamics of the yellow rice borer Tryporza itcertulas (Walker) and its damage to the rice plant MS thesis University of the Philippines at Ios Bafios Laguna Philippines 92 p

235 Uichanco L B 1911 Pests Pages 55-74 it Rice Philipp Agric 11-200

IRPS No 123 January 1987 41

236 van der Laan P A 195 1 Life history of the rice seedling fly Contrib Gen Agric Res Stn Boor Indonesia 118 15 p

237 van der Laan 11A 1959 Correlation between rainfall in the dry season and the occurrence of the white rice borer (Scirpolhaga innotala WIk) in lava Entomol Fxp Appl 212-20

238 Van Driesche R G 1983 Meaning of percent parasitismin studies of insect parasitism Fnviron Entomol 121 I1- 122

239 Venugopal M S and A litsingcr 1984 Fffect ofcarbofuran on rice growth Prot Ecol 7313-317

240 Vietnam Directorate of Rural Affairs 1962 Annual work progress report on crop improvement program of rice sugarcane vegetable and field crops (lor the period from July 1961 to June 1962) (11) 299 p (Rev Appl Entomol A51526-527)

241 Wan M T K 1972 Observations on rice leaf- and planthoppers in Sarawak (Malaysian Borneo) Malay Agric 148308-335

242 West Africa Rice I)esclopnient Association 1978 Annual research report Vol 2 Statistics Monrovia Liberia

243 West Africa Rice Development Association 1979 Annual research report p47 Monrovia Liberia

244 Williams ) F and CS lofgren 1981 Fli I illy l-468 anew bait toxicant for control of the red imported lire at Fla Entomol 64472477

245 Williams I) 1 1970 The mealybugs OFlomoptera Coccoidea Iscudococcidac) of sugar-cane rice and sorghum Bull Entomol Res 0109-188

246 Williams D) 1 1982 Pulinaria icervi (Signoret) and its allies ott sugar cane and other grasses Btll Entoinol Res 72111-117

247 Wirjosuiardjo S 1976 Study ofchenmical cont rot of insect pests ott upland rice Imiu Pertanian (Jogjakarta) 1323-329

248 Wirjosuhardjo S and N Wirarta 1976 Chemical control of the white grub tIolorithiatlltriin upland rice in M uneng Irobolinggo Indonesia) [in Indonesian] Imut Pertanian (Jogjakarta) 1330-332

249 Yano K T Miyake and V F Eastop 1983 The biology and economic importance of rice aphids (Henuiptera Aphid idae) a review Bull Entomol Res 73539-566

250 Yelp D and P Rhodes 19(4 lhe success of drilled dryland rice iii the Fiji Islands International Rice Commission Working Party on Rice Production and Protection Manila Philippines 3-10 March1964 Food and Agriculture Organization Rome Italy 13 p

251 Young W R 1981 Fifty-five years of research on the sorghum shootfly Insect Sci Appi 23-9

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No 101The economics of hybrid rice production in (hina No 113 Boron toxicity in rice No 102 Rice ratooning No 114 Energy analysis rice production systems and rice research No 103 Growth and development of the (Jeep water rice plant No 115 Production risk and optimal fertilizer rates an application of the

No 104 Faridpur acomputer-assisted instruction nmodel forrainfed lowland random coefficient model No 116 Consumer demand for rice grain quality in Thailand Indonesia andrice

No 105 A readingand listening comprehcnsion test inEnglish for non native the Philippines speakers applying for tiraining at IRIRI No 117 Morphological changes in rice panicle development a review of

No 106 Rice grassy stunt virus 2 a new strain of rice grassy stunt in tire literature

Philippines No 118 IRRI-Korca collaborative project for the development of cold-No 107 Physical losses and quality deterioration in rice plstprodnCtion tolerant lines through anthei culture

systemis No 119 Problen soils as potenial areas for adverse soils-tolerant rice No 108 (olptblication ofIRRI materials a survey of translators and varieties in South and Ssuthcasi Asia

publishers No 120 Changes isinall-farm iLtireliing techii)hgy in Thailand and [he No 109 Classification of Philippine rainfall patterns Philippines No 110 Contributions of modern rice varieties to rutrition in Asia No 121 Landforms and modern rice varieties No Ill Changes in rice breeding in 10 Asian countries 1965-84 No 122 Yield stability and modern rice technology No 112 Design parameters affecting the performance of the IRRI-designed

axial-flow pump

Page 16: UPLANDRICE. INSECTl--1i M T -E!R.EOLOU t RTANCE ,AND …delphacid.s3.amazonaws.com › 6958.pdf · The significance of abiotic factors in upland rice insect ecology will be apparent

16 IRPS No 123 January 1987

which lays its eggs hign on tile foliage (199) L acuta and L solonlonlesis oviposit at ground level the former on litter and the latter loose on the ground These egg laying habis explain theirtenvironmental prefcrencecs(214) In Japan the postdiapause adults of L chinensis migrate from moun-tainous areas to upland ricefields during floweiing (1 14)

Pentatornid bugs prevail in Latin America (8182 195) and in Africa pentatomid and coreid bugs arc equally cited (6 82) l)amage is often characterstic of the species (5)rcoris ventralis and Mnida spp are the common

pentatomid seed bugs of Philippine upland rice but they prefer lEchinochva spp grass seed to rice Vearaviritda occurs worldwide and hoften more abundant in the uplands(54) We saw few reports of seed bugs on upland rice but see(d bugs apparently opelrate cqually between by land and upland rice Good dispersal is necessary ior species that call feed for only 2-3 xon a crop and must first find those crops at ripening stages

Other pct species feed n flowers In Mato Grosso Brazil (194) and Santa Cruz Bolivia the panicle weevil Aeo-hariuia amllitarsis attacks rice during flowering (68 233)The larva e rice sten horers adutlts feed at the bases of spikelets caIusing empty grains Also inLatin America thecydnid -lkiituttatralusisa seed bug as anad tilt and nymph (194)

A number of beetles ecd on poilen - coccinellids lialrotia Aulac lhora and Monohpl-a (55) - but becatise riCe is self-pollinating and fcrtiliation occurs before pikelets open they pose no threat to yield

IST ECOLOGY

We now look at characteiistics of the life cycles of upland rice insect pests to lea rn hew each plays an adaptive role in upland rice ecology

Life history strategies Orzasativadoes not tolerate drodgh well so upland rice is highly seasonal normally grown in the wettest months of the year Usually rice is present for less than half of the year The rice-fiee fallow poses serious problems of survival to upland rice insects which have evolved att least four mechanisms to ovrcolne tle cvcl ic-l lack of a hostI) polyphagy 2) longevity 3) dormancy and 4) vagility (dispersal)

Poldt)agy lolyphagy is defined as having hosts of at least two botanical families oligophagy of more than one gents and ionophagy of only one ventus (40) - inour case Orvza Most wild rices however are illi tiC and theieiirc are normally far removed from upland rice hahitat othewise more would be alternate hosts for uplanJ iice insects Thte highly mornophagous rice pests shySciriol)haita hiceritlas Nilal)arvaiahiens Iand Nealo--tettiyvircsctws in Asia and Aalirpha secparatella and 1ioj)sis lngicornis inAfrica - occur on upland rice but

are more abundant in lowland culture They can specialize in rice because they are highly vagile and can attack all growth stages By specializing in rice monophagous rice pests can outcompete related but polyphagous speciesIntensive andI extensive lowland rice culture has favored monophagous species Riqwla alhinella (Cramer) the onlymonophagous species in Latin America is rarely reported on upland ice (I1)

Because of its limited temporal and spatial existence upland rice has not favored the evolution of specialized species Clhang(44)shows evidence that upland rice has only recently been cultured by man from the lowlands thus pests have had little time to fully adapt to the crop All but the lowland-adapted insect pest species have alternate planthosts to rice Root pests with limited mobility are highly polyphagous Pests that specialize in one growth stage eg secdling maggots seed bugs are oligophagous or polyphagous

Most alternat hosts of rice pests are annual grasseswhich also are seasonal Upla_rice is also a host but not the preferred one of the most highly polypliagous spcciessoil pests grass hoppers armvwarrs and Vearaviirielda

The d ry season poses igreat obstacle to ani upland riceinsects survival and only sonic perennial plant hosts are suitable food during the off-seasoin Plolyphagy allows upland rice insects to suLrx-ixe during thC rainy season when plants- both annual and perennial -areactivelygrowing and therefore more nutritious

Loiugevitr The ability of aiinsect to live 2-3 mo without undergoing dormancy helps species sich as root-feedingwhite grbs (as larvae) and secd-feeding bugs (as adtilt) to survive unfavorable periods White grubs arc not highly fecund and their life strategy is to improve survivorship of limited progeny Their subterranean habitat hides them from manv natural enemies lBtut root tissue is not highly nutritious so they ltilst eat great quantities and need a largeinsect biomass to digest it Other soil-inhabiting rootshyfeeding insects have short life cycles but are more fecund

Seed pests need greater longcvi vas adults to locate ahost at early grain dcclopment arnd give sedentary offspring time to develop Few groups of upland rice insects live longin active development Polyphagy and longevity only increase their ability to survive during the rainy seasonOther mechanisms are needed to survive the dry season

orantc In the tropics the nonrice season is the dry season while intemperate areas tile off-+season is winter Many upland rice insects undergo dormancy inlsummer (aestivation) or winter (hibernation) Dormancy is simplyinactivity during winters with prolonged temperatures below 10 C insects find shelter arnd cease movement Activity is resu med with warmer t empcera tu res A deep state of irnactivity termed diapauise occurs in sole species that prepare physiologically (havxing low metabolism and convcrting gonad s to fatty fuel reserves) for the unfavorable season (148) They enter diapaUse inanticipation of unshy

IRPS No 123 January 1987 17

favorable weather and resume activity after favorable Armyworms skippers and locusts actively travel hundreds weather begins usually in response to photoperiod or of kilometers in air fronts such as intertropical convergencethrough a delayed response to a stimulus such as heavy zones to descend onto upland rice fields far from their rainfall (237) Not all individuals of a population have the breeding grounds (31) Several generations can build ip on same threshold for environmental responses for entering or alternate hosts in tile grass family before rice is plantedterminating diapause -- this is seen as a mechanism to These large insects are active fliers Smaller insects such as prevent total mortality if a false cue appears such as a thrips aphids and planthoppers passively migrate long sudden warm spell in winter or rain in the middle of the dry distances by the wind season (154) Otherspecies ean readily travel tens of kilometers - stem

Probably more upland rice insects undL go dormancy borers leaffolders leafhoppes cutworms spittle bugs leaf than we realize Candidates are Atherigona spp root beetles seed bugs ants and termites aphids root weevils crickets leaf beetles thrips and seed The least dispersive species are white grabs mealybugSbugs Insects known to entervarious states of aestivation are seedling maggots crickets weevils and gall midgesleptocorisaspp adults (199) Maliarphascparatellalarvae Each species has evolved a unique set of attributes to (9) Scirpophaga innotata larvae (117) Chilo partelhtsv enable it to survive and adapt to the changing uplandlarvae (21) ScirpopThagainccetulas (179) white grub larvae environments Several evolutionary avenues lead to fitness (134) Patangasuccincia adults (6 82) and Schistocerca (Table 2) but there is no reason to believe one set of gregariaeggs (82) attributes is better than another (224)

Insects known to enter hibernation are LodeIphaDstriatellus nymphs (124) Chtio Tvlpressalis larivae (125) DroughtSesaia inferens larvae (146) but not S calavisis(2) Well-drained rainfed upland soils are subject to drought ifIDiaraea saccharalis larvae (75) Scirpophaga- tcerlttas rains do not fall within 2-3 wk Prolonged drought followedlarae(5 lacchrli s lae (75)saSlOhagasshoperseggs (82)and felo pidas athias pOpae (232) by favorable rains stimulates armyworm and locust outshybreaks but mealybtigs root aphids and thrips become

Lcpmotorisa oratorisand L acuta adults aggregate on numerous onlyafter dry spells for two reasons First heavy trees sugarcane or other shady moist sites L paluwanmsis rain normally kills soft-bodied foliage-feeding thrips and confined to Sulu and Palawan Philippines lives on the mealybugs and soil-inhabiting root aphids and root mealyshygrass Brachiariamutica (Forsk) Stapf in the absence of bugs Drought eliminates this source of mortality Second rice The adults can live for 5 mo When aestivating their the rice plant responds to drought by breaking down rate of metabolism drops and they do not feed They will proteins into soluble N compounds which enter the phloem however puncture plant tissues of their aestivation host in and are taken up by these sap-feeding insects (245) Greater search of water This behavior has led to errors of the host nutrition therefore leads to greater survivorship and range of rice bugs (I) Aestivation quickly terminates with fecundity rain probably meaning aestivation is not deep Small rice area

Some stem borers aestivate as mature larvae in rice Except in the highly mechanized large landholdings of stubble or in straw stacked as livestock feed However Latin America upland ricefieldstypically are patchy within moriality increases proportionally with the duration of the a highly diverse flora The small fields result in higher ratios dry season even if the stubble remains undisturbed If the of perimeter to area than with the typically larger lowland stubble is plowed for a crop following rice few stem borers ricefields Upland ricefields themselves may be intercroppedsurvive Rainfall terminates dormancy but it normally with cereals legumes or root crops Mixed intercroppingtakes several weeks for moths to develop and emerge with a wide variety of species also occ in tribal slash-andshy

Last-instar white grub larvae tunnel 1-2 m deep in the soil burn agriculture (50 74) to construct pupal chambers in which they aestivate The Upland ricefields tend to be small because of the labor larvae are very sensitive to dryness so pupal chambers are needed to clear land in slash-and-burn areas and to till land sealed to coierve body moisture Aestivation terminates to minimize weeds Rice requires more tillage than does with the first heavy rains but as in stem borer 3-4 wk pass maize because rice competes less well with weeds Uplandbefore adults develop rice is more a subsistence than a cash crop and small areas

Insects may respond differently in each region to cold (I ha) can feed a family temperature or drought Biotypes or loal populations may Small fields favor some insect pests such as some ant evolve (56) Local populations of Scirpophagaincertulasor species that can forage from more permanent field borders Patangasuccincta may have become adapted to prevailing Trees provide food and mating sites for white grubs (134) as conditions well as aestivation sites for LetTocorisa spp (199) Field

iagiliti The ability to disperse combined with high border grasses plus tIhose within a ricefield provide alternate fecundity orshort life cycles enable some upland rice pests to food for all but the highly monophagous upland rice better exploit the temporarily favorable upland habitat colonizers and a habitat for natural enemies

0 z

Table 2 Life history patterns of upland rice insects or insect groups

Host range Longevity Dormancy Vagility FertilityMonophagy Oligophagy Polyphagy Low Medium High Yes No Low Dispersive Migratory Low Medium High(Oryza spp) (grasses sedges) (angiosperms) (lt3 wk) (gt2 mo) (lt1 kin) (lt50 eggs) (gt500 eggs)

Ants xTermites x x x

Mole crickets x x x xx x xField crickets x xx

White grubs x x x xx xRoot aphids x x xxMealybugs x

x x x xxSeedling maggots x

x x x x xLeaf beetles

x xx xArmyworms x x xx x

Thrips x x xx xChilo spp x x xx xMaliarpha x x x x

xLocusts x x Xx x xGall midges x x xxiVephotettL iirescens x x xx x x xLeprocorisaspp x

x xLeaftolders x x x

Sogatellafurcifera x

xx x x x x x x

00

IRPS No 123 January 1Q87 19

All this appeals to favor insect pest buildup but the notwithstanding the release of stored nutrients in response contrary is true Loevinsohn (138) showed that insect pest to drought that temporarily benefits philoem feeders populations respond exponentially with the proportion of land devoted to rice up to about 75 of the area then the M POR ANCE I I P1A 1) RWI P1St S

rate of response declines In most upland rico areas tie rice crop occupies less than 50i of the area- therefore the Visits to upland ricefields normally reveal 1kw insect pests potential for population buildup is low This relationship Loevinsohn (138) and Locvinsohn ct al (139) compared may not hold for seed pests that can locate small ricefields insect abundance using annual light trap atchcs in upland

Another outcome of the highly diverse flora and phical rainfed lowland and irrigated lowland sites in the Philipshyenvironment of upland rice isthateach upland ricearca has pines Although only one upland site was studied data a unique composition of insect pests showed equal or fewer insects at the upland site than at the

For example in Tanauan Batangas Philippines the rainfcd lowland sites and fewer than at the irrigated sites main upland rice pests are rice leaffolder(RLF) armvworm (Table 3) M Lltiple regression analysis showed that the key Aythntnaseparata WBPH ants Soli psis gentitataand factors explaining insect abundance among a set of cropping white grub Leucopholis irrorata Sixty kilometers away in intensity variables were the number of rice crops grown per Real Quezon the main pests are seedling maggot Athe- year followed by area devoted to rice Table 3 shows that rigona orae flea beetle (Thacitcmnra hasalis thrips the upland rice site had only I rice crop perym rand that rice Stenchaetothrips hiurnmis and rice bug Leptocorisa was planted in only 20Vi of the area Fertiliter and oratorius In a third Philippine upland rice site in Claveria insecticide use had little bea ring on the trend The armters at Misamis Oriental Mindanao the main pests are ants the upland site applied 60 kg N ha higher than at most Solenopsis gemtinata seedling maggot Alieriona orvzae irrigated sites white grubs Holotrichia in)iUaiuoan t and Lucopholiv Insect occurrence at the upland site was omparable to irrorata stem borers Sesamia ifiArelr (hio auriciliusC that at the Cagayan rainfed lowland site parlicularly in 1982stpessallis Scirpophaga incertulas S innoiata and when drought prevented many farmers fron planting(only

4c ona chrrsurapella root aphids Tetraneura niriah- 40(of the rice area eventually was planted that year) M ore dunrinalisand Rhupaosiphunm riahbdminalis lcaffolders favorable weather occurred in 1980 and 1981 which Marasniapataltis and Chaphalocrocis Medinalis army- supported greater rice areas and consequently more insects worm MIthinna separata and rice bug Leltucorisa The most intensive site for insect pests was South Cotabato oratoriusThe great diversity of insect problems is typical Mindanao where farmers plant 23-25 rice crops per year fer upland rice and makes control efforts more difficult with irrigation The Zaragoa Nutva Ecija site also had particularly regarding breeding for insect resistance high pest abundance and at this site the fields were highly

asynchronous Asynchrony was the third most important Low yield potential Iactor in explaining rice insect abundance Most upland rice is grown on low-nutrient and mineral- Rice cropping intensity in upland areas is bound to toxic soils which when combined with eiratic rainfall remain low Only one rainid upland crop is possible per make upland rice a highly riskycrop for the farmerto invest year (Farmers in Claveria Misamis Oriental Mindanao in costly soil amendments and land management Often Philippines tried asecond upland ricecrop i 1984a ycarof upland rice isgrown far from markets making inputs even favorable rains but no harvest was possible b-cause of more expensive Farmers growing a subsistence crop hosi- drought blast and birds) Rice area isaso bound to remain tate to invest in costly inputs to raise yields low because of the high labor and power rcquirelmenlts to

Even having varieties that could double existing yield prepare the land (because of weeds)and because upland rice potential under present management levels would probably wvill not compete with lowland rice as a cash crop Some not prompt many farmers to use purchased insecticide upland rice sites with prolonged rainy seasons howeverare unless tile risk of crop failure were reduced such as it was planted asynchronously such as the Clacra site with irrigation systems in lowland rice The seasonal abundance of Upland rice insect pests

The Tanauan Batangas site is atypical for upland rice attracted toa light trap wasgrapied for crop year 1980-81 in The soils and rainfall pattern are favorable and because of Tanauan FBatangas (Fig I 2) W13P11 was he most its nearness to Manila farmers have cash resources ftm common rice hopper and its population peaked at the sales of vegetables to purchase tertilier They applied an midgrowth stage as it does in lowland rice D-Vac suction average of 60 kg Nha after particle initiation and obtained samples showed an earlier peak than that from a light trap 25-35 tha yields The high fertility isone reason why RLF indicating emigration W13PI usually emigrate after the armyworms and WBPH developed into large numbers vegetative stage Populations in 1980 reached levels that Farmers however used no insecticide on upland rice caused patches of slight yellowing in some fields WBPH although they did on vegetables probably cause hopperburn in Batangas They immigrate

The poor growing conditions fOr rice generally mean year-round as shown by low levels in the light trap BPH poor growth and low fecundity for insect pests fIeding on it however were scarce in the light trap and not collected with

Table 3 Abundance of rice insect pests and factors in rice cropping intensity of Philippine upland rainfed lowland and irrigated lowland locations where light traps were operated daily for atleast I yr -I

0Rainfed Rainfed lowland Irrigated lowlandupland

Pangasinan Cagayan lloilo Santa Maria Nueva Ecija South Cotabato Tanauan Caaringayan Caaringayan Bangag Bangag Bangag Buray

LagunaCale

Batangas Manaoag 1980-81 Santa Motiica 1982-83 Cabanatuan Zaragoza Avancena NaniamanSolana 1981-82 1982-83 Oton

1980-81 1979-80 Oton 1981-82 1981-82 Koronadal Koronadal1980-81 1979-80 1979-80 1983 1983

Insect (kerosene light trapNephotcttixcatch nolyr)virescens 650 1240 500 1800 330 210 3400 1600 160N nigropictus 210 - 360 18400 5500 2900- 140 90 20 - shy 1700 -Reciliadorsalis 710 - 3200 1700180 890 630 140 170 2400 800Nilaparratalugens 130 690 - - 4200 4600300 520 750 180 60 3300 1600 740 310Sogatellafiircifera 1700 31000 13000 8200310 210 3300 610 270 980 1600 6300 -Cyrrorhinus - 12000 6700600 540 3200 290 160 330 120 410Scirpophaga spp 50 410 620 - - 21000 12000150 180 220 270 30 1000Cnaphalocrocismedinalis 70 70 540 610 270 11000 6100200 140 100 6 60 250Chilo suppressalis 0 60 60 1900 300 200- - 30 20 0 6 3 160 -C auricilius - 0 00 - - 20 3 60 20 3 0Sesania inferens 0 - 80 60 3 80 10 51 -Cropping intensity _ -

Rice crops (noyr) 10 10 12 10 10 10 13Rice area (7c) 17 19 20 20 25 2320 85 85 60 70 40 85 85 90Fertilizer (kg Nha) 60 20 85 80 70 7020 0 0 0 30 30Insecticide (no of 40 60 40 300 04 04 0 03 300 13 13 30 50 40 33 33applicationscrop)

__________

IRPS No 123 January 1987 21

Light trapped Insects Drytnd __ 2(notrap-nght) Suctioned Insects (nom )

50 -)o 9 jO40- SOW ffurciWO Light trp

D-Vc suction

20 to 4

20

Egg porasitizahion (M) 75 60

300 SOOampReOftwciffr

45 30shy

15

-insects (notrop-night) 4 Nileporvata Iuen5

Egg parastization()

Light trapped insects (no trop-nght) Suchoned insects (no m 2 )

6 8 Cytorh~nuls hvidipennii 0

4 05 2

50 0 Insects (noltrop-night)40-ui Ipanh pe g

Moohase 1 Seasonal abundance measured by light trap

Rainfall(etod) and D-Vac suction sampler of whitebacked

to ~ hopper Nilaparvata lugens and lea ffolder60)-t Cnaphalocrocis mnedinalis andplthpe g20 L I II I I I parasites and mtrid predator Ortorhjntus livi-Apr May Jun Jul Aug Sep Oct Nov Dec JaIn Feb Mar dipennis set against cropping pattern daily1 9 80o 19 8 1rainfall and moon phase Tanauan Batangas

1980-81

the D- Vac Cyrtorhinus build up on the crop coterminous are collected in light traps during full moon (109)with WBPH buildup Cyrtorhinus disperses during the Because upland rice shares most of the lowland rice pestsrainy season RLF not readily collected in light traps was and those pest species are generally less impotant to uplandprevalent only during the rice crop rice than lowland rice there is a general belief that insect

Nephotettix virescens was more abundant than Nnigro- pests are less important onpicLus in the light trap collections and on the crop GLH upland than lowland rice Arcview of limited data on upland rice shows an averag of population peaked at the reproductive stage as did zigzag 10-21 yield loss from insect pests on a par with that ofleafhopper (ZLH) Crtorhinus tracked the populations of lowland uee determined by the same methodology (132) planthoppers and leafhoppers Both GLH species were (Table 4) McGuire and Crandall (144) made a similarcollected year-round in the light trap indicating immigra- estimate for Central America Cramer (51) howevertion from nearby irrigated rice areas Possibly because of the estimated 35 yieldlow populations loss in South America based on ainsect number did not correlate with similar pest complex - stem borers and seed bugs shymoon phase although reports indicate that more hoppers attacking irrigated rice in North America This estimate did

22 IRPS No 123 January 1987

Light trapped insects D~~nd (no trap-night )

St plht Suctined insects (nom 2

)

D-Voc suction 1520

1o

0 Lih trap 5 ~105 o5

00 Eggparositizotion()5040 NephOtgtha virneens

30 20

I010 A-Insects (notrop -night)

4 Nephatelh nigropccent i

2 o I vV^J - -- ^ VA____ f _

Light trapped insects (no trap-night) Suctioned insects (no rnE

10 -1208 ReCdO doRsol

6 15

4 10 052 iwxALAiamp 0

Light trapped insects (notrap-night) 2Suc oned nsects (nom )

8 CyrtorhmnusI6o nn]10 i0075

4 05

2 0 0

025

2 Seasonal abundance measured by light trap Roinfall (mmd) Moon phase and D-Vac suction sampler of green leafhoppersNephotettix virescens and N ngropicrus and I zigzag leafhopper Recilia dorsalis their egg 60

parasites and micid predator Cyrtorhinus livi-dipennis set against cropping pattern dailyrainfall and moon phase Tanauan Batangas1980-81

40

5 Lf Apr May Jun

1980 Jul Aug Sp Oct Nov I

-aMar

1981

not account for soil pests and was not based on field data in the absence of pests Carbofuran granules are phytotonicAkinsola (I1) cited mainly data from irrigated rice areas in to rice (239) The prevailing hypothesis is that carbofuranAfrica but estimated yield losses of upland rice between 15 prevents soil bacteria from consuming fertilizer The phytoshyand 30 from insects However losses in Thailand over a tonic effect is through greater availability of fertilizer to the4-yr period ranged from I to 13 averaging 5(106) plant Denitrifying bacteria are known to be more active inYield loss data however should be interpreted with aerobic than anaerobic soils therefore it may not be merecaution (184) The Philippine data were all derived from coincidence that the highest recorded yield losses (22-69)trials on farmers fields under farmers normal agronomic occurred in those trials with 12-30 kg basal Nha On themanagement practices except for Bukidnon and Capiz other hand contro with carbofuran granules may not be aswhere researchers used fertilizer Batangas farmers are good as a seed treatment against seed pests and perhapsatypical because ofhigh levels of fertilizer used The trials in yields would have been higher if a systemic seed treatmentBrazil were carried out at an experiment station but insecticide had been used conditions were similar to those of local farmers The 1979 and 1980 trials in Batangas had two full-

Carbofuran granules were used in every trial as a broad- protection treatments one using carbofuran granules andspectrum insecticide applied basally with the seed An the other using bendiocarb WP as a seed treatment Plotsobjection to using insecticides to measure yield losses isthat protected with bendiocarb yielded higher than those proshyinsecticidescandirectlyorindirectlystimulateplantgrowth tected with carbofuran granules The other trials in the

Table 4 Yield losses to insect pests and yield responses to insecticide in upland rice in the Philippines Brazil and Upper Volta

Yield (tha) Yield loss (5) Location Year Cultivar Protected

with Protected

without Check Total Vegetative Reproductive Ripening

Basal N (kgha) Reference

carbofuran carbofuran Protected No with carbofuran

carbofuran

Philippines Pili Camarines Sur Tanauan Batangas

Pangantucan Bukidnon

Dumarao t-apiz

BrazilGoias Minas Ge~ais

Upper Volta6

1975 1974 1974 1975 1975 1976 1977 1978

1979

1980

1979 1980 1979

1977 1978 1979

Bursiging puti Kinanda Dagge Kinanda Dagge Dagge Dagge Dagge UPL Ri5 Dagge UPL Ri5 Dagge UPL RiS UPL Ri5 UPL Ri5 UPL RiS

IAC47 IAC47 IAC47

14 33 11 36 35 31 30 27 37 26 40 30 40 42 46 35

19 21 09

--

-34 24 24 27 --

29 43 31 42 -

47 25

---

10 28 05 27 27 30 26 29 33 28 33 29 43 23 36 11

15 15 06

29 15 55 25 23

3 13 -7 11

-7 18

3 -7 45 22 69

24 28 35

--

21 -11 -20

4 -

-3

23 7

-2 -23 56

-

-

-__0

6 --

-

-23 14

5 0 8 2

22

-

--

0

-

--18 11

3 -10

4 10

7

--7 -

14 -

-1 6

10 -5

8 9 7

-

--

0 0 0 0 0 0 0 0 0 0 0

30 30 30

12 12 12

99 99

100 100 101 101 102 102 102 102 103 103 104 104 104

66 66 66

Farako-Ba

Mean

1977 1978

IRAT10 IRATIO

38 66

--

40 43

-5 35

21

-

10

--

--

--

- 242 243

Z

tJ

-w

24 IRPS No 123 January 1987

Philippines which had two treatments offering protection In the 9 Philippine trials where yield losses were measuredwith and without carbofuran granules relied on further by growth stage 10 of the yield loss (21) occurred at thecontrol with foliar spray and not on seed dressings The vegetative stage and 517 occurred at each of the reproshycarbofuran-protected plots had an 18yield los compared ductive and ripening stages Ibis result is similar to thewith only 10 with foliar sprayS This yield difference is pattern of yield loss in lowland rice measured by successiveprobably due to differences in protection against sown-seed treatments where the crop is unprotected at each growthpests (foliar spray do not control sown-seed pests) rather stage but proticted during the other growth stagesthan phytotonic effects became basal fertilizer wa used in However the insect complexes r sponsible for vegetativeonly 2 of the 10 comparisons stage yield losses in upland rice are entirely diftfrent than in

The recorded yield losses were not related to yield lowland rice The key vegetative stage pests in lowland ricepotential (Fig 3) One might expect higheryielding crops to arc aquatic whereas those in upland rice are sown--seedhave higher percentage of yield losses but this ws not tile (sown in soil) root and seedling pests This dift rence maycase One reason is that insect damage exacerbates plant explain why agricul torists have overlooked tile ilmportanccinjury from drought stress low yielding fields from d ought of upland rice insect pests Many reports of upland rice pestsstress would show disproportionate differences between in Asia ha Ve focused on major lowland species (46 108)insect-protected and -unprotected treatments Shoot and It isalso surprising that tei vegctative stage of upland rice root pests would be more responsible for losses from would record the highest yield losses becatuseseeral studiesdrought stress than sown-scd pests A second reason have shown that upland rice can readily recover from foliageconcerns seed pests which would have a greater impact on reinoval during that stage (117 152 165 188) Upland ricelow tillering low yielding varieties such as )aggc Low poscsses a hitgh obilitV to compensate for ca rly loss oftillering varieties nnot fill in the space created by removal foliage and will even be stinulated by foliage removal toof seeds oweer 1) R i 5 is high yielding because it tillersP1 produce a higher viel than plants ithout any foliageactively which can close the canopy to provide greater weed removed (63 192 231) [ie reason for this apparentcontrol discrepancy hovce is that yield loss at the vegetative

stage does not octUr frol leat arca loss but fron pestsYield loss () removing sown seed and feeding on devcloping tillers and roots -- damage that cannot be readily conpLtnsatCd for

70 - Measuring yield losses in upland rice cnvironrnents is further complicated by a report from Brazil that greater yield losses occurred in response to better insect control60 - =-09 as Insecticide seed treatments resuled in denser plant stands

(n=21) which in turn created a selectively more favorable environshyment for blast (69) This resuIt was parlticurla rlxexacerbated50 when cirbofuran granules were used in seed furrows Aside

ffrom a greater plant stand ca rbofuran iay have auigmented the N lecl and stimulated blast If so ca rbofuran should be40 - replaced with another insecticide or the seeding or fertilizerI rates decreased

Other methods to determine the impact of upland rice30 shy pests wvcrc to interview farners or to develop single speciesbull bull correlations of population levels and yield loss Feneira (66)2 NIM)st (75(1) Golas farmers reporteddid both in Brazil

20- insect losses in ufpland rice and 251 said those lossesS avcragcd 37(1 [his result is consistent with tie yield losses 0 0 measured by field trials in the same statc(Table4) For every

I0 4 of seedlings removC d by leaf-cutting ants a I 7yield loss is predicted Similarly Ferreira (66) found that ai

0 - average of 53 thrips per panicle before panicle initiation meant 3 times more unfillh d grains

-IO0 I 1 1 I I-101 0 2 3 4 5 6 7

Yield (tho) C)N [ROt IM FI IODS 3 Correlation between upland rice yield and yield lols caused by Upland rice insect pests may be controlled by culturalinsects measured in 21 trials in the Philippines Ivory Coast andBrazil genetic biological and chemical methods

IRPS No 123 January 1987 25

Cultural control dry season plowing will have little effect on them Plowing Crop husbandry to reduce upland insect pest populations when the soil is wet from the first monsoon rains will expose can be classified into practices effective in a single field and soil insects that tunnel close to the soil surface to predatory those effective only at the community level Cultural birds chickens dogs and even man (field crickets nole controls should come first in a pest management program crickets and white grubs a re delicacies in man tra ditional They have broad stable effects as pests have little pos- diets) sibility of overcoming them through selection of biotyps The greatest degree i1 inIscct control fron tillage comes (138) On the other hand some cultu ral controls work only from plowing soon after rice ha rvest when most soil insects with high labor or power inputs and connunity-wilt lie close to the soil surface anrd stern borers are in the rice methods require coordination of iany farniters stuble Whitegrubsare mature and tillage exposesthem to

Pantin time The luxury of being able to shift the time of predators Tirnelv control will prtec the crop following pla ntring upland rice is open onl hr la rniers in areas with a rice and lowr the populationIfor succeedinig years Plowingprolonged rainifall pa ttern lI copping intensity or underthe ice stulble helps it decompose killing sten borer rnechanied land preparatiot Gcnerally planlting as soon larvae inside as the rFiinv season begins will lower populatiois of 1110st [requent tillage destroys ait nests bit not termites insect pests Insect populationsarelo aftcra dlvseoll or Tillage and clean culture howes er rcliove food for winter fallow delayed plaiting lets them build tip on termites which then mtay attack a young rice crop Scarab alIternate hosts that gi vigorotsly with the first seasonial beetles also prefer to miposit in eceitly tilled fields Noshyratiris This first flush iof weds cros lItxuttriartlv with iceCult is dibbling faors the survival oftillace Clre Such rineralied N released during the dr season aid made stils i aels tnd insects thtt pttpate or lay eggs- in the soil

available to the plants with the rains It iliso igrtmitcally Zero tilla hio eerl conserves predators ofthe GIH (37) advanttgeous for the rice ciop to tap this trlLItlaI fertility nt it mcttoi lo lessen the impact o0 soil insects and grow vigorotsly to compete with wrus and to tolerate brotdcasting seed or planting in ftltrrows is prefc ired over pest infestation dihlirig in hills Crickets are large enotlgh to eat seedlings in

Larly plantngs terd to escape seedling maggolts Upland a hill atrd tillering normnally ca iriot fill in tlhe missing space rice in Bitangas is planted in Ma and ine itli the Iirst Oftcn dibbll rd is not well covered and iseasily found by rains Fairners are highly mtotiated to plant early tocscape foraging insets ioIdents and birds drlitght and typlotns ricar harct Arn Atrgtst planting Plan uit sii Incrcatsini sced density protects tile cropwould be evcely attacked by It riim orivzo At tti againrst seed and seelling pests especially ants particularly other Philippiie tipland sites Rea l Qteot antd (laelit if the crop will germinate and ermerge quickly M isainis Oriental seedlig mnaggots arnrige ric boe t se hr trftinll itI India rice intcrclopped witi cotton or farnes have a longer growing season and plnit 2 rin or pigeon pci had lower 6IIt and WBIlII poptrlations than mote after the ortset of the rainy season liaaiangas ftrmners rice alone (211) NfailC arid Urplald ir_ 2re coilltori in the plant ca rly becia use ofa shorter raiinfall period arnd the desi re Philippines he chatngC in pest status folii iniercropping is to lliarvet before the typhoor sea son peaks ltighly locatioll-specific (133) cmianarid the net elfc be either

Early plantings cornbind with early-iatiuring varictics nil bcnelficial ortdetriential (189) For example BIatangias provide higher crp tolerartce for white grubs Aestiatior farmers intercrip Itro in their ipland ricefields A liornshyterminates with the first rains but 5-6 moare ineeded for the nrii iotd-Ahis i (I iiiacius) which norlially feeds resting larvae to develop into beetles emerge and lay eggs on taro ill feed or miarbv rice plants Intercropping and for larvae to reach the danriigii g ird-instar stage probably would not afft0 seed pests A crluanion crop (134 In lataigas this normually iseiough tiite for harvest planted ith rice and haing gleater tolerarnce for root

It Brazil howecver delayed planting is a suggested dariage or being toxic nayact isi trapcrop to root-feeditig cultural control for the lesser cornstalk borer and insect pests Acial-fei i ig pests voultId be most affected by pests favored l drought (66 195) Cropping internsity in intrer ropping Shoot aphids leaf beetles sten borers and Brail is low and delayed plraing places the rice crop in the hopplers would be likely ca ntidates to note effcts On the niost stable rainfall period reducing likelihood old rought othlt halnd the corlipalior crop stch~ sitniaie Iiiv be more

In rilniy arca s rinmirsare roi atl to plant other crops benehited than rice from initecropping ( 137) before rice and althNgh it woultl be desirabh to avoid 11echi Weeuling luring the first month after clop infestations by planting rice cuatl lariners lack the labor cstablishent will force pcts such is anyvworris which and time to do so ol a low priority crp prefer grassy cetds ontor tle vorling rice Soil iampsecs will

77age Uplild rice Stils ae light textured and Call be mvnye to the rice crop Cleani citrltirc also lay lore territes plowed duiring the d rv seasor notl only to grin time for to attack I u cetincti inrice ceds i ay he le optinal planting with lie fitst Inilsoon riiains but also to desiccate solition 13) soil insects arid weeds such is ntsedge that foim rhizomes Irtlizr Itneatsing N Iciiliier oin hlwland rice has Htowever becta use most soil insects lie below die plow layer fAtwvored Irigher populains of pliithoppeiv Imlhoppers

26 IRPS No 123 January 1987

and leaffolde rs However because of the need to temper fertilizer use on upland rice o prevent blast fIrtilizer rates will be lower than is comonia r for lowlad rice causing relatixely less hopper and IaIfTfoide r population buildup However inupland rice areas without a history of blast such Is Batangas farmers apply high rates of N and conscqtrently the crop has high infcstation levels of WBPI]and RIF The fhrmrcrs leec insect buildup by splitting the fertili11r in0to two to three applicatiois bit do not applV fertilizer at the vegetati estage because this prodrces a tall crop which illeasily lodge during typhoons Greater crop fertility leads to greater insect survival (better nutrition) larger insects (to cat no re foliage) arid greater fec titd t (to lav nmre eggs) (128 145 171) Not all insect species respondequally hovr Increasing rates (if N l anrid K increased Dialraa s-acchralis and (iaeiurema flea beetle but reduced thrips (66) Zn rcdLuced deadheart density from the lesser cornIstalk borer

Fnldingi Although most Upland rice areas do nIot hawxcaccess to wa ter floodiig the fields is rRrrnriierdcd in areas of Brazil or root bug and white gib control ( 195)

Crop rotatioti Tillage before planting a crop after rc harvest will control stern boreis and other pests renain n the stubble arid will unearth soil pests Planting a non-gratriiaceous crop after rice is recommncrded for termite control (66) Crop rotation is less nttdcd in upland rice culture than in lowlands because atcrop-free period is assured even after harvest of crops following rice

t1ulchilt iryl plastic irli irig designed for weedcoitrol ii upland rice inJapan resulted in ntr highanri liy infestation of (ihilo sulytresvais (93 9-1 95) NIulched Upland rice grew more luxuriantlv and attracted more (Avipositing mothis than u lched rice Also the rice plant and stein borers imatoured 10rC quickly and a partialgeneratiori af vouing ista iNoverwintcred in the stubbleLotili--malitrt varieti Quickly maturing crops reduce the tmbCr of pCst generations that Calt build up Rapid crop establishment olan ca rl v-natu ring vaitvty if)pa 11icLa r will lessen white gib and stem borer damiage White grub larvae intle last iistar are highly destIruCtive so aii early-iiatri ng variety will escape serious root loss Stem borers build 1ip slowlyx in rice to ecormic abundant in a late-Maturing crop anrd caiuse whi tlieads Numibers of somespecies increase exponeitially with each succeeding genera-ti so a xarietyexhat atures I trowh earier thian another will hac less insect damage Early-naturing vaiicties however will not lessen seed pest daiage either ii the soil or oina standing crop

Strwhoi hAIJttam Inrscts tcd irig on thlie aerial portions of rice plats disperse from carly to late plantedfields AS iiiibCirs increase exponentially with eachinsect

gtscnrattm e eiglbrig f rmc~s1(uil upp wh rc generation pests build tIP where neighboring fihrTC e9r1sstagger their plantings II Batangas farmers plant uplandrice within I iro use 120-d s-arieties and plant at the start of the rainy season Ini Misarnis Oriental laricrs stagger their

plantings up to 3 mo apart and use a set of atieties that mature in 4-7 mo Insect pest danage particularly from stem borers is greater in Misarmis Oriental Also seedling maggots damage rice there because rainfall favors frequent planting of maic In Batangas rice escapes seedling maggots bCca usC of distinct drs season and synchron0us early planting with the onset of ains

Therefore combinations of cultural practices ea rlyplanting synchronous planing crop rotation and earlyshymaturing vaneties protect the rice crop against most insect pests

Ilmant resistance Beca tse upland rice has few specialized insect pests and many insect types attack the crop worldwide regional breeding and strong internatiornal cooperation are required

Sown-sced pests arc economically controlled by insectishycide seed treatment and should the relore receive lowxpriori ty inbreeding objectives

Root pests are normally cost ly to control with insecticides but finding resista t sonres mayvbe di fficult Tanaka (229) for exa -ple fi led to find varieties resistant to root aphids Selecting for large root bioniass perhaps should be the strategy for pests sLuch as Whlite grubs that renliove roots Fainter(174) howcvcr cites varietiesof crops tolerant ofor resistant t(i root pests Some sorghun varieties are high in cyanide so rice sorghun might incorporate broadshyspcctruri chemical resistance

Seedling vigor and drought tolerance are high prioritybreeding objectives so tupland rices should have large r root systernsand be able to tolerate ligherlevels of root loss Sonie root loss inolder plants is beneficial if it stimulates new root developnrut and therefore crihances intake of soil nutrients (53) The rice seedling inaggot rtiLotia spp may be

controlled thirough resistant aittics A hi rgc breeding effort on sorghniu has had fair success inideveloping arreties esrstait to Asoccata Rodani (251) A field trial comparing 10 upland ricc varieties showed differences aniong varieties va ring from II to 4(- damaged tilleis (215) Hovver resistarce to 1Iill-0r gtia species may rot cross ox-rti otier

Shiraki (210) reported that upland i-ice stens wcrc harder than lows-land rice stemsperhaps becaIusc (f a higher silicoricontent and thus were more resistant to yellow stern borer hc tortia Iow tillering (f uplaid iice xarneties prodtres a

higher percentage of infested tillers than would occor in higher tillering lowland iice under a siiilar egg density The corollary therefore is to select for high tilleriig upland rices

Stemii bor amcstheh main targes dobrecders inAfrica (571 204 )but nticl oifth e sc reen irig is dotie unde r lo wland co d t osrm s wih A h r~ o l pe i s n acorditions (11 96 218) As with lthrinrnwspecies mayhave to be dealt with independently however initial results show cross resistance between African and Asian species(92)

In Brazil screening Ior insect resistance focttses oil lou I pest problems Varieties tolerant of lasmolwalpus ligi-sellus and resistant to )iatraea saccharalis have been identified (70 153) Current work also emphasies 7iraca limativetntris and Thmialspis (=Iosis) flavouicta (66) These insect pests Ilt onlV citise IIigh yield losses tihey also ire difficult to control chemically ofr by other ilealls

ManV upland rice vaieties aic early maturing and are resistant to pests such as gall midge (I Xl ) Early mIturity is often a highly desirable trait to esuilpe pest buildup but it is not true resista nce bca usc ilthose va reties are plinted late they will be itsdamaged as ain early planted susc ible variety

P~erhaps tile cosiiiopolitall of all upland rice pestsimost are the sced pests (ripenling stagc) which pose a particular problem beta use scuds re ised IsIO for hitnit4ans Ayltoxic substatice introd tcld geticticllyilmay also be toxic to nal iniless it caln he placed old in thfe IdhuffSelectmi fo hairiness or lotng wns kill mIlct sthilrohleiiis dutilig threshing pa rtictfarl maitial threshinfl [hireshers ill complain of itcfimin atid itteascd (fust levels

Attempts itIRRI to fitid tcistatncc to the rice bug plorisa have ilotbeCnl prmitisilg lIIetIattonlids iorc

through the lenilna or palca hillhairs sedstVIor seeds wilh awns that deter bugs promote complaints from threshers Seed bugs liptocorisa and Stcnocoris enter the seed through the opening between tile lemna and palC but varieties with narrow openings are diifficult to mill

Breeding priorities should concentrate ott inotCI gruLtLIps that are most difficult to c nttrol by other meains Steil) horers appear to be the most widespread group that sigiiificantly dalag s rice aind is hard to manage by other Ilnca us

Biological control Very fe stutdic of mitatt~l eneties of rice pests in upland envirollnletits wee in tile wefouiid literature Therefore have not compiled a list of species Ve present a list of natural enemies iOtid in upland ricefields tuic latialln Batangas for some conitno iOldiar pest species ([itble 5)

Muich ofl thc lollowinle itfor ittiol cotlliCs rol Stutdies in the lowlatnds or roi crops otithcr thlin tice

Qu(ailtr o 11ll t11t( Nat trrtf lii hcoeits ight Ile used agaitist ile social insects tilt and termites Sollie predator specice live ill theirf nests n ilnic theirnllt ip-pearanIlCe aMRidbehavior IJtrouMdCing a mote aggressise ant species to displace pest ant codonies is a possibility (169) Patihogens as potential natural encnies have reccived little atteition (I13) Natural ncnuics asa group however hac not been toIl iporta it in Iegulating aintshown be and ternite numbers

The nocturnal behavior aid subteitmien habitat of lole crickets and field crickets protect them from many ntllral enemies There are only limited records of natural cnelics for these groups of pests some pathogens (87) but mainly

IRPS No 123 JattuIny I 271987

parasites - scelionids on eggsand sphecids on ilymphs and adults Sphecid wasps Larra carutmaria (SImitI hI L huzwuensis Rohwer and lsaltiuhta Willi ams speciali in mole crickets pa ralyzing them nd dragging the ninto thcir nests as food lot their voting ()tier sphCcid species Lins aurulenoa (Iabricius) lot maiac (Ashmead ) 1 lhshylssellaus (Smith) and Ml lal)oriosu (Snithf specialie in field clickets

White grUbsO the other hand hae Icti scriuns enuigh to have been extensively studied for cotttroi 1W nati al enemies( 112 140 141) Scoliid parasites C(ajsoMcispp have beetn intrtduced itto tilePhilippites for control of Ielxl ]udisiro la The rcsearchcts w Intttrdced themle during ill era claimed that the white gttionthreak as controlled (142) howevcr it isnormal for pest epidemics to subside and there was no diect evidence to indictte that tfe parasite was responsible A tmore promisitg titctllod might he to identify iso ltcttass prt~f tcartd apply hiactelaln fingal pathogens to tile soil Ihe lalattese beetle in North America has hcci coitrolled hv tite conmnleciall aailable hactcrial prcpa ration itacihlpu pav ofr milksdiscas (71) Spote prtyefott the soil itloted ilrtier reitiain actic fr Veas

Root aphid sardtnctly bugs readilslic tilib to predatots and parlsites once their tetndine ants are controlled The larvac and adtilts of a coccinellid beetle 1Scynms sp prey on llriadlouais in slash and burn upland rice Baits treated with insecticide may be used to control tending ants by killing their ourig (244)

Root-tkcditig blugs false wireoris wireorms and root wccils aic r ofsuch little ifilpoltatice that thenomttll role of liatural Cticniics hias tiot beei asscssed A possibility hoevrl tiglt he tit ItaIIitic niaCIMtodcs lintt1g11en Itese soil-itlhlfbitine mCtitodcs W5uhtfhave to be imiass Produiced

large miltibcr of niatltrll cnemics ircknown ft CltIoriS IC IpblCeii is thtt clttvortnlls ofonlie the crop soott alter land ptel)tritiOti a1tithe bcgit iniig of the raittv

Citsoln hfien tiatllral leniiiy populations aric lo Colltloil WOtuld iii ole mass prodtticing andirclcaslng key spccies

le egg stage is the fpirt (olthe Cedlitgt Ialot life cycle tI hich to f)ctis ftor tittlril elctit cottntrol hplraites o1 pleditors ttsit elleggs highly expoIsYCl I Sgg pailsites or predators stolld probahly e ittass andtbe produced released Littlh i toruialioll is a ailable ott what species to trV o ho to Iniss podLice thiCIi A etiloplhid wasp Icmihlaol sp effctiicly regushy

lates the leatf lcloalapolm2a spioanoiiitr heat andc rice antid should be collsctvcd

leaf beetles whCllose e am] eggs ate laid in the grutindhirO ould be motC difficutlt to control by niatulral enelV Mnll ipuhltiOn But species sUici IS c_gsthispid beetles ith

and larVle oilleascs ate vtilcratble to iparaites and pledators

Natural enemisc of tlirips otl rice are little known

28 IRPS No 123 January 1987

Table S Parasites recorded from upland rice insect pests in Tanauan Batangas Philippines 1977-84

Order Family Species Host stage

I lymttenoptera Braconidae Leaffolders Cnaphalocrocis Marasinia spy

Cardiochilesphilipplinensis Ashmead a Larva

Cotesia angustillasis(Gahan)Cotesia or cypris (Nixon)

Larva Larva

Cotesia or taeniaticornis(Nixon) Larva Cilonusinotakatae Munakata Chelonits SppAlacrocetrts philippinensis (Ashtmead)

Egg to larva Egg to larva Larva

Iclineumonidac Orgilts spp Isclutojppa hireator(Fabricius)

Larva Larva

Itoplectis naraigav(Ashmcad) Larva Tenelhtcha philippiensis (Ashmead) Larva Tenelchha stangli (Ashmead) a

irichomma cnaphalocrosis Uchida Larva Larva

Bethylidac Elasmidac

Goniozms or trianguli r Kieffer lasmtusalhopictus Crawford

Larva Larva

Diptera

Encyrtidac Chalcididac

Tachinidac

Elasmtus spp Copidosomopsisnacohiae (Fidy) Brach nteria evcarinata Gahan Biachyieria lasus (Walker) Ztgohothria ciliata (Wulp)

Larva Egg to larva Pupa Pupa Larva

Argyrophylx nigrotihialis(Blaranov) Larva

l-ymenoptera Braconidac Yellow stern borer Scirpopiaga incertulas

Braco chinensisSzeplegeti Larva CIthelotis spp Chelonus niunakatae Munakata Cotesi flaripes Cameron Stenoracon nicerilli (Bingham) a

Larva Egg to larva Larva Larva

Ichneumonidac Tropobracon schoenobii (Viereck) Amatroinorpiaaccepta metathoracica(Ashmiead)Frihorus sinicus (I lolmgren)

Larva Larva Larva

Ischnojoppaluteator (Fabricius) Larva Isotima nr dammeratiti Rohwer Larva Tenelucha philippinensis (Ashmead) a

Larva Tenehicha stangli (Ashmcad)Trichomma enaphalocrosis Uchida

Larva Lara

Chalcididac Eulophidac Trichogranmmatidae

Xanthopimnpla stemmator (Thunberg) Charops hrachYpterumn Gupta and Malteswary Brachtm eria sppTetrasticttus schoenohii Ferriere Trichorigranm chihunis Ishi

Larva to pupa Larva Pupa Egg Egg

Diptera

Scelionidac Pteromalidae Tachinidae

Tt ichograintnta aponiicuitt (Ashniead) Telenomits rowani (Gahan) a

Trichoinalopsis apattelctena (Crawford) Jeirhaea spp

Egg Fgg Egg pupa Larva

Z4gobothria ciliata (Wulp) Larva

Ilymenoptera lecneumonirlac Gol fringed borer (hilo auricilius

Trichomm enoptalocrosisUchida Larva

Ieioenoptera Braconidae

Aantholimlh steiniator (Thtinberg) a

Pink stet Ioret Sesamia inferens StoInacoil nicrilhi (Bingh am)

Loarva to pupa

larva

Diptera Ilyinenoptera

Armvtnyorms Mythimna separata Spodoptera mauritia acronyetoidesTachinidae Zvgohothriaatropiora( Robinea i -Desvoid y)Eulophidae It-phctrtiscitapadoe (Ashimead)

Larva Larva

(halcididae Braconidac

Brachpin ria spip (oesia spp

Larva larva

IHyntenoptera I)iptera

Rice skipper Ielop ida (halcididae Tachinidae

math ias aidgreen iortedci ciliarNIelan itis eda Brat lhincria sp iii matuqitatt Cameron Irgyrolhirlaxi trotlhiais(marumtv)a1

ismtne Larva Larva

Diptera Ilymenoptera

Tachnidae lelineutionidae

fuown se tilo per Mois frugalisIrgtrophrlax 1igrothiahiv 3 a rat ov) anthopimpa pmnctata (Iabricius)

Larva Larva to pupa

aMost dominant

IRPS No 123 January 1987 29

Normally predatory thrips prey on herbivorous thrips strepsiptera on both Spiders dominate tilenymphaladultNatural enemies of stem bugs --Scolinoplara and Tibraa predators Spittle bugs move to rice from pasture grasses - are in order of importance egg par~isites and predators and biocontrol would be successful only ifcarried out before fungal pathogens and nymphal adult parasites Egg tiley immigrate into ricefields parasites could either be introduced fron other areas or Biological control tactics for hemipterous seed bugs mass produced and released ould follow those outlined for stein bugs

Armyworms are normally lwld in hcck by the activities Coml)arison of environnewtns Natural enemies are of egg and larval parasites When these natural enemies fail perhaps faced with even greaterconstraints than insect pestsusually because of drought armyworns become epidemic to survive ind reproduce in upland environments TheyIt may be worthwhile therefore to release parasites during have adapted ani survived by having wide host ranges the the rainy months following a drought Virus diseases of ability to aestivate or hibernate over unfavorable seasonslarvae would beanotheravnue to exploreforarnyworms and the ability to disperse or by other mechanisms

Parasites have ben traditionallyconsidered for biolegical Comparing the most dominant and therefore the most control attempts against stein borers usually by rekasing adapted species of natural enemies collected in various exotic species( 166) This approach has not met with success environments in the Philippines we can understand how and new avenues should be explored The role of predators different rice environments are arid pathogens is little inderstood and deserves greater Anagqrms olptahiisa mymarid planthopper egg parasiteattention Egg parasitism rates are normally reasonably is cquallyadapted to all Philippine rice environments (Tablehigh but should be supplemented with effective egg 6) It is selective for planthoppers and even parasitizes eggspredators Orthopterans are potential egg predators of the n-iize planthopper Peregrinitsmaidis itt upland areas Metioche vittaticollis (Sta ) Anavipha nifiwnnis (Saus- Alavohis also parasitizes maize planthopper eggs but is sure) and Conocephahs hongipnni (de i-laan) feed on most prevalent on BPH in the irrigated wetlands (onashystem borer eggs etioche and Anaxil)ha specialize in eggs tocerus spp like A optalhilis on BPH and WBPI Iattacks with no hair such as Chilo spp The omnivorous (ono- the eggs of GLH in ell rie environments It is also found cephahsspecializes in eggs cov red with hairs it is a proven parasitizing the eggs ef the white rice licahopper (oimna egg predator of Scirlouphaga spp but has the discouraging spetra Oirosita naias a trichogrammatid egg parasitehabit of eating rice guin (65) attacks BIPH and WBPH in the wetlands and has no other

Grasshoppers have egg (scelionid) and nymphalad ult host A related species 0 aesopi specializes in G1-H in (nemestrinid tachinid and sarcophagid) v irasites Among wetland environments Dryinid strepsipt-an and pipunshypathogens protozoans have been recorded most frequently culid nymphal and adult parasites of planthoppers and Control of locust species by natural enemies could con- leafhoppers also specialize either in hopper species or centrate on their habitual breeding areas environments Among the dryinids Pseuhdgonatoipsflavi-

Leaffolders have rich complexes of natural enemies some feimur prefers the lowlands and BPH while P nuch has no adapted to upland rice (22 27) ranging from egg predators environmental prefirences but is adapted only to WBPH (gryllids coccinellids) to larval parasites (braconids ichncu- The main d ryinid species on GLH aptogonatop)usspp is monids) to larval prcdators (ants and carabids) to larval most dominant in rainfed lowlands and also attacks other pathogeiis (vira and fungal diseases) Efforts to augment leafhoppers within Amnrasca Cicadulinaand Bachaltuathese natural enemies should focus on the egg predators Strepsiptera are more dominant on plarithoppers than (rearing releasing)and viral or fungal pathogens (culturing lealhoppers The Elenchus species in the Philippines attack spraying) all species of planthoppers within the genera Sogawilla

Parasitization of the large conspicuous larvae of Sogatodels NilaparsataHarualia and Opiconsiva Elenshyskippers green horned caterpillars and other polyphagous cusrvasuntaisuiis adapted to the major rice environments Lepidoptera by tachinids and chalcids is normally low ELg but prefers BPH in rainfed arid WBPIH in wetland environshyparasites and predators are perhaps the key to their control ments Ilalictoph(utsmtnroeiand HVpectrus attack G LH

Gall midges are parasitized by pterormalids and platy- and COfimravcctra respectivelygasterids that aestivate within their larval host between Pipunculids are mainly parasitic oii lealhoppers Each seasons Shoot aphids are preyed upon by coccinellids species has a unique environmental preference Tomosshysyrphids and chrysopids varireIa oraetoraprefers Ne7 otetltix nigropictrs and

Planthoppers arid leafhoppers have egg parasites and Pilunwulhtsjavanensisattacks Dehoceplhahls spp which predators as well as nymphialad nlt parasitesand predators are more abundant in upland than irrigated lowland their rich complex of natural enemies includes fungal environments T suhvirescensattacks Nephotettix virescens pathogens The principal egg parasites are myrnarids and and prefers the lowlands Pilpuiulus nttillattls is an trichogramniatids Egg predators are norinally mirid plant upland rice parasite arid attacks only Nephotetix speciesbugs that suck out the yolk Nymphal ad nIt parasites are More than 17 recorded species of leaffolder parasites in dryinids on planthoppers pipunculids on lealhoppers and the Philippines attack species of C(naplwocrocis and

30 IRPS No 123 January 1987

Table 6 Dominant natural enemies of rice pests in upland rainfed lowland and irrigated lowland rice environments of the Philippines

Natural enemy

Host stage Species

Brown planthop-Parasite Egg

MymaridaeAnagrus optabilis(Perkins) Anagrusflaveolus Waterhouse

TrichogrammatidacOligosita naiasGirault

NymphAdult Dryinidac

Pseudogotatopuslaiifemttr Esaki et Hashimoto Elenchidae

Elenchus yasurnatsui Kifune and Hirashima

Environment

Upand Rainfed lowland Irrigated lowland

Ailaparata lugens

x x X x

x

x x

x X X Whitebacked planthoppers Sogatella furcifera and Sogatodes pusanusEgg

MymaridaeAnagrus optabilis(Perkins)

TrichogrammatidacOl~gositanaias Girault

NymphAdult Dryinidae

Pseudogonatopusnudus Perkins Elenchidae

Elenchus yasutatsui Kifune et Hirashima

Parasite Egg

MymaridaeGonatocerus spp

TrichogrammatidaeOligositaaesopi Girault

NymphAdult Pipunculidac

Tomosvaryella subviresccns Loew Tomosvaryella oryzaetora (Koizumi) Pipunclusmutillatus (de Meijere)

DryinidaeIlaplogonatopusspp

Egg Copidosomopsisnacoleiae (Eady)

Larva BraLonidae

Cardiochiles philipphnensisAshnead lchneumonidac

Trichiomnia cnaphalocrosisUchida Temelucha stangli (Ashmead)

Pupa Chalcididae

BrachymeriaexcarinataGahan Brachymerialasus (Walker) Brachymeria nr tachardiaegroup

Parasite Egg

Scelionidae Telenomus rowani (Gahan)

EulophidacTettasticusschoenobhi Ferriere

Larva Braconidae

Stenobraconnicevillei (Bingham) Iclhncurnonidac

Temelucha phiiippinensis(Ashinead)Temelucha stangli (Ashmead)

x x x

x x

X x X

x x Green leafhopper Nephotettix virescens

X x x

x x

X X X X

X

X Rice leaffolders CnaphalocrocisMarasmia

x X x

X

X X

x x x x x X x x x

Yellow stem borer Scirpophaga incertulas

X X x

x

x

x X XAinauromorphaaccepta metathoracica(Ashmead) x

continued on next page

IRPS No 123 January 1987 31

Table 6 cntinued

Natural enemy Environment

Host stage Species Upland Rainfed lowland Irrigated lowland

Larva-pupa Gold fringed borer Chilo attriciiuslchneumonidae

Xanthopimpla stemmator (Thunberg) x aPupa x

Pteromalidae Trichoinalopsis apanteloctena (Crawford) ax

Pink stem borerb Sesamnia inferensParasite

Larva Braconidae

Stenobracon nicevillei (Bingham) x b b Armyworms Mvthinna SpodopteraLarva

Tachinidac Zygobothriaarroptvora (Robineau-Desvoidy) x x

Braconidac Cotesia spp

X Skipper Pelopidas mathias and green horned caterpillar Melanitis leda ismeneF-gg

TrichogramrnatidacTrichogrannaspp x

Scelionidae Undetermined Scelionid

x Larva

Tachinidae Argyroph lax nitrotibialis (Baranov) X x

Pteromalidae Trichomnalopsis apanteloctena (Crawford)

X Predator

CoenagrionidaeAgriocnenis femina femina Brauer x xIschnura senegalensis tambur

Miridae CrtorhinsliidipennisReuter x x

Nabidac Stcnonabis tagalicus (Stal) x

VeliidaeAlicrovelia douglasi atrolineata (Bergroth) x

Coccinellidae Micraspis crocea (Mulsant) X x x

Vespidae Eumenes campaniformis (Fabricius) x

Formicidac Diaecamna spp xOdontoponera transversa (F Smith) x

Carabidae Chaenius spp OphioneaW- aJasciataSchmidt-Goebel

x

Ophionea ishii ishii Habu x x X

Gryllidav Anaxipta longipennis (Saussur-) x xMetioche ittaticollis (Sial) x x x

Tettigoniidae Conocephalus longipennis (de Haan) x x x

aNo rearing made on rainfed lowland bNo rearing made on rainfed lowland or Irrigated lowland

MrasmiaAmong the Laval parasites the braconid Cardio- lowlands and parasitizes oriental maize borer OsIriniachilesphilijpphensis is most adapted to an upland environ- furnacalis (Guenee) larvae The leaffolder pupal parasitesment It also parasitizes Hydelepta indicata a common not only occur equally in all environments but also haveleaffolder of legumes The ichneumonids Trichomma cna- wvide host ranges- parasitizing species of OsIrinia Marucaphalocrosis and Temehcha stangli occur more in the Hydelepta and Honiona - in maize and legumeslowlands T stangli is most adapted to irrigated lowlands The principal egg parasite of yeltow stem borer (YSB)and also parasitizes Chilo and Scirpophaga stem borer Telenomus rowani is widely adapted to all environmentslarvae T cnaphalocrosis is most prevalent in rainfed and has an unusual alternate host -Tabanus eggs (25)

32 IRPS No 123 January 1987

Tetrastictu schoenobii specializes only in YSB and occurs mainly in irrigated lowlands -- further proof that YSB is most adapted to the wetter environments

Larval parasites of YSB are highly environment-specific The braconid Stenohracon nicevillei occurs only in upland rice but also attacks the pink stem borer (PSB) Tieinchwa phiiiplPwnsis an ichneumonid occurs in upland rice and also parasitizes the larvae of (7hiu stqpresaisinatro-morpha accepta melaihoracica like 7 stangi is mnost adapted to the irrigated lowlands

The dark-headed stem borer(l)HSB) (ih pochit3sus was not collected in all rice environments and the gold fringed borer Chilo auricilius nd PSB were collected only in upland rice Xaantiholiplastenrnaloran ichneumonid larval parasite appears to be widely adapted whereas TrihomaslopisapanitI)cteiiaa pteromalid pupal parasite occurs mostly in wetlands and also parasitizes larvae of Chilo suppressais and Pelhuidas mathias

Armyworms Myit hinin and Spodujipera have two main larval parasites - a tachinid Zrgohothriaatropivora in rainfed environments and a braconid Cotesia spp in irrigated lowlands

The butterfly pests Pelopidas and 4elanitishave tricho-grammatid egg parasites found only in irrigated lowland environments

Two larval parasites show environmental preferences --Argrroph av nigrotihialis a tachinid in rainfed habitats and Trichonialopsisapantelkcnela in irrigated ricefields A limgrotibialisalsoattacks the sweet potato hornwormAgrius convolvtdi and T apanteloctena parasitizes DH S1B

Predators as a group show more distinct environmental preferences The aquatic damselflies Coenagrionidae are most abundant in the lowlands Cvriorhinus is also most adapted to the lowlands The nabid Stenonabis tagalicus occurs only in the uplands and isalso prevalent on legumes The coccinellid lady beetles wasps and ants are more prevalent in upland areas AlicrasIpis cit-ea larvae prey ona variety of aphids attacking legumes and maize Etunienes campanonia vespid mud wasp makes nests in trees and is therefore most adapted to the more botanically diverse uplands Soil-dwelling ants cannot tolerate flooding

Upland environments are habitats to arboreal carabid beetles Three species of Chlaeniusbeetles prey on leaffolder larvae and are perhaps more important than parasites (27)

The gryllids and tettigoniids are widely adapted egg predators not only on rice but also on maize and legumes They feed on eggs of most insects that are laid on leaves

The spidercommunity of upland rice environments isrich in species At one site (Batangas Philippines) 31 species have been recorded (23) Of the 176 spider species recorded in Philippine ricefields about one-half (82) occur in upland rice The spiderspecies of upland rice environments overlap more with those of rainfed lowland than with those of irrigated lowland rice (24) The same study showed that of the three environments irrigated rice has (ie greatest spider species diversity foiiowed by rainfed lowland and upland

However of the 10 most prevalent ricefield spiders 9 were abundant in upland rice showing wide adaptation All three spider guilds -- orb-web space-web and huting spiders shywere prevalent in upland ricefields There were however differences in environmental prelerences for some spider species

Of the orb-wcavers Tetragniathamanlihulatawas partishycila rlv abundant 7 japonica was low and Ltucatge dehcorata was absent in upland rice

As in other environments Aiypeiwnaifirtosana was the most dominant space-web spider however its relative numbers were lowerthan for lowland sites Among hunting spiders Ox|opes javanus and Lcosa leatcost fgla were more abundant and Lvcosa lpseudoanmnlata was less abundant in upland compared with lowland rice envishyroniments

Extensive rearing of rice insect pests in upland rainfed lowland and irrigated sites over an 8-yr period in the Philippines provided insight into the effectiveness of parasites as natural enemies (Table 7 8) Most of toe data are from large sample sizes taken over at least one crop which overcomes the pitfalls expressed by Van Driesche (238)

VSB egg parasitization was surprisingly similar across all environments Larval parasitization was highly dynamic among sites and even years within the same site but showed somewhat lowerlevels in uplandsthan in irrigated wetlands This would have been predicted for egg parasites as well since YSB is adapted to the lowlands and so must be its parasites This isevidence that Telenomus rowani is as good at dispersing as its main hosts Trowani clings onto the anal tufts of female YSB and parasitizes eggs as they are laid This phoresy may explain its wide adaptation The more specialized lowland YSB larval parasites therefore appear more effective than their upland counterparts with more alternate hosts

Low collection levels of PSB larvae especially in the wetlands make comparisons between environments difficult Low levels of larval parasifization were recorded in both rainfed environments Except in the Cagayan rainfed wetland site in 1980 low levels ef larval parasitization were evident at irrigated and rainfed sitesalike No differences by environment were evident Higher levels of parasitization occurred in Sarawak during a 1967 outbreak also from tachinids Rothschild (197) indicated that higher parasishytization occurred in upland rice areas

Skipper and green horned caterpillar larval parasitization rates were generally low and showed no environmental effect htighest activity occurred in Cagayan in 1980 (48 parasitization) Although larval parasitization levels of RLF were variable year to year activity appeared greater in the irrigated lowlands than in the uplands Again the highest levels were recorded in Cagayan in 1980 Cagayan has extensive grasslands and fallow around the rice areas and a relatively long rainfall period Rainfall in 1930 was good for crop growth and apparently for increase of larval parasites

IRPS No 123 January 1987 33

Table 7 Parasitization rates of rice planthoppers and green leafhopper in upland rainfed lowland and irrigated lowland environments Philipshypines 1976-840

Environment and site Year Egg parasitization () Nymphaladult parasitization ()

BPH W13P GLH BPII WBPH GLIH

UplandBatangas

Rainfed lowland

1976 1979 1980 1982

15 17 12

2(47)8(39)

4(51)

9(1800)14 (212)

12(304)

8 (101)6 (127)

9(116)

Iloilo

Cagayan

1976 1977 1978 1981 1980 1981 1982

0 (69)

11

44 48 40

-

-

46 55 44

0 (471)

11

52 56 49

0 (200)9 (214)

10 (1259)8(235)6 (900) 3(800)

-7 (462)

11 (270)3 (313)5 (900) 1 (800)

4 (600)10 (725)10 (362)12(870)5 (900) 3 (800)

Pangasinan

Irrigatedlowland

1984 1976 1978 1979 1982

13 (436) 13(379) 14 (457)

6(14)7 (197)

17(198) 11 (486) 18 (275)

3 (81)10 (274)3 (116)

16(871) 19(88)

28(211)9 (764)5 (647)7 (1204) 8(305)

Laguna 1977 11 7 21 1978 31 1979 14 4 4 28 1980 7 8 20 1981 1983

56 7(278)

26 16 (1650)

16 27(1921)

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R I Apostol unpublished 1lo1lo and Pangasinan 136 Cagayan BCanapi unpublished Liuna 175 I C Pantua unpublished 118 234 43 Carino and Shepard unpublished BPH = brown planthopper Nila parvara htgens W131Pi =whitebacked planthopper Sogatelia furcifera GLH = green leafhopper Nephotettrix virescens Figiires in parentheses are sample sizes

Tauher et al (230) reported that internal external and genetic factors influence the seasonal activity of parasitoids

Hopper egg parasites but not nymphaladult parasites were also parlicularly abundant at the Cagayan site possibly the egg parasites have more alternative hopper prey in the weedy areas than nymphaladult paisites

The seasonal dynamics of hopper egg parasites can be seen for upland rice in Batangas(Fig I 2) and for rainfed lowland lice in Pangasinan( 135)and Cagayan( 105) Levels of BPHWBPI-L and GLH egg parasitization were corn-parable in upland and irrigated lowland environments Except for some high rates of paraMsitization early in the season the parasitization rate tended to be steady in all envilonments However hopper nymphaladult parasi-tizution levels were more similar in the rainfed environments and lower than in irrigated rice

Overall parasites do not appear highly effective by themselves against upland or even lowland pests Among natural enemies parasites have attracted the most attention because they can be more readily assesscd

Predators and pathogens lend themselves better to management practices Predators the most important group of natural enemies in riccfields are difficult to quantify On the other hand pathogens appear less im-portant but can be readily cultured and disseminated

The natural enemy commulnity of upland rice is rich in species and differs significantly from that of lowland rice

These beneficial organisms must be conserved by applying insecticides judiciously particularly sprays Seed or soil placement of chemicals minimizes exposure to natural enemies The strategy to derive the greatest benefit from natural enemies isto allow the greatest number of beneficial species to thrive Then perhaps at least one species will be effective against each pest at any tine overcoming the variable abundance of each species within and between years

Programs to introduce exotic species or mass produce indigenous ones are ambitious and expensive because they require trained people on a sustained not ad hoc basis

Chemical control Insecticides offer rapid and efficient control of upland insect pests (190) but-should be used only after other control measures have been considered Insecticides are rarely used on upland rice because of cost (32) Upland rice yields are normally too low to justify the expenditure Also most upland rice is grown as a cash crop Spraying upland rice is more difficult than lowland rice because water is less accessible Government subsidy programs to provide pestishycides to tribal peoples for upland rice production have created problems of toxicity to humans because isolated people have had no experience to be able to handle pesticides safely

However ins-eticide use has beenjustified when a crop is

0Z

Table 8 Parasitization rates of lepidopterous pests in upland rainfed lowland and irrigated lowland environments Philippines 1976-84a

Parasitization (5)

Environment and site Year YSB GFB PSB Earcutting Swarming Skipper Greenborned RLF Brown larva larva caterpillar caterpillar caterpillar caterpillar larva semilooperEgg Larva larva larva larva larva larva

UplandBatangas 1977 - 0 (57) - 0 (63) - 0 (71) - 0 (106) 11(142) 2 (101)1978 43 (4697) 8(451) 5 (38) 3 (40) 6 (61) 2(58) 15 (57) 13 (129) 13 (1286) 4 (30)1979 59(4135) 12(320) 13 (11) 5(28) 9(45) 6(46) 7(23) 8(160) 9(1152) 8(26)

Rainfed lowland Iloilo 1976 - 0 (10) 0(10) 0 (7) 0 (19) 2 13) 15 (180) 5 (76)1978 0 (54) 0 (3) 0 (17) 9 (64) 4 (78) 3 (73) 0 (121) 0 (191)1979 64 (2350) 16 (84) 6 (17) 0(6) 18 (68) 25 (104)

1981 56 (1322) 18(304) 0(5) 12(51) 18(138)Pangasinan 1976 60 (7 7 )b 62(82) 0(111) 5 (39)

1978 58 (53)b 5(51) Cagayan 1979 59(713) 8 (101) 6 (34) 0(7)

1980 0(11) 4 (28) 15 (286)

62 ( 48(177) 61(140)1981 0(5) 12 (2) 13(45) 14 (92)

Irrigatedlowland Laguna 1974 62 8

1974-75 50 41 1975 43 72 1978 96 (623) 21(24) 33 (60)1979 83(2017) 18(110) 8(62) 11 (89) 22(400)1980 78 (1625) 20(75) 0 (16) 4 (72)1983 60

9 (80) 7 (270) 30

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R F Apostol unpublished Iloilo and Pangasnan 136 Cagayan B Canapi unpublished Laguna 175 P CPantua unpublished 118 234 143 Carino and Shepard unpublished YSB = yellow stem borer ScirpophagaincertulasGFB = gold fringed borer Chilo auricilius PSB = pink stem borerSesamia inferens earcutting caterpillar Alythimna separata swarming caterpillar Spcdoptera mauritia acronyctoides skipper Pelopidas mathias reen horned caterpillar Melanitls ledaismene RLF = rice leaffoldersCnaphalocrocismedinalis and Marasmiaspp brown semilooper Mocisfrugalis Figures in parentheses are sample sizes Egg masses

IRPS No 123 January 1987 35

Table 9 Effect of seeding rate and insecticide on plant density and Yield losses from insect pests in upland rice often areyield of Dagge upland rice Batangas Philippines 1976a comparable to those of lowland rainfed or irrigated rice

Seeding rate Insecticide Dosage Plant density Yield (0-Y30o) but usually from different pests The principal (kg aiha) seed treatment (kg aiha) (noIra-row) (tha) groups are soil pests feeding on sown seed and rootsat 14 DE followed by seedling pests These groups have largely gone

50 None - 35 a 20 a unnoticed leading researchers to conclude that stem borers50 Carbofuran F 10 38 ab 23 a and seed bugs are the most important pests (17) Yield loss 10 beCabofuranF100 None - 4551 c 23 a trials that pinpoint major pests need to includc seed

100 Dieldrin WP 10 52 cd 22 a treatments Granules applied in the furrow may not give150 None - 59 d 22 a adequate protection from many seed pests

aAv of 8 fields DE = days after crop emergence In a column means The floristic and pedological diversity of upland rice followed by a common letter are not significantly different at the environments is matched by the diversity of arthropods5level by D)MRT F = flowable Wi = wettable powder both pests and their natural enemies Irrigation has homoshy

genized lowland rice environments Flooded soils tend tothreatened by epidemics of locusts or armyworms New have the same properties and to eliminate soil pests The technology stressing efficiency has minimized insecticide spatial and temporal dominance of cultivated rice has cost benefited specialist pests

Seed treatment is normally inexpensive and can be Upland rice has no specialist arthropod species The economicallyjustified in -nanycases(66) Treating seed with monophagous pests on upland rice prefer the lowlands systemic insecticide (216) but not chlorinated hydrocarbons where rice evolved Rice in the uplands exists only because (247) is most effective in controlling seedling maggots of man and even though lowland rice predates upland riceFoliar sprays have to be repeated to obtain control and specialist species evolved Wheat maize and sorghum do often are washed off by frequent early season rains not have monophagous pests leading us to conclude that Granular insecticides are effective but required dosages are there is no evolutionary advantage for inssects to have too expensive to justify except under heavy infestation narrow host ranges in upland graminaceae(247) When directed against first-instar larvae banding Pests have adapted to the uplands not only by having granules in seed furrows (177) can be effective against white wider plant host ranges but also by having long life cyclesgrubs at low dosages (025 kg aii ha) (134) Seed treatments undergoing dormancy or being dispersive Each pest has a are not promising for white grub control (248) unique life history strategy suited to highly unpredictable

Seed treatment is also effective ainst ants and may be environments In agivenyear someinsect specics an better cheaper than increasing the seeding rate In Batangas exploit upland rice as a food source but no pest is a senious treating 50 kg seed ha was as effective as increasing the one every year on upland rice seeding rate to 100 kgha and using untreated seeds (Table The low yielding potential of upland rice means less 9) In Brazil seed treatments control termites and other soil quantity and lower quality of food for pests to exploit but insects including the lessercornstalk borer(66) Domiciano also less incentive for farmers to undertake control (58) however found seed treatments inconsistent in their The most effective control would be early and synchroshyeffect against high populations of lesser corn stalk borer nous planting of early-maturing upland rice varieties to

Baiting can be an inexpensive way of controlling ants prevent insect pest buildup Breeding for insect resistance seedling maggot flies mole crickets and field crickets Baits should initially focus on stem borers because they are verymade of locally available material and impregnated with difficult to control by other means Biocontrol efforts insecticide can be sparingly distributed taking adantage of should first stress conservation of natural enemies preshythe pests ability to disperse and encounter bait sites dators and pathogens rather than parasites Efficient

The new low-volume hand-held controlled-droplet chemical control includes seed treatment or baiting for seed sprayers may offer an advantage over high-volume knap- seedling and some root pests and using controlled dropletsack sprayes for controlling foliar pests Upland rice sprayers for spot treatments against foliar pests and seed canopies are more open than those in lowland rice and bugs when economic thresholds are reached better droplet penetration should result Controlled droplet applicators would be ideal for seedbugs and defoliators

CONCLUSION

Upland rice is attacked by a wider array of insect pests than is lowland rice mainly beca use of the addition of soil pests Generally population levels of lowland rice pests are lower in upland than lowland rice leading many people to conclude that insect pests are not important on upland rice

36 IRPS No 123 January 1987

REFERENCES CITED

1Abraham C C and K S R Mony 1978 Occurrence ofLeptocorisa acuta Fabr (Coreidae Hemiptera) as a pest of nutmeg trees J Bombay Nat ttistSoc 74553

2 Agyen-Sampong M 1975 HlierogrlphusdiaganensisKrauss (Orthop-tera Acrididae) a new pest of rice in Northern Ghana Ghana J Agric Sci 8249-253

3 Agyen-Samnpong M 1976 Varietal resistance and other inle-ratcd pest control components as factors of rice varietal improvement in West Africa Prcsented at the Second varietal improvement serinar 14-18 Sep 1976 Bouak Ivory Coast West Africa Rice Development Association Monrovia Liberia 18 p

4 Agyen-Sampong M 1977 Outbreak of Ihccrorrdnisorae in rice seedlings near Rokupr Sierra Leone lit Rice Res Newsl 2(6)16-17

5 Agyen-Sampong M 1980 Dirty panicles and rice yield reduction caused by hugs IntRice Res Newsl 5(l)11-12

6 Agyen-Sampong M 1982 The major pest problems of irrigated upland and mangrove swanip rice ecosystems ir the humid tropical Guinea savanna and Salel climate zones Pages 318-328 in Pro-ceedings training course cn concepts techniques and application of irtegrated pest management in rice inWest Africa 10-20 Jan 1982 Fendall Liberia West Africa Rice l)evelopioent Association Monrovia Liberia 505 p

7 Akibo-Betts V I and S A Raymundo 1978 Aphids as rice pests in Sierra Leone lit Rice Res Newsl 3(6)15-16

8 Akinsola F A 1975 Present status of different rice stemborers in parts of Nigeria iicL Entomo Newsl 328

9 Akinsola FA 1979 Ihe biology and ecology of rice stemborers in Nigeria Ph 1) thesis University ol lbadan Nigeria 217 p

10 Akinsola E A 1979 Guidelines of integrated management of insect pests of upland rice in West Africa Presented at the Seminar on integrated management of rice diseases and insect pests at Bobo-Dioulasso Upper Volta 17-22 Sep 1979 West Africa Rice Develop-ment Association Monrovia Liberia

II Akinsola IA 1984 Insect pests of upland rice in Africa Pages 301-305 in An overview of upland rice research Proceedings of the 1982 lIouak Iwsrv Coast upland rice workshop International Rice Research Institute 13O Box 933 Manila Philippines

12 Alam M S V TJohn and Kaung Zan 1985 Insect pests and diseases of rice in Africa Pages 67-82 in Rice improvement in Eastern Central and Southern Africa proceedings of the international rice workshop at Iusaka Zambia 9-19 April 1984

13 Alticri M A A van Schoonhovcn and J D Doll 1977 The ecological role of weeds in insect pest management systems a review illustiated with bean (Phaseohsvulgaris L)cropping systems PANS (Pest Artie News Summ) 23195-205

14 Anmante F 1967 The leaf-cutting ant Atia capiguaraa pest of pastures [in Portuguese] Biol6gico (Sao Paulo) 33113-120 (Rev Appl Entomol A56207)

15Andrade A C and D Puzzi 1951 Preliminary results of experi-merits on the control of Sculptcoris iastantus on sugarcane [in Portuguese] Bioulgico (Sao Paulo) 1744-49 (Rev Appl Entomol A41421422)

16 Appert J 1970 Maliarplia separaelha (rice stem borer) new observations of entomological problems of rice in Madagascar [in French] Agron Trop (Paris) 25329-334

17 Ariadeau Mand Z Harahap 1986 Relevant upland rice breeding objectives Pages 189-197 in Progress in upland rice research Proceedings of the 1985 Jakarta conference International Rice Research Institute 1O Box 933 Manila Philippines

18Asar S and Kisimoto 1975 Seasonal occurrence of the smaller brown planthopper in upland rice [in Japanese] Proc Kanto-Tosan Plant Prot Soc 2228

19 Bakker W 1974 Characterization and ecological aspects of rice yellow mottle virus in Kenya Meded Landbouwhogesh Wage-ningen 152 p

20 Balut F F 1970 White grubs attacking rice Oryza ativa L [in Portuguese] Biol6gico (So Paulo) 36321-322

21 Banerjee S N and L M Pramanik 1967 The lepidopterous stalk borers of rice and their life cycles in the tropics pages 103-124 in The major insect pests ofthe rice plant Proceedings ofa symposium at the International Rice Research Institute September 1964 The Johns Hopkins Press Baltimore Maryland USA

22 Barrion A T and J A Litsinger 1980 Ants a natural enemy of cliaiarac in dryland rice IntRice Res Newsl

5(4)22-23 23 Barrion A T and J A litsinger 1981 The spider fauna of

Philippine rice agroecosystem 1Dryland Philipp Entomnol 5139-166

24 Barrion A T and 1 A Iitsinger 1984 The spider fauna of Philippine rice agroccosystcms II Welland Philipp Eiitomol 611-37

25 Barrion A T1 and J A Litsinger 1984 lihanus (Diptera Tabanidae) eggs an alternative host of rice stei borer (SI3) egg parasite Telenoonis digns (Iliymenoptcra Scelionidae) lit Rice Res Newvsl 9(6)19

26 Barrion A T and J A Litsinger 1985 Identilication of rice leaffolders by wing markings Int Rice Res Newsl 10( 1)24

27 Barrion A T and J A Litsinger 1985 Chctiosspp (Coleoptera Carabidae) a leaffolder ILF)predator lit Rice Res Newsl 10(1)21

28 Bertels A 1970 Insect pests of rice and theircontrol [in Portuguese] Pelotas Brazil Instituto de Expcrimentacao Agropccuararias do Sul Circular 43 24 p

29 Bonzi S M 1982 Ctilo (flisilineus J de Joannis (Lepidoptera Pyralidae) stem borer of irrigated cereals in Upper Volta LioFrench] Agron Trop Riu Rizic Cult Vivrieres Trop 37207-209

30 Bourke T V T L Fenner J N LStihick G Baker EHassan D F OSullivan and C S li 1977 Insect pest survey for the year ending 30th June 1969 Papta New Guinea )epartment of Agrishyculture Stock and Fisheries Port Moresby 57 p

31 Bowden 1 1973 Migration of pests in the tropics Meded Rijksfac Landbouwwet Gent 38785-796

32 Breniere J 1973 The main rice parasites in West Africa and their control Pages 27-42 in Proceedings seminar on plant protection for the rice crop West Africa Rice Development Association Monrovia Liberia

33 Breniere J 1976 The principal insect pests of rice in West Africa and their control West Africa Rice Development Association Monrovia Liberia 52 p

34 Brown ES 1962 The African arinyworn Spoiolptera eXI)Ina (Walker) (Lepidoptera Noctuidae) a review of the literature Cornshymonwealth Institute of Entomology London UK57 p

35 Brown F B 1968 Upland rice in Latin America FAO-International Rice Commission 12th Session of the Working Party on Rice Production and Protection 2-14 Sept Peradeniya Sri lanka Food and Agriculture Organization Rome Italy 6 p

36 Buyckx F J 1 1962 Disease and insect pests of crops inCongo Rwanda and Burundi [iil French] Publications dc LInstilut National pour lestude Agrononiquedu Congo M Weissenbruck S A Brussels Belgium 708 p

37 Cariiio F 0 1981 Role of natural enentics inpopulation suppression and pest management of green rice lealhoppers Ph D thesis University ofthe Philippines at Los Bafitos PhilippinesLaguna 136 -

38 Carpente A J 1983 The development of rice cultivation and extension of vegetable productin in Zanzibar Final Report Vol II1 October 1983 Food and Agricultme Organization Ronte Italy

39 Carroll C P and S1 Risch 1983 Tropical annual cropping systems ant ecology Environ Manage 751-57

4(0 Cates R G 198 1 Host plant predictability and the feeding patterns of monophagous oligophagous and polyphagous insect herbivores Oecologia (Berl) 48319-326

IRPS No 123 January 1987 37

41 Cendcafia S M and F B Caora 1967 Insect pests of rice in the 62 Ene 1 C 1981 I)iiorldhol)ierusihilis Slater (licteroplera Philippines Pages 591-616 in The najor insect pests ofthe rice plant lygacidue Illissinac) a new rice pest front Nigeria lit Rice Res Proceedings of a symposium at the International Rice Research Newsl N3)15 Institute September 1964 The Johns Hopkins Press Baltimore 63 Fugenio T S 1965 rhe effect oi e-vCgetation on rice production Maryland USA 729 p and the possibility of overcoming it by leaf pruning MS thesis

42 Centro Ihternacional de Agricklittira Tropical 1984 Upland rice in University ol the Philippines at Los Baijos ILaguna Ihilippins 69 p Iatin Anerica Pages 93-119 hi An os ervies of uplaind rice research 04 Farrow R A 1975 The Alrican nmigratory locust ii is main Proceedings of the 1982 Boinakamp Ivory Coast upland rice workshop outbreak area of the middle Niger qlinlitatise studies of solitary International Rice Research Institute P0 Blox 933 Manila populations in relation to environmental factors Locusla No II Philippines 198 p

43 Chandra G 1979 Taxonomy ad hn riononlics ofthe imect parasites of 65 Fennal R G 1943 Food crop investigation Windward and rice Icalloppers and planthoppers in the Pihilippines and their Ieeward Islands First report Report onl work dunring the periodiiiportairce iii natural biol)gical cointrol Philipp Fntoltol 4119-139 April 1942-April 1943 Imperial College of Tropical Agriculture

44 Chang T T 1976 The originii colut ion enlIivation dissemination Trinidad 16 p (Rev of Appl Fintornol A3222-25) and diersificition of Asian and African rices -uphytica 25425-44 1 66 Ferreira F 1983 Integrated pest ritaageiret of upland rice [ill

45 Chatierjee P B 1979 Rice leaffolder attacks in India lit Rice Res Portuguese English summary] Pages 323-341 in Upland riceculttre Newsl 4(3)21 factors affecting production E Ferreira T Yamada and F-

46 Cheng Cbien-pan 19(4 Upland rice in Taiwan International Rice Malmolta eds Institutoda Potassae Foslato Institnto Internacional Cnmmission Wiorking Party oit Rice Production ard Protection I()th da Potassa Piracicaha Sao Paulo lraid Meeting 3-10 March 1964 Manila Philippines Food and Agricul- 67 Ferreira E arid F dl Silva Martins 1979 Occurrence of ture Organiaion Rome Italy 5 p Sogatohs orvziola (Muir) inupland rice in Goias Ilrail lit Rice

47 Chcrrett 1 M and 1) 1Peregrine 1976 A review of the status of Rcs Ncwssl 4(4)18 leaf-cutting ants and their control Ann Appl Biol 84124-128 68 Ferreira E and 1F da Silva Martiris 1984 Insect pestof rice ill

48 Chipeta F M 1974 Pests of rice in Malawi Pages 90-95 in Brazil and their control Jill Portugucsej FMIlRAtPA-CNPAF Proceedings Sith eastern Africa cereals research conference 10-15 Goiania Brmiil 67 p March 1974 DRB Mialda ed Ministry of Agriculture lilongwe 69 Ferreira FI F ca Silha Miartins S S Nero and 1 It P de Malawi Carvallio 1982 M tltiplc regression analysis todeterimine the effect of

49 Chu Yau-i 1979 The d atiaging period of the rice seedling fly insect populatiois ott upland rice yield [inIPortuguese Pesqui (Atlterigotiavxigu Stcin) (NItiscide Diptera) oil cori) at Fast Java Agropecu Bras 17671-675 Indonesia Plant Prot Bill 21403413 70 Ferreira F 1F da Silva Martins and F 1PZimmermann 1979

50 Conklin H C 1957 IHaninoo agriculture a report on aln integral Rice varietal resistance to tle lesser corn stalk borer [in Port uguese] systent of shi fti rig cult ivationi in the Pltilippincs Food and Agriculture Pesqui Agropccu Biras 14317-321 Organization Rome Italy 209 p 71 Fleming W F 1968 Biological control of lie hapanese beetle U S

51 Cramer H II 1967 Plant proection and world crop production Dep Agric RLs Sers [ech Bull 1383 78 p Pilaniensclhutz-Nachr 201-524 72 Food and Agriculture Organiation of the United Nations 1982

52 l)ani R C and Majunidar 1978 Prelininary observations on tie Country report surmimary Pages 27-37 in Report on the first incidence of root aphids on differeit rice cultnrcs Sci Cult 4488-89 FAOIUN)IPhail and regional training coiuirse in iniproved ctltural

53 Davidson R I 1979 lffects of root feeding on foliage yield Pages practices for upland rice 5-23 Oct 1981 Northern Region Agrculturd117-120 in T K Ci osb and R B Pottinger eds Proceedings of the Developririt Center Chiang Mai Food and Agricuilture Organiiashy2nd Australian conlerence on grassland invertebrate ecology Govt tion Rome Italy 37 n Printer Wellington New Zealand 73 Franco do Ariaral S 195 1 Residual action of insecticides in turfy

54 l)ean G 1W 1976 Rice insect pests inlaos Int Rice Res Newsl soil--the results of a field experiment [in Portuguese] Biol6gico (Sao 1(2)15 Paulo) 17209-214 (Rev Appl Entomol A41423)

55 I)ean G J W 1978 Insect pests of rice in Laos PANS 24(3)280-289 74 Freeman ) 1970 Report onithe Iban london School of -coroiics 56 )ingle H 1984 Behavior genes and lifc histories conplex adapta- Monographs on Social Anthropology No 41 Athlane Press

tions to uncertain environments Pages 169-194 in A new ecology London UK 317 p novel approaches to interactive systems P W Price C N Slobod- 75 Fuchs T W1 A Harding and 1W Smithlr 1979 Induction and chikoff and W S Gaud cdsl ohn Wiley New York USA termination oldiapause in the sugarcane borer Ann Flitonlol Soc

57 Dobelmann-JP and M Falais 1968 Upland rice cultivation ill tie Am 72271-274 North-western part of Madagascar in 1966-1967 [in French English 76 (hbriel B1) 1975 A review tf the major insect pests of some tpland summary] Trop Agron 2313-21 crops in tlc Philippines Pages 86-111 in Ad Htoc panel of experts on

58 Doniiciano N L 1979 Chemical control of tie lesser corn stalk pest disease arnd weed problems in some rainfed crops Bangkok (lolsonno~paloIgnowslhts) (Zeller 1848) (I-epid(ptera Ihycitid ae) Thailand Food aid Agriculture Organization Roire Italy

on rice in Ponta Gressa Parara 1977-1978 [in Portuguese] Pages 77 Garg D K and N K Shal 1983 White grub outbreak on rainfed 291-303 in Anais 1 Reuniao de tecnicos cmiirizicultura do estado de dryland rice in Uttar Pradeslh Int Rice Res Ncwsl 81(4)16-17 Sao Paulo-Campinas Cati Brazil 78 Garg I) K and IP Tandon 1982 Major insect pests of rice on hilly

59 Dunsmore 1 R 1970 Investigations on tlie varieties pests and tracts of Uttar Pradesh India lilt Rice Res Newsl 7(11)11-12diseases of upland rice (hill padi) in Sarawak Malaysia Int Rice 79 Garrity 1) P 1983 Upland rice research an international bibliograpy Comm Newsl 1929-35 1975-1982 Iternational Rice Research hnstitnte PO Box 933

60 Elias 1R1965 The new decline of rice seedlings is not caused by Manila Philippines 121 p parasites [in Portuguese] Biol6gico (Sao Paulo) 313-6 80 Ghauri M S K 1982 A new subspecies of Iinorh)rone bull

61 Empresa Brasiliera de Pesquisa Agropecuaria 1984 Upland rice itt cornlus Slater (Hemiptera Iygaeidae) front Papua New Guinea on Brazil Pages 121-134 it Av overview of upland rice research rice and carpet grass Ihll lEntoniol Res 72133-137 Proceedings ofthe 1982 Blouak Ivory Coast Upland rice workshop 81 Gonzales F 0 PArregoces R I ihernandef and OT Parada International Rice Research Institute PO Box 933 Manila 1983 Insect and imite pests of rice arid their control in Latin America Philippines [in Spanish] la Federacion Nacional de Arrocerts (Fedearroz)

Bogota Colombia 60 n

38 IRPS No 123 January 1987

82 Grist D H and R J A W Lever 1969 Pests of rice Longmans London UK520 p

83 Hale P R 1979 Two subterranean pests of upland rice in Papua New Guirnca IntRice Res Newsl 4(2)16-17

84 Hale P R and J T Hale 1975 Insect pests of rice in the New Guinea Islands Rice Entomol Newsl 34

85 Harris W V 1969 Termites as pests of crops and trees Common-wealth Institute of Entomology London UK 41 p

86 Hattori 1 1968 A method to distinguish between grass leaf roller (Ctaplha rocis medinalisGuenee) and the rice leaf roller (Susurtia exigua Butler) [in Japanese] Plant Prot (Japan) 23167-170

87 Henry I E and 1A Onsager 1984 Experimental control of the mormon cricket Iahrosshnph by NJsena locustae(Aficrospora Microsporidae) a protozoan parasite of grasshoppers (Ort Acri-didae) Entomophaga 27197-201

88 Hidaka 1 P Vungsilabulr and SKadkao 1974 Studies on ecology and control of the rice gall midge in Thailand Tech Bull 6 TropicalAgricultural Research Center Tokyo Japan 113 p

89 Hincklcy A 1) 1963 Ecology and control of rice planthoppers in Fiji Bull Entomrol Res 53467 481

90 Hirao J 1982 Ecology and chtInical control of the rice leaffolder Jpn Pestic Inf 4114-17

91 Ho DT 1984 Sten borer incifience in rice ecosystems in KenyaEast Africa Pages 307-320 in A overview of upland rice research Proceedings of the 1982 Bouac Ivory Coast upland rice workshop International Rice Research Institute PO Box 933 Manila Philippines

92 Ho )T J 1Njokah and J GKibuka 1983 Studies on rice sten borers in Kenya with emphasis on ifaliarpiaipiraellRag Insect Sci Appl 465-73

93 Ichihara I T Ito A NIatsuo and M Sawada 1971 Chemical control of the rice striped stem borer in upland rice grown in plastic mulch [iii Japanese] Proc Kanto-Tosan Plant Prot Soc 1882

94 IclhiharaIand A Matsuo 1969 Damage caused by rice striped stett borer to upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1686-87

95 Ichhara I atid A MIatsun11970 Conside ration oft lie extraordinary prevalence of rice striped stem borer in upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1778-79

96 Institut de Recherches Agrolnliqes Iropicale et des Cultures Vivrieres 1978 Insect resistance Pages 12-13 in Upland rice 1978 atnual report Bouak Ivory Coast 27 p

97 International Institute for Tropical Agriculture 1983 Aphids itt rice roots I ITA Res Briefs 4(1-3)4-5

98 International Institite fi Tropical Agriculture 1983 Hopperburn in rice caused by brown plathopper IITA Res Briefs 4(l-3)6-7

99 International Rice Research Institute 1975 Annual report for 1974 PO Box 933 Manila Philippines p 216

100 International Rice Restarch Institute 1976 Annual repiort for 1975 PO Box 933 Manila Philippines p 228

101 International Rice Research Institute 1979 Cropping systems program Component technology development and evaluation Page 404 i Annual report for 1978 11O Box 933 Manila Philippines

102 International Rice Research Institute Department of Entomology 1981) Insecticid evaluation 1979 Tables 73 74 PO Box 933Manila Philipes

103 International Rice Research Institute Department of Entonmology 1981 Insecticide evalomtion 1980 Table 81 POBox 933 Manila Philippines

104 International Rice Research Institute 1981 Report of a meeting of Asian rice-based cropping systems entomologists Eleventh cropping systems working group meeting 18-22 May 1981 PO Box 933 Manila Philippines p 132 133 144

105 International Rice Research Institute 1984 Cropping systems program Pages 368-373 in Annual report for 1983 PO Box 933 Manila Philippines

106 International Rice Research Institutc 1984 Upland rice in Asia Pages 45-68 in An overview of upland rice research Proceedings of the 1982 Bouakamp Ivory Coast upland rice workiiop loternational Rice Research Institute 11O Box 933 Manila Philippines

107 Irshad M 1977 Grasshoppers in rice in Pakistan lit Rice Res Newsl 2(3)8-9

108 Ito R and TOno 1964 Upland lice its culture and varieties in Japan lIternational Rice Cormmission loth Meeting of the Working Party on Rice Production and Protection 3-10 March 1964 Manila Philippines Food and Agriculture Organization Rome Italy 12 p

109 Jeffrey M I and V A Dyck 1983 Effect of moonlight on lightshytrap catches of brown planthopper lit Rice Res Newsl 8(6) 13

110 Jennings P R and T Alicia Pineda 1983 Management ofthe rice plnthoppers Sogatodsorivi(coIl in Latin America Presented at the meeting on stable plant resistance FAO and lIndocsian I)inectorate of Food Crops Protection Bali lndonesia 29 Novcniber-2 l)ccemshyber 9 p

IllJimenez Gomez J A 1979 Rice insect pests i lla ios ()riCntales lil Spanish] Pages 99-103 in Rice ciltivation Instituto (olotibiatto Agropecuario Compndio No 29

112 Jongeleen F JJ Soesanto and A Mockidjo 1979 Studies oti pathogens of two species of Scarabacidac froinIndonesia Z Angew Enlomnol 87412-417

113 Jouvenaz 1) PC S lofgren and W A Baiks 1981 Biological control of imported fire antsa review of cirrent knowledge Bull Entomol Soc Am 27203-208

114 Kainoh Y K Shiniiu S MarUani Y Taiaki 1980 lost finding behavior of tie rice bu lplotori+JchinvittjI)allas (liciniptera Corcidac) with special reference to diet pattern ol aggregation and feeding on the rice plant Appl Entomol Zool 15225-233

115 Kakkar K 0 P Sharma and P I Sharma 1977 Note on Ihteronyrchus liodresRedt (Coleopicra I)ynastidae) as asporadic pest of paddy and maie inHimachal Pradesh Corr Agric 156-57

116 Kalode M131 79 Major insect pest problems and their control in upland rice (Unpublishcd mineogriph CWI oJNaaiat lni Chandaratat (198 1)original not seen)

117 Kalshoven L i 98 I The pests of crops in I ldoniesia (translated by P A vatn der Laan) 1 T Ichtiar B3aru-vati Iloeve lakarta Indonesia 701 p

118 Kamal N Q 1981 Suppression of whitebacked planto ner Sogatella fircaifra (Horvath) aLId rice lealfolder (nhal -cis licdb lis (Guenee) populations by natural cnclmie hlI) tiesis Grcgorio Araneta University Foundation Victoncta Park Metto Manila Philippines 179 Ip

119 Karim A N M Reaul 1975 A species ofliealy bug Trioni i70s ceres new oii rice in the Philippines Rice Entoniol Newsl 247

120 Katanyukul Wand C Chandaratat 1981 Insect pestsof uplaid rice and their control Report firstFAOUNlFi Tliailand training course on improved cultural practices for tplad rice Chiang Mai Thailand 5-23 Oct 1981 Food and Agriculture Organiation Rome Italy 22 p

121 Kern F 1955 lhletwioa(I)uninemts) bhhit Biturm(Coleoptera) a new pest of sugarcane inVenezuela [inSpanish with Etnglish sunmary] Agron Trop (Maracay) 527-29 (Rev Appl Eitoniol A4572)

122 Khan K M and S Ghai 1974 White grubs ald their control in India Pesticides (1Bombay) 819-25

123 Kiritani K 1975 Integrated pest management of rice pests Pages 66-70 itt Proceedings platning workshop ott cooperative hIeld research in pest managementP SMotooka ed East-West Center Honololu Hawaii USA

124 Kisimoto R 1967 Ecology of insect vectors forecasting and chemical control Pages 243-255 in The virus diseases of tle rice plant Proceedings of a symposium atthe International Rice Research Institute April 1967 The Johns Hopkins Press altimore Maryshyland USA

125 Kisimoto R and V A Dyck 1976 Climate and rice insects Pages 367-391 in Climate and rice International Rice Research Institute PO Box 933 Manila Philippines

126 Kureha S S Kobayashi SAsari and T Ishihara 1977 Life history and control of the mole cricket Grrllotalpa africana Palisot de Beauvois [in Japanese English translation 7520 1RRI library] Proc Kanto-Tosan Plant Prot Soc 21192-197

127 Kushwaha K S 1981 The black beetle as a pest of rice in Haryana India Int Rice Res Newsl 6(4)14

128 Lawani S M 1982 A review of tile effects of various agronomic practices on cereal stem borers Trop Pest Manage 28266-276

129 Leuck D B 1966 Biology of the lesser cornstalk borer in South Georgia J Econ Entomol 59 797-801

130 lever R J A W 1955 Insect pests of the rice plant in Malaya Int Rice Comm Newsl 1612-21

131 Li Ching-Sing and Shin-Foon Chiu 1951 The study ofthe rice gall midge Iachydiplosis orvza Wood Mason [in Chinese with English summary] 1Taiwan Agric Res 21-13

132 Iitsinger J A 1984 Assesament of the need-based application for rice Proceedings TechnologyTransfer Workshop 15-16 March 1984 International Rice Research Institute 1PO Box 933 Manila Philippines 6 p

133 Litsinger J A 1984 Environmental diversity and insect pest abundance with reference to the Pacific Pages 285-307 in Proceedings sub-regional training course on methods of controlling diseases insects and other pests of plants in the South Pacific 4-20 Oct 1982 Vaini Kingdom of Tonga Nukualofa

134 Litsinger J A RF Apostol and M BObusan 1983 White grub Leucopholis irrorata (Coleoptera Scarabaeidae) pest status popula-tion dynamics and chemical conllol in arice-maize cropping pattern in the Philippines J Econ Entoinol 761133-1138

135 Litsinger J A C de la Cruz F Raymundo A T Barrion M D Lumaban J P Bandong M S Venugopal F Paragna and E Balete 1982 Insect pests and insecticide reponse of dry-seeded bunded rice Pages 179-193 in Report of workshop on cropping systems research in Asia International Rice Research Institute Los Bafos Philippine

136 Litsinger J A M 1) Luinaban J P Bandong A T Barrion W Katanyukul and N Panda 1978 Insect pest management in rainfcd lowland rice agroecosystms of tropical Asia International Rice Research Conference 17-21 April 1978 International Rice Research Institute Los Bafios Philippines

137 Litsinger J A and K Moody 1976 Integrated pest management in multiple cropping systems Pages 293-317 in R I Papendick P A Sanchez and G B Triplett eds Multiple cropping American Society of Agronomy Madison Wisconsin USA

138 Loevinsohn M E 1984 The ecology and control of rice pests in relation to the intensity and synchrony of cultivation Ph D thesis University of London 36) p

139 Loevinsohn M E J A Litsinger N Panda M A Austria R F Apostol C Andrion and J Bandong 1982 A basis for predicting rice insect pest abundance in cropping systems Paper presented at the 1982 International rice research conference on rice and cropping systems 19-23 April 1982 International Rice Research Institute Los Bafios Iaguna Philippines 25 p

140 Lopez A W 1930 Repot of the Entomology Department Pages 145-172 in Annual report of the Philippine Sugar Association 1929-30 Manila

141 Lopez AW 1931 Entomology Department Pages 227-266 in Annual report of the director or research 1930-31 Philippine Sugar Association Manila

142 Lopez A W 1932 Entomology Deparment Pages 252-279 in Annual report of the Philippine Sugar Association 1931-32 Manila

143 Lungibill P and G G Ainslee 1917 The lessercornstalk borer US Dep Agric Agric Res Serv Tech Bull 539 27 p

IRPS No 123 January 1987 39

144 McGuire J U Jr and BS Crandall 1967 Survey of insect pests and plant diseases ofselected food crops of Mexico Central America and Panama U SDepartment of Agriculture International Agriculshytural Development Service Agricultuial Researrh Service October 1967 11 p

145 McNeill Sand T R ESouthwood 1978 The role of nitrogen in the development of insectplant relations Pages 77-9 in lliochemical aspects of plant and animal coevolution J B Ilarbornu ed Academic Press London UK

146 Maebara H 1955 Spatial distribution of lastinstiarlanaeof purplish stem borer (Sesamia iqferens Walker) hibernating in an upland rice field Bull Kagoshima Univ Fac Agric 438-41

147 Mallick E H and N G Hajra 1977 Response of rice plant to defoliation treatments at different developmental stages Indian Agric 2151-56

148 Mansingh A 1971 Physiological classification of dormancies in insects Canl Entomol 103983-10)9

149 Martell C G 1974 Catalogue of plant feeding insects of Mexico [in Spanish] Fitofilo 27(69)1-176

150 Martinez 0 J 1982 Principal rice pests in Ilanos orientalis [in Spanish] P~ages 80-94 in Curso deact nalization de conocimientos en arroz Villaniencis Instituto (olonmbiano Agropecuario

151 Martins J F da Silva F Ferrcira A S Prabhu F 1 P Zimmermann 1980 Use of pesticides for the control of the main subterranean upland rice pests esqui Agropecu Bras 1553-u

152 Martins I F da Silva F Fe eira rod BdamSilveira Pinheiro I Q2 Damage simulation caused by defoliators of upland rice [in Portuguese with English summary] Pesqui Agropecu Bras 171113-1119

153 Martins 1F da Silva N(iyuen van Tan and 1Ida Silveira Pinheiro 1981 Stem borer resistance in upland rice associated with plant morphological characters [in Portuuce Iesqui Agropecu Bras 16187-192

154 Masaki S 1980 Sumnmer diapause Ann Rev Entomiol 251-25 155 Masuzawa T 11 Suwa and F Nakasoji 1983 l)ifkleences of

oviposition preferencc and survival rate of two skipper butterflies Parnara guttata and PIchlpidai iathiais (Iepidoptera HIesperiidac) on rice plant and cogon grass New Entomol 3247-56

156 Maurya D M and C P Vaish 1984 Upland rice in India Pages 69-92 in An overview oftpland rice research Proceedings ofthe 1982 Bouakamp Ivory Coast upland rice workshop International Rice Research Institute IO Box 933 Mnila Philippines

157 Menon M G R R N Katiyai S Kuar and S S Misra 1970 Termites attacking the paddy crop Sci Cult 36294-295

158 Miyahara Y and 1 Nishikubo 1967 Behavior of young nymphs of the red rice root aphid Iholahsipiun ri liabhdontinalisSasaki in relation to movement from the leaf to the roots of upland rice [in Japanese] Pioc Assoc Plant Prot Kyushu 1363-66

159 Mo ljoa Tjien 1952 White grub of upland rice (Holotrichiahelleri Brsk) [in Indonesian with English summary] Contrib Agric Res Stn Bogor No 13134 p

160 Mochida 0 and T Suryana 1979 The occurrence of the brown planthopper and ragged stunt on lowland and upland rice ill Central Java and logyakarta in tile late of January 1979 Entomnolgy Survey Report No 15 Central Research Institute of Agriculture (CRIA) Sukamandi Indonesia I February 1979 9 p

161Monet C 1979 Varietal resistance to upland rice pests [inI French] Pages 35-39 in Annual report 1977 Institut des Savanes Department des Cultures Vivrieres Bouak6 Ivory Coast

162 Monte 0 1942 A caterpillar ofrice-fields[in Portuguese] Biol6gico (Sao Paulo) 8161-163 (Rev Appl Entomol A31414)

163 Munroe I) D 1979 Pest management strategy for hill rice in four resettlement areas in Sarawak Malaysia Pages 89-102 i1 Guidelines for integrated control of rice insect pests Food and Agriculture Organization Plant Production and Protection Paper 14 Rome Italy 115 p

40 IRPS No 123 January 1987

164 Nakasuji F M Ishii and Jen-Zon Ito 1984 Rice skippers in Taiwan and their life histories ZZrC (Taipei) Tech Bull 82 10 p

165 Navas I) 1976 Fall armyworum in rice Pages 99-106 il Ecological animal control by habitat management Proc Tall Timbers Con-ference 28 Feb-I Mar 1974 Fall Timbers Res Stn Misc Publ 6 Tallahassee Horida USA

166 Nickel 1 1 1961 Biological control of rice stem borer A feasibility study Tcch Bull 2 International Rice Rcarch Institute PO Box 933 Manila Philippines IIl p

167 Nigan P M and R A Vernia 1985 Insect pests of uplaild rice in Uttar Pradesh Int Rice Res Newsl 10(6)22

168 Nilakhe S S 1985 -cological observations on spittle bugs with emphasis on their occirence in rice Pcsqtui Agropecu Bras 20407-414

169 Nixon G EJ 195 1 lie association of ats with aphids and coccids Commonwealth Iusti lcte of Entonology Iondon UK 36 p

170 Nurdin F an Ishak Manti 1985 Study oit seedling fly of upland rice Interlational Upland Rice Moitoring Tour Report 26 lebruary-2 MarLI Sukaranti West Sumatra

171 Oka 1 N 1979 Ciltural conttol of the brown planthopper Pages 357-369 in Brown planthopper threat to rice production in Asia International Rice Research Institutc 1p0 Box 933 Manila Philippines

172 Ohni M and A Vilnaii 1975 Rice insect pests of Italy biology of Sipha grcria (Kaltenbach) and its control [in Italian] Riso (Milan) 2459-71

173 Otanes F Q atnd 1 I Sison 194 1 Pests of rice Philipp J Agric 12211-259

174 Painter R I 1951 Insect icsistance in crop plants University Press of Kansits Lawrence Kansas USA 520 p

175 Pantua P C and 1A litsinger 1981 Arthropod abundance in continuous vs biannual rice cropping systeis Int Rice Res Ncwsl 6(2)20-21

176 Pathak M ) and V A l)yck 1975 Sttudics ott insect pests of upland rice Pages 186-191 in Major research in upland rice International Rice Research Institute PO Box 933 Ilanila Philippines

177 Patil B R B M IHasabe and 1) S Ajri 198 I Chemical control of white gritb lIditriihinhorraia Fabr intfesting groundntut paddy atnd sorghum Indian 1Entomol 43232-236

178 Pawar A 1) 1974 (ollection and identification of insect pests of rice rlerminal repirt lInternational Rice Research Institute PO Box 933 Manila Philippines 72 p (tnimco)

179 IerrajU A aid A S Reedy 1965 Observations ott diapausc itt tie steti borer ohrice Rice News Teller (lai-Mar)9-13

180 Pont A C 1972 A review of the Oriental species of theri ona Rondani (I)iptera Muscidac) of econtomic importance Pages 27-102 in Control of sorglium shoot fly M G lomtani and W R Young eds Oxford atd IH1 Ptthl Co New I)clhi India 324 p

181 Prasad S C 1PChand aid M 1PSingh 1983 Incidence of rice gall midge in the upland plateau region of Hihar Oryza 20235-236

182 Rai P S 198 0 Nisaga simpex damage to rice in the hill tracts of South India Int Rice kles News[ 5(I)12

183 Rathorc Y S 1969 Gonocplhahli dcprssunt Fabr as itpredator of (tilo zonwIlhis (Swinhoc) and other moths in field cage at Patnagar Uttar Pradesh Indian 1Entotnol 3127

184 Reed W 1983 Crop losses causcd by insect pists in the developing world Proceedings 10th Internatial Congress of Plant Protection lrighton UK 174-80

185 Reedy M V 1981 Ateactogasterinhens(Walker)a new root-grub pest of tipland paddy and inaize Curr Sci 501076

186 Reveno M A 1968 Catalogue of the insect pests of economic crops in Colombia [in Spaiish] Asociacion Latino Americana de Entomo-logia (AIAE) 129-33

187 Reyes G M 1957 Rice dwarf disease in the Philippines Food and Agriculture Organiiation Plant Prot Bull 617-19

188 Rice S E A A Grigarick and M 0 Way 1982 Effect of leaf and panicle feeding byarmyworm (Lepidoptera Noctuidae) larvae on rice grain yield Environ Entomol 7593-595

189 Risch S J 1) Andow and M A Altieri 1983 Agioccosystem diversity arnd pest control data tentative conclusions and nizw research dirctions Environ Entomol 12625-629

190 Rivera C T 1956 Three organic insecticides in thecontrol of insects affecting upland rice Philipp Agric 39465-472

191 Roffrey 1 1972 Locusta outbreaks in the Philippines Acrida 1177-188

192 Rojas R 1) 1941 The effect of pruning at varying ages upon the yield of Ramairio Phipp Agric 29851-860

193 Roonwal M i 1979 Ternite life and termite control in tropical South Asia Scientilic Pnbl odhpr India 139 p

194 Rosetto C1 F da Silva Martins N C Schmidt and L EAzzini 1978 Damage caused by pasturespittle bugs 4Dvoi)lavopictaandDJ schach) on rice lBragantia Nota 6 (Matcli)25-27

195 Rosztto C1- S S Neto I- Amantc I) Iink 1Garcia-Vieira D M de SoLza N V Bansa t and A M de Olivcira 1971 Rice pests of Brazil [in Portuguese] Pages 149-238 inrFA()-International Rice Commission 2nd meeting of the Rice Committee for the Americas 6-11 l)ecember 1971 Pelotas Rio Grande do Sul Brazil

196 Rothschild G H L 1969 Obseratiois on the armyworm Spodoptera matritia acronrctoiiles Gn (Lep Noctuidae) Bull Entomol Res 59143-160

197 Rothschild G 1- L 1974 Parasites of the armworm Spolopiera mauritiaacroncitoihes in Malaysia Fntornopliaga 19293-299

198 Sachan i N S K Gangwar and 1N Katujar 1980 Field crickets damaging cages in North Eastern IHill Region Indian J Entomol 42526-527

199 Sands 1) P A 1977 The biology and ecology of Lepocorisa (Hemiptera Alydidac) in Papua New Guinea Res Bull 18 Departshyment of Primary Industry Port Moresby Papua New Guinea 104 p

200 Sarawak 1968 titomology division Pages 97-106 in Report of the research branch of the Department of Altriculture 1966 Kuching (Rev AIppl Eintomol A56603)

201 Satpathy JM M S Das and K Naik 1977 Effect of multiple and mixed cropping on the incidence of scine important pestsI Entomol Res 178-85

202 Sauer IHF G 1939 Notes Ott Elasmpnlpualtsligniosellths Zeller (Lep Pyr) a serious pest ofcereals in Sao Paolo [in Portuguese] Arq Inst Biol 10199-206 (Rev Appl Entotnol A28378)

203 Satinders 1L A 1 SKing and C L Vargas S 1983 Crop pests of Central Anerica with t bibliography [in Spanish] Centro Agroshynotnico Tropical de Investigacion y Ensenanza Turrialba Costa Rica (CATIE) Tec Bol 9 90 p

204 Sauphanor B 1985 Some factors of upland rice tolerttce to stem borers in West Africa Inseu Sci Appl 6429434

205 Sen P atnd S Chakravorty 1976 Stentflics in Haritighata Nadia West Bengal India [it Rice Res Newsl 1(12) 15

206 Senapai B and 1M Satpathy 1983 The seedling fly Atherigona orrzae Malloch is a pest of rice in Orissa (India) J Entomol Res 777-79

207 Shhjahan M S Alatn M Ah nad and BN Islam 1983 A new scarabaeid beetle pest of dryland rice in Bangladesh Int Rice Res Ncwsl 8(I)16-17

208 Sharma G L S C Modgol and R C Gautam 1979 Effect of depth of submergence ott incidence of bacterial blight and white grub infestation in transplanted rice Int Rice Res Newsl 4(6)12-13

209 Sharma S 1985 Scarabaeid beetle outbreak itt upland rice Int Rice Res Newsl 1(1(l)24

210 Shiraki T 1917 The paddy borer Schoenobhits incertulas Walker [achaku Agric Exp Stn Formosa 256 p

211 Silva A de Brito and 11P Magalhaes 1981 Insect pests of rice [in Portuguese] Circular Tecnica No 22 EMBRAPA Centro de Pesquisa t ecuaria de Tropico Unido (CPAJU) Belem Brazil 14 p

212 Singh S A 1979 Emergence of rice leaf butterfly as a rice pest in Manipur India int Rice Res Newsl 4(4)16-17

213 SinghS A 1983 Changing trends of insect pestsof rice in Manipur Int Rice Res Newsl 8(f)12

214 Siwi S S and P H van Doesburg 1984 Lenovorisa Latreille in Indonesia (Heteroptera Coreidae Alydinae) Zool Meded (Leiden) 58117-129

215 Soejitno J 1977 Reaction of some upland rice varieties toseedling fly eltherikonaexigua Stein [in Indonesian] Siri Hana I Penyakit 630-35 Lembga I osat Fenclilian Pertanian Bogor Indonesia

216 Socnardi 1967 Insect pests of rice in Indonesia Pages675-683 in The major inse pests of thu rice plant Proceedings of a symposium at the Internatital Rice Rec Wi itt S pternber 1964 The Johns Hopkins Press Baltimore Maryland USA

217 Soto P E A T Petcz and I W Buddenhagen 1976 Survey of insect pests and diseases of rice in different ecological zones in Nigeria Rice Entomol Ncwsl 435-46

218 Soto 11E and Z Ziddiqi 1976 Screening for resistance to African rice insect pests Varietal improsetnent sCmi n r Bouak Ivory Coast 13-17 Sept 1976 West Africa Rice l)evelopment Association Monrovia Iiberia

219 Squire F A 1972 Entomological problems in Bolivia PANS (PestArtic News Sumr) 18249-268

220 Stapely 1 II 1978 The rice leaf roller in the Solomon Islands Int Rice Res Newsl 3(l)13

221 Suenaga H 1937 On aphids attacking the roots of rice Oyo-I)oubts Zasshi 9150-152 [in Japanese] (Rev Appl Entomol A25745)

222 Suenaga H 1952 On the seasonal abundance of three subterranean aphids on upland rice Bull Kyushu Agric Exp Sti 1249-262

223 Sutynso Yas and Dandi Soekaria 1964 Beberapa hana yang ttenperang tana man padi tanah kering [in Indonesian] Rapat kerja Padi Tanah Kering 6 sd 8 Juily 1964 Kaliurang 9 p

224 Tallamy I) W and R F Denno 1983 Alternative life history patterns in risky environments in example from lace bugs Pages 129-147 in Insect life history patterns R F l)cnio and If Dingle eds Spring-Verlag 225 p

225 Tamura 1 1939 Observations on some insect pests of field crops [in Japanese] 1 Plant Prot 26414-420 (Rev Appl EntomolA27-556)

226 Tamura and Suzuki 1964 Influences of the early-transplanting practice in rice culture upon development of the rice stem borer Chilo siql)ressalisWalker Bull Ilokuriku Natil Agric Exp Stn 761-94

227 Tan J P 1924 The rice root aphid Drowhi ihirsoia Philipp Agric 13277-288

228 Tanaka 1 1957 Taxonomy and distribution of some subterranean aphids injurious to upland rice in Japan with description of a new species [in Japanese English summary] Botyu-Kagaku 22168-176

229 Tanaka T 1961 The rice root aphids their ecology and control Spec [full College of Agriculture Utsunomiya University Japan F3 p

230 Tauber M IC A Tauber R Nechols and 1Obrycki 1983 Seasonal activity of parasitoids control by external internal and genetic factors Pages 87-108 itt I)iapause and life cycle strategies in insects V K Brown and I Ilodek eds )r W Junk The Hague Netherlands

231 Taylor W F 1972 Effects of artificial defoliation (simulating pest damage) on varieties of upland rice Exp Agric 879-83

232 feotia T P S and S Natud 1966 Biotoomics of the rice skipper Parnaraniathiis Fabricius (Lepidoptera Hesperiidac) Indian J Entomol 28181-186

233 Teran F 0 1971 Rice pests of Santa Cruz [in Spanish Ministerio de Agricultura Sita Cruz Bolivia Bull 45 8 p

234 Than Itun 1976 Population dynamics of the yellow rice borer Tryporza itcertulas (Walker) and its damage to the rice plant MS thesis University of the Philippines at Ios Bafios Laguna Philippines 92 p

235 Uichanco L B 1911 Pests Pages 55-74 it Rice Philipp Agric 11-200

IRPS No 123 January 1987 41

236 van der Laan P A 195 1 Life history of the rice seedling fly Contrib Gen Agric Res Stn Boor Indonesia 118 15 p

237 van der Laan 11A 1959 Correlation between rainfall in the dry season and the occurrence of the white rice borer (Scirpolhaga innotala WIk) in lava Entomol Fxp Appl 212-20

238 Van Driesche R G 1983 Meaning of percent parasitismin studies of insect parasitism Fnviron Entomol 121 I1- 122

239 Venugopal M S and A litsingcr 1984 Fffect ofcarbofuran on rice growth Prot Ecol 7313-317

240 Vietnam Directorate of Rural Affairs 1962 Annual work progress report on crop improvement program of rice sugarcane vegetable and field crops (lor the period from July 1961 to June 1962) (11) 299 p (Rev Appl Entomol A51526-527)

241 Wan M T K 1972 Observations on rice leaf- and planthoppers in Sarawak (Malaysian Borneo) Malay Agric 148308-335

242 West Africa Rice I)esclopnient Association 1978 Annual research report Vol 2 Statistics Monrovia Liberia

243 West Africa Rice Development Association 1979 Annual research report p47 Monrovia Liberia

244 Williams ) F and CS lofgren 1981 Fli I illy l-468 anew bait toxicant for control of the red imported lire at Fla Entomol 64472477

245 Williams I) 1 1970 The mealybugs OFlomoptera Coccoidea Iscudococcidac) of sugar-cane rice and sorghum Bull Entomol Res 0109-188

246 Williams D) 1 1982 Pulinaria icervi (Signoret) and its allies ott sugar cane and other grasses Btll Entoinol Res 72111-117

247 Wirjosuiardjo S 1976 Study ofchenmical cont rot of insect pests ott upland rice Imiu Pertanian (Jogjakarta) 1323-329

248 Wirjosuhardjo S and N Wirarta 1976 Chemical control of the white grub tIolorithiatlltriin upland rice in M uneng Irobolinggo Indonesia) [in Indonesian] Imut Pertanian (Jogjakarta) 1330-332

249 Yano K T Miyake and V F Eastop 1983 The biology and economic importance of rice aphids (Henuiptera Aphid idae) a review Bull Entomol Res 73539-566

250 Yelp D and P Rhodes 19(4 lhe success of drilled dryland rice iii the Fiji Islands International Rice Commission Working Party on Rice Production and Protection Manila Philippines 3-10 March1964 Food and Agriculture Organization Rome Italy 13 p

251 Young W R 1981 Fifty-five years of research on the sorghum shootfly Insect Sci Appi 23-9

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ISSN 05-3862Other papers in this series TITLES OF NUMBERS 1-45 ARE LISTED ON THE LAST PAGE OF NO46 THOSE OF NUMBERS 46-70 ARE ON THE LAST PAGE OF NO 71-80AND THOSE OF NUMBERS 71-1O0ARE ON

THE LAST PAGE OF NO 101-122

No 101The economics of hybrid rice production in (hina No 113 Boron toxicity in rice No 102 Rice ratooning No 114 Energy analysis rice production systems and rice research No 103 Growth and development of the (Jeep water rice plant No 115 Production risk and optimal fertilizer rates an application of the

No 104 Faridpur acomputer-assisted instruction nmodel forrainfed lowland random coefficient model No 116 Consumer demand for rice grain quality in Thailand Indonesia andrice

No 105 A readingand listening comprehcnsion test inEnglish for non native the Philippines speakers applying for tiraining at IRIRI No 117 Morphological changes in rice panicle development a review of

No 106 Rice grassy stunt virus 2 a new strain of rice grassy stunt in tire literature

Philippines No 118 IRRI-Korca collaborative project for the development of cold-No 107 Physical losses and quality deterioration in rice plstprodnCtion tolerant lines through anthei culture

systemis No 119 Problen soils as potenial areas for adverse soils-tolerant rice No 108 (olptblication ofIRRI materials a survey of translators and varieties in South and Ssuthcasi Asia

publishers No 120 Changes isinall-farm iLtireliing techii)hgy in Thailand and [he No 109 Classification of Philippine rainfall patterns Philippines No 110 Contributions of modern rice varieties to rutrition in Asia No 121 Landforms and modern rice varieties No Ill Changes in rice breeding in 10 Asian countries 1965-84 No 122 Yield stability and modern rice technology No 112 Design parameters affecting the performance of the IRRI-designed

axial-flow pump

Page 17: UPLANDRICE. INSECTl--1i M T -E!R.EOLOU t RTANCE ,AND …delphacid.s3.amazonaws.com › 6958.pdf · The significance of abiotic factors in upland rice insect ecology will be apparent

IRPS No 123 January 1987 17

favorable weather and resume activity after favorable Armyworms skippers and locusts actively travel hundreds weather begins usually in response to photoperiod or of kilometers in air fronts such as intertropical convergencethrough a delayed response to a stimulus such as heavy zones to descend onto upland rice fields far from their rainfall (237) Not all individuals of a population have the breeding grounds (31) Several generations can build ip on same threshold for environmental responses for entering or alternate hosts in tile grass family before rice is plantedterminating diapause -- this is seen as a mechanism to These large insects are active fliers Smaller insects such as prevent total mortality if a false cue appears such as a thrips aphids and planthoppers passively migrate long sudden warm spell in winter or rain in the middle of the dry distances by the wind season (154) Otherspecies ean readily travel tens of kilometers - stem

Probably more upland rice insects undL go dormancy borers leaffolders leafhoppes cutworms spittle bugs leaf than we realize Candidates are Atherigona spp root beetles seed bugs ants and termites aphids root weevils crickets leaf beetles thrips and seed The least dispersive species are white grabs mealybugSbugs Insects known to entervarious states of aestivation are seedling maggots crickets weevils and gall midgesleptocorisaspp adults (199) Maliarphascparatellalarvae Each species has evolved a unique set of attributes to (9) Scirpophaga innotata larvae (117) Chilo partelhtsv enable it to survive and adapt to the changing uplandlarvae (21) ScirpopThagainccetulas (179) white grub larvae environments Several evolutionary avenues lead to fitness (134) Patangasuccincia adults (6 82) and Schistocerca (Table 2) but there is no reason to believe one set of gregariaeggs (82) attributes is better than another (224)

Insects known to enter hibernation are LodeIphaDstriatellus nymphs (124) Chtio Tvlpressalis larivae (125) DroughtSesaia inferens larvae (146) but not S calavisis(2) Well-drained rainfed upland soils are subject to drought ifIDiaraea saccharalis larvae (75) Scirpophaga- tcerlttas rains do not fall within 2-3 wk Prolonged drought followedlarae(5 lacchrli s lae (75)saSlOhagasshoperseggs (82)and felo pidas athias pOpae (232) by favorable rains stimulates armyworm and locust outshybreaks but mealybtigs root aphids and thrips become

Lcpmotorisa oratorisand L acuta adults aggregate on numerous onlyafter dry spells for two reasons First heavy trees sugarcane or other shady moist sites L paluwanmsis rain normally kills soft-bodied foliage-feeding thrips and confined to Sulu and Palawan Philippines lives on the mealybugs and soil-inhabiting root aphids and root mealyshygrass Brachiariamutica (Forsk) Stapf in the absence of bugs Drought eliminates this source of mortality Second rice The adults can live for 5 mo When aestivating their the rice plant responds to drought by breaking down rate of metabolism drops and they do not feed They will proteins into soluble N compounds which enter the phloem however puncture plant tissues of their aestivation host in and are taken up by these sap-feeding insects (245) Greater search of water This behavior has led to errors of the host nutrition therefore leads to greater survivorship and range of rice bugs (I) Aestivation quickly terminates with fecundity rain probably meaning aestivation is not deep Small rice area

Some stem borers aestivate as mature larvae in rice Except in the highly mechanized large landholdings of stubble or in straw stacked as livestock feed However Latin America upland ricefieldstypically are patchy within moriality increases proportionally with the duration of the a highly diverse flora The small fields result in higher ratios dry season even if the stubble remains undisturbed If the of perimeter to area than with the typically larger lowland stubble is plowed for a crop following rice few stem borers ricefields Upland ricefields themselves may be intercroppedsurvive Rainfall terminates dormancy but it normally with cereals legumes or root crops Mixed intercroppingtakes several weeks for moths to develop and emerge with a wide variety of species also occ in tribal slash-andshy

Last-instar white grub larvae tunnel 1-2 m deep in the soil burn agriculture (50 74) to construct pupal chambers in which they aestivate The Upland ricefields tend to be small because of the labor larvae are very sensitive to dryness so pupal chambers are needed to clear land in slash-and-burn areas and to till land sealed to coierve body moisture Aestivation terminates to minimize weeds Rice requires more tillage than does with the first heavy rains but as in stem borer 3-4 wk pass maize because rice competes less well with weeds Uplandbefore adults develop rice is more a subsistence than a cash crop and small areas

Insects may respond differently in each region to cold (I ha) can feed a family temperature or drought Biotypes or loal populations may Small fields favor some insect pests such as some ant evolve (56) Local populations of Scirpophagaincertulasor species that can forage from more permanent field borders Patangasuccincta may have become adapted to prevailing Trees provide food and mating sites for white grubs (134) as conditions well as aestivation sites for LetTocorisa spp (199) Field

iagiliti The ability to disperse combined with high border grasses plus tIhose within a ricefield provide alternate fecundity orshort life cycles enable some upland rice pests to food for all but the highly monophagous upland rice better exploit the temporarily favorable upland habitat colonizers and a habitat for natural enemies

0 z

Table 2 Life history patterns of upland rice insects or insect groups

Host range Longevity Dormancy Vagility FertilityMonophagy Oligophagy Polyphagy Low Medium High Yes No Low Dispersive Migratory Low Medium High(Oryza spp) (grasses sedges) (angiosperms) (lt3 wk) (gt2 mo) (lt1 kin) (lt50 eggs) (gt500 eggs)

Ants xTermites x x x

Mole crickets x x x xx x xField crickets x xx

White grubs x x x xx xRoot aphids x x xxMealybugs x

x x x xxSeedling maggots x

x x x x xLeaf beetles

x xx xArmyworms x x xx x

Thrips x x xx xChilo spp x x xx xMaliarpha x x x x

xLocusts x x Xx x xGall midges x x xxiVephotettL iirescens x x xx x x xLeprocorisaspp x

x xLeaftolders x x x

Sogatellafurcifera x

xx x x x x x x

00

IRPS No 123 January 1Q87 19

All this appeals to favor insect pest buildup but the notwithstanding the release of stored nutrients in response contrary is true Loevinsohn (138) showed that insect pest to drought that temporarily benefits philoem feeders populations respond exponentially with the proportion of land devoted to rice up to about 75 of the area then the M POR ANCE I I P1A 1) RWI P1St S

rate of response declines In most upland rico areas tie rice crop occupies less than 50i of the area- therefore the Visits to upland ricefields normally reveal 1kw insect pests potential for population buildup is low This relationship Loevinsohn (138) and Locvinsohn ct al (139) compared may not hold for seed pests that can locate small ricefields insect abundance using annual light trap atchcs in upland

Another outcome of the highly diverse flora and phical rainfed lowland and irrigated lowland sites in the Philipshyenvironment of upland rice isthateach upland ricearca has pines Although only one upland site was studied data a unique composition of insect pests showed equal or fewer insects at the upland site than at the

For example in Tanauan Batangas Philippines the rainfcd lowland sites and fewer than at the irrigated sites main upland rice pests are rice leaffolder(RLF) armvworm (Table 3) M Lltiple regression analysis showed that the key Aythntnaseparata WBPH ants Soli psis gentitataand factors explaining insect abundance among a set of cropping white grub Leucopholis irrorata Sixty kilometers away in intensity variables were the number of rice crops grown per Real Quezon the main pests are seedling maggot Athe- year followed by area devoted to rice Table 3 shows that rigona orae flea beetle (Thacitcmnra hasalis thrips the upland rice site had only I rice crop perym rand that rice Stenchaetothrips hiurnmis and rice bug Leptocorisa was planted in only 20Vi of the area Fertiliter and oratorius In a third Philippine upland rice site in Claveria insecticide use had little bea ring on the trend The armters at Misamis Oriental Mindanao the main pests are ants the upland site applied 60 kg N ha higher than at most Solenopsis gemtinata seedling maggot Alieriona orvzae irrigated sites white grubs Holotrichia in)iUaiuoan t and Lucopholiv Insect occurrence at the upland site was omparable to irrorata stem borers Sesamia ifiArelr (hio auriciliusC that at the Cagayan rainfed lowland site parlicularly in 1982stpessallis Scirpophaga incertulas S innoiata and when drought prevented many farmers fron planting(only

4c ona chrrsurapella root aphids Tetraneura niriah- 40(of the rice area eventually was planted that year) M ore dunrinalisand Rhupaosiphunm riahbdminalis lcaffolders favorable weather occurred in 1980 and 1981 which Marasniapataltis and Chaphalocrocis Medinalis army- supported greater rice areas and consequently more insects worm MIthinna separata and rice bug Leltucorisa The most intensive site for insect pests was South Cotabato oratoriusThe great diversity of insect problems is typical Mindanao where farmers plant 23-25 rice crops per year fer upland rice and makes control efforts more difficult with irrigation The Zaragoa Nutva Ecija site also had particularly regarding breeding for insect resistance high pest abundance and at this site the fields were highly

asynchronous Asynchrony was the third most important Low yield potential Iactor in explaining rice insect abundance Most upland rice is grown on low-nutrient and mineral- Rice cropping intensity in upland areas is bound to toxic soils which when combined with eiratic rainfall remain low Only one rainid upland crop is possible per make upland rice a highly riskycrop for the farmerto invest year (Farmers in Claveria Misamis Oriental Mindanao in costly soil amendments and land management Often Philippines tried asecond upland ricecrop i 1984a ycarof upland rice isgrown far from markets making inputs even favorable rains but no harvest was possible b-cause of more expensive Farmers growing a subsistence crop hosi- drought blast and birds) Rice area isaso bound to remain tate to invest in costly inputs to raise yields low because of the high labor and power rcquirelmenlts to

Even having varieties that could double existing yield prepare the land (because of weeds)and because upland rice potential under present management levels would probably wvill not compete with lowland rice as a cash crop Some not prompt many farmers to use purchased insecticide upland rice sites with prolonged rainy seasons howeverare unless tile risk of crop failure were reduced such as it was planted asynchronously such as the Clacra site with irrigation systems in lowland rice The seasonal abundance of Upland rice insect pests

The Tanauan Batangas site is atypical for upland rice attracted toa light trap wasgrapied for crop year 1980-81 in The soils and rainfall pattern are favorable and because of Tanauan FBatangas (Fig I 2) W13P11 was he most its nearness to Manila farmers have cash resources ftm common rice hopper and its population peaked at the sales of vegetables to purchase tertilier They applied an midgrowth stage as it does in lowland rice D-Vac suction average of 60 kg Nha after particle initiation and obtained samples showed an earlier peak than that from a light trap 25-35 tha yields The high fertility isone reason why RLF indicating emigration W13PI usually emigrate after the armyworms and WBPH developed into large numbers vegetative stage Populations in 1980 reached levels that Farmers however used no insecticide on upland rice caused patches of slight yellowing in some fields WBPH although they did on vegetables probably cause hopperburn in Batangas They immigrate

The poor growing conditions fOr rice generally mean year-round as shown by low levels in the light trap BPH poor growth and low fecundity for insect pests fIeding on it however were scarce in the light trap and not collected with

Table 3 Abundance of rice insect pests and factors in rice cropping intensity of Philippine upland rainfed lowland and irrigated lowland locations where light traps were operated daily for atleast I yr -I

0Rainfed Rainfed lowland Irrigated lowlandupland

Pangasinan Cagayan lloilo Santa Maria Nueva Ecija South Cotabato Tanauan Caaringayan Caaringayan Bangag Bangag Bangag Buray

LagunaCale

Batangas Manaoag 1980-81 Santa Motiica 1982-83 Cabanatuan Zaragoza Avancena NaniamanSolana 1981-82 1982-83 Oton

1980-81 1979-80 Oton 1981-82 1981-82 Koronadal Koronadal1980-81 1979-80 1979-80 1983 1983

Insect (kerosene light trapNephotcttixcatch nolyr)virescens 650 1240 500 1800 330 210 3400 1600 160N nigropictus 210 - 360 18400 5500 2900- 140 90 20 - shy 1700 -Reciliadorsalis 710 - 3200 1700180 890 630 140 170 2400 800Nilaparratalugens 130 690 - - 4200 4600300 520 750 180 60 3300 1600 740 310Sogatellafiircifera 1700 31000 13000 8200310 210 3300 610 270 980 1600 6300 -Cyrrorhinus - 12000 6700600 540 3200 290 160 330 120 410Scirpophaga spp 50 410 620 - - 21000 12000150 180 220 270 30 1000Cnaphalocrocismedinalis 70 70 540 610 270 11000 6100200 140 100 6 60 250Chilo suppressalis 0 60 60 1900 300 200- - 30 20 0 6 3 160 -C auricilius - 0 00 - - 20 3 60 20 3 0Sesania inferens 0 - 80 60 3 80 10 51 -Cropping intensity _ -

Rice crops (noyr) 10 10 12 10 10 10 13Rice area (7c) 17 19 20 20 25 2320 85 85 60 70 40 85 85 90Fertilizer (kg Nha) 60 20 85 80 70 7020 0 0 0 30 30Insecticide (no of 40 60 40 300 04 04 0 03 300 13 13 30 50 40 33 33applicationscrop)

__________

IRPS No 123 January 1987 21

Light trapped Insects Drytnd __ 2(notrap-nght) Suctioned Insects (nom )

50 -)o 9 jO40- SOW ffurciWO Light trp

D-Vc suction

20 to 4

20

Egg porasitizahion (M) 75 60

300 SOOampReOftwciffr

45 30shy

15

-insects (notrop-night) 4 Nileporvata Iuen5

Egg parastization()

Light trapped insects (no trop-nght) Suchoned insects (no m 2 )

6 8 Cytorh~nuls hvidipennii 0

4 05 2

50 0 Insects (noltrop-night)40-ui Ipanh pe g

Moohase 1 Seasonal abundance measured by light trap

Rainfall(etod) and D-Vac suction sampler of whitebacked

to ~ hopper Nilaparvata lugens and lea ffolder60)-t Cnaphalocrocis mnedinalis andplthpe g20 L I II I I I parasites and mtrid predator Ortorhjntus livi-Apr May Jun Jul Aug Sep Oct Nov Dec JaIn Feb Mar dipennis set against cropping pattern daily1 9 80o 19 8 1rainfall and moon phase Tanauan Batangas

1980-81

the D- Vac Cyrtorhinus build up on the crop coterminous are collected in light traps during full moon (109)with WBPH buildup Cyrtorhinus disperses during the Because upland rice shares most of the lowland rice pestsrainy season RLF not readily collected in light traps was and those pest species are generally less impotant to uplandprevalent only during the rice crop rice than lowland rice there is a general belief that insect

Nephotettix virescens was more abundant than Nnigro- pests are less important onpicLus in the light trap collections and on the crop GLH upland than lowland rice Arcview of limited data on upland rice shows an averag of population peaked at the reproductive stage as did zigzag 10-21 yield loss from insect pests on a par with that ofleafhopper (ZLH) Crtorhinus tracked the populations of lowland uee determined by the same methodology (132) planthoppers and leafhoppers Both GLH species were (Table 4) McGuire and Crandall (144) made a similarcollected year-round in the light trap indicating immigra- estimate for Central America Cramer (51) howevertion from nearby irrigated rice areas Possibly because of the estimated 35 yieldlow populations loss in South America based on ainsect number did not correlate with similar pest complex - stem borers and seed bugs shymoon phase although reports indicate that more hoppers attacking irrigated rice in North America This estimate did

22 IRPS No 123 January 1987

Light trapped insects D~~nd (no trap-night )

St plht Suctined insects (nom 2

)

D-Voc suction 1520

1o

0 Lih trap 5 ~105 o5

00 Eggparositizotion()5040 NephOtgtha virneens

30 20

I010 A-Insects (notrop -night)

4 Nephatelh nigropccent i

2 o I vV^J - -- ^ VA____ f _

Light trapped insects (no trap-night) Suctioned insects (no rnE

10 -1208 ReCdO doRsol

6 15

4 10 052 iwxALAiamp 0

Light trapped insects (notrap-night) 2Suc oned nsects (nom )

8 CyrtorhmnusI6o nn]10 i0075

4 05

2 0 0

025

2 Seasonal abundance measured by light trap Roinfall (mmd) Moon phase and D-Vac suction sampler of green leafhoppersNephotettix virescens and N ngropicrus and I zigzag leafhopper Recilia dorsalis their egg 60

parasites and micid predator Cyrtorhinus livi-dipennis set against cropping pattern dailyrainfall and moon phase Tanauan Batangas1980-81

40

5 Lf Apr May Jun

1980 Jul Aug Sp Oct Nov I

-aMar

1981

not account for soil pests and was not based on field data in the absence of pests Carbofuran granules are phytotonicAkinsola (I1) cited mainly data from irrigated rice areas in to rice (239) The prevailing hypothesis is that carbofuranAfrica but estimated yield losses of upland rice between 15 prevents soil bacteria from consuming fertilizer The phytoshyand 30 from insects However losses in Thailand over a tonic effect is through greater availability of fertilizer to the4-yr period ranged from I to 13 averaging 5(106) plant Denitrifying bacteria are known to be more active inYield loss data however should be interpreted with aerobic than anaerobic soils therefore it may not be merecaution (184) The Philippine data were all derived from coincidence that the highest recorded yield losses (22-69)trials on farmers fields under farmers normal agronomic occurred in those trials with 12-30 kg basal Nha On themanagement practices except for Bukidnon and Capiz other hand contro with carbofuran granules may not be aswhere researchers used fertilizer Batangas farmers are good as a seed treatment against seed pests and perhapsatypical because ofhigh levels of fertilizer used The trials in yields would have been higher if a systemic seed treatmentBrazil were carried out at an experiment station but insecticide had been used conditions were similar to those of local farmers The 1979 and 1980 trials in Batangas had two full-

Carbofuran granules were used in every trial as a broad- protection treatments one using carbofuran granules andspectrum insecticide applied basally with the seed An the other using bendiocarb WP as a seed treatment Plotsobjection to using insecticides to measure yield losses isthat protected with bendiocarb yielded higher than those proshyinsecticidescandirectlyorindirectlystimulateplantgrowth tected with carbofuran granules The other trials in the

Table 4 Yield losses to insect pests and yield responses to insecticide in upland rice in the Philippines Brazil and Upper Volta

Yield (tha) Yield loss (5) Location Year Cultivar Protected

with Protected

without Check Total Vegetative Reproductive Ripening

Basal N (kgha) Reference

carbofuran carbofuran Protected No with carbofuran

carbofuran

Philippines Pili Camarines Sur Tanauan Batangas

Pangantucan Bukidnon

Dumarao t-apiz

BrazilGoias Minas Ge~ais

Upper Volta6

1975 1974 1974 1975 1975 1976 1977 1978

1979

1980

1979 1980 1979

1977 1978 1979

Bursiging puti Kinanda Dagge Kinanda Dagge Dagge Dagge Dagge UPL Ri5 Dagge UPL Ri5 Dagge UPL RiS UPL Ri5 UPL Ri5 UPL RiS

IAC47 IAC47 IAC47

14 33 11 36 35 31 30 27 37 26 40 30 40 42 46 35

19 21 09

--

-34 24 24 27 --

29 43 31 42 -

47 25

---

10 28 05 27 27 30 26 29 33 28 33 29 43 23 36 11

15 15 06

29 15 55 25 23

3 13 -7 11

-7 18

3 -7 45 22 69

24 28 35

--

21 -11 -20

4 -

-3

23 7

-2 -23 56

-

-

-__0

6 --

-

-23 14

5 0 8 2

22

-

--

0

-

--18 11

3 -10

4 10

7

--7 -

14 -

-1 6

10 -5

8 9 7

-

--

0 0 0 0 0 0 0 0 0 0 0

30 30 30

12 12 12

99 99

100 100 101 101 102 102 102 102 103 103 104 104 104

66 66 66

Farako-Ba

Mean

1977 1978

IRAT10 IRATIO

38 66

--

40 43

-5 35

21

-

10

--

--

--

- 242 243

Z

tJ

-w

24 IRPS No 123 January 1987

Philippines which had two treatments offering protection In the 9 Philippine trials where yield losses were measuredwith and without carbofuran granules relied on further by growth stage 10 of the yield loss (21) occurred at thecontrol with foliar spray and not on seed dressings The vegetative stage and 517 occurred at each of the reproshycarbofuran-protected plots had an 18yield los compared ductive and ripening stages Ibis result is similar to thewith only 10 with foliar sprayS This yield difference is pattern of yield loss in lowland rice measured by successiveprobably due to differences in protection against sown-seed treatments where the crop is unprotected at each growthpests (foliar spray do not control sown-seed pests) rather stage but proticted during the other growth stagesthan phytotonic effects became basal fertilizer wa used in However the insect complexes r sponsible for vegetativeonly 2 of the 10 comparisons stage yield losses in upland rice are entirely diftfrent than in

The recorded yield losses were not related to yield lowland rice The key vegetative stage pests in lowland ricepotential (Fig 3) One might expect higheryielding crops to arc aquatic whereas those in upland rice are sown--seedhave higher percentage of yield losses but this ws not tile (sown in soil) root and seedling pests This dift rence maycase One reason is that insect damage exacerbates plant explain why agricul torists have overlooked tile ilmportanccinjury from drought stress low yielding fields from d ought of upland rice insect pests Many reports of upland rice pestsstress would show disproportionate differences between in Asia ha Ve focused on major lowland species (46 108)insect-protected and -unprotected treatments Shoot and It isalso surprising that tei vegctative stage of upland rice root pests would be more responsible for losses from would record the highest yield losses becatuseseeral studiesdrought stress than sown-scd pests A second reason have shown that upland rice can readily recover from foliageconcerns seed pests which would have a greater impact on reinoval during that stage (117 152 165 188) Upland ricelow tillering low yielding varieties such as )aggc Low poscsses a hitgh obilitV to compensate for ca rly loss oftillering varieties nnot fill in the space created by removal foliage and will even be stinulated by foliage removal toof seeds oweer 1) R i 5 is high yielding because it tillersP1 produce a higher viel than plants ithout any foliageactively which can close the canopy to provide greater weed removed (63 192 231) [ie reason for this apparentcontrol discrepancy hovce is that yield loss at the vegetative

stage does not octUr frol leat arca loss but fron pestsYield loss () removing sown seed and feeding on devcloping tillers and roots -- damage that cannot be readily conpLtnsatCd for

70 - Measuring yield losses in upland rice cnvironrnents is further complicated by a report from Brazil that greater yield losses occurred in response to better insect control60 - =-09 as Insecticide seed treatments resuled in denser plant stands

(n=21) which in turn created a selectively more favorable environshyment for blast (69) This resuIt was parlticurla rlxexacerbated50 when cirbofuran granules were used in seed furrows Aside

ffrom a greater plant stand ca rbofuran iay have auigmented the N lecl and stimulated blast If so ca rbofuran should be40 - replaced with another insecticide or the seeding or fertilizerI rates decreased

Other methods to determine the impact of upland rice30 shy pests wvcrc to interview farners or to develop single speciesbull bull correlations of population levels and yield loss Feneira (66)2 NIM)st (75(1) Golas farmers reporteddid both in Brazil

20- insect losses in ufpland rice and 251 said those lossesS avcragcd 37(1 [his result is consistent with tie yield losses 0 0 measured by field trials in the same statc(Table4) For every

I0 4 of seedlings removC d by leaf-cutting ants a I 7yield loss is predicted Similarly Ferreira (66) found that ai

0 - average of 53 thrips per panicle before panicle initiation meant 3 times more unfillh d grains

-IO0 I 1 1 I I-101 0 2 3 4 5 6 7

Yield (tho) C)N [ROt IM FI IODS 3 Correlation between upland rice yield and yield lols caused by Upland rice insect pests may be controlled by culturalinsects measured in 21 trials in the Philippines Ivory Coast andBrazil genetic biological and chemical methods

IRPS No 123 January 1987 25

Cultural control dry season plowing will have little effect on them Plowing Crop husbandry to reduce upland insect pest populations when the soil is wet from the first monsoon rains will expose can be classified into practices effective in a single field and soil insects that tunnel close to the soil surface to predatory those effective only at the community level Cultural birds chickens dogs and even man (field crickets nole controls should come first in a pest management program crickets and white grubs a re delicacies in man tra ditional They have broad stable effects as pests have little pos- diets) sibility of overcoming them through selection of biotyps The greatest degree i1 inIscct control fron tillage comes (138) On the other hand some cultu ral controls work only from plowing soon after rice ha rvest when most soil insects with high labor or power inputs and connunity-wilt lie close to the soil surface anrd stern borers are in the rice methods require coordination of iany farniters stuble Whitegrubsare mature and tillage exposesthem to

Pantin time The luxury of being able to shift the time of predators Tirnelv control will prtec the crop following pla ntring upland rice is open onl hr la rniers in areas with a rice and lowr the populationIfor succeedinig years Plowingprolonged rainifall pa ttern lI copping intensity or underthe ice stulble helps it decompose killing sten borer rnechanied land preparatiot Gcnerally planlting as soon larvae inside as the rFiinv season begins will lower populatiois of 1110st [requent tillage destroys ait nests bit not termites insect pests Insect populationsarelo aftcra dlvseoll or Tillage and clean culture howes er rcliove food for winter fallow delayed plaiting lets them build tip on termites which then mtay attack a young rice crop Scarab alIternate hosts that gi vigorotsly with the first seasonial beetles also prefer to miposit in eceitly tilled fields Noshyratiris This first flush iof weds cros lItxuttriartlv with iceCult is dibbling faors the survival oftillace Clre Such rineralied N released during the dr season aid made stils i aels tnd insects thtt pttpate or lay eggs- in the soil

available to the plants with the rains It iliso igrtmitcally Zero tilla hio eerl conserves predators ofthe GIH (37) advanttgeous for the rice ciop to tap this trlLItlaI fertility nt it mcttoi lo lessen the impact o0 soil insects and grow vigorotsly to compete with wrus and to tolerate brotdcasting seed or planting in ftltrrows is prefc ired over pest infestation dihlirig in hills Crickets are large enotlgh to eat seedlings in

Larly plantngs terd to escape seedling maggolts Upland a hill atrd tillering normnally ca iriot fill in tlhe missing space rice in Bitangas is planted in Ma and ine itli the Iirst Oftcn dibbll rd is not well covered and iseasily found by rains Fairners are highly mtotiated to plant early tocscape foraging insets ioIdents and birds drlitght and typlotns ricar harct Arn Atrgtst planting Plan uit sii Incrcatsini sced density protects tile cropwould be evcely attacked by It riim orivzo At tti againrst seed and seelling pests especially ants particularly other Philippiie tipland sites Rea l Qteot antd (laelit if the crop will germinate and ermerge quickly M isainis Oriental seedlig mnaggots arnrige ric boe t se hr trftinll itI India rice intcrclopped witi cotton or farnes have a longer growing season and plnit 2 rin or pigeon pci had lower 6IIt and WBIlII poptrlations than mote after the ortset of the rainy season liaaiangas ftrmners rice alone (211) NfailC arid Urplald ir_ 2re coilltori in the plant ca rly becia use ofa shorter raiinfall period arnd the desi re Philippines he chatngC in pest status folii iniercropping is to lliarvet before the typhoor sea son peaks ltighly locatioll-specific (133) cmianarid the net elfc be either

Early plantings cornbind with early-iatiuring varictics nil bcnelficial ortdetriential (189) For example BIatangias provide higher crp tolerartce for white grubs Aestiatior farmers intercrip Itro in their ipland ricefields A liornshyterminates with the first rains but 5-6 moare ineeded for the nrii iotd-Ahis i (I iiiacius) which norlially feeds resting larvae to develop into beetles emerge and lay eggs on taro ill feed or miarbv rice plants Intercropping and for larvae to reach the danriigii g ird-instar stage probably would not afft0 seed pests A crluanion crop (134 In lataigas this normually iseiough tiite for harvest planted ith rice and haing gleater tolerarnce for root

It Brazil howecver delayed planting is a suggested dariage or being toxic nayact isi trapcrop to root-feeditig cultural control for the lesser cornstalk borer and insect pests Acial-fei i ig pests voultId be most affected by pests favored l drought (66 195) Cropping internsity in intrer ropping Shoot aphids leaf beetles sten borers and Brail is low and delayed plraing places the rice crop in the hopplers would be likely ca ntidates to note effcts On the niost stable rainfall period reducing likelihood old rought othlt halnd the corlipalior crop stch~ sitniaie Iiiv be more

In rilniy arca s rinmirsare roi atl to plant other crops benehited than rice from initecropping ( 137) before rice and althNgh it woultl be desirabh to avoid 11echi Weeuling luring the first month after clop infestations by planting rice cuatl lariners lack the labor cstablishent will force pcts such is anyvworris which and time to do so ol a low priority crp prefer grassy cetds ontor tle vorling rice Soil iampsecs will

77age Uplild rice Stils ae light textured and Call be mvnye to the rice crop Cleani citrltirc also lay lore territes plowed duiring the d rv seasor notl only to grin time for to attack I u cetincti inrice ceds i ay he le optinal planting with lie fitst Inilsoon riiains but also to desiccate solition 13) soil insects arid weeds such is ntsedge that foim rhizomes Irtlizr Itneatsing N Iciiliier oin hlwland rice has Htowever becta use most soil insects lie below die plow layer fAtwvored Irigher populains of pliithoppeiv Imlhoppers

26 IRPS No 123 January 1987

and leaffolde rs However because of the need to temper fertilizer use on upland rice o prevent blast fIrtilizer rates will be lower than is comonia r for lowlad rice causing relatixely less hopper and IaIfTfoide r population buildup However inupland rice areas without a history of blast such Is Batangas farmers apply high rates of N and conscqtrently the crop has high infcstation levels of WBPI]and RIF The fhrmrcrs leec insect buildup by splitting the fertili11r in0to two to three applicatiois bit do not applV fertilizer at the vegetati estage because this prodrces a tall crop which illeasily lodge during typhoons Greater crop fertility leads to greater insect survival (better nutrition) larger insects (to cat no re foliage) arid greater fec titd t (to lav nmre eggs) (128 145 171) Not all insect species respondequally hovr Increasing rates (if N l anrid K increased Dialraa s-acchralis and (iaeiurema flea beetle but reduced thrips (66) Zn rcdLuced deadheart density from the lesser cornIstalk borer

Fnldingi Although most Upland rice areas do nIot hawxcaccess to wa ter floodiig the fields is rRrrnriierdcd in areas of Brazil or root bug and white gib control ( 195)

Crop rotatioti Tillage before planting a crop after rc harvest will control stern boreis and other pests renain n the stubble arid will unearth soil pests Planting a non-gratriiaceous crop after rice is recommncrded for termite control (66) Crop rotation is less nttdcd in upland rice culture than in lowlands because atcrop-free period is assured even after harvest of crops following rice

t1ulchilt iryl plastic irli irig designed for weedcoitrol ii upland rice inJapan resulted in ntr highanri liy infestation of (ihilo sulytresvais (93 9-1 95) NIulched Upland rice grew more luxuriantlv and attracted more (Avipositing mothis than u lched rice Also the rice plant and stein borers imatoured 10rC quickly and a partialgeneratiori af vouing ista iNoverwintcred in the stubbleLotili--malitrt varieti Quickly maturing crops reduce the tmbCr of pCst generations that Calt build up Rapid crop establishment olan ca rl v-natu ring vaitvty if)pa 11icLa r will lessen white gib and stem borer damiage White grub larvae intle last iistar are highly destIruCtive so aii early-iiatri ng variety will escape serious root loss Stem borers build 1ip slowlyx in rice to ecormic abundant in a late-Maturing crop anrd caiuse whi tlieads Numibers of somespecies increase exponeitially with each succeeding genera-ti so a xarietyexhat atures I trowh earier thian another will hac less insect damage Early-naturing vaiicties however will not lessen seed pest daiage either ii the soil or oina standing crop

Strwhoi hAIJttam Inrscts tcd irig on thlie aerial portions of rice plats disperse from carly to late plantedfields AS iiiibCirs increase exponentially with eachinsect

gtscnrattm e eiglbrig f rmc~s1(uil upp wh rc generation pests build tIP where neighboring fihrTC e9r1sstagger their plantings II Batangas farmers plant uplandrice within I iro use 120-d s-arieties and plant at the start of the rainy season Ini Misarnis Oriental laricrs stagger their

plantings up to 3 mo apart and use a set of atieties that mature in 4-7 mo Insect pest danage particularly from stem borers is greater in Misarmis Oriental Also seedling maggots damage rice there because rainfall favors frequent planting of maic In Batangas rice escapes seedling maggots bCca usC of distinct drs season and synchron0us early planting with the onset of ains

Therefore combinations of cultural practices ea rlyplanting synchronous planing crop rotation and earlyshymaturing vaneties protect the rice crop against most insect pests

Ilmant resistance Beca tse upland rice has few specialized insect pests and many insect types attack the crop worldwide regional breeding and strong internatiornal cooperation are required

Sown-sced pests arc economically controlled by insectishycide seed treatment and should the relore receive lowxpriori ty inbreeding objectives

Root pests are normally cost ly to control with insecticides but finding resista t sonres mayvbe di fficult Tanaka (229) for exa -ple fi led to find varieties resistant to root aphids Selecting for large root bioniass perhaps should be the strategy for pests sLuch as Whlite grubs that renliove roots Fainter(174) howcvcr cites varietiesof crops tolerant ofor resistant t(i root pests Some sorghun varieties are high in cyanide so rice sorghun might incorporate broadshyspcctruri chemical resistance

Seedling vigor and drought tolerance are high prioritybreeding objectives so tupland rices should have large r root systernsand be able to tolerate ligherlevels of root loss Sonie root loss inolder plants is beneficial if it stimulates new root developnrut and therefore crihances intake of soil nutrients (53) The rice seedling inaggot rtiLotia spp may be

controlled thirough resistant aittics A hi rgc breeding effort on sorghniu has had fair success inideveloping arreties esrstait to Asoccata Rodani (251) A field trial comparing 10 upland ricc varieties showed differences aniong varieties va ring from II to 4(- damaged tilleis (215) Hovver resistarce to 1Iill-0r gtia species may rot cross ox-rti otier

Shiraki (210) reported that upland i-ice stens wcrc harder than lows-land rice stemsperhaps becaIusc (f a higher silicoricontent and thus were more resistant to yellow stern borer hc tortia Iow tillering (f uplaid iice xarneties prodtres a

higher percentage of infested tillers than would occor in higher tillering lowland iice under a siiilar egg density The corollary therefore is to select for high tilleriig upland rices

Stemii bor amcstheh main targes dobrecders inAfrica (571 204 )but nticl oifth e sc reen irig is dotie unde r lo wland co d t osrm s wih A h r~ o l pe i s n acorditions (11 96 218) As with lthrinrnwspecies mayhave to be dealt with independently however initial results show cross resistance between African and Asian species(92)

In Brazil screening Ior insect resistance focttses oil lou I pest problems Varieties tolerant of lasmolwalpus ligi-sellus and resistant to )iatraea saccharalis have been identified (70 153) Current work also emphasies 7iraca limativetntris and Thmialspis (=Iosis) flavouicta (66) These insect pests Ilt onlV citise IIigh yield losses tihey also ire difficult to control chemically ofr by other ilealls

ManV upland rice vaieties aic early maturing and are resistant to pests such as gall midge (I Xl ) Early mIturity is often a highly desirable trait to esuilpe pest buildup but it is not true resista nce bca usc ilthose va reties are plinted late they will be itsdamaged as ain early planted susc ible variety

P~erhaps tile cosiiiopolitall of all upland rice pestsimost are the sced pests (ripenling stagc) which pose a particular problem beta use scuds re ised IsIO for hitnit4ans Ayltoxic substatice introd tcld geticticllyilmay also be toxic to nal iniless it caln he placed old in thfe IdhuffSelectmi fo hairiness or lotng wns kill mIlct sthilrohleiiis dutilig threshing pa rtictfarl maitial threshinfl [hireshers ill complain of itcfimin atid itteascd (fust levels

Attempts itIRRI to fitid tcistatncc to the rice bug plorisa have ilotbeCnl prmitisilg lIIetIattonlids iorc

through the lenilna or palca hillhairs sedstVIor seeds wilh awns that deter bugs promote complaints from threshers Seed bugs liptocorisa and Stcnocoris enter the seed through the opening between tile lemna and palC but varieties with narrow openings are diifficult to mill

Breeding priorities should concentrate ott inotCI gruLtLIps that are most difficult to c nttrol by other meains Steil) horers appear to be the most widespread group that sigiiificantly dalag s rice aind is hard to manage by other Ilnca us

Biological control Very fe stutdic of mitatt~l eneties of rice pests in upland envirollnletits wee in tile wefouiid literature Therefore have not compiled a list of species Ve present a list of natural enemies iOtid in upland ricefields tuic latialln Batangas for some conitno iOldiar pest species ([itble 5)

Muich ofl thc lollowinle itfor ittiol cotlliCs rol Stutdies in the lowlatnds or roi crops otithcr thlin tice

Qu(ailtr o 11ll t11t( Nat trrtf lii hcoeits ight Ile used agaitist ile social insects tilt and termites Sollie predator specice live ill theirf nests n ilnic theirnllt ip-pearanIlCe aMRidbehavior IJtrouMdCing a mote aggressise ant species to displace pest ant codonies is a possibility (169) Patihogens as potential natural encnies have reccived little atteition (I13) Natural ncnuics asa group however hac not been toIl iporta it in Iegulating aintshown be and ternite numbers

The nocturnal behavior aid subteitmien habitat of lole crickets and field crickets protect them from many ntllral enemies There are only limited records of natural cnelics for these groups of pests some pathogens (87) but mainly

IRPS No 123 JattuIny I 271987

parasites - scelionids on eggsand sphecids on ilymphs and adults Sphecid wasps Larra carutmaria (SImitI hI L huzwuensis Rohwer and lsaltiuhta Willi ams speciali in mole crickets pa ralyzing them nd dragging the ninto thcir nests as food lot their voting ()tier sphCcid species Lins aurulenoa (Iabricius) lot maiac (Ashmead ) 1 lhshylssellaus (Smith) and Ml lal)oriosu (Snithf specialie in field clickets

White grUbsO the other hand hae Icti scriuns enuigh to have been extensively studied for cotttroi 1W nati al enemies( 112 140 141) Scoliid parasites C(ajsoMcispp have beetn intrtduced itto tilePhilippites for control of Ielxl ]udisiro la The rcsearchcts w Intttrdced themle during ill era claimed that the white gttionthreak as controlled (142) howevcr it isnormal for pest epidemics to subside and there was no diect evidence to indictte that tfe parasite was responsible A tmore promisitg titctllod might he to identify iso ltcttass prt~f tcartd apply hiactelaln fingal pathogens to tile soil Ihe lalattese beetle in North America has hcci coitrolled hv tite conmnleciall aailable hactcrial prcpa ration itacihlpu pav ofr milksdiscas (71) Spote prtyefott the soil itloted ilrtier reitiain actic fr Veas

Root aphid sardtnctly bugs readilslic tilib to predatots and parlsites once their tetndine ants are controlled The larvac and adtilts of a coccinellid beetle 1Scynms sp prey on llriadlouais in slash and burn upland rice Baits treated with insecticide may be used to control tending ants by killing their ourig (244)

Root-tkcditig blugs false wireoris wireorms and root wccils aic r ofsuch little ifilpoltatice that thenomttll role of liatural Cticniics hias tiot beei asscssed A possibility hoevrl tiglt he tit ItaIIitic niaCIMtodcs lintt1g11en Itese soil-itlhlfbitine mCtitodcs W5uhtfhave to be imiass Produiced

large miltibcr of niatltrll cnemics ircknown ft CltIoriS IC IpblCeii is thtt clttvortnlls ofonlie the crop soott alter land ptel)tritiOti a1tithe bcgit iniig of the raittv

Citsoln hfien tiatllral leniiiy populations aric lo Colltloil WOtuld iii ole mass prodtticing andirclcaslng key spccies

le egg stage is the fpirt (olthe Cedlitgt Ialot life cycle tI hich to f)ctis ftor tittlril elctit cottntrol hplraites o1 pleditors ttsit elleggs highly expoIsYCl I Sgg pailsites or predators stolld probahly e ittass andtbe produced released Littlh i toruialioll is a ailable ott what species to trV o ho to Iniss podLice thiCIi A etiloplhid wasp Icmihlaol sp effctiicly regushy

lates the leatf lcloalapolm2a spioanoiiitr heat andc rice antid should be collsctvcd

leaf beetles whCllose e am] eggs ate laid in the grutindhirO ould be motC difficutlt to control by niatulral enelV Mnll ipuhltiOn But species sUici IS c_gsthispid beetles ith

and larVle oilleascs ate vtilcratble to iparaites and pledators

Natural enemisc of tlirips otl rice are little known

28 IRPS No 123 January 1987

Table S Parasites recorded from upland rice insect pests in Tanauan Batangas Philippines 1977-84

Order Family Species Host stage

I lymttenoptera Braconidae Leaffolders Cnaphalocrocis Marasinia spy

Cardiochilesphilipplinensis Ashmead a Larva

Cotesia angustillasis(Gahan)Cotesia or cypris (Nixon)

Larva Larva

Cotesia or taeniaticornis(Nixon) Larva Cilonusinotakatae Munakata Chelonits SppAlacrocetrts philippinensis (Ashtmead)

Egg to larva Egg to larva Larva

Iclineumonidac Orgilts spp Isclutojppa hireator(Fabricius)

Larva Larva

Itoplectis naraigav(Ashmcad) Larva Tenelhtcha philippiensis (Ashmead) Larva Tenelchha stangli (Ashmead) a

irichomma cnaphalocrosis Uchida Larva Larva

Bethylidac Elasmidac

Goniozms or trianguli r Kieffer lasmtusalhopictus Crawford

Larva Larva

Diptera

Encyrtidac Chalcididac

Tachinidac

Elasmtus spp Copidosomopsisnacohiae (Fidy) Brach nteria evcarinata Gahan Biachyieria lasus (Walker) Ztgohothria ciliata (Wulp)

Larva Egg to larva Pupa Pupa Larva

Argyrophylx nigrotihialis(Blaranov) Larva

l-ymenoptera Braconidac Yellow stern borer Scirpopiaga incertulas

Braco chinensisSzeplegeti Larva CIthelotis spp Chelonus niunakatae Munakata Cotesi flaripes Cameron Stenoracon nicerilli (Bingham) a

Larva Egg to larva Larva Larva

Ichneumonidac Tropobracon schoenobii (Viereck) Amatroinorpiaaccepta metathoracica(Ashmiead)Frihorus sinicus (I lolmgren)

Larva Larva Larva

Ischnojoppaluteator (Fabricius) Larva Isotima nr dammeratiti Rohwer Larva Tenelucha philippinensis (Ashmead) a

Larva Tenehicha stangli (Ashmcad)Trichomma enaphalocrosis Uchida

Larva Lara

Chalcididac Eulophidac Trichogranmmatidae

Xanthopimnpla stemmator (Thunberg) Charops hrachYpterumn Gupta and Malteswary Brachtm eria sppTetrasticttus schoenohii Ferriere Trichorigranm chihunis Ishi

Larva to pupa Larva Pupa Egg Egg

Diptera

Scelionidac Pteromalidae Tachinidae

Tt ichograintnta aponiicuitt (Ashniead) Telenomits rowani (Gahan) a

Trichoinalopsis apattelctena (Crawford) Jeirhaea spp

Egg Fgg Egg pupa Larva

Z4gobothria ciliata (Wulp) Larva

Ilymenoptera lecneumonirlac Gol fringed borer (hilo auricilius

Trichomm enoptalocrosisUchida Larva

Ieioenoptera Braconidae

Aantholimlh steiniator (Thtinberg) a

Pink stet Ioret Sesamia inferens StoInacoil nicrilhi (Bingh am)

Loarva to pupa

larva

Diptera Ilyinenoptera

Armvtnyorms Mythimna separata Spodoptera mauritia acronyetoidesTachinidae Zvgohothriaatropiora( Robinea i -Desvoid y)Eulophidae It-phctrtiscitapadoe (Ashimead)

Larva Larva

(halcididae Braconidac

Brachpin ria spip (oesia spp

Larva larva

IHyntenoptera I)iptera

Rice skipper Ielop ida (halcididae Tachinidae

math ias aidgreen iortedci ciliarNIelan itis eda Brat lhincria sp iii matuqitatt Cameron Irgyrolhirlaxi trotlhiais(marumtv)a1

ismtne Larva Larva

Diptera Ilymenoptera

Tachnidae lelineutionidae

fuown se tilo per Mois frugalisIrgtrophrlax 1igrothiahiv 3 a rat ov) anthopimpa pmnctata (Iabricius)

Larva Larva to pupa

aMost dominant

IRPS No 123 January 1987 29

Normally predatory thrips prey on herbivorous thrips strepsiptera on both Spiders dominate tilenymphaladultNatural enemies of stem bugs --Scolinoplara and Tibraa predators Spittle bugs move to rice from pasture grasses - are in order of importance egg par~isites and predators and biocontrol would be successful only ifcarried out before fungal pathogens and nymphal adult parasites Egg tiley immigrate into ricefields parasites could either be introduced fron other areas or Biological control tactics for hemipterous seed bugs mass produced and released ould follow those outlined for stein bugs

Armyworms are normally lwld in hcck by the activities Coml)arison of environnewtns Natural enemies are of egg and larval parasites When these natural enemies fail perhaps faced with even greaterconstraints than insect pestsusually because of drought armyworns become epidemic to survive ind reproduce in upland environments TheyIt may be worthwhile therefore to release parasites during have adapted ani survived by having wide host ranges the the rainy months following a drought Virus diseases of ability to aestivate or hibernate over unfavorable seasonslarvae would beanotheravnue to exploreforarnyworms and the ability to disperse or by other mechanisms

Parasites have ben traditionallyconsidered for biolegical Comparing the most dominant and therefore the most control attempts against stein borers usually by rekasing adapted species of natural enemies collected in various exotic species( 166) This approach has not met with success environments in the Philippines we can understand how and new avenues should be explored The role of predators different rice environments are arid pathogens is little inderstood and deserves greater Anagqrms olptahiisa mymarid planthopper egg parasiteattention Egg parasitism rates are normally reasonably is cquallyadapted to all Philippine rice environments (Tablehigh but should be supplemented with effective egg 6) It is selective for planthoppers and even parasitizes eggspredators Orthopterans are potential egg predators of the n-iize planthopper Peregrinitsmaidis itt upland areas Metioche vittaticollis (Sta ) Anavipha nifiwnnis (Saus- Alavohis also parasitizes maize planthopper eggs but is sure) and Conocephahs hongipnni (de i-laan) feed on most prevalent on BPH in the irrigated wetlands (onashystem borer eggs etioche and Anaxil)ha specialize in eggs tocerus spp like A optalhilis on BPH and WBPI Iattacks with no hair such as Chilo spp The omnivorous (ono- the eggs of GLH in ell rie environments It is also found cephahsspecializes in eggs cov red with hairs it is a proven parasitizing the eggs ef the white rice licahopper (oimna egg predator of Scirlouphaga spp but has the discouraging spetra Oirosita naias a trichogrammatid egg parasitehabit of eating rice guin (65) attacks BIPH and WBPH in the wetlands and has no other

Grasshoppers have egg (scelionid) and nymphalad ult host A related species 0 aesopi specializes in G1-H in (nemestrinid tachinid and sarcophagid) v irasites Among wetland environments Dryinid strepsipt-an and pipunshypathogens protozoans have been recorded most frequently culid nymphal and adult parasites of planthoppers and Control of locust species by natural enemies could con- leafhoppers also specialize either in hopper species or centrate on their habitual breeding areas environments Among the dryinids Pseuhdgonatoipsflavi-

Leaffolders have rich complexes of natural enemies some feimur prefers the lowlands and BPH while P nuch has no adapted to upland rice (22 27) ranging from egg predators environmental prefirences but is adapted only to WBPH (gryllids coccinellids) to larval parasites (braconids ichncu- The main d ryinid species on GLH aptogonatop)usspp is monids) to larval prcdators (ants and carabids) to larval most dominant in rainfed lowlands and also attacks other pathogeiis (vira and fungal diseases) Efforts to augment leafhoppers within Amnrasca Cicadulinaand Bachaltuathese natural enemies should focus on the egg predators Strepsiptera are more dominant on plarithoppers than (rearing releasing)and viral or fungal pathogens (culturing lealhoppers The Elenchus species in the Philippines attack spraying) all species of planthoppers within the genera Sogawilla

Parasitization of the large conspicuous larvae of Sogatodels NilaparsataHarualia and Opiconsiva Elenshyskippers green horned caterpillars and other polyphagous cusrvasuntaisuiis adapted to the major rice environments Lepidoptera by tachinids and chalcids is normally low ELg but prefers BPH in rainfed arid WBPIH in wetland environshyparasites and predators are perhaps the key to their control ments Ilalictoph(utsmtnroeiand HVpectrus attack G LH

Gall midges are parasitized by pterormalids and platy- and COfimravcctra respectivelygasterids that aestivate within their larval host between Pipunculids are mainly parasitic oii lealhoppers Each seasons Shoot aphids are preyed upon by coccinellids species has a unique environmental preference Tomosshysyrphids and chrysopids varireIa oraetoraprefers Ne7 otetltix nigropictrs and

Planthoppers arid leafhoppers have egg parasites and Pilunwulhtsjavanensisattacks Dehoceplhahls spp which predators as well as nymphialad nlt parasitesand predators are more abundant in upland than irrigated lowland their rich complex of natural enemies includes fungal environments T suhvirescensattacks Nephotettix virescens pathogens The principal egg parasites are myrnarids and and prefers the lowlands Pilpuiulus nttillattls is an trichogramniatids Egg predators are norinally mirid plant upland rice parasite arid attacks only Nephotetix speciesbugs that suck out the yolk Nymphal ad nIt parasites are More than 17 recorded species of leaffolder parasites in dryinids on planthoppers pipunculids on lealhoppers and the Philippines attack species of C(naplwocrocis and

30 IRPS No 123 January 1987

Table 6 Dominant natural enemies of rice pests in upland rainfed lowland and irrigated lowland rice environments of the Philippines

Natural enemy

Host stage Species

Brown planthop-Parasite Egg

MymaridaeAnagrus optabilis(Perkins) Anagrusflaveolus Waterhouse

TrichogrammatidacOligosita naiasGirault

NymphAdult Dryinidac

Pseudogotatopuslaiifemttr Esaki et Hashimoto Elenchidae

Elenchus yasurnatsui Kifune and Hirashima

Environment

Upand Rainfed lowland Irrigated lowland

Ailaparata lugens

x x X x

x

x x

x X X Whitebacked planthoppers Sogatella furcifera and Sogatodes pusanusEgg

MymaridaeAnagrus optabilis(Perkins)

TrichogrammatidacOl~gositanaias Girault

NymphAdult Dryinidae

Pseudogonatopusnudus Perkins Elenchidae

Elenchus yasutatsui Kifune et Hirashima

Parasite Egg

MymaridaeGonatocerus spp

TrichogrammatidaeOligositaaesopi Girault

NymphAdult Pipunculidac

Tomosvaryella subviresccns Loew Tomosvaryella oryzaetora (Koizumi) Pipunclusmutillatus (de Meijere)

DryinidaeIlaplogonatopusspp

Egg Copidosomopsisnacoleiae (Eady)

Larva BraLonidae

Cardiochiles philipphnensisAshnead lchneumonidac

Trichiomnia cnaphalocrosisUchida Temelucha stangli (Ashmead)

Pupa Chalcididae

BrachymeriaexcarinataGahan Brachymerialasus (Walker) Brachymeria nr tachardiaegroup

Parasite Egg

Scelionidae Telenomus rowani (Gahan)

EulophidacTettasticusschoenobhi Ferriere

Larva Braconidae

Stenobraconnicevillei (Bingham) Iclhncurnonidac

Temelucha phiiippinensis(Ashinead)Temelucha stangli (Ashmead)

x x x

x x

X x X

x x Green leafhopper Nephotettix virescens

X x x

x x

X X X X

X

X Rice leaffolders CnaphalocrocisMarasmia

x X x

X

X X

x x x x x X x x x

Yellow stem borer Scirpophaga incertulas

X X x

x

x

x X XAinauromorphaaccepta metathoracica(Ashmead) x

continued on next page

IRPS No 123 January 1987 31

Table 6 cntinued

Natural enemy Environment

Host stage Species Upland Rainfed lowland Irrigated lowland

Larva-pupa Gold fringed borer Chilo attriciiuslchneumonidae

Xanthopimpla stemmator (Thunberg) x aPupa x

Pteromalidae Trichoinalopsis apanteloctena (Crawford) ax

Pink stem borerb Sesamnia inferensParasite

Larva Braconidae

Stenobracon nicevillei (Bingham) x b b Armyworms Mvthinna SpodopteraLarva

Tachinidac Zygobothriaarroptvora (Robineau-Desvoidy) x x

Braconidac Cotesia spp

X Skipper Pelopidas mathias and green horned caterpillar Melanitis leda ismeneF-gg

TrichogramrnatidacTrichogrannaspp x

Scelionidae Undetermined Scelionid

x Larva

Tachinidae Argyroph lax nitrotibialis (Baranov) X x

Pteromalidae Trichomnalopsis apanteloctena (Crawford)

X Predator

CoenagrionidaeAgriocnenis femina femina Brauer x xIschnura senegalensis tambur

Miridae CrtorhinsliidipennisReuter x x

Nabidac Stcnonabis tagalicus (Stal) x

VeliidaeAlicrovelia douglasi atrolineata (Bergroth) x

Coccinellidae Micraspis crocea (Mulsant) X x x

Vespidae Eumenes campaniformis (Fabricius) x

Formicidac Diaecamna spp xOdontoponera transversa (F Smith) x

Carabidae Chaenius spp OphioneaW- aJasciataSchmidt-Goebel

x

Ophionea ishii ishii Habu x x X

Gryllidav Anaxipta longipennis (Saussur-) x xMetioche ittaticollis (Sial) x x x

Tettigoniidae Conocephalus longipennis (de Haan) x x x

aNo rearing made on rainfed lowland bNo rearing made on rainfed lowland or Irrigated lowland

MrasmiaAmong the Laval parasites the braconid Cardio- lowlands and parasitizes oriental maize borer OsIriniachilesphilijpphensis is most adapted to an upland environ- furnacalis (Guenee) larvae The leaffolder pupal parasitesment It also parasitizes Hydelepta indicata a common not only occur equally in all environments but also haveleaffolder of legumes The ichneumonids Trichomma cna- wvide host ranges- parasitizing species of OsIrinia Marucaphalocrosis and Temehcha stangli occur more in the Hydelepta and Honiona - in maize and legumeslowlands T stangli is most adapted to irrigated lowlands The principal egg parasite of yeltow stem borer (YSB)and also parasitizes Chilo and Scirpophaga stem borer Telenomus rowani is widely adapted to all environmentslarvae T cnaphalocrosis is most prevalent in rainfed and has an unusual alternate host -Tabanus eggs (25)

32 IRPS No 123 January 1987

Tetrastictu schoenobii specializes only in YSB and occurs mainly in irrigated lowlands -- further proof that YSB is most adapted to the wetter environments

Larval parasites of YSB are highly environment-specific The braconid Stenohracon nicevillei occurs only in upland rice but also attacks the pink stem borer (PSB) Tieinchwa phiiiplPwnsis an ichneumonid occurs in upland rice and also parasitizes the larvae of (7hiu stqpresaisinatro-morpha accepta melaihoracica like 7 stangi is mnost adapted to the irrigated lowlands

The dark-headed stem borer(l)HSB) (ih pochit3sus was not collected in all rice environments and the gold fringed borer Chilo auricilius nd PSB were collected only in upland rice Xaantiholiplastenrnaloran ichneumonid larval parasite appears to be widely adapted whereas TrihomaslopisapanitI)cteiiaa pteromalid pupal parasite occurs mostly in wetlands and also parasitizes larvae of Chilo suppressais and Pelhuidas mathias

Armyworms Myit hinin and Spodujipera have two main larval parasites - a tachinid Zrgohothriaatropivora in rainfed environments and a braconid Cotesia spp in irrigated lowlands

The butterfly pests Pelopidas and 4elanitishave tricho-grammatid egg parasites found only in irrigated lowland environments

Two larval parasites show environmental preferences --Argrroph av nigrotihialis a tachinid in rainfed habitats and Trichonialopsisapantelkcnela in irrigated ricefields A limgrotibialisalsoattacks the sweet potato hornwormAgrius convolvtdi and T apanteloctena parasitizes DH S1B

Predators as a group show more distinct environmental preferences The aquatic damselflies Coenagrionidae are most abundant in the lowlands Cvriorhinus is also most adapted to the lowlands The nabid Stenonabis tagalicus occurs only in the uplands and isalso prevalent on legumes The coccinellid lady beetles wasps and ants are more prevalent in upland areas AlicrasIpis cit-ea larvae prey ona variety of aphids attacking legumes and maize Etunienes campanonia vespid mud wasp makes nests in trees and is therefore most adapted to the more botanically diverse uplands Soil-dwelling ants cannot tolerate flooding

Upland environments are habitats to arboreal carabid beetles Three species of Chlaeniusbeetles prey on leaffolder larvae and are perhaps more important than parasites (27)

The gryllids and tettigoniids are widely adapted egg predators not only on rice but also on maize and legumes They feed on eggs of most insects that are laid on leaves

The spidercommunity of upland rice environments isrich in species At one site (Batangas Philippines) 31 species have been recorded (23) Of the 176 spider species recorded in Philippine ricefields about one-half (82) occur in upland rice The spiderspecies of upland rice environments overlap more with those of rainfed lowland than with those of irrigated lowland rice (24) The same study showed that of the three environments irrigated rice has (ie greatest spider species diversity foiiowed by rainfed lowland and upland

However of the 10 most prevalent ricefield spiders 9 were abundant in upland rice showing wide adaptation All three spider guilds -- orb-web space-web and huting spiders shywere prevalent in upland ricefields There were however differences in environmental prelerences for some spider species

Of the orb-wcavers Tetragniathamanlihulatawas partishycila rlv abundant 7 japonica was low and Ltucatge dehcorata was absent in upland rice

As in other environments Aiypeiwnaifirtosana was the most dominant space-web spider however its relative numbers were lowerthan for lowland sites Among hunting spiders Ox|opes javanus and Lcosa leatcost fgla were more abundant and Lvcosa lpseudoanmnlata was less abundant in upland compared with lowland rice envishyroniments

Extensive rearing of rice insect pests in upland rainfed lowland and irrigated sites over an 8-yr period in the Philippines provided insight into the effectiveness of parasites as natural enemies (Table 7 8) Most of toe data are from large sample sizes taken over at least one crop which overcomes the pitfalls expressed by Van Driesche (238)

VSB egg parasitization was surprisingly similar across all environments Larval parasitization was highly dynamic among sites and even years within the same site but showed somewhat lowerlevels in uplandsthan in irrigated wetlands This would have been predicted for egg parasites as well since YSB is adapted to the lowlands and so must be its parasites This isevidence that Telenomus rowani is as good at dispersing as its main hosts Trowani clings onto the anal tufts of female YSB and parasitizes eggs as they are laid This phoresy may explain its wide adaptation The more specialized lowland YSB larval parasites therefore appear more effective than their upland counterparts with more alternate hosts

Low collection levels of PSB larvae especially in the wetlands make comparisons between environments difficult Low levels of larval parasifization were recorded in both rainfed environments Except in the Cagayan rainfed wetland site in 1980 low levels ef larval parasitization were evident at irrigated and rainfed sitesalike No differences by environment were evident Higher levels of parasitization occurred in Sarawak during a 1967 outbreak also from tachinids Rothschild (197) indicated that higher parasishytization occurred in upland rice areas

Skipper and green horned caterpillar larval parasitization rates were generally low and showed no environmental effect htighest activity occurred in Cagayan in 1980 (48 parasitization) Although larval parasitization levels of RLF were variable year to year activity appeared greater in the irrigated lowlands than in the uplands Again the highest levels were recorded in Cagayan in 1980 Cagayan has extensive grasslands and fallow around the rice areas and a relatively long rainfall period Rainfall in 1930 was good for crop growth and apparently for increase of larval parasites

IRPS No 123 January 1987 33

Table 7 Parasitization rates of rice planthoppers and green leafhopper in upland rainfed lowland and irrigated lowland environments Philipshypines 1976-840

Environment and site Year Egg parasitization () Nymphaladult parasitization ()

BPH W13P GLH BPII WBPH GLIH

UplandBatangas

Rainfed lowland

1976 1979 1980 1982

15 17 12

2(47)8(39)

4(51)

9(1800)14 (212)

12(304)

8 (101)6 (127)

9(116)

Iloilo

Cagayan

1976 1977 1978 1981 1980 1981 1982

0 (69)

11

44 48 40

-

-

46 55 44

0 (471)

11

52 56 49

0 (200)9 (214)

10 (1259)8(235)6 (900) 3(800)

-7 (462)

11 (270)3 (313)5 (900) 1 (800)

4 (600)10 (725)10 (362)12(870)5 (900) 3 (800)

Pangasinan

Irrigatedlowland

1984 1976 1978 1979 1982

13 (436) 13(379) 14 (457)

6(14)7 (197)

17(198) 11 (486) 18 (275)

3 (81)10 (274)3 (116)

16(871) 19(88)

28(211)9 (764)5 (647)7 (1204) 8(305)

Laguna 1977 11 7 21 1978 31 1979 14 4 4 28 1980 7 8 20 1981 1983

56 7(278)

26 16 (1650)

16 27(1921)

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R I Apostol unpublished 1lo1lo and Pangasinan 136 Cagayan BCanapi unpublished Liuna 175 I C Pantua unpublished 118 234 43 Carino and Shepard unpublished BPH = brown planthopper Nila parvara htgens W131Pi =whitebacked planthopper Sogatelia furcifera GLH = green leafhopper Nephotettrix virescens Figiires in parentheses are sample sizes

Tauher et al (230) reported that internal external and genetic factors influence the seasonal activity of parasitoids

Hopper egg parasites but not nymphaladult parasites were also parlicularly abundant at the Cagayan site possibly the egg parasites have more alternative hopper prey in the weedy areas than nymphaladult paisites

The seasonal dynamics of hopper egg parasites can be seen for upland rice in Batangas(Fig I 2) and for rainfed lowland lice in Pangasinan( 135)and Cagayan( 105) Levels of BPHWBPI-L and GLH egg parasitization were corn-parable in upland and irrigated lowland environments Except for some high rates of paraMsitization early in the season the parasitization rate tended to be steady in all envilonments However hopper nymphaladult parasi-tizution levels were more similar in the rainfed environments and lower than in irrigated rice

Overall parasites do not appear highly effective by themselves against upland or even lowland pests Among natural enemies parasites have attracted the most attention because they can be more readily assesscd

Predators and pathogens lend themselves better to management practices Predators the most important group of natural enemies in riccfields are difficult to quantify On the other hand pathogens appear less im-portant but can be readily cultured and disseminated

The natural enemy commulnity of upland rice is rich in species and differs significantly from that of lowland rice

These beneficial organisms must be conserved by applying insecticides judiciously particularly sprays Seed or soil placement of chemicals minimizes exposure to natural enemies The strategy to derive the greatest benefit from natural enemies isto allow the greatest number of beneficial species to thrive Then perhaps at least one species will be effective against each pest at any tine overcoming the variable abundance of each species within and between years

Programs to introduce exotic species or mass produce indigenous ones are ambitious and expensive because they require trained people on a sustained not ad hoc basis

Chemical control Insecticides offer rapid and efficient control of upland insect pests (190) but-should be used only after other control measures have been considered Insecticides are rarely used on upland rice because of cost (32) Upland rice yields are normally too low to justify the expenditure Also most upland rice is grown as a cash crop Spraying upland rice is more difficult than lowland rice because water is less accessible Government subsidy programs to provide pestishycides to tribal peoples for upland rice production have created problems of toxicity to humans because isolated people have had no experience to be able to handle pesticides safely

However ins-eticide use has beenjustified when a crop is

0Z

Table 8 Parasitization rates of lepidopterous pests in upland rainfed lowland and irrigated lowland environments Philippines 1976-84a

Parasitization (5)

Environment and site Year YSB GFB PSB Earcutting Swarming Skipper Greenborned RLF Brown larva larva caterpillar caterpillar caterpillar caterpillar larva semilooperEgg Larva larva larva larva larva larva

UplandBatangas 1977 - 0 (57) - 0 (63) - 0 (71) - 0 (106) 11(142) 2 (101)1978 43 (4697) 8(451) 5 (38) 3 (40) 6 (61) 2(58) 15 (57) 13 (129) 13 (1286) 4 (30)1979 59(4135) 12(320) 13 (11) 5(28) 9(45) 6(46) 7(23) 8(160) 9(1152) 8(26)

Rainfed lowland Iloilo 1976 - 0 (10) 0(10) 0 (7) 0 (19) 2 13) 15 (180) 5 (76)1978 0 (54) 0 (3) 0 (17) 9 (64) 4 (78) 3 (73) 0 (121) 0 (191)1979 64 (2350) 16 (84) 6 (17) 0(6) 18 (68) 25 (104)

1981 56 (1322) 18(304) 0(5) 12(51) 18(138)Pangasinan 1976 60 (7 7 )b 62(82) 0(111) 5 (39)

1978 58 (53)b 5(51) Cagayan 1979 59(713) 8 (101) 6 (34) 0(7)

1980 0(11) 4 (28) 15 (286)

62 ( 48(177) 61(140)1981 0(5) 12 (2) 13(45) 14 (92)

Irrigatedlowland Laguna 1974 62 8

1974-75 50 41 1975 43 72 1978 96 (623) 21(24) 33 (60)1979 83(2017) 18(110) 8(62) 11 (89) 22(400)1980 78 (1625) 20(75) 0 (16) 4 (72)1983 60

9 (80) 7 (270) 30

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R F Apostol unpublished Iloilo and Pangasnan 136 Cagayan B Canapi unpublished Laguna 175 P CPantua unpublished 118 234 143 Carino and Shepard unpublished YSB = yellow stem borer ScirpophagaincertulasGFB = gold fringed borer Chilo auricilius PSB = pink stem borerSesamia inferens earcutting caterpillar Alythimna separata swarming caterpillar Spcdoptera mauritia acronyctoides skipper Pelopidas mathias reen horned caterpillar Melanitls ledaismene RLF = rice leaffoldersCnaphalocrocismedinalis and Marasmiaspp brown semilooper Mocisfrugalis Figures in parentheses are sample sizes Egg masses

IRPS No 123 January 1987 35

Table 9 Effect of seeding rate and insecticide on plant density and Yield losses from insect pests in upland rice often areyield of Dagge upland rice Batangas Philippines 1976a comparable to those of lowland rainfed or irrigated rice

Seeding rate Insecticide Dosage Plant density Yield (0-Y30o) but usually from different pests The principal (kg aiha) seed treatment (kg aiha) (noIra-row) (tha) groups are soil pests feeding on sown seed and rootsat 14 DE followed by seedling pests These groups have largely gone

50 None - 35 a 20 a unnoticed leading researchers to conclude that stem borers50 Carbofuran F 10 38 ab 23 a and seed bugs are the most important pests (17) Yield loss 10 beCabofuranF100 None - 4551 c 23 a trials that pinpoint major pests need to includc seed

100 Dieldrin WP 10 52 cd 22 a treatments Granules applied in the furrow may not give150 None - 59 d 22 a adequate protection from many seed pests

aAv of 8 fields DE = days after crop emergence In a column means The floristic and pedological diversity of upland rice followed by a common letter are not significantly different at the environments is matched by the diversity of arthropods5level by D)MRT F = flowable Wi = wettable powder both pests and their natural enemies Irrigation has homoshy

genized lowland rice environments Flooded soils tend tothreatened by epidemics of locusts or armyworms New have the same properties and to eliminate soil pests The technology stressing efficiency has minimized insecticide spatial and temporal dominance of cultivated rice has cost benefited specialist pests

Seed treatment is normally inexpensive and can be Upland rice has no specialist arthropod species The economicallyjustified in -nanycases(66) Treating seed with monophagous pests on upland rice prefer the lowlands systemic insecticide (216) but not chlorinated hydrocarbons where rice evolved Rice in the uplands exists only because (247) is most effective in controlling seedling maggots of man and even though lowland rice predates upland riceFoliar sprays have to be repeated to obtain control and specialist species evolved Wheat maize and sorghum do often are washed off by frequent early season rains not have monophagous pests leading us to conclude that Granular insecticides are effective but required dosages are there is no evolutionary advantage for inssects to have too expensive to justify except under heavy infestation narrow host ranges in upland graminaceae(247) When directed against first-instar larvae banding Pests have adapted to the uplands not only by having granules in seed furrows (177) can be effective against white wider plant host ranges but also by having long life cyclesgrubs at low dosages (025 kg aii ha) (134) Seed treatments undergoing dormancy or being dispersive Each pest has a are not promising for white grub control (248) unique life history strategy suited to highly unpredictable

Seed treatment is also effective ainst ants and may be environments In agivenyear someinsect specics an better cheaper than increasing the seeding rate In Batangas exploit upland rice as a food source but no pest is a senious treating 50 kg seed ha was as effective as increasing the one every year on upland rice seeding rate to 100 kgha and using untreated seeds (Table The low yielding potential of upland rice means less 9) In Brazil seed treatments control termites and other soil quantity and lower quality of food for pests to exploit but insects including the lessercornstalk borer(66) Domiciano also less incentive for farmers to undertake control (58) however found seed treatments inconsistent in their The most effective control would be early and synchroshyeffect against high populations of lesser corn stalk borer nous planting of early-maturing upland rice varieties to

Baiting can be an inexpensive way of controlling ants prevent insect pest buildup Breeding for insect resistance seedling maggot flies mole crickets and field crickets Baits should initially focus on stem borers because they are verymade of locally available material and impregnated with difficult to control by other means Biocontrol efforts insecticide can be sparingly distributed taking adantage of should first stress conservation of natural enemies preshythe pests ability to disperse and encounter bait sites dators and pathogens rather than parasites Efficient

The new low-volume hand-held controlled-droplet chemical control includes seed treatment or baiting for seed sprayers may offer an advantage over high-volume knap- seedling and some root pests and using controlled dropletsack sprayes for controlling foliar pests Upland rice sprayers for spot treatments against foliar pests and seed canopies are more open than those in lowland rice and bugs when economic thresholds are reached better droplet penetration should result Controlled droplet applicators would be ideal for seedbugs and defoliators

CONCLUSION

Upland rice is attacked by a wider array of insect pests than is lowland rice mainly beca use of the addition of soil pests Generally population levels of lowland rice pests are lower in upland than lowland rice leading many people to conclude that insect pests are not important on upland rice

36 IRPS No 123 January 1987

REFERENCES CITED

1Abraham C C and K S R Mony 1978 Occurrence ofLeptocorisa acuta Fabr (Coreidae Hemiptera) as a pest of nutmeg trees J Bombay Nat ttistSoc 74553

2 Agyen-Sampong M 1975 HlierogrlphusdiaganensisKrauss (Orthop-tera Acrididae) a new pest of rice in Northern Ghana Ghana J Agric Sci 8249-253

3 Agyen-Samnpong M 1976 Varietal resistance and other inle-ratcd pest control components as factors of rice varietal improvement in West Africa Prcsented at the Second varietal improvement serinar 14-18 Sep 1976 Bouak Ivory Coast West Africa Rice Development Association Monrovia Liberia 18 p

4 Agyen-Sampong M 1977 Outbreak of Ihccrorrdnisorae in rice seedlings near Rokupr Sierra Leone lit Rice Res Newsl 2(6)16-17

5 Agyen-Sampong M 1980 Dirty panicles and rice yield reduction caused by hugs IntRice Res Newsl 5(l)11-12

6 Agyen-Sampong M 1982 The major pest problems of irrigated upland and mangrove swanip rice ecosystems ir the humid tropical Guinea savanna and Salel climate zones Pages 318-328 in Pro-ceedings training course cn concepts techniques and application of irtegrated pest management in rice inWest Africa 10-20 Jan 1982 Fendall Liberia West Africa Rice l)evelopioent Association Monrovia Liberia 505 p

7 Akibo-Betts V I and S A Raymundo 1978 Aphids as rice pests in Sierra Leone lit Rice Res Newsl 3(6)15-16

8 Akinsola F A 1975 Present status of different rice stemborers in parts of Nigeria iicL Entomo Newsl 328

9 Akinsola FA 1979 Ihe biology and ecology of rice stemborers in Nigeria Ph 1) thesis University ol lbadan Nigeria 217 p

10 Akinsola E A 1979 Guidelines of integrated management of insect pests of upland rice in West Africa Presented at the Seminar on integrated management of rice diseases and insect pests at Bobo-Dioulasso Upper Volta 17-22 Sep 1979 West Africa Rice Develop-ment Association Monrovia Liberia

II Akinsola IA 1984 Insect pests of upland rice in Africa Pages 301-305 in An overview of upland rice research Proceedings of the 1982 lIouak Iwsrv Coast upland rice workshop International Rice Research Institute 13O Box 933 Manila Philippines

12 Alam M S V TJohn and Kaung Zan 1985 Insect pests and diseases of rice in Africa Pages 67-82 in Rice improvement in Eastern Central and Southern Africa proceedings of the international rice workshop at Iusaka Zambia 9-19 April 1984

13 Alticri M A A van Schoonhovcn and J D Doll 1977 The ecological role of weeds in insect pest management systems a review illustiated with bean (Phaseohsvulgaris L)cropping systems PANS (Pest Artie News Summ) 23195-205

14 Anmante F 1967 The leaf-cutting ant Atia capiguaraa pest of pastures [in Portuguese] Biol6gico (Sao Paulo) 33113-120 (Rev Appl Entomol A56207)

15Andrade A C and D Puzzi 1951 Preliminary results of experi-merits on the control of Sculptcoris iastantus on sugarcane [in Portuguese] Bioulgico (Sao Paulo) 1744-49 (Rev Appl Entomol A41421422)

16 Appert J 1970 Maliarplia separaelha (rice stem borer) new observations of entomological problems of rice in Madagascar [in French] Agron Trop (Paris) 25329-334

17 Ariadeau Mand Z Harahap 1986 Relevant upland rice breeding objectives Pages 189-197 in Progress in upland rice research Proceedings of the 1985 Jakarta conference International Rice Research Institute 1O Box 933 Manila Philippines

18Asar S and Kisimoto 1975 Seasonal occurrence of the smaller brown planthopper in upland rice [in Japanese] Proc Kanto-Tosan Plant Prot Soc 2228

19 Bakker W 1974 Characterization and ecological aspects of rice yellow mottle virus in Kenya Meded Landbouwhogesh Wage-ningen 152 p

20 Balut F F 1970 White grubs attacking rice Oryza ativa L [in Portuguese] Biol6gico (So Paulo) 36321-322

21 Banerjee S N and L M Pramanik 1967 The lepidopterous stalk borers of rice and their life cycles in the tropics pages 103-124 in The major insect pests ofthe rice plant Proceedings ofa symposium at the International Rice Research Institute September 1964 The Johns Hopkins Press Baltimore Maryland USA

22 Barrion A T and J A Litsinger 1980 Ants a natural enemy of cliaiarac in dryland rice IntRice Res Newsl

5(4)22-23 23 Barrion A T and J A litsinger 1981 The spider fauna of

Philippine rice agroecosystem 1Dryland Philipp Entomnol 5139-166

24 Barrion A T and 1 A Iitsinger 1984 The spider fauna of Philippine rice agroccosystcms II Welland Philipp Eiitomol 611-37

25 Barrion A T1 and J A Litsinger 1984 lihanus (Diptera Tabanidae) eggs an alternative host of rice stei borer (SI3) egg parasite Telenoonis digns (Iliymenoptcra Scelionidae) lit Rice Res Newvsl 9(6)19

26 Barrion A T and J A Litsinger 1985 Identilication of rice leaffolders by wing markings Int Rice Res Newsl 10( 1)24

27 Barrion A T and J A Litsinger 1985 Chctiosspp (Coleoptera Carabidae) a leaffolder ILF)predator lit Rice Res Newsl 10(1)21

28 Bertels A 1970 Insect pests of rice and theircontrol [in Portuguese] Pelotas Brazil Instituto de Expcrimentacao Agropccuararias do Sul Circular 43 24 p

29 Bonzi S M 1982 Ctilo (flisilineus J de Joannis (Lepidoptera Pyralidae) stem borer of irrigated cereals in Upper Volta LioFrench] Agron Trop Riu Rizic Cult Vivrieres Trop 37207-209

30 Bourke T V T L Fenner J N LStihick G Baker EHassan D F OSullivan and C S li 1977 Insect pest survey for the year ending 30th June 1969 Papta New Guinea )epartment of Agrishyculture Stock and Fisheries Port Moresby 57 p

31 Bowden 1 1973 Migration of pests in the tropics Meded Rijksfac Landbouwwet Gent 38785-796

32 Breniere J 1973 The main rice parasites in West Africa and their control Pages 27-42 in Proceedings seminar on plant protection for the rice crop West Africa Rice Development Association Monrovia Liberia

33 Breniere J 1976 The principal insect pests of rice in West Africa and their control West Africa Rice Development Association Monrovia Liberia 52 p

34 Brown ES 1962 The African arinyworn Spoiolptera eXI)Ina (Walker) (Lepidoptera Noctuidae) a review of the literature Cornshymonwealth Institute of Entomology London UK57 p

35 Brown F B 1968 Upland rice in Latin America FAO-International Rice Commission 12th Session of the Working Party on Rice Production and Protection 2-14 Sept Peradeniya Sri lanka Food and Agriculture Organization Rome Italy 6 p

36 Buyckx F J 1 1962 Disease and insect pests of crops inCongo Rwanda and Burundi [iil French] Publications dc LInstilut National pour lestude Agrononiquedu Congo M Weissenbruck S A Brussels Belgium 708 p

37 Cariiio F 0 1981 Role of natural enentics inpopulation suppression and pest management of green rice lealhoppers Ph D thesis University ofthe Philippines at Los Bafitos PhilippinesLaguna 136 -

38 Carpente A J 1983 The development of rice cultivation and extension of vegetable productin in Zanzibar Final Report Vol II1 October 1983 Food and Agricultme Organization Ronte Italy

39 Carroll C P and S1 Risch 1983 Tropical annual cropping systems ant ecology Environ Manage 751-57

4(0 Cates R G 198 1 Host plant predictability and the feeding patterns of monophagous oligophagous and polyphagous insect herbivores Oecologia (Berl) 48319-326

IRPS No 123 January 1987 37

41 Cendcafia S M and F B Caora 1967 Insect pests of rice in the 62 Ene 1 C 1981 I)iiorldhol)ierusihilis Slater (licteroplera Philippines Pages 591-616 in The najor insect pests ofthe rice plant lygacidue Illissinac) a new rice pest front Nigeria lit Rice Res Proceedings of a symposium at the International Rice Research Newsl N3)15 Institute September 1964 The Johns Hopkins Press Baltimore 63 Fugenio T S 1965 rhe effect oi e-vCgetation on rice production Maryland USA 729 p and the possibility of overcoming it by leaf pruning MS thesis

42 Centro Ihternacional de Agricklittira Tropical 1984 Upland rice in University ol the Philippines at Los Baijos ILaguna Ihilippins 69 p Iatin Anerica Pages 93-119 hi An os ervies of uplaind rice research 04 Farrow R A 1975 The Alrican nmigratory locust ii is main Proceedings of the 1982 Boinakamp Ivory Coast upland rice workshop outbreak area of the middle Niger qlinlitatise studies of solitary International Rice Research Institute P0 Blox 933 Manila populations in relation to environmental factors Locusla No II Philippines 198 p

43 Chandra G 1979 Taxonomy ad hn riononlics ofthe imect parasites of 65 Fennal R G 1943 Food crop investigation Windward and rice Icalloppers and planthoppers in the Pihilippines and their Ieeward Islands First report Report onl work dunring the periodiiiportairce iii natural biol)gical cointrol Philipp Fntoltol 4119-139 April 1942-April 1943 Imperial College of Tropical Agriculture

44 Chang T T 1976 The originii colut ion enlIivation dissemination Trinidad 16 p (Rev of Appl Fintornol A3222-25) and diersificition of Asian and African rices -uphytica 25425-44 1 66 Ferreira F 1983 Integrated pest ritaageiret of upland rice [ill

45 Chatierjee P B 1979 Rice leaffolder attacks in India lit Rice Res Portuguese English summary] Pages 323-341 in Upland riceculttre Newsl 4(3)21 factors affecting production E Ferreira T Yamada and F-

46 Cheng Cbien-pan 19(4 Upland rice in Taiwan International Rice Malmolta eds Institutoda Potassae Foslato Institnto Internacional Cnmmission Wiorking Party oit Rice Production ard Protection I()th da Potassa Piracicaha Sao Paulo lraid Meeting 3-10 March 1964 Manila Philippines Food and Agricul- 67 Ferreira E arid F dl Silva Martins 1979 Occurrence of ture Organiaion Rome Italy 5 p Sogatohs orvziola (Muir) inupland rice in Goias Ilrail lit Rice

47 Chcrrett 1 M and 1) 1Peregrine 1976 A review of the status of Rcs Ncwssl 4(4)18 leaf-cutting ants and their control Ann Appl Biol 84124-128 68 Ferreira E and 1F da Silva Martiris 1984 Insect pestof rice ill

48 Chipeta F M 1974 Pests of rice in Malawi Pages 90-95 in Brazil and their control Jill Portugucsej FMIlRAtPA-CNPAF Proceedings Sith eastern Africa cereals research conference 10-15 Goiania Brmiil 67 p March 1974 DRB Mialda ed Ministry of Agriculture lilongwe 69 Ferreira FI F ca Silha Miartins S S Nero and 1 It P de Malawi Carvallio 1982 M tltiplc regression analysis todeterimine the effect of

49 Chu Yau-i 1979 The d atiaging period of the rice seedling fly insect populatiois ott upland rice yield [inIPortuguese Pesqui (Atlterigotiavxigu Stcin) (NItiscide Diptera) oil cori) at Fast Java Agropecu Bras 17671-675 Indonesia Plant Prot Bill 21403413 70 Ferreira F 1F da Silva Martins and F 1PZimmermann 1979

50 Conklin H C 1957 IHaninoo agriculture a report on aln integral Rice varietal resistance to tle lesser corn stalk borer [in Port uguese] systent of shi fti rig cult ivationi in the Pltilippincs Food and Agriculture Pesqui Agropccu Biras 14317-321 Organization Rome Italy 209 p 71 Fleming W F 1968 Biological control of lie hapanese beetle U S

51 Cramer H II 1967 Plant proection and world crop production Dep Agric RLs Sers [ech Bull 1383 78 p Pilaniensclhutz-Nachr 201-524 72 Food and Agriculture Organiation of the United Nations 1982

52 l)ani R C and Majunidar 1978 Prelininary observations on tie Country report surmimary Pages 27-37 in Report on the first incidence of root aphids on differeit rice cultnrcs Sci Cult 4488-89 FAOIUN)IPhail and regional training coiuirse in iniproved ctltural

53 Davidson R I 1979 lffects of root feeding on foliage yield Pages practices for upland rice 5-23 Oct 1981 Northern Region Agrculturd117-120 in T K Ci osb and R B Pottinger eds Proceedings of the Developririt Center Chiang Mai Food and Agricuilture Organiiashy2nd Australian conlerence on grassland invertebrate ecology Govt tion Rome Italy 37 n Printer Wellington New Zealand 73 Franco do Ariaral S 195 1 Residual action of insecticides in turfy

54 l)ean G 1W 1976 Rice insect pests inlaos Int Rice Res Newsl soil--the results of a field experiment [in Portuguese] Biol6gico (Sao 1(2)15 Paulo) 17209-214 (Rev Appl Entomol A41423)

55 I)ean G J W 1978 Insect pests of rice in Laos PANS 24(3)280-289 74 Freeman ) 1970 Report onithe Iban london School of -coroiics 56 )ingle H 1984 Behavior genes and lifc histories conplex adapta- Monographs on Social Anthropology No 41 Athlane Press

tions to uncertain environments Pages 169-194 in A new ecology London UK 317 p novel approaches to interactive systems P W Price C N Slobod- 75 Fuchs T W1 A Harding and 1W Smithlr 1979 Induction and chikoff and W S Gaud cdsl ohn Wiley New York USA termination oldiapause in the sugarcane borer Ann Flitonlol Soc

57 Dobelmann-JP and M Falais 1968 Upland rice cultivation ill tie Am 72271-274 North-western part of Madagascar in 1966-1967 [in French English 76 (hbriel B1) 1975 A review tf the major insect pests of some tpland summary] Trop Agron 2313-21 crops in tlc Philippines Pages 86-111 in Ad Htoc panel of experts on

58 Doniiciano N L 1979 Chemical control of tie lesser corn stalk pest disease arnd weed problems in some rainfed crops Bangkok (lolsonno~paloIgnowslhts) (Zeller 1848) (I-epid(ptera Ihycitid ae) Thailand Food aid Agriculture Organization Roire Italy

on rice in Ponta Gressa Parara 1977-1978 [in Portuguese] Pages 77 Garg D K and N K Shal 1983 White grub outbreak on rainfed 291-303 in Anais 1 Reuniao de tecnicos cmiirizicultura do estado de dryland rice in Uttar Pradeslh Int Rice Res Ncwsl 81(4)16-17 Sao Paulo-Campinas Cati Brazil 78 Garg I) K and IP Tandon 1982 Major insect pests of rice on hilly

59 Dunsmore 1 R 1970 Investigations on tlie varieties pests and tracts of Uttar Pradesh India lilt Rice Res Newsl 7(11)11-12diseases of upland rice (hill padi) in Sarawak Malaysia Int Rice 79 Garrity 1) P 1983 Upland rice research an international bibliograpy Comm Newsl 1929-35 1975-1982 Iternational Rice Research hnstitnte PO Box 933

60 Elias 1R1965 The new decline of rice seedlings is not caused by Manila Philippines 121 p parasites [in Portuguese] Biol6gico (Sao Paulo) 313-6 80 Ghauri M S K 1982 A new subspecies of Iinorh)rone bull

61 Empresa Brasiliera de Pesquisa Agropecuaria 1984 Upland rice itt cornlus Slater (Hemiptera Iygaeidae) front Papua New Guinea on Brazil Pages 121-134 it Av overview of upland rice research rice and carpet grass Ihll lEntoniol Res 72133-137 Proceedings ofthe 1982 Blouak Ivory Coast Upland rice workshop 81 Gonzales F 0 PArregoces R I ihernandef and OT Parada International Rice Research Institute PO Box 933 Manila 1983 Insect and imite pests of rice arid their control in Latin America Philippines [in Spanish] la Federacion Nacional de Arrocerts (Fedearroz)

Bogota Colombia 60 n

38 IRPS No 123 January 1987

82 Grist D H and R J A W Lever 1969 Pests of rice Longmans London UK520 p

83 Hale P R 1979 Two subterranean pests of upland rice in Papua New Guirnca IntRice Res Newsl 4(2)16-17

84 Hale P R and J T Hale 1975 Insect pests of rice in the New Guinea Islands Rice Entomol Newsl 34

85 Harris W V 1969 Termites as pests of crops and trees Common-wealth Institute of Entomology London UK 41 p

86 Hattori 1 1968 A method to distinguish between grass leaf roller (Ctaplha rocis medinalisGuenee) and the rice leaf roller (Susurtia exigua Butler) [in Japanese] Plant Prot (Japan) 23167-170

87 Henry I E and 1A Onsager 1984 Experimental control of the mormon cricket Iahrosshnph by NJsena locustae(Aficrospora Microsporidae) a protozoan parasite of grasshoppers (Ort Acri-didae) Entomophaga 27197-201

88 Hidaka 1 P Vungsilabulr and SKadkao 1974 Studies on ecology and control of the rice gall midge in Thailand Tech Bull 6 TropicalAgricultural Research Center Tokyo Japan 113 p

89 Hincklcy A 1) 1963 Ecology and control of rice planthoppers in Fiji Bull Entomrol Res 53467 481

90 Hirao J 1982 Ecology and chtInical control of the rice leaffolder Jpn Pestic Inf 4114-17

91 Ho DT 1984 Sten borer incifience in rice ecosystems in KenyaEast Africa Pages 307-320 in A overview of upland rice research Proceedings of the 1982 Bouac Ivory Coast upland rice workshop International Rice Research Institute PO Box 933 Manila Philippines

92 Ho )T J 1Njokah and J GKibuka 1983 Studies on rice sten borers in Kenya with emphasis on ifaliarpiaipiraellRag Insect Sci Appl 465-73

93 Ichihara I T Ito A NIatsuo and M Sawada 1971 Chemical control of the rice striped stem borer in upland rice grown in plastic mulch [iii Japanese] Proc Kanto-Tosan Plant Prot Soc 1882

94 IclhiharaIand A Matsuo 1969 Damage caused by rice striped stett borer to upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1686-87

95 Ichhara I atid A MIatsun11970 Conside ration oft lie extraordinary prevalence of rice striped stem borer in upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1778-79

96 Institut de Recherches Agrolnliqes Iropicale et des Cultures Vivrieres 1978 Insect resistance Pages 12-13 in Upland rice 1978 atnual report Bouak Ivory Coast 27 p

97 International Institute for Tropical Agriculture 1983 Aphids itt rice roots I ITA Res Briefs 4(1-3)4-5

98 International Institite fi Tropical Agriculture 1983 Hopperburn in rice caused by brown plathopper IITA Res Briefs 4(l-3)6-7

99 International Rice Research Institute 1975 Annual report for 1974 PO Box 933 Manila Philippines p 216

100 International Rice Restarch Institute 1976 Annual repiort for 1975 PO Box 933 Manila Philippines p 228

101 International Rice Research Institute 1979 Cropping systems program Component technology development and evaluation Page 404 i Annual report for 1978 11O Box 933 Manila Philippines

102 International Rice Research Institute Department of Entomology 1981) Insecticid evaluation 1979 Tables 73 74 PO Box 933Manila Philipes

103 International Rice Research Institute Department of Entonmology 1981 Insecticide evalomtion 1980 Table 81 POBox 933 Manila Philippines

104 International Rice Research Institute 1981 Report of a meeting of Asian rice-based cropping systems entomologists Eleventh cropping systems working group meeting 18-22 May 1981 PO Box 933 Manila Philippines p 132 133 144

105 International Rice Research Institute 1984 Cropping systems program Pages 368-373 in Annual report for 1983 PO Box 933 Manila Philippines

106 International Rice Research Institutc 1984 Upland rice in Asia Pages 45-68 in An overview of upland rice research Proceedings of the 1982 Bouakamp Ivory Coast upland rice workiiop loternational Rice Research Institute 11O Box 933 Manila Philippines

107 Irshad M 1977 Grasshoppers in rice in Pakistan lit Rice Res Newsl 2(3)8-9

108 Ito R and TOno 1964 Upland lice its culture and varieties in Japan lIternational Rice Cormmission loth Meeting of the Working Party on Rice Production and Protection 3-10 March 1964 Manila Philippines Food and Agriculture Organization Rome Italy 12 p

109 Jeffrey M I and V A Dyck 1983 Effect of moonlight on lightshytrap catches of brown planthopper lit Rice Res Newsl 8(6) 13

110 Jennings P R and T Alicia Pineda 1983 Management ofthe rice plnthoppers Sogatodsorivi(coIl in Latin America Presented at the meeting on stable plant resistance FAO and lIndocsian I)inectorate of Food Crops Protection Bali lndonesia 29 Novcniber-2 l)ccemshyber 9 p

IllJimenez Gomez J A 1979 Rice insect pests i lla ios ()riCntales lil Spanish] Pages 99-103 in Rice ciltivation Instituto (olotibiatto Agropecuario Compndio No 29

112 Jongeleen F JJ Soesanto and A Mockidjo 1979 Studies oti pathogens of two species of Scarabacidac froinIndonesia Z Angew Enlomnol 87412-417

113 Jouvenaz 1) PC S lofgren and W A Baiks 1981 Biological control of imported fire antsa review of cirrent knowledge Bull Entomol Soc Am 27203-208

114 Kainoh Y K Shiniiu S MarUani Y Taiaki 1980 lost finding behavior of tie rice bu lplotori+JchinvittjI)allas (liciniptera Corcidac) with special reference to diet pattern ol aggregation and feeding on the rice plant Appl Entomol Zool 15225-233

115 Kakkar K 0 P Sharma and P I Sharma 1977 Note on Ihteronyrchus liodresRedt (Coleopicra I)ynastidae) as asporadic pest of paddy and maie inHimachal Pradesh Corr Agric 156-57

116 Kalode M131 79 Major insect pest problems and their control in upland rice (Unpublishcd mineogriph CWI oJNaaiat lni Chandaratat (198 1)original not seen)

117 Kalshoven L i 98 I The pests of crops in I ldoniesia (translated by P A vatn der Laan) 1 T Ichtiar B3aru-vati Iloeve lakarta Indonesia 701 p

118 Kamal N Q 1981 Suppression of whitebacked planto ner Sogatella fircaifra (Horvath) aLId rice lealfolder (nhal -cis licdb lis (Guenee) populations by natural cnclmie hlI) tiesis Grcgorio Araneta University Foundation Victoncta Park Metto Manila Philippines 179 Ip

119 Karim A N M Reaul 1975 A species ofliealy bug Trioni i70s ceres new oii rice in the Philippines Rice Entoniol Newsl 247

120 Katanyukul Wand C Chandaratat 1981 Insect pestsof uplaid rice and their control Report firstFAOUNlFi Tliailand training course on improved cultural practices for tplad rice Chiang Mai Thailand 5-23 Oct 1981 Food and Agriculture Organiation Rome Italy 22 p

121 Kern F 1955 lhletwioa(I)uninemts) bhhit Biturm(Coleoptera) a new pest of sugarcane inVenezuela [inSpanish with Etnglish sunmary] Agron Trop (Maracay) 527-29 (Rev Appl Eitoniol A4572)

122 Khan K M and S Ghai 1974 White grubs ald their control in India Pesticides (1Bombay) 819-25

123 Kiritani K 1975 Integrated pest management of rice pests Pages 66-70 itt Proceedings platning workshop ott cooperative hIeld research in pest managementP SMotooka ed East-West Center Honololu Hawaii USA

124 Kisimoto R 1967 Ecology of insect vectors forecasting and chemical control Pages 243-255 in The virus diseases of tle rice plant Proceedings of a symposium atthe International Rice Research Institute April 1967 The Johns Hopkins Press altimore Maryshyland USA

125 Kisimoto R and V A Dyck 1976 Climate and rice insects Pages 367-391 in Climate and rice International Rice Research Institute PO Box 933 Manila Philippines

126 Kureha S S Kobayashi SAsari and T Ishihara 1977 Life history and control of the mole cricket Grrllotalpa africana Palisot de Beauvois [in Japanese English translation 7520 1RRI library] Proc Kanto-Tosan Plant Prot Soc 21192-197

127 Kushwaha K S 1981 The black beetle as a pest of rice in Haryana India Int Rice Res Newsl 6(4)14

128 Lawani S M 1982 A review of tile effects of various agronomic practices on cereal stem borers Trop Pest Manage 28266-276

129 Leuck D B 1966 Biology of the lesser cornstalk borer in South Georgia J Econ Entomol 59 797-801

130 lever R J A W 1955 Insect pests of the rice plant in Malaya Int Rice Comm Newsl 1612-21

131 Li Ching-Sing and Shin-Foon Chiu 1951 The study ofthe rice gall midge Iachydiplosis orvza Wood Mason [in Chinese with English summary] 1Taiwan Agric Res 21-13

132 Iitsinger J A 1984 Assesament of the need-based application for rice Proceedings TechnologyTransfer Workshop 15-16 March 1984 International Rice Research Institute 1PO Box 933 Manila Philippines 6 p

133 Litsinger J A 1984 Environmental diversity and insect pest abundance with reference to the Pacific Pages 285-307 in Proceedings sub-regional training course on methods of controlling diseases insects and other pests of plants in the South Pacific 4-20 Oct 1982 Vaini Kingdom of Tonga Nukualofa

134 Litsinger J A RF Apostol and M BObusan 1983 White grub Leucopholis irrorata (Coleoptera Scarabaeidae) pest status popula-tion dynamics and chemical conllol in arice-maize cropping pattern in the Philippines J Econ Entoinol 761133-1138

135 Litsinger J A C de la Cruz F Raymundo A T Barrion M D Lumaban J P Bandong M S Venugopal F Paragna and E Balete 1982 Insect pests and insecticide reponse of dry-seeded bunded rice Pages 179-193 in Report of workshop on cropping systems research in Asia International Rice Research Institute Los Bafos Philippine

136 Litsinger J A M 1) Luinaban J P Bandong A T Barrion W Katanyukul and N Panda 1978 Insect pest management in rainfcd lowland rice agroecosystms of tropical Asia International Rice Research Conference 17-21 April 1978 International Rice Research Institute Los Bafios Philippines

137 Litsinger J A and K Moody 1976 Integrated pest management in multiple cropping systems Pages 293-317 in R I Papendick P A Sanchez and G B Triplett eds Multiple cropping American Society of Agronomy Madison Wisconsin USA

138 Loevinsohn M E 1984 The ecology and control of rice pests in relation to the intensity and synchrony of cultivation Ph D thesis University of London 36) p

139 Loevinsohn M E J A Litsinger N Panda M A Austria R F Apostol C Andrion and J Bandong 1982 A basis for predicting rice insect pest abundance in cropping systems Paper presented at the 1982 International rice research conference on rice and cropping systems 19-23 April 1982 International Rice Research Institute Los Bafios Iaguna Philippines 25 p

140 Lopez A W 1930 Repot of the Entomology Department Pages 145-172 in Annual report of the Philippine Sugar Association 1929-30 Manila

141 Lopez AW 1931 Entomology Department Pages 227-266 in Annual report of the director or research 1930-31 Philippine Sugar Association Manila

142 Lopez A W 1932 Entomology Deparment Pages 252-279 in Annual report of the Philippine Sugar Association 1931-32 Manila

143 Lungibill P and G G Ainslee 1917 The lessercornstalk borer US Dep Agric Agric Res Serv Tech Bull 539 27 p

IRPS No 123 January 1987 39

144 McGuire J U Jr and BS Crandall 1967 Survey of insect pests and plant diseases ofselected food crops of Mexico Central America and Panama U SDepartment of Agriculture International Agriculshytural Development Service Agricultuial Researrh Service October 1967 11 p

145 McNeill Sand T R ESouthwood 1978 The role of nitrogen in the development of insectplant relations Pages 77-9 in lliochemical aspects of plant and animal coevolution J B Ilarbornu ed Academic Press London UK

146 Maebara H 1955 Spatial distribution of lastinstiarlanaeof purplish stem borer (Sesamia iqferens Walker) hibernating in an upland rice field Bull Kagoshima Univ Fac Agric 438-41

147 Mallick E H and N G Hajra 1977 Response of rice plant to defoliation treatments at different developmental stages Indian Agric 2151-56

148 Mansingh A 1971 Physiological classification of dormancies in insects Canl Entomol 103983-10)9

149 Martell C G 1974 Catalogue of plant feeding insects of Mexico [in Spanish] Fitofilo 27(69)1-176

150 Martinez 0 J 1982 Principal rice pests in Ilanos orientalis [in Spanish] P~ages 80-94 in Curso deact nalization de conocimientos en arroz Villaniencis Instituto (olonmbiano Agropecuario

151 Martins J F da Silva F Ferrcira A S Prabhu F 1 P Zimmermann 1980 Use of pesticides for the control of the main subterranean upland rice pests esqui Agropecu Bras 1553-u

152 Martins I F da Silva F Fe eira rod BdamSilveira Pinheiro I Q2 Damage simulation caused by defoliators of upland rice [in Portuguese with English summary] Pesqui Agropecu Bras 171113-1119

153 Martins 1F da Silva N(iyuen van Tan and 1Ida Silveira Pinheiro 1981 Stem borer resistance in upland rice associated with plant morphological characters [in Portuuce Iesqui Agropecu Bras 16187-192

154 Masaki S 1980 Sumnmer diapause Ann Rev Entomiol 251-25 155 Masuzawa T 11 Suwa and F Nakasoji 1983 l)ifkleences of

oviposition preferencc and survival rate of two skipper butterflies Parnara guttata and PIchlpidai iathiais (Iepidoptera HIesperiidac) on rice plant and cogon grass New Entomol 3247-56

156 Maurya D M and C P Vaish 1984 Upland rice in India Pages 69-92 in An overview oftpland rice research Proceedings ofthe 1982 Bouakamp Ivory Coast upland rice workshop International Rice Research Institute IO Box 933 Mnila Philippines

157 Menon M G R R N Katiyai S Kuar and S S Misra 1970 Termites attacking the paddy crop Sci Cult 36294-295

158 Miyahara Y and 1 Nishikubo 1967 Behavior of young nymphs of the red rice root aphid Iholahsipiun ri liabhdontinalisSasaki in relation to movement from the leaf to the roots of upland rice [in Japanese] Pioc Assoc Plant Prot Kyushu 1363-66

159 Mo ljoa Tjien 1952 White grub of upland rice (Holotrichiahelleri Brsk) [in Indonesian with English summary] Contrib Agric Res Stn Bogor No 13134 p

160 Mochida 0 and T Suryana 1979 The occurrence of the brown planthopper and ragged stunt on lowland and upland rice ill Central Java and logyakarta in tile late of January 1979 Entomnolgy Survey Report No 15 Central Research Institute of Agriculture (CRIA) Sukamandi Indonesia I February 1979 9 p

161Monet C 1979 Varietal resistance to upland rice pests [inI French] Pages 35-39 in Annual report 1977 Institut des Savanes Department des Cultures Vivrieres Bouak6 Ivory Coast

162 Monte 0 1942 A caterpillar ofrice-fields[in Portuguese] Biol6gico (Sao Paulo) 8161-163 (Rev Appl Entomol A31414)

163 Munroe I) D 1979 Pest management strategy for hill rice in four resettlement areas in Sarawak Malaysia Pages 89-102 i1 Guidelines for integrated control of rice insect pests Food and Agriculture Organization Plant Production and Protection Paper 14 Rome Italy 115 p

40 IRPS No 123 January 1987

164 Nakasuji F M Ishii and Jen-Zon Ito 1984 Rice skippers in Taiwan and their life histories ZZrC (Taipei) Tech Bull 82 10 p

165 Navas I) 1976 Fall armyworum in rice Pages 99-106 il Ecological animal control by habitat management Proc Tall Timbers Con-ference 28 Feb-I Mar 1974 Fall Timbers Res Stn Misc Publ 6 Tallahassee Horida USA

166 Nickel 1 1 1961 Biological control of rice stem borer A feasibility study Tcch Bull 2 International Rice Rcarch Institute PO Box 933 Manila Philippines IIl p

167 Nigan P M and R A Vernia 1985 Insect pests of uplaild rice in Uttar Pradesh Int Rice Res Newsl 10(6)22

168 Nilakhe S S 1985 -cological observations on spittle bugs with emphasis on their occirence in rice Pcsqtui Agropecu Bras 20407-414

169 Nixon G EJ 195 1 lie association of ats with aphids and coccids Commonwealth Iusti lcte of Entonology Iondon UK 36 p

170 Nurdin F an Ishak Manti 1985 Study oit seedling fly of upland rice Interlational Upland Rice Moitoring Tour Report 26 lebruary-2 MarLI Sukaranti West Sumatra

171 Oka 1 N 1979 Ciltural conttol of the brown planthopper Pages 357-369 in Brown planthopper threat to rice production in Asia International Rice Research Institutc 1p0 Box 933 Manila Philippines

172 Ohni M and A Vilnaii 1975 Rice insect pests of Italy biology of Sipha grcria (Kaltenbach) and its control [in Italian] Riso (Milan) 2459-71

173 Otanes F Q atnd 1 I Sison 194 1 Pests of rice Philipp J Agric 12211-259

174 Painter R I 1951 Insect icsistance in crop plants University Press of Kansits Lawrence Kansas USA 520 p

175 Pantua P C and 1A litsinger 1981 Arthropod abundance in continuous vs biannual rice cropping systeis Int Rice Res Ncwsl 6(2)20-21

176 Pathak M ) and V A l)yck 1975 Sttudics ott insect pests of upland rice Pages 186-191 in Major research in upland rice International Rice Research Institute PO Box 933 Ilanila Philippines

177 Patil B R B M IHasabe and 1) S Ajri 198 I Chemical control of white gritb lIditriihinhorraia Fabr intfesting groundntut paddy atnd sorghum Indian 1Entomol 43232-236

178 Pawar A 1) 1974 (ollection and identification of insect pests of rice rlerminal repirt lInternational Rice Research Institute PO Box 933 Manila Philippines 72 p (tnimco)

179 IerrajU A aid A S Reedy 1965 Observations ott diapausc itt tie steti borer ohrice Rice News Teller (lai-Mar)9-13

180 Pont A C 1972 A review of the Oriental species of theri ona Rondani (I)iptera Muscidac) of econtomic importance Pages 27-102 in Control of sorglium shoot fly M G lomtani and W R Young eds Oxford atd IH1 Ptthl Co New I)clhi India 324 p

181 Prasad S C 1PChand aid M 1PSingh 1983 Incidence of rice gall midge in the upland plateau region of Hihar Oryza 20235-236

182 Rai P S 198 0 Nisaga simpex damage to rice in the hill tracts of South India Int Rice kles News[ 5(I)12

183 Rathorc Y S 1969 Gonocplhahli dcprssunt Fabr as itpredator of (tilo zonwIlhis (Swinhoc) and other moths in field cage at Patnagar Uttar Pradesh Indian 1Entotnol 3127

184 Reed W 1983 Crop losses causcd by insect pists in the developing world Proceedings 10th Internatial Congress of Plant Protection lrighton UK 174-80

185 Reedy M V 1981 Ateactogasterinhens(Walker)a new root-grub pest of tipland paddy and inaize Curr Sci 501076

186 Reveno M A 1968 Catalogue of the insect pests of economic crops in Colombia [in Spaiish] Asociacion Latino Americana de Entomo-logia (AIAE) 129-33

187 Reyes G M 1957 Rice dwarf disease in the Philippines Food and Agriculture Organiiation Plant Prot Bull 617-19

188 Rice S E A A Grigarick and M 0 Way 1982 Effect of leaf and panicle feeding byarmyworm (Lepidoptera Noctuidae) larvae on rice grain yield Environ Entomol 7593-595

189 Risch S J 1) Andow and M A Altieri 1983 Agioccosystem diversity arnd pest control data tentative conclusions and nizw research dirctions Environ Entomol 12625-629

190 Rivera C T 1956 Three organic insecticides in thecontrol of insects affecting upland rice Philipp Agric 39465-472

191 Roffrey 1 1972 Locusta outbreaks in the Philippines Acrida 1177-188

192 Rojas R 1) 1941 The effect of pruning at varying ages upon the yield of Ramairio Phipp Agric 29851-860

193 Roonwal M i 1979 Ternite life and termite control in tropical South Asia Scientilic Pnbl odhpr India 139 p

194 Rosetto C1 F da Silva Martins N C Schmidt and L EAzzini 1978 Damage caused by pasturespittle bugs 4Dvoi)lavopictaandDJ schach) on rice lBragantia Nota 6 (Matcli)25-27

195 Rosztto C1- S S Neto I- Amantc I) Iink 1Garcia-Vieira D M de SoLza N V Bansa t and A M de Olivcira 1971 Rice pests of Brazil [in Portuguese] Pages 149-238 inrFA()-International Rice Commission 2nd meeting of the Rice Committee for the Americas 6-11 l)ecember 1971 Pelotas Rio Grande do Sul Brazil

196 Rothschild G H L 1969 Obseratiois on the armyworm Spodoptera matritia acronrctoiiles Gn (Lep Noctuidae) Bull Entomol Res 59143-160

197 Rothschild G 1- L 1974 Parasites of the armworm Spolopiera mauritiaacroncitoihes in Malaysia Fntornopliaga 19293-299

198 Sachan i N S K Gangwar and 1N Katujar 1980 Field crickets damaging cages in North Eastern IHill Region Indian J Entomol 42526-527

199 Sands 1) P A 1977 The biology and ecology of Lepocorisa (Hemiptera Alydidac) in Papua New Guinea Res Bull 18 Departshyment of Primary Industry Port Moresby Papua New Guinea 104 p

200 Sarawak 1968 titomology division Pages 97-106 in Report of the research branch of the Department of Altriculture 1966 Kuching (Rev AIppl Eintomol A56603)

201 Satpathy JM M S Das and K Naik 1977 Effect of multiple and mixed cropping on the incidence of scine important pestsI Entomol Res 178-85

202 Sauer IHF G 1939 Notes Ott Elasmpnlpualtsligniosellths Zeller (Lep Pyr) a serious pest ofcereals in Sao Paolo [in Portuguese] Arq Inst Biol 10199-206 (Rev Appl Entotnol A28378)

203 Satinders 1L A 1 SKing and C L Vargas S 1983 Crop pests of Central Anerica with t bibliography [in Spanish] Centro Agroshynotnico Tropical de Investigacion y Ensenanza Turrialba Costa Rica (CATIE) Tec Bol 9 90 p

204 Sauphanor B 1985 Some factors of upland rice tolerttce to stem borers in West Africa Inseu Sci Appl 6429434

205 Sen P atnd S Chakravorty 1976 Stentflics in Haritighata Nadia West Bengal India [it Rice Res Newsl 1(12) 15

206 Senapai B and 1M Satpathy 1983 The seedling fly Atherigona orrzae Malloch is a pest of rice in Orissa (India) J Entomol Res 777-79

207 Shhjahan M S Alatn M Ah nad and BN Islam 1983 A new scarabaeid beetle pest of dryland rice in Bangladesh Int Rice Res Ncwsl 8(I)16-17

208 Sharma G L S C Modgol and R C Gautam 1979 Effect of depth of submergence ott incidence of bacterial blight and white grub infestation in transplanted rice Int Rice Res Newsl 4(6)12-13

209 Sharma S 1985 Scarabaeid beetle outbreak itt upland rice Int Rice Res Newsl 1(1(l)24

210 Shiraki T 1917 The paddy borer Schoenobhits incertulas Walker [achaku Agric Exp Stn Formosa 256 p

211 Silva A de Brito and 11P Magalhaes 1981 Insect pests of rice [in Portuguese] Circular Tecnica No 22 EMBRAPA Centro de Pesquisa t ecuaria de Tropico Unido (CPAJU) Belem Brazil 14 p

212 Singh S A 1979 Emergence of rice leaf butterfly as a rice pest in Manipur India int Rice Res Newsl 4(4)16-17

213 SinghS A 1983 Changing trends of insect pestsof rice in Manipur Int Rice Res Newsl 8(f)12

214 Siwi S S and P H van Doesburg 1984 Lenovorisa Latreille in Indonesia (Heteroptera Coreidae Alydinae) Zool Meded (Leiden) 58117-129

215 Soejitno J 1977 Reaction of some upland rice varieties toseedling fly eltherikonaexigua Stein [in Indonesian] Siri Hana I Penyakit 630-35 Lembga I osat Fenclilian Pertanian Bogor Indonesia

216 Socnardi 1967 Insect pests of rice in Indonesia Pages675-683 in The major inse pests of thu rice plant Proceedings of a symposium at the Internatital Rice Rec Wi itt S pternber 1964 The Johns Hopkins Press Baltimore Maryland USA

217 Soto P E A T Petcz and I W Buddenhagen 1976 Survey of insect pests and diseases of rice in different ecological zones in Nigeria Rice Entomol Ncwsl 435-46

218 Soto 11E and Z Ziddiqi 1976 Screening for resistance to African rice insect pests Varietal improsetnent sCmi n r Bouak Ivory Coast 13-17 Sept 1976 West Africa Rice l)evelopment Association Monrovia Iiberia

219 Squire F A 1972 Entomological problems in Bolivia PANS (PestArtic News Sumr) 18249-268

220 Stapely 1 II 1978 The rice leaf roller in the Solomon Islands Int Rice Res Newsl 3(l)13

221 Suenaga H 1937 On aphids attacking the roots of rice Oyo-I)oubts Zasshi 9150-152 [in Japanese] (Rev Appl Entomol A25745)

222 Suenaga H 1952 On the seasonal abundance of three subterranean aphids on upland rice Bull Kyushu Agric Exp Sti 1249-262

223 Sutynso Yas and Dandi Soekaria 1964 Beberapa hana yang ttenperang tana man padi tanah kering [in Indonesian] Rapat kerja Padi Tanah Kering 6 sd 8 Juily 1964 Kaliurang 9 p

224 Tallamy I) W and R F Denno 1983 Alternative life history patterns in risky environments in example from lace bugs Pages 129-147 in Insect life history patterns R F l)cnio and If Dingle eds Spring-Verlag 225 p

225 Tamura 1 1939 Observations on some insect pests of field crops [in Japanese] 1 Plant Prot 26414-420 (Rev Appl EntomolA27-556)

226 Tamura and Suzuki 1964 Influences of the early-transplanting practice in rice culture upon development of the rice stem borer Chilo siql)ressalisWalker Bull Ilokuriku Natil Agric Exp Stn 761-94

227 Tan J P 1924 The rice root aphid Drowhi ihirsoia Philipp Agric 13277-288

228 Tanaka 1 1957 Taxonomy and distribution of some subterranean aphids injurious to upland rice in Japan with description of a new species [in Japanese English summary] Botyu-Kagaku 22168-176

229 Tanaka T 1961 The rice root aphids their ecology and control Spec [full College of Agriculture Utsunomiya University Japan F3 p

230 Tauber M IC A Tauber R Nechols and 1Obrycki 1983 Seasonal activity of parasitoids control by external internal and genetic factors Pages 87-108 itt I)iapause and life cycle strategies in insects V K Brown and I Ilodek eds )r W Junk The Hague Netherlands

231 Taylor W F 1972 Effects of artificial defoliation (simulating pest damage) on varieties of upland rice Exp Agric 879-83

232 feotia T P S and S Natud 1966 Biotoomics of the rice skipper Parnaraniathiis Fabricius (Lepidoptera Hesperiidac) Indian J Entomol 28181-186

233 Teran F 0 1971 Rice pests of Santa Cruz [in Spanish Ministerio de Agricultura Sita Cruz Bolivia Bull 45 8 p

234 Than Itun 1976 Population dynamics of the yellow rice borer Tryporza itcertulas (Walker) and its damage to the rice plant MS thesis University of the Philippines at Ios Bafios Laguna Philippines 92 p

235 Uichanco L B 1911 Pests Pages 55-74 it Rice Philipp Agric 11-200

IRPS No 123 January 1987 41

236 van der Laan P A 195 1 Life history of the rice seedling fly Contrib Gen Agric Res Stn Boor Indonesia 118 15 p

237 van der Laan 11A 1959 Correlation between rainfall in the dry season and the occurrence of the white rice borer (Scirpolhaga innotala WIk) in lava Entomol Fxp Appl 212-20

238 Van Driesche R G 1983 Meaning of percent parasitismin studies of insect parasitism Fnviron Entomol 121 I1- 122

239 Venugopal M S and A litsingcr 1984 Fffect ofcarbofuran on rice growth Prot Ecol 7313-317

240 Vietnam Directorate of Rural Affairs 1962 Annual work progress report on crop improvement program of rice sugarcane vegetable and field crops (lor the period from July 1961 to June 1962) (11) 299 p (Rev Appl Entomol A51526-527)

241 Wan M T K 1972 Observations on rice leaf- and planthoppers in Sarawak (Malaysian Borneo) Malay Agric 148308-335

242 West Africa Rice I)esclopnient Association 1978 Annual research report Vol 2 Statistics Monrovia Liberia

243 West Africa Rice Development Association 1979 Annual research report p47 Monrovia Liberia

244 Williams ) F and CS lofgren 1981 Fli I illy l-468 anew bait toxicant for control of the red imported lire at Fla Entomol 64472477

245 Williams I) 1 1970 The mealybugs OFlomoptera Coccoidea Iscudococcidac) of sugar-cane rice and sorghum Bull Entomol Res 0109-188

246 Williams D) 1 1982 Pulinaria icervi (Signoret) and its allies ott sugar cane and other grasses Btll Entoinol Res 72111-117

247 Wirjosuiardjo S 1976 Study ofchenmical cont rot of insect pests ott upland rice Imiu Pertanian (Jogjakarta) 1323-329

248 Wirjosuhardjo S and N Wirarta 1976 Chemical control of the white grub tIolorithiatlltriin upland rice in M uneng Irobolinggo Indonesia) [in Indonesian] Imut Pertanian (Jogjakarta) 1330-332

249 Yano K T Miyake and V F Eastop 1983 The biology and economic importance of rice aphids (Henuiptera Aphid idae) a review Bull Entomol Res 73539-566

250 Yelp D and P Rhodes 19(4 lhe success of drilled dryland rice iii the Fiji Islands International Rice Commission Working Party on Rice Production and Protection Manila Philippines 3-10 March1964 Food and Agriculture Organization Rome Italy 13 p

251 Young W R 1981 Fifty-five years of research on the sorghum shootfly Insect Sci Appi 23-9

IRR Irch Institute Co EN CAS DE NON REMISE RENYOVER A

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ISSN 05-3862Other papers in this series TITLES OF NUMBERS 1-45 ARE LISTED ON THE LAST PAGE OF NO46 THOSE OF NUMBERS 46-70 ARE ON THE LAST PAGE OF NO 71-80AND THOSE OF NUMBERS 71-1O0ARE ON

THE LAST PAGE OF NO 101-122

No 101The economics of hybrid rice production in (hina No 113 Boron toxicity in rice No 102 Rice ratooning No 114 Energy analysis rice production systems and rice research No 103 Growth and development of the (Jeep water rice plant No 115 Production risk and optimal fertilizer rates an application of the

No 104 Faridpur acomputer-assisted instruction nmodel forrainfed lowland random coefficient model No 116 Consumer demand for rice grain quality in Thailand Indonesia andrice

No 105 A readingand listening comprehcnsion test inEnglish for non native the Philippines speakers applying for tiraining at IRIRI No 117 Morphological changes in rice panicle development a review of

No 106 Rice grassy stunt virus 2 a new strain of rice grassy stunt in tire literature

Philippines No 118 IRRI-Korca collaborative project for the development of cold-No 107 Physical losses and quality deterioration in rice plstprodnCtion tolerant lines through anthei culture

systemis No 119 Problen soils as potenial areas for adverse soils-tolerant rice No 108 (olptblication ofIRRI materials a survey of translators and varieties in South and Ssuthcasi Asia

publishers No 120 Changes isinall-farm iLtireliing techii)hgy in Thailand and [he No 109 Classification of Philippine rainfall patterns Philippines No 110 Contributions of modern rice varieties to rutrition in Asia No 121 Landforms and modern rice varieties No Ill Changes in rice breeding in 10 Asian countries 1965-84 No 122 Yield stability and modern rice technology No 112 Design parameters affecting the performance of the IRRI-designed

axial-flow pump

Page 18: UPLANDRICE. INSECTl--1i M T -E!R.EOLOU t RTANCE ,AND …delphacid.s3.amazonaws.com › 6958.pdf · The significance of abiotic factors in upland rice insect ecology will be apparent

0 z

Table 2 Life history patterns of upland rice insects or insect groups

Host range Longevity Dormancy Vagility FertilityMonophagy Oligophagy Polyphagy Low Medium High Yes No Low Dispersive Migratory Low Medium High(Oryza spp) (grasses sedges) (angiosperms) (lt3 wk) (gt2 mo) (lt1 kin) (lt50 eggs) (gt500 eggs)

Ants xTermites x x x

Mole crickets x x x xx x xField crickets x xx

White grubs x x x xx xRoot aphids x x xxMealybugs x

x x x xxSeedling maggots x

x x x x xLeaf beetles

x xx xArmyworms x x xx x

Thrips x x xx xChilo spp x x xx xMaliarpha x x x x

xLocusts x x Xx x xGall midges x x xxiVephotettL iirescens x x xx x x xLeprocorisaspp x

x xLeaftolders x x x

Sogatellafurcifera x

xx x x x x x x

00

IRPS No 123 January 1Q87 19

All this appeals to favor insect pest buildup but the notwithstanding the release of stored nutrients in response contrary is true Loevinsohn (138) showed that insect pest to drought that temporarily benefits philoem feeders populations respond exponentially with the proportion of land devoted to rice up to about 75 of the area then the M POR ANCE I I P1A 1) RWI P1St S

rate of response declines In most upland rico areas tie rice crop occupies less than 50i of the area- therefore the Visits to upland ricefields normally reveal 1kw insect pests potential for population buildup is low This relationship Loevinsohn (138) and Locvinsohn ct al (139) compared may not hold for seed pests that can locate small ricefields insect abundance using annual light trap atchcs in upland

Another outcome of the highly diverse flora and phical rainfed lowland and irrigated lowland sites in the Philipshyenvironment of upland rice isthateach upland ricearca has pines Although only one upland site was studied data a unique composition of insect pests showed equal or fewer insects at the upland site than at the

For example in Tanauan Batangas Philippines the rainfcd lowland sites and fewer than at the irrigated sites main upland rice pests are rice leaffolder(RLF) armvworm (Table 3) M Lltiple regression analysis showed that the key Aythntnaseparata WBPH ants Soli psis gentitataand factors explaining insect abundance among a set of cropping white grub Leucopholis irrorata Sixty kilometers away in intensity variables were the number of rice crops grown per Real Quezon the main pests are seedling maggot Athe- year followed by area devoted to rice Table 3 shows that rigona orae flea beetle (Thacitcmnra hasalis thrips the upland rice site had only I rice crop perym rand that rice Stenchaetothrips hiurnmis and rice bug Leptocorisa was planted in only 20Vi of the area Fertiliter and oratorius In a third Philippine upland rice site in Claveria insecticide use had little bea ring on the trend The armters at Misamis Oriental Mindanao the main pests are ants the upland site applied 60 kg N ha higher than at most Solenopsis gemtinata seedling maggot Alieriona orvzae irrigated sites white grubs Holotrichia in)iUaiuoan t and Lucopholiv Insect occurrence at the upland site was omparable to irrorata stem borers Sesamia ifiArelr (hio auriciliusC that at the Cagayan rainfed lowland site parlicularly in 1982stpessallis Scirpophaga incertulas S innoiata and when drought prevented many farmers fron planting(only

4c ona chrrsurapella root aphids Tetraneura niriah- 40(of the rice area eventually was planted that year) M ore dunrinalisand Rhupaosiphunm riahbdminalis lcaffolders favorable weather occurred in 1980 and 1981 which Marasniapataltis and Chaphalocrocis Medinalis army- supported greater rice areas and consequently more insects worm MIthinna separata and rice bug Leltucorisa The most intensive site for insect pests was South Cotabato oratoriusThe great diversity of insect problems is typical Mindanao where farmers plant 23-25 rice crops per year fer upland rice and makes control efforts more difficult with irrigation The Zaragoa Nutva Ecija site also had particularly regarding breeding for insect resistance high pest abundance and at this site the fields were highly

asynchronous Asynchrony was the third most important Low yield potential Iactor in explaining rice insect abundance Most upland rice is grown on low-nutrient and mineral- Rice cropping intensity in upland areas is bound to toxic soils which when combined with eiratic rainfall remain low Only one rainid upland crop is possible per make upland rice a highly riskycrop for the farmerto invest year (Farmers in Claveria Misamis Oriental Mindanao in costly soil amendments and land management Often Philippines tried asecond upland ricecrop i 1984a ycarof upland rice isgrown far from markets making inputs even favorable rains but no harvest was possible b-cause of more expensive Farmers growing a subsistence crop hosi- drought blast and birds) Rice area isaso bound to remain tate to invest in costly inputs to raise yields low because of the high labor and power rcquirelmenlts to

Even having varieties that could double existing yield prepare the land (because of weeds)and because upland rice potential under present management levels would probably wvill not compete with lowland rice as a cash crop Some not prompt many farmers to use purchased insecticide upland rice sites with prolonged rainy seasons howeverare unless tile risk of crop failure were reduced such as it was planted asynchronously such as the Clacra site with irrigation systems in lowland rice The seasonal abundance of Upland rice insect pests

The Tanauan Batangas site is atypical for upland rice attracted toa light trap wasgrapied for crop year 1980-81 in The soils and rainfall pattern are favorable and because of Tanauan FBatangas (Fig I 2) W13P11 was he most its nearness to Manila farmers have cash resources ftm common rice hopper and its population peaked at the sales of vegetables to purchase tertilier They applied an midgrowth stage as it does in lowland rice D-Vac suction average of 60 kg Nha after particle initiation and obtained samples showed an earlier peak than that from a light trap 25-35 tha yields The high fertility isone reason why RLF indicating emigration W13PI usually emigrate after the armyworms and WBPH developed into large numbers vegetative stage Populations in 1980 reached levels that Farmers however used no insecticide on upland rice caused patches of slight yellowing in some fields WBPH although they did on vegetables probably cause hopperburn in Batangas They immigrate

The poor growing conditions fOr rice generally mean year-round as shown by low levels in the light trap BPH poor growth and low fecundity for insect pests fIeding on it however were scarce in the light trap and not collected with

Table 3 Abundance of rice insect pests and factors in rice cropping intensity of Philippine upland rainfed lowland and irrigated lowland locations where light traps were operated daily for atleast I yr -I

0Rainfed Rainfed lowland Irrigated lowlandupland

Pangasinan Cagayan lloilo Santa Maria Nueva Ecija South Cotabato Tanauan Caaringayan Caaringayan Bangag Bangag Bangag Buray

LagunaCale

Batangas Manaoag 1980-81 Santa Motiica 1982-83 Cabanatuan Zaragoza Avancena NaniamanSolana 1981-82 1982-83 Oton

1980-81 1979-80 Oton 1981-82 1981-82 Koronadal Koronadal1980-81 1979-80 1979-80 1983 1983

Insect (kerosene light trapNephotcttixcatch nolyr)virescens 650 1240 500 1800 330 210 3400 1600 160N nigropictus 210 - 360 18400 5500 2900- 140 90 20 - shy 1700 -Reciliadorsalis 710 - 3200 1700180 890 630 140 170 2400 800Nilaparratalugens 130 690 - - 4200 4600300 520 750 180 60 3300 1600 740 310Sogatellafiircifera 1700 31000 13000 8200310 210 3300 610 270 980 1600 6300 -Cyrrorhinus - 12000 6700600 540 3200 290 160 330 120 410Scirpophaga spp 50 410 620 - - 21000 12000150 180 220 270 30 1000Cnaphalocrocismedinalis 70 70 540 610 270 11000 6100200 140 100 6 60 250Chilo suppressalis 0 60 60 1900 300 200- - 30 20 0 6 3 160 -C auricilius - 0 00 - - 20 3 60 20 3 0Sesania inferens 0 - 80 60 3 80 10 51 -Cropping intensity _ -

Rice crops (noyr) 10 10 12 10 10 10 13Rice area (7c) 17 19 20 20 25 2320 85 85 60 70 40 85 85 90Fertilizer (kg Nha) 60 20 85 80 70 7020 0 0 0 30 30Insecticide (no of 40 60 40 300 04 04 0 03 300 13 13 30 50 40 33 33applicationscrop)

__________

IRPS No 123 January 1987 21

Light trapped Insects Drytnd __ 2(notrap-nght) Suctioned Insects (nom )

50 -)o 9 jO40- SOW ffurciWO Light trp

D-Vc suction

20 to 4

20

Egg porasitizahion (M) 75 60

300 SOOampReOftwciffr

45 30shy

15

-insects (notrop-night) 4 Nileporvata Iuen5

Egg parastization()

Light trapped insects (no trop-nght) Suchoned insects (no m 2 )

6 8 Cytorh~nuls hvidipennii 0

4 05 2

50 0 Insects (noltrop-night)40-ui Ipanh pe g

Moohase 1 Seasonal abundance measured by light trap

Rainfall(etod) and D-Vac suction sampler of whitebacked

to ~ hopper Nilaparvata lugens and lea ffolder60)-t Cnaphalocrocis mnedinalis andplthpe g20 L I II I I I parasites and mtrid predator Ortorhjntus livi-Apr May Jun Jul Aug Sep Oct Nov Dec JaIn Feb Mar dipennis set against cropping pattern daily1 9 80o 19 8 1rainfall and moon phase Tanauan Batangas

1980-81

the D- Vac Cyrtorhinus build up on the crop coterminous are collected in light traps during full moon (109)with WBPH buildup Cyrtorhinus disperses during the Because upland rice shares most of the lowland rice pestsrainy season RLF not readily collected in light traps was and those pest species are generally less impotant to uplandprevalent only during the rice crop rice than lowland rice there is a general belief that insect

Nephotettix virescens was more abundant than Nnigro- pests are less important onpicLus in the light trap collections and on the crop GLH upland than lowland rice Arcview of limited data on upland rice shows an averag of population peaked at the reproductive stage as did zigzag 10-21 yield loss from insect pests on a par with that ofleafhopper (ZLH) Crtorhinus tracked the populations of lowland uee determined by the same methodology (132) planthoppers and leafhoppers Both GLH species were (Table 4) McGuire and Crandall (144) made a similarcollected year-round in the light trap indicating immigra- estimate for Central America Cramer (51) howevertion from nearby irrigated rice areas Possibly because of the estimated 35 yieldlow populations loss in South America based on ainsect number did not correlate with similar pest complex - stem borers and seed bugs shymoon phase although reports indicate that more hoppers attacking irrigated rice in North America This estimate did

22 IRPS No 123 January 1987

Light trapped insects D~~nd (no trap-night )

St plht Suctined insects (nom 2

)

D-Voc suction 1520

1o

0 Lih trap 5 ~105 o5

00 Eggparositizotion()5040 NephOtgtha virneens

30 20

I010 A-Insects (notrop -night)

4 Nephatelh nigropccent i

2 o I vV^J - -- ^ VA____ f _

Light trapped insects (no trap-night) Suctioned insects (no rnE

10 -1208 ReCdO doRsol

6 15

4 10 052 iwxALAiamp 0

Light trapped insects (notrap-night) 2Suc oned nsects (nom )

8 CyrtorhmnusI6o nn]10 i0075

4 05

2 0 0

025

2 Seasonal abundance measured by light trap Roinfall (mmd) Moon phase and D-Vac suction sampler of green leafhoppersNephotettix virescens and N ngropicrus and I zigzag leafhopper Recilia dorsalis their egg 60

parasites and micid predator Cyrtorhinus livi-dipennis set against cropping pattern dailyrainfall and moon phase Tanauan Batangas1980-81

40

5 Lf Apr May Jun

1980 Jul Aug Sp Oct Nov I

-aMar

1981

not account for soil pests and was not based on field data in the absence of pests Carbofuran granules are phytotonicAkinsola (I1) cited mainly data from irrigated rice areas in to rice (239) The prevailing hypothesis is that carbofuranAfrica but estimated yield losses of upland rice between 15 prevents soil bacteria from consuming fertilizer The phytoshyand 30 from insects However losses in Thailand over a tonic effect is through greater availability of fertilizer to the4-yr period ranged from I to 13 averaging 5(106) plant Denitrifying bacteria are known to be more active inYield loss data however should be interpreted with aerobic than anaerobic soils therefore it may not be merecaution (184) The Philippine data were all derived from coincidence that the highest recorded yield losses (22-69)trials on farmers fields under farmers normal agronomic occurred in those trials with 12-30 kg basal Nha On themanagement practices except for Bukidnon and Capiz other hand contro with carbofuran granules may not be aswhere researchers used fertilizer Batangas farmers are good as a seed treatment against seed pests and perhapsatypical because ofhigh levels of fertilizer used The trials in yields would have been higher if a systemic seed treatmentBrazil were carried out at an experiment station but insecticide had been used conditions were similar to those of local farmers The 1979 and 1980 trials in Batangas had two full-

Carbofuran granules were used in every trial as a broad- protection treatments one using carbofuran granules andspectrum insecticide applied basally with the seed An the other using bendiocarb WP as a seed treatment Plotsobjection to using insecticides to measure yield losses isthat protected with bendiocarb yielded higher than those proshyinsecticidescandirectlyorindirectlystimulateplantgrowth tected with carbofuran granules The other trials in the

Table 4 Yield losses to insect pests and yield responses to insecticide in upland rice in the Philippines Brazil and Upper Volta

Yield (tha) Yield loss (5) Location Year Cultivar Protected

with Protected

without Check Total Vegetative Reproductive Ripening

Basal N (kgha) Reference

carbofuran carbofuran Protected No with carbofuran

carbofuran

Philippines Pili Camarines Sur Tanauan Batangas

Pangantucan Bukidnon

Dumarao t-apiz

BrazilGoias Minas Ge~ais

Upper Volta6

1975 1974 1974 1975 1975 1976 1977 1978

1979

1980

1979 1980 1979

1977 1978 1979

Bursiging puti Kinanda Dagge Kinanda Dagge Dagge Dagge Dagge UPL Ri5 Dagge UPL Ri5 Dagge UPL RiS UPL Ri5 UPL Ri5 UPL RiS

IAC47 IAC47 IAC47

14 33 11 36 35 31 30 27 37 26 40 30 40 42 46 35

19 21 09

--

-34 24 24 27 --

29 43 31 42 -

47 25

---

10 28 05 27 27 30 26 29 33 28 33 29 43 23 36 11

15 15 06

29 15 55 25 23

3 13 -7 11

-7 18

3 -7 45 22 69

24 28 35

--

21 -11 -20

4 -

-3

23 7

-2 -23 56

-

-

-__0

6 --

-

-23 14

5 0 8 2

22

-

--

0

-

--18 11

3 -10

4 10

7

--7 -

14 -

-1 6

10 -5

8 9 7

-

--

0 0 0 0 0 0 0 0 0 0 0

30 30 30

12 12 12

99 99

100 100 101 101 102 102 102 102 103 103 104 104 104

66 66 66

Farako-Ba

Mean

1977 1978

IRAT10 IRATIO

38 66

--

40 43

-5 35

21

-

10

--

--

--

- 242 243

Z

tJ

-w

24 IRPS No 123 January 1987

Philippines which had two treatments offering protection In the 9 Philippine trials where yield losses were measuredwith and without carbofuran granules relied on further by growth stage 10 of the yield loss (21) occurred at thecontrol with foliar spray and not on seed dressings The vegetative stage and 517 occurred at each of the reproshycarbofuran-protected plots had an 18yield los compared ductive and ripening stages Ibis result is similar to thewith only 10 with foliar sprayS This yield difference is pattern of yield loss in lowland rice measured by successiveprobably due to differences in protection against sown-seed treatments where the crop is unprotected at each growthpests (foliar spray do not control sown-seed pests) rather stage but proticted during the other growth stagesthan phytotonic effects became basal fertilizer wa used in However the insect complexes r sponsible for vegetativeonly 2 of the 10 comparisons stage yield losses in upland rice are entirely diftfrent than in

The recorded yield losses were not related to yield lowland rice The key vegetative stage pests in lowland ricepotential (Fig 3) One might expect higheryielding crops to arc aquatic whereas those in upland rice are sown--seedhave higher percentage of yield losses but this ws not tile (sown in soil) root and seedling pests This dift rence maycase One reason is that insect damage exacerbates plant explain why agricul torists have overlooked tile ilmportanccinjury from drought stress low yielding fields from d ought of upland rice insect pests Many reports of upland rice pestsstress would show disproportionate differences between in Asia ha Ve focused on major lowland species (46 108)insect-protected and -unprotected treatments Shoot and It isalso surprising that tei vegctative stage of upland rice root pests would be more responsible for losses from would record the highest yield losses becatuseseeral studiesdrought stress than sown-scd pests A second reason have shown that upland rice can readily recover from foliageconcerns seed pests which would have a greater impact on reinoval during that stage (117 152 165 188) Upland ricelow tillering low yielding varieties such as )aggc Low poscsses a hitgh obilitV to compensate for ca rly loss oftillering varieties nnot fill in the space created by removal foliage and will even be stinulated by foliage removal toof seeds oweer 1) R i 5 is high yielding because it tillersP1 produce a higher viel than plants ithout any foliageactively which can close the canopy to provide greater weed removed (63 192 231) [ie reason for this apparentcontrol discrepancy hovce is that yield loss at the vegetative

stage does not octUr frol leat arca loss but fron pestsYield loss () removing sown seed and feeding on devcloping tillers and roots -- damage that cannot be readily conpLtnsatCd for

70 - Measuring yield losses in upland rice cnvironrnents is further complicated by a report from Brazil that greater yield losses occurred in response to better insect control60 - =-09 as Insecticide seed treatments resuled in denser plant stands

(n=21) which in turn created a selectively more favorable environshyment for blast (69) This resuIt was parlticurla rlxexacerbated50 when cirbofuran granules were used in seed furrows Aside

ffrom a greater plant stand ca rbofuran iay have auigmented the N lecl and stimulated blast If so ca rbofuran should be40 - replaced with another insecticide or the seeding or fertilizerI rates decreased

Other methods to determine the impact of upland rice30 shy pests wvcrc to interview farners or to develop single speciesbull bull correlations of population levels and yield loss Feneira (66)2 NIM)st (75(1) Golas farmers reporteddid both in Brazil

20- insect losses in ufpland rice and 251 said those lossesS avcragcd 37(1 [his result is consistent with tie yield losses 0 0 measured by field trials in the same statc(Table4) For every

I0 4 of seedlings removC d by leaf-cutting ants a I 7yield loss is predicted Similarly Ferreira (66) found that ai

0 - average of 53 thrips per panicle before panicle initiation meant 3 times more unfillh d grains

-IO0 I 1 1 I I-101 0 2 3 4 5 6 7

Yield (tho) C)N [ROt IM FI IODS 3 Correlation between upland rice yield and yield lols caused by Upland rice insect pests may be controlled by culturalinsects measured in 21 trials in the Philippines Ivory Coast andBrazil genetic biological and chemical methods

IRPS No 123 January 1987 25

Cultural control dry season plowing will have little effect on them Plowing Crop husbandry to reduce upland insect pest populations when the soil is wet from the first monsoon rains will expose can be classified into practices effective in a single field and soil insects that tunnel close to the soil surface to predatory those effective only at the community level Cultural birds chickens dogs and even man (field crickets nole controls should come first in a pest management program crickets and white grubs a re delicacies in man tra ditional They have broad stable effects as pests have little pos- diets) sibility of overcoming them through selection of biotyps The greatest degree i1 inIscct control fron tillage comes (138) On the other hand some cultu ral controls work only from plowing soon after rice ha rvest when most soil insects with high labor or power inputs and connunity-wilt lie close to the soil surface anrd stern borers are in the rice methods require coordination of iany farniters stuble Whitegrubsare mature and tillage exposesthem to

Pantin time The luxury of being able to shift the time of predators Tirnelv control will prtec the crop following pla ntring upland rice is open onl hr la rniers in areas with a rice and lowr the populationIfor succeedinig years Plowingprolonged rainifall pa ttern lI copping intensity or underthe ice stulble helps it decompose killing sten borer rnechanied land preparatiot Gcnerally planlting as soon larvae inside as the rFiinv season begins will lower populatiois of 1110st [requent tillage destroys ait nests bit not termites insect pests Insect populationsarelo aftcra dlvseoll or Tillage and clean culture howes er rcliove food for winter fallow delayed plaiting lets them build tip on termites which then mtay attack a young rice crop Scarab alIternate hosts that gi vigorotsly with the first seasonial beetles also prefer to miposit in eceitly tilled fields Noshyratiris This first flush iof weds cros lItxuttriartlv with iceCult is dibbling faors the survival oftillace Clre Such rineralied N released during the dr season aid made stils i aels tnd insects thtt pttpate or lay eggs- in the soil

available to the plants with the rains It iliso igrtmitcally Zero tilla hio eerl conserves predators ofthe GIH (37) advanttgeous for the rice ciop to tap this trlLItlaI fertility nt it mcttoi lo lessen the impact o0 soil insects and grow vigorotsly to compete with wrus and to tolerate brotdcasting seed or planting in ftltrrows is prefc ired over pest infestation dihlirig in hills Crickets are large enotlgh to eat seedlings in

Larly plantngs terd to escape seedling maggolts Upland a hill atrd tillering normnally ca iriot fill in tlhe missing space rice in Bitangas is planted in Ma and ine itli the Iirst Oftcn dibbll rd is not well covered and iseasily found by rains Fairners are highly mtotiated to plant early tocscape foraging insets ioIdents and birds drlitght and typlotns ricar harct Arn Atrgtst planting Plan uit sii Incrcatsini sced density protects tile cropwould be evcely attacked by It riim orivzo At tti againrst seed and seelling pests especially ants particularly other Philippiie tipland sites Rea l Qteot antd (laelit if the crop will germinate and ermerge quickly M isainis Oriental seedlig mnaggots arnrige ric boe t se hr trftinll itI India rice intcrclopped witi cotton or farnes have a longer growing season and plnit 2 rin or pigeon pci had lower 6IIt and WBIlII poptrlations than mote after the ortset of the rainy season liaaiangas ftrmners rice alone (211) NfailC arid Urplald ir_ 2re coilltori in the plant ca rly becia use ofa shorter raiinfall period arnd the desi re Philippines he chatngC in pest status folii iniercropping is to lliarvet before the typhoor sea son peaks ltighly locatioll-specific (133) cmianarid the net elfc be either

Early plantings cornbind with early-iatiuring varictics nil bcnelficial ortdetriential (189) For example BIatangias provide higher crp tolerartce for white grubs Aestiatior farmers intercrip Itro in their ipland ricefields A liornshyterminates with the first rains but 5-6 moare ineeded for the nrii iotd-Ahis i (I iiiacius) which norlially feeds resting larvae to develop into beetles emerge and lay eggs on taro ill feed or miarbv rice plants Intercropping and for larvae to reach the danriigii g ird-instar stage probably would not afft0 seed pests A crluanion crop (134 In lataigas this normually iseiough tiite for harvest planted ith rice and haing gleater tolerarnce for root

It Brazil howecver delayed planting is a suggested dariage or being toxic nayact isi trapcrop to root-feeditig cultural control for the lesser cornstalk borer and insect pests Acial-fei i ig pests voultId be most affected by pests favored l drought (66 195) Cropping internsity in intrer ropping Shoot aphids leaf beetles sten borers and Brail is low and delayed plraing places the rice crop in the hopplers would be likely ca ntidates to note effcts On the niost stable rainfall period reducing likelihood old rought othlt halnd the corlipalior crop stch~ sitniaie Iiiv be more

In rilniy arca s rinmirsare roi atl to plant other crops benehited than rice from initecropping ( 137) before rice and althNgh it woultl be desirabh to avoid 11echi Weeuling luring the first month after clop infestations by planting rice cuatl lariners lack the labor cstablishent will force pcts such is anyvworris which and time to do so ol a low priority crp prefer grassy cetds ontor tle vorling rice Soil iampsecs will

77age Uplild rice Stils ae light textured and Call be mvnye to the rice crop Cleani citrltirc also lay lore territes plowed duiring the d rv seasor notl only to grin time for to attack I u cetincti inrice ceds i ay he le optinal planting with lie fitst Inilsoon riiains but also to desiccate solition 13) soil insects arid weeds such is ntsedge that foim rhizomes Irtlizr Itneatsing N Iciiliier oin hlwland rice has Htowever becta use most soil insects lie below die plow layer fAtwvored Irigher populains of pliithoppeiv Imlhoppers

26 IRPS No 123 January 1987

and leaffolde rs However because of the need to temper fertilizer use on upland rice o prevent blast fIrtilizer rates will be lower than is comonia r for lowlad rice causing relatixely less hopper and IaIfTfoide r population buildup However inupland rice areas without a history of blast such Is Batangas farmers apply high rates of N and conscqtrently the crop has high infcstation levels of WBPI]and RIF The fhrmrcrs leec insect buildup by splitting the fertili11r in0to two to three applicatiois bit do not applV fertilizer at the vegetati estage because this prodrces a tall crop which illeasily lodge during typhoons Greater crop fertility leads to greater insect survival (better nutrition) larger insects (to cat no re foliage) arid greater fec titd t (to lav nmre eggs) (128 145 171) Not all insect species respondequally hovr Increasing rates (if N l anrid K increased Dialraa s-acchralis and (iaeiurema flea beetle but reduced thrips (66) Zn rcdLuced deadheart density from the lesser cornIstalk borer

Fnldingi Although most Upland rice areas do nIot hawxcaccess to wa ter floodiig the fields is rRrrnriierdcd in areas of Brazil or root bug and white gib control ( 195)

Crop rotatioti Tillage before planting a crop after rc harvest will control stern boreis and other pests renain n the stubble arid will unearth soil pests Planting a non-gratriiaceous crop after rice is recommncrded for termite control (66) Crop rotation is less nttdcd in upland rice culture than in lowlands because atcrop-free period is assured even after harvest of crops following rice

t1ulchilt iryl plastic irli irig designed for weedcoitrol ii upland rice inJapan resulted in ntr highanri liy infestation of (ihilo sulytresvais (93 9-1 95) NIulched Upland rice grew more luxuriantlv and attracted more (Avipositing mothis than u lched rice Also the rice plant and stein borers imatoured 10rC quickly and a partialgeneratiori af vouing ista iNoverwintcred in the stubbleLotili--malitrt varieti Quickly maturing crops reduce the tmbCr of pCst generations that Calt build up Rapid crop establishment olan ca rl v-natu ring vaitvty if)pa 11icLa r will lessen white gib and stem borer damiage White grub larvae intle last iistar are highly destIruCtive so aii early-iiatri ng variety will escape serious root loss Stem borers build 1ip slowlyx in rice to ecormic abundant in a late-Maturing crop anrd caiuse whi tlieads Numibers of somespecies increase exponeitially with each succeeding genera-ti so a xarietyexhat atures I trowh earier thian another will hac less insect damage Early-naturing vaiicties however will not lessen seed pest daiage either ii the soil or oina standing crop

Strwhoi hAIJttam Inrscts tcd irig on thlie aerial portions of rice plats disperse from carly to late plantedfields AS iiiibCirs increase exponentially with eachinsect

gtscnrattm e eiglbrig f rmc~s1(uil upp wh rc generation pests build tIP where neighboring fihrTC e9r1sstagger their plantings II Batangas farmers plant uplandrice within I iro use 120-d s-arieties and plant at the start of the rainy season Ini Misarnis Oriental laricrs stagger their

plantings up to 3 mo apart and use a set of atieties that mature in 4-7 mo Insect pest danage particularly from stem borers is greater in Misarmis Oriental Also seedling maggots damage rice there because rainfall favors frequent planting of maic In Batangas rice escapes seedling maggots bCca usC of distinct drs season and synchron0us early planting with the onset of ains

Therefore combinations of cultural practices ea rlyplanting synchronous planing crop rotation and earlyshymaturing vaneties protect the rice crop against most insect pests

Ilmant resistance Beca tse upland rice has few specialized insect pests and many insect types attack the crop worldwide regional breeding and strong internatiornal cooperation are required

Sown-sced pests arc economically controlled by insectishycide seed treatment and should the relore receive lowxpriori ty inbreeding objectives

Root pests are normally cost ly to control with insecticides but finding resista t sonres mayvbe di fficult Tanaka (229) for exa -ple fi led to find varieties resistant to root aphids Selecting for large root bioniass perhaps should be the strategy for pests sLuch as Whlite grubs that renliove roots Fainter(174) howcvcr cites varietiesof crops tolerant ofor resistant t(i root pests Some sorghun varieties are high in cyanide so rice sorghun might incorporate broadshyspcctruri chemical resistance

Seedling vigor and drought tolerance are high prioritybreeding objectives so tupland rices should have large r root systernsand be able to tolerate ligherlevels of root loss Sonie root loss inolder plants is beneficial if it stimulates new root developnrut and therefore crihances intake of soil nutrients (53) The rice seedling inaggot rtiLotia spp may be

controlled thirough resistant aittics A hi rgc breeding effort on sorghniu has had fair success inideveloping arreties esrstait to Asoccata Rodani (251) A field trial comparing 10 upland ricc varieties showed differences aniong varieties va ring from II to 4(- damaged tilleis (215) Hovver resistarce to 1Iill-0r gtia species may rot cross ox-rti otier

Shiraki (210) reported that upland i-ice stens wcrc harder than lows-land rice stemsperhaps becaIusc (f a higher silicoricontent and thus were more resistant to yellow stern borer hc tortia Iow tillering (f uplaid iice xarneties prodtres a

higher percentage of infested tillers than would occor in higher tillering lowland iice under a siiilar egg density The corollary therefore is to select for high tilleriig upland rices

Stemii bor amcstheh main targes dobrecders inAfrica (571 204 )but nticl oifth e sc reen irig is dotie unde r lo wland co d t osrm s wih A h r~ o l pe i s n acorditions (11 96 218) As with lthrinrnwspecies mayhave to be dealt with independently however initial results show cross resistance between African and Asian species(92)

In Brazil screening Ior insect resistance focttses oil lou I pest problems Varieties tolerant of lasmolwalpus ligi-sellus and resistant to )iatraea saccharalis have been identified (70 153) Current work also emphasies 7iraca limativetntris and Thmialspis (=Iosis) flavouicta (66) These insect pests Ilt onlV citise IIigh yield losses tihey also ire difficult to control chemically ofr by other ilealls

ManV upland rice vaieties aic early maturing and are resistant to pests such as gall midge (I Xl ) Early mIturity is often a highly desirable trait to esuilpe pest buildup but it is not true resista nce bca usc ilthose va reties are plinted late they will be itsdamaged as ain early planted susc ible variety

P~erhaps tile cosiiiopolitall of all upland rice pestsimost are the sced pests (ripenling stagc) which pose a particular problem beta use scuds re ised IsIO for hitnit4ans Ayltoxic substatice introd tcld geticticllyilmay also be toxic to nal iniless it caln he placed old in thfe IdhuffSelectmi fo hairiness or lotng wns kill mIlct sthilrohleiiis dutilig threshing pa rtictfarl maitial threshinfl [hireshers ill complain of itcfimin atid itteascd (fust levels

Attempts itIRRI to fitid tcistatncc to the rice bug plorisa have ilotbeCnl prmitisilg lIIetIattonlids iorc

through the lenilna or palca hillhairs sedstVIor seeds wilh awns that deter bugs promote complaints from threshers Seed bugs liptocorisa and Stcnocoris enter the seed through the opening between tile lemna and palC but varieties with narrow openings are diifficult to mill

Breeding priorities should concentrate ott inotCI gruLtLIps that are most difficult to c nttrol by other meains Steil) horers appear to be the most widespread group that sigiiificantly dalag s rice aind is hard to manage by other Ilnca us

Biological control Very fe stutdic of mitatt~l eneties of rice pests in upland envirollnletits wee in tile wefouiid literature Therefore have not compiled a list of species Ve present a list of natural enemies iOtid in upland ricefields tuic latialln Batangas for some conitno iOldiar pest species ([itble 5)

Muich ofl thc lollowinle itfor ittiol cotlliCs rol Stutdies in the lowlatnds or roi crops otithcr thlin tice

Qu(ailtr o 11ll t11t( Nat trrtf lii hcoeits ight Ile used agaitist ile social insects tilt and termites Sollie predator specice live ill theirf nests n ilnic theirnllt ip-pearanIlCe aMRidbehavior IJtrouMdCing a mote aggressise ant species to displace pest ant codonies is a possibility (169) Patihogens as potential natural encnies have reccived little atteition (I13) Natural ncnuics asa group however hac not been toIl iporta it in Iegulating aintshown be and ternite numbers

The nocturnal behavior aid subteitmien habitat of lole crickets and field crickets protect them from many ntllral enemies There are only limited records of natural cnelics for these groups of pests some pathogens (87) but mainly

IRPS No 123 JattuIny I 271987

parasites - scelionids on eggsand sphecids on ilymphs and adults Sphecid wasps Larra carutmaria (SImitI hI L huzwuensis Rohwer and lsaltiuhta Willi ams speciali in mole crickets pa ralyzing them nd dragging the ninto thcir nests as food lot their voting ()tier sphCcid species Lins aurulenoa (Iabricius) lot maiac (Ashmead ) 1 lhshylssellaus (Smith) and Ml lal)oriosu (Snithf specialie in field clickets

White grUbsO the other hand hae Icti scriuns enuigh to have been extensively studied for cotttroi 1W nati al enemies( 112 140 141) Scoliid parasites C(ajsoMcispp have beetn intrtduced itto tilePhilippites for control of Ielxl ]udisiro la The rcsearchcts w Intttrdced themle during ill era claimed that the white gttionthreak as controlled (142) howevcr it isnormal for pest epidemics to subside and there was no diect evidence to indictte that tfe parasite was responsible A tmore promisitg titctllod might he to identify iso ltcttass prt~f tcartd apply hiactelaln fingal pathogens to tile soil Ihe lalattese beetle in North America has hcci coitrolled hv tite conmnleciall aailable hactcrial prcpa ration itacihlpu pav ofr milksdiscas (71) Spote prtyefott the soil itloted ilrtier reitiain actic fr Veas

Root aphid sardtnctly bugs readilslic tilib to predatots and parlsites once their tetndine ants are controlled The larvac and adtilts of a coccinellid beetle 1Scynms sp prey on llriadlouais in slash and burn upland rice Baits treated with insecticide may be used to control tending ants by killing their ourig (244)

Root-tkcditig blugs false wireoris wireorms and root wccils aic r ofsuch little ifilpoltatice that thenomttll role of liatural Cticniics hias tiot beei asscssed A possibility hoevrl tiglt he tit ItaIIitic niaCIMtodcs lintt1g11en Itese soil-itlhlfbitine mCtitodcs W5uhtfhave to be imiass Produiced

large miltibcr of niatltrll cnemics ircknown ft CltIoriS IC IpblCeii is thtt clttvortnlls ofonlie the crop soott alter land ptel)tritiOti a1tithe bcgit iniig of the raittv

Citsoln hfien tiatllral leniiiy populations aric lo Colltloil WOtuld iii ole mass prodtticing andirclcaslng key spccies

le egg stage is the fpirt (olthe Cedlitgt Ialot life cycle tI hich to f)ctis ftor tittlril elctit cottntrol hplraites o1 pleditors ttsit elleggs highly expoIsYCl I Sgg pailsites or predators stolld probahly e ittass andtbe produced released Littlh i toruialioll is a ailable ott what species to trV o ho to Iniss podLice thiCIi A etiloplhid wasp Icmihlaol sp effctiicly regushy

lates the leatf lcloalapolm2a spioanoiiitr heat andc rice antid should be collsctvcd

leaf beetles whCllose e am] eggs ate laid in the grutindhirO ould be motC difficutlt to control by niatulral enelV Mnll ipuhltiOn But species sUici IS c_gsthispid beetles ith

and larVle oilleascs ate vtilcratble to iparaites and pledators

Natural enemisc of tlirips otl rice are little known

28 IRPS No 123 January 1987

Table S Parasites recorded from upland rice insect pests in Tanauan Batangas Philippines 1977-84

Order Family Species Host stage

I lymttenoptera Braconidae Leaffolders Cnaphalocrocis Marasinia spy

Cardiochilesphilipplinensis Ashmead a Larva

Cotesia angustillasis(Gahan)Cotesia or cypris (Nixon)

Larva Larva

Cotesia or taeniaticornis(Nixon) Larva Cilonusinotakatae Munakata Chelonits SppAlacrocetrts philippinensis (Ashtmead)

Egg to larva Egg to larva Larva

Iclineumonidac Orgilts spp Isclutojppa hireator(Fabricius)

Larva Larva

Itoplectis naraigav(Ashmcad) Larva Tenelhtcha philippiensis (Ashmead) Larva Tenelchha stangli (Ashmead) a

irichomma cnaphalocrosis Uchida Larva Larva

Bethylidac Elasmidac

Goniozms or trianguli r Kieffer lasmtusalhopictus Crawford

Larva Larva

Diptera

Encyrtidac Chalcididac

Tachinidac

Elasmtus spp Copidosomopsisnacohiae (Fidy) Brach nteria evcarinata Gahan Biachyieria lasus (Walker) Ztgohothria ciliata (Wulp)

Larva Egg to larva Pupa Pupa Larva

Argyrophylx nigrotihialis(Blaranov) Larva

l-ymenoptera Braconidac Yellow stern borer Scirpopiaga incertulas

Braco chinensisSzeplegeti Larva CIthelotis spp Chelonus niunakatae Munakata Cotesi flaripes Cameron Stenoracon nicerilli (Bingham) a

Larva Egg to larva Larva Larva

Ichneumonidac Tropobracon schoenobii (Viereck) Amatroinorpiaaccepta metathoracica(Ashmiead)Frihorus sinicus (I lolmgren)

Larva Larva Larva

Ischnojoppaluteator (Fabricius) Larva Isotima nr dammeratiti Rohwer Larva Tenelucha philippinensis (Ashmead) a

Larva Tenehicha stangli (Ashmcad)Trichomma enaphalocrosis Uchida

Larva Lara

Chalcididac Eulophidac Trichogranmmatidae

Xanthopimnpla stemmator (Thunberg) Charops hrachYpterumn Gupta and Malteswary Brachtm eria sppTetrasticttus schoenohii Ferriere Trichorigranm chihunis Ishi

Larva to pupa Larva Pupa Egg Egg

Diptera

Scelionidac Pteromalidae Tachinidae

Tt ichograintnta aponiicuitt (Ashniead) Telenomits rowani (Gahan) a

Trichoinalopsis apattelctena (Crawford) Jeirhaea spp

Egg Fgg Egg pupa Larva

Z4gobothria ciliata (Wulp) Larva

Ilymenoptera lecneumonirlac Gol fringed borer (hilo auricilius

Trichomm enoptalocrosisUchida Larva

Ieioenoptera Braconidae

Aantholimlh steiniator (Thtinberg) a

Pink stet Ioret Sesamia inferens StoInacoil nicrilhi (Bingh am)

Loarva to pupa

larva

Diptera Ilyinenoptera

Armvtnyorms Mythimna separata Spodoptera mauritia acronyetoidesTachinidae Zvgohothriaatropiora( Robinea i -Desvoid y)Eulophidae It-phctrtiscitapadoe (Ashimead)

Larva Larva

(halcididae Braconidac

Brachpin ria spip (oesia spp

Larva larva

IHyntenoptera I)iptera

Rice skipper Ielop ida (halcididae Tachinidae

math ias aidgreen iortedci ciliarNIelan itis eda Brat lhincria sp iii matuqitatt Cameron Irgyrolhirlaxi trotlhiais(marumtv)a1

ismtne Larva Larva

Diptera Ilymenoptera

Tachnidae lelineutionidae

fuown se tilo per Mois frugalisIrgtrophrlax 1igrothiahiv 3 a rat ov) anthopimpa pmnctata (Iabricius)

Larva Larva to pupa

aMost dominant

IRPS No 123 January 1987 29

Normally predatory thrips prey on herbivorous thrips strepsiptera on both Spiders dominate tilenymphaladultNatural enemies of stem bugs --Scolinoplara and Tibraa predators Spittle bugs move to rice from pasture grasses - are in order of importance egg par~isites and predators and biocontrol would be successful only ifcarried out before fungal pathogens and nymphal adult parasites Egg tiley immigrate into ricefields parasites could either be introduced fron other areas or Biological control tactics for hemipterous seed bugs mass produced and released ould follow those outlined for stein bugs

Armyworms are normally lwld in hcck by the activities Coml)arison of environnewtns Natural enemies are of egg and larval parasites When these natural enemies fail perhaps faced with even greaterconstraints than insect pestsusually because of drought armyworns become epidemic to survive ind reproduce in upland environments TheyIt may be worthwhile therefore to release parasites during have adapted ani survived by having wide host ranges the the rainy months following a drought Virus diseases of ability to aestivate or hibernate over unfavorable seasonslarvae would beanotheravnue to exploreforarnyworms and the ability to disperse or by other mechanisms

Parasites have ben traditionallyconsidered for biolegical Comparing the most dominant and therefore the most control attempts against stein borers usually by rekasing adapted species of natural enemies collected in various exotic species( 166) This approach has not met with success environments in the Philippines we can understand how and new avenues should be explored The role of predators different rice environments are arid pathogens is little inderstood and deserves greater Anagqrms olptahiisa mymarid planthopper egg parasiteattention Egg parasitism rates are normally reasonably is cquallyadapted to all Philippine rice environments (Tablehigh but should be supplemented with effective egg 6) It is selective for planthoppers and even parasitizes eggspredators Orthopterans are potential egg predators of the n-iize planthopper Peregrinitsmaidis itt upland areas Metioche vittaticollis (Sta ) Anavipha nifiwnnis (Saus- Alavohis also parasitizes maize planthopper eggs but is sure) and Conocephahs hongipnni (de i-laan) feed on most prevalent on BPH in the irrigated wetlands (onashystem borer eggs etioche and Anaxil)ha specialize in eggs tocerus spp like A optalhilis on BPH and WBPI Iattacks with no hair such as Chilo spp The omnivorous (ono- the eggs of GLH in ell rie environments It is also found cephahsspecializes in eggs cov red with hairs it is a proven parasitizing the eggs ef the white rice licahopper (oimna egg predator of Scirlouphaga spp but has the discouraging spetra Oirosita naias a trichogrammatid egg parasitehabit of eating rice guin (65) attacks BIPH and WBPH in the wetlands and has no other

Grasshoppers have egg (scelionid) and nymphalad ult host A related species 0 aesopi specializes in G1-H in (nemestrinid tachinid and sarcophagid) v irasites Among wetland environments Dryinid strepsipt-an and pipunshypathogens protozoans have been recorded most frequently culid nymphal and adult parasites of planthoppers and Control of locust species by natural enemies could con- leafhoppers also specialize either in hopper species or centrate on their habitual breeding areas environments Among the dryinids Pseuhdgonatoipsflavi-

Leaffolders have rich complexes of natural enemies some feimur prefers the lowlands and BPH while P nuch has no adapted to upland rice (22 27) ranging from egg predators environmental prefirences but is adapted only to WBPH (gryllids coccinellids) to larval parasites (braconids ichncu- The main d ryinid species on GLH aptogonatop)usspp is monids) to larval prcdators (ants and carabids) to larval most dominant in rainfed lowlands and also attacks other pathogeiis (vira and fungal diseases) Efforts to augment leafhoppers within Amnrasca Cicadulinaand Bachaltuathese natural enemies should focus on the egg predators Strepsiptera are more dominant on plarithoppers than (rearing releasing)and viral or fungal pathogens (culturing lealhoppers The Elenchus species in the Philippines attack spraying) all species of planthoppers within the genera Sogawilla

Parasitization of the large conspicuous larvae of Sogatodels NilaparsataHarualia and Opiconsiva Elenshyskippers green horned caterpillars and other polyphagous cusrvasuntaisuiis adapted to the major rice environments Lepidoptera by tachinids and chalcids is normally low ELg but prefers BPH in rainfed arid WBPIH in wetland environshyparasites and predators are perhaps the key to their control ments Ilalictoph(utsmtnroeiand HVpectrus attack G LH

Gall midges are parasitized by pterormalids and platy- and COfimravcctra respectivelygasterids that aestivate within their larval host between Pipunculids are mainly parasitic oii lealhoppers Each seasons Shoot aphids are preyed upon by coccinellids species has a unique environmental preference Tomosshysyrphids and chrysopids varireIa oraetoraprefers Ne7 otetltix nigropictrs and

Planthoppers arid leafhoppers have egg parasites and Pilunwulhtsjavanensisattacks Dehoceplhahls spp which predators as well as nymphialad nlt parasitesand predators are more abundant in upland than irrigated lowland their rich complex of natural enemies includes fungal environments T suhvirescensattacks Nephotettix virescens pathogens The principal egg parasites are myrnarids and and prefers the lowlands Pilpuiulus nttillattls is an trichogramniatids Egg predators are norinally mirid plant upland rice parasite arid attacks only Nephotetix speciesbugs that suck out the yolk Nymphal ad nIt parasites are More than 17 recorded species of leaffolder parasites in dryinids on planthoppers pipunculids on lealhoppers and the Philippines attack species of C(naplwocrocis and

30 IRPS No 123 January 1987

Table 6 Dominant natural enemies of rice pests in upland rainfed lowland and irrigated lowland rice environments of the Philippines

Natural enemy

Host stage Species

Brown planthop-Parasite Egg

MymaridaeAnagrus optabilis(Perkins) Anagrusflaveolus Waterhouse

TrichogrammatidacOligosita naiasGirault

NymphAdult Dryinidac

Pseudogotatopuslaiifemttr Esaki et Hashimoto Elenchidae

Elenchus yasurnatsui Kifune and Hirashima

Environment

Upand Rainfed lowland Irrigated lowland

Ailaparata lugens

x x X x

x

x x

x X X Whitebacked planthoppers Sogatella furcifera and Sogatodes pusanusEgg

MymaridaeAnagrus optabilis(Perkins)

TrichogrammatidacOl~gositanaias Girault

NymphAdult Dryinidae

Pseudogonatopusnudus Perkins Elenchidae

Elenchus yasutatsui Kifune et Hirashima

Parasite Egg

MymaridaeGonatocerus spp

TrichogrammatidaeOligositaaesopi Girault

NymphAdult Pipunculidac

Tomosvaryella subviresccns Loew Tomosvaryella oryzaetora (Koizumi) Pipunclusmutillatus (de Meijere)

DryinidaeIlaplogonatopusspp

Egg Copidosomopsisnacoleiae (Eady)

Larva BraLonidae

Cardiochiles philipphnensisAshnead lchneumonidac

Trichiomnia cnaphalocrosisUchida Temelucha stangli (Ashmead)

Pupa Chalcididae

BrachymeriaexcarinataGahan Brachymerialasus (Walker) Brachymeria nr tachardiaegroup

Parasite Egg

Scelionidae Telenomus rowani (Gahan)

EulophidacTettasticusschoenobhi Ferriere

Larva Braconidae

Stenobraconnicevillei (Bingham) Iclhncurnonidac

Temelucha phiiippinensis(Ashinead)Temelucha stangli (Ashmead)

x x x

x x

X x X

x x Green leafhopper Nephotettix virescens

X x x

x x

X X X X

X

X Rice leaffolders CnaphalocrocisMarasmia

x X x

X

X X

x x x x x X x x x

Yellow stem borer Scirpophaga incertulas

X X x

x

x

x X XAinauromorphaaccepta metathoracica(Ashmead) x

continued on next page

IRPS No 123 January 1987 31

Table 6 cntinued

Natural enemy Environment

Host stage Species Upland Rainfed lowland Irrigated lowland

Larva-pupa Gold fringed borer Chilo attriciiuslchneumonidae

Xanthopimpla stemmator (Thunberg) x aPupa x

Pteromalidae Trichoinalopsis apanteloctena (Crawford) ax

Pink stem borerb Sesamnia inferensParasite

Larva Braconidae

Stenobracon nicevillei (Bingham) x b b Armyworms Mvthinna SpodopteraLarva

Tachinidac Zygobothriaarroptvora (Robineau-Desvoidy) x x

Braconidac Cotesia spp

X Skipper Pelopidas mathias and green horned caterpillar Melanitis leda ismeneF-gg

TrichogramrnatidacTrichogrannaspp x

Scelionidae Undetermined Scelionid

x Larva

Tachinidae Argyroph lax nitrotibialis (Baranov) X x

Pteromalidae Trichomnalopsis apanteloctena (Crawford)

X Predator

CoenagrionidaeAgriocnenis femina femina Brauer x xIschnura senegalensis tambur

Miridae CrtorhinsliidipennisReuter x x

Nabidac Stcnonabis tagalicus (Stal) x

VeliidaeAlicrovelia douglasi atrolineata (Bergroth) x

Coccinellidae Micraspis crocea (Mulsant) X x x

Vespidae Eumenes campaniformis (Fabricius) x

Formicidac Diaecamna spp xOdontoponera transversa (F Smith) x

Carabidae Chaenius spp OphioneaW- aJasciataSchmidt-Goebel

x

Ophionea ishii ishii Habu x x X

Gryllidav Anaxipta longipennis (Saussur-) x xMetioche ittaticollis (Sial) x x x

Tettigoniidae Conocephalus longipennis (de Haan) x x x

aNo rearing made on rainfed lowland bNo rearing made on rainfed lowland or Irrigated lowland

MrasmiaAmong the Laval parasites the braconid Cardio- lowlands and parasitizes oriental maize borer OsIriniachilesphilijpphensis is most adapted to an upland environ- furnacalis (Guenee) larvae The leaffolder pupal parasitesment It also parasitizes Hydelepta indicata a common not only occur equally in all environments but also haveleaffolder of legumes The ichneumonids Trichomma cna- wvide host ranges- parasitizing species of OsIrinia Marucaphalocrosis and Temehcha stangli occur more in the Hydelepta and Honiona - in maize and legumeslowlands T stangli is most adapted to irrigated lowlands The principal egg parasite of yeltow stem borer (YSB)and also parasitizes Chilo and Scirpophaga stem borer Telenomus rowani is widely adapted to all environmentslarvae T cnaphalocrosis is most prevalent in rainfed and has an unusual alternate host -Tabanus eggs (25)

32 IRPS No 123 January 1987

Tetrastictu schoenobii specializes only in YSB and occurs mainly in irrigated lowlands -- further proof that YSB is most adapted to the wetter environments

Larval parasites of YSB are highly environment-specific The braconid Stenohracon nicevillei occurs only in upland rice but also attacks the pink stem borer (PSB) Tieinchwa phiiiplPwnsis an ichneumonid occurs in upland rice and also parasitizes the larvae of (7hiu stqpresaisinatro-morpha accepta melaihoracica like 7 stangi is mnost adapted to the irrigated lowlands

The dark-headed stem borer(l)HSB) (ih pochit3sus was not collected in all rice environments and the gold fringed borer Chilo auricilius nd PSB were collected only in upland rice Xaantiholiplastenrnaloran ichneumonid larval parasite appears to be widely adapted whereas TrihomaslopisapanitI)cteiiaa pteromalid pupal parasite occurs mostly in wetlands and also parasitizes larvae of Chilo suppressais and Pelhuidas mathias

Armyworms Myit hinin and Spodujipera have two main larval parasites - a tachinid Zrgohothriaatropivora in rainfed environments and a braconid Cotesia spp in irrigated lowlands

The butterfly pests Pelopidas and 4elanitishave tricho-grammatid egg parasites found only in irrigated lowland environments

Two larval parasites show environmental preferences --Argrroph av nigrotihialis a tachinid in rainfed habitats and Trichonialopsisapantelkcnela in irrigated ricefields A limgrotibialisalsoattacks the sweet potato hornwormAgrius convolvtdi and T apanteloctena parasitizes DH S1B

Predators as a group show more distinct environmental preferences The aquatic damselflies Coenagrionidae are most abundant in the lowlands Cvriorhinus is also most adapted to the lowlands The nabid Stenonabis tagalicus occurs only in the uplands and isalso prevalent on legumes The coccinellid lady beetles wasps and ants are more prevalent in upland areas AlicrasIpis cit-ea larvae prey ona variety of aphids attacking legumes and maize Etunienes campanonia vespid mud wasp makes nests in trees and is therefore most adapted to the more botanically diverse uplands Soil-dwelling ants cannot tolerate flooding

Upland environments are habitats to arboreal carabid beetles Three species of Chlaeniusbeetles prey on leaffolder larvae and are perhaps more important than parasites (27)

The gryllids and tettigoniids are widely adapted egg predators not only on rice but also on maize and legumes They feed on eggs of most insects that are laid on leaves

The spidercommunity of upland rice environments isrich in species At one site (Batangas Philippines) 31 species have been recorded (23) Of the 176 spider species recorded in Philippine ricefields about one-half (82) occur in upland rice The spiderspecies of upland rice environments overlap more with those of rainfed lowland than with those of irrigated lowland rice (24) The same study showed that of the three environments irrigated rice has (ie greatest spider species diversity foiiowed by rainfed lowland and upland

However of the 10 most prevalent ricefield spiders 9 were abundant in upland rice showing wide adaptation All three spider guilds -- orb-web space-web and huting spiders shywere prevalent in upland ricefields There were however differences in environmental prelerences for some spider species

Of the orb-wcavers Tetragniathamanlihulatawas partishycila rlv abundant 7 japonica was low and Ltucatge dehcorata was absent in upland rice

As in other environments Aiypeiwnaifirtosana was the most dominant space-web spider however its relative numbers were lowerthan for lowland sites Among hunting spiders Ox|opes javanus and Lcosa leatcost fgla were more abundant and Lvcosa lpseudoanmnlata was less abundant in upland compared with lowland rice envishyroniments

Extensive rearing of rice insect pests in upland rainfed lowland and irrigated sites over an 8-yr period in the Philippines provided insight into the effectiveness of parasites as natural enemies (Table 7 8) Most of toe data are from large sample sizes taken over at least one crop which overcomes the pitfalls expressed by Van Driesche (238)

VSB egg parasitization was surprisingly similar across all environments Larval parasitization was highly dynamic among sites and even years within the same site but showed somewhat lowerlevels in uplandsthan in irrigated wetlands This would have been predicted for egg parasites as well since YSB is adapted to the lowlands and so must be its parasites This isevidence that Telenomus rowani is as good at dispersing as its main hosts Trowani clings onto the anal tufts of female YSB and parasitizes eggs as they are laid This phoresy may explain its wide adaptation The more specialized lowland YSB larval parasites therefore appear more effective than their upland counterparts with more alternate hosts

Low collection levels of PSB larvae especially in the wetlands make comparisons between environments difficult Low levels of larval parasifization were recorded in both rainfed environments Except in the Cagayan rainfed wetland site in 1980 low levels ef larval parasitization were evident at irrigated and rainfed sitesalike No differences by environment were evident Higher levels of parasitization occurred in Sarawak during a 1967 outbreak also from tachinids Rothschild (197) indicated that higher parasishytization occurred in upland rice areas

Skipper and green horned caterpillar larval parasitization rates were generally low and showed no environmental effect htighest activity occurred in Cagayan in 1980 (48 parasitization) Although larval parasitization levels of RLF were variable year to year activity appeared greater in the irrigated lowlands than in the uplands Again the highest levels were recorded in Cagayan in 1980 Cagayan has extensive grasslands and fallow around the rice areas and a relatively long rainfall period Rainfall in 1930 was good for crop growth and apparently for increase of larval parasites

IRPS No 123 January 1987 33

Table 7 Parasitization rates of rice planthoppers and green leafhopper in upland rainfed lowland and irrigated lowland environments Philipshypines 1976-840

Environment and site Year Egg parasitization () Nymphaladult parasitization ()

BPH W13P GLH BPII WBPH GLIH

UplandBatangas

Rainfed lowland

1976 1979 1980 1982

15 17 12

2(47)8(39)

4(51)

9(1800)14 (212)

12(304)

8 (101)6 (127)

9(116)

Iloilo

Cagayan

1976 1977 1978 1981 1980 1981 1982

0 (69)

11

44 48 40

-

-

46 55 44

0 (471)

11

52 56 49

0 (200)9 (214)

10 (1259)8(235)6 (900) 3(800)

-7 (462)

11 (270)3 (313)5 (900) 1 (800)

4 (600)10 (725)10 (362)12(870)5 (900) 3 (800)

Pangasinan

Irrigatedlowland

1984 1976 1978 1979 1982

13 (436) 13(379) 14 (457)

6(14)7 (197)

17(198) 11 (486) 18 (275)

3 (81)10 (274)3 (116)

16(871) 19(88)

28(211)9 (764)5 (647)7 (1204) 8(305)

Laguna 1977 11 7 21 1978 31 1979 14 4 4 28 1980 7 8 20 1981 1983

56 7(278)

26 16 (1650)

16 27(1921)

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R I Apostol unpublished 1lo1lo and Pangasinan 136 Cagayan BCanapi unpublished Liuna 175 I C Pantua unpublished 118 234 43 Carino and Shepard unpublished BPH = brown planthopper Nila parvara htgens W131Pi =whitebacked planthopper Sogatelia furcifera GLH = green leafhopper Nephotettrix virescens Figiires in parentheses are sample sizes

Tauher et al (230) reported that internal external and genetic factors influence the seasonal activity of parasitoids

Hopper egg parasites but not nymphaladult parasites were also parlicularly abundant at the Cagayan site possibly the egg parasites have more alternative hopper prey in the weedy areas than nymphaladult paisites

The seasonal dynamics of hopper egg parasites can be seen for upland rice in Batangas(Fig I 2) and for rainfed lowland lice in Pangasinan( 135)and Cagayan( 105) Levels of BPHWBPI-L and GLH egg parasitization were corn-parable in upland and irrigated lowland environments Except for some high rates of paraMsitization early in the season the parasitization rate tended to be steady in all envilonments However hopper nymphaladult parasi-tizution levels were more similar in the rainfed environments and lower than in irrigated rice

Overall parasites do not appear highly effective by themselves against upland or even lowland pests Among natural enemies parasites have attracted the most attention because they can be more readily assesscd

Predators and pathogens lend themselves better to management practices Predators the most important group of natural enemies in riccfields are difficult to quantify On the other hand pathogens appear less im-portant but can be readily cultured and disseminated

The natural enemy commulnity of upland rice is rich in species and differs significantly from that of lowland rice

These beneficial organisms must be conserved by applying insecticides judiciously particularly sprays Seed or soil placement of chemicals minimizes exposure to natural enemies The strategy to derive the greatest benefit from natural enemies isto allow the greatest number of beneficial species to thrive Then perhaps at least one species will be effective against each pest at any tine overcoming the variable abundance of each species within and between years

Programs to introduce exotic species or mass produce indigenous ones are ambitious and expensive because they require trained people on a sustained not ad hoc basis

Chemical control Insecticides offer rapid and efficient control of upland insect pests (190) but-should be used only after other control measures have been considered Insecticides are rarely used on upland rice because of cost (32) Upland rice yields are normally too low to justify the expenditure Also most upland rice is grown as a cash crop Spraying upland rice is more difficult than lowland rice because water is less accessible Government subsidy programs to provide pestishycides to tribal peoples for upland rice production have created problems of toxicity to humans because isolated people have had no experience to be able to handle pesticides safely

However ins-eticide use has beenjustified when a crop is

0Z

Table 8 Parasitization rates of lepidopterous pests in upland rainfed lowland and irrigated lowland environments Philippines 1976-84a

Parasitization (5)

Environment and site Year YSB GFB PSB Earcutting Swarming Skipper Greenborned RLF Brown larva larva caterpillar caterpillar caterpillar caterpillar larva semilooperEgg Larva larva larva larva larva larva

UplandBatangas 1977 - 0 (57) - 0 (63) - 0 (71) - 0 (106) 11(142) 2 (101)1978 43 (4697) 8(451) 5 (38) 3 (40) 6 (61) 2(58) 15 (57) 13 (129) 13 (1286) 4 (30)1979 59(4135) 12(320) 13 (11) 5(28) 9(45) 6(46) 7(23) 8(160) 9(1152) 8(26)

Rainfed lowland Iloilo 1976 - 0 (10) 0(10) 0 (7) 0 (19) 2 13) 15 (180) 5 (76)1978 0 (54) 0 (3) 0 (17) 9 (64) 4 (78) 3 (73) 0 (121) 0 (191)1979 64 (2350) 16 (84) 6 (17) 0(6) 18 (68) 25 (104)

1981 56 (1322) 18(304) 0(5) 12(51) 18(138)Pangasinan 1976 60 (7 7 )b 62(82) 0(111) 5 (39)

1978 58 (53)b 5(51) Cagayan 1979 59(713) 8 (101) 6 (34) 0(7)

1980 0(11) 4 (28) 15 (286)

62 ( 48(177) 61(140)1981 0(5) 12 (2) 13(45) 14 (92)

Irrigatedlowland Laguna 1974 62 8

1974-75 50 41 1975 43 72 1978 96 (623) 21(24) 33 (60)1979 83(2017) 18(110) 8(62) 11 (89) 22(400)1980 78 (1625) 20(75) 0 (16) 4 (72)1983 60

9 (80) 7 (270) 30

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R F Apostol unpublished Iloilo and Pangasnan 136 Cagayan B Canapi unpublished Laguna 175 P CPantua unpublished 118 234 143 Carino and Shepard unpublished YSB = yellow stem borer ScirpophagaincertulasGFB = gold fringed borer Chilo auricilius PSB = pink stem borerSesamia inferens earcutting caterpillar Alythimna separata swarming caterpillar Spcdoptera mauritia acronyctoides skipper Pelopidas mathias reen horned caterpillar Melanitls ledaismene RLF = rice leaffoldersCnaphalocrocismedinalis and Marasmiaspp brown semilooper Mocisfrugalis Figures in parentheses are sample sizes Egg masses

IRPS No 123 January 1987 35

Table 9 Effect of seeding rate and insecticide on plant density and Yield losses from insect pests in upland rice often areyield of Dagge upland rice Batangas Philippines 1976a comparable to those of lowland rainfed or irrigated rice

Seeding rate Insecticide Dosage Plant density Yield (0-Y30o) but usually from different pests The principal (kg aiha) seed treatment (kg aiha) (noIra-row) (tha) groups are soil pests feeding on sown seed and rootsat 14 DE followed by seedling pests These groups have largely gone

50 None - 35 a 20 a unnoticed leading researchers to conclude that stem borers50 Carbofuran F 10 38 ab 23 a and seed bugs are the most important pests (17) Yield loss 10 beCabofuranF100 None - 4551 c 23 a trials that pinpoint major pests need to includc seed

100 Dieldrin WP 10 52 cd 22 a treatments Granules applied in the furrow may not give150 None - 59 d 22 a adequate protection from many seed pests

aAv of 8 fields DE = days after crop emergence In a column means The floristic and pedological diversity of upland rice followed by a common letter are not significantly different at the environments is matched by the diversity of arthropods5level by D)MRT F = flowable Wi = wettable powder both pests and their natural enemies Irrigation has homoshy

genized lowland rice environments Flooded soils tend tothreatened by epidemics of locusts or armyworms New have the same properties and to eliminate soil pests The technology stressing efficiency has minimized insecticide spatial and temporal dominance of cultivated rice has cost benefited specialist pests

Seed treatment is normally inexpensive and can be Upland rice has no specialist arthropod species The economicallyjustified in -nanycases(66) Treating seed with monophagous pests on upland rice prefer the lowlands systemic insecticide (216) but not chlorinated hydrocarbons where rice evolved Rice in the uplands exists only because (247) is most effective in controlling seedling maggots of man and even though lowland rice predates upland riceFoliar sprays have to be repeated to obtain control and specialist species evolved Wheat maize and sorghum do often are washed off by frequent early season rains not have monophagous pests leading us to conclude that Granular insecticides are effective but required dosages are there is no evolutionary advantage for inssects to have too expensive to justify except under heavy infestation narrow host ranges in upland graminaceae(247) When directed against first-instar larvae banding Pests have adapted to the uplands not only by having granules in seed furrows (177) can be effective against white wider plant host ranges but also by having long life cyclesgrubs at low dosages (025 kg aii ha) (134) Seed treatments undergoing dormancy or being dispersive Each pest has a are not promising for white grub control (248) unique life history strategy suited to highly unpredictable

Seed treatment is also effective ainst ants and may be environments In agivenyear someinsect specics an better cheaper than increasing the seeding rate In Batangas exploit upland rice as a food source but no pest is a senious treating 50 kg seed ha was as effective as increasing the one every year on upland rice seeding rate to 100 kgha and using untreated seeds (Table The low yielding potential of upland rice means less 9) In Brazil seed treatments control termites and other soil quantity and lower quality of food for pests to exploit but insects including the lessercornstalk borer(66) Domiciano also less incentive for farmers to undertake control (58) however found seed treatments inconsistent in their The most effective control would be early and synchroshyeffect against high populations of lesser corn stalk borer nous planting of early-maturing upland rice varieties to

Baiting can be an inexpensive way of controlling ants prevent insect pest buildup Breeding for insect resistance seedling maggot flies mole crickets and field crickets Baits should initially focus on stem borers because they are verymade of locally available material and impregnated with difficult to control by other means Biocontrol efforts insecticide can be sparingly distributed taking adantage of should first stress conservation of natural enemies preshythe pests ability to disperse and encounter bait sites dators and pathogens rather than parasites Efficient

The new low-volume hand-held controlled-droplet chemical control includes seed treatment or baiting for seed sprayers may offer an advantage over high-volume knap- seedling and some root pests and using controlled dropletsack sprayes for controlling foliar pests Upland rice sprayers for spot treatments against foliar pests and seed canopies are more open than those in lowland rice and bugs when economic thresholds are reached better droplet penetration should result Controlled droplet applicators would be ideal for seedbugs and defoliators

CONCLUSION

Upland rice is attacked by a wider array of insect pests than is lowland rice mainly beca use of the addition of soil pests Generally population levels of lowland rice pests are lower in upland than lowland rice leading many people to conclude that insect pests are not important on upland rice

36 IRPS No 123 January 1987

REFERENCES CITED

1Abraham C C and K S R Mony 1978 Occurrence ofLeptocorisa acuta Fabr (Coreidae Hemiptera) as a pest of nutmeg trees J Bombay Nat ttistSoc 74553

2 Agyen-Sampong M 1975 HlierogrlphusdiaganensisKrauss (Orthop-tera Acrididae) a new pest of rice in Northern Ghana Ghana J Agric Sci 8249-253

3 Agyen-Samnpong M 1976 Varietal resistance and other inle-ratcd pest control components as factors of rice varietal improvement in West Africa Prcsented at the Second varietal improvement serinar 14-18 Sep 1976 Bouak Ivory Coast West Africa Rice Development Association Monrovia Liberia 18 p

4 Agyen-Sampong M 1977 Outbreak of Ihccrorrdnisorae in rice seedlings near Rokupr Sierra Leone lit Rice Res Newsl 2(6)16-17

5 Agyen-Sampong M 1980 Dirty panicles and rice yield reduction caused by hugs IntRice Res Newsl 5(l)11-12

6 Agyen-Sampong M 1982 The major pest problems of irrigated upland and mangrove swanip rice ecosystems ir the humid tropical Guinea savanna and Salel climate zones Pages 318-328 in Pro-ceedings training course cn concepts techniques and application of irtegrated pest management in rice inWest Africa 10-20 Jan 1982 Fendall Liberia West Africa Rice l)evelopioent Association Monrovia Liberia 505 p

7 Akibo-Betts V I and S A Raymundo 1978 Aphids as rice pests in Sierra Leone lit Rice Res Newsl 3(6)15-16

8 Akinsola F A 1975 Present status of different rice stemborers in parts of Nigeria iicL Entomo Newsl 328

9 Akinsola FA 1979 Ihe biology and ecology of rice stemborers in Nigeria Ph 1) thesis University ol lbadan Nigeria 217 p

10 Akinsola E A 1979 Guidelines of integrated management of insect pests of upland rice in West Africa Presented at the Seminar on integrated management of rice diseases and insect pests at Bobo-Dioulasso Upper Volta 17-22 Sep 1979 West Africa Rice Develop-ment Association Monrovia Liberia

II Akinsola IA 1984 Insect pests of upland rice in Africa Pages 301-305 in An overview of upland rice research Proceedings of the 1982 lIouak Iwsrv Coast upland rice workshop International Rice Research Institute 13O Box 933 Manila Philippines

12 Alam M S V TJohn and Kaung Zan 1985 Insect pests and diseases of rice in Africa Pages 67-82 in Rice improvement in Eastern Central and Southern Africa proceedings of the international rice workshop at Iusaka Zambia 9-19 April 1984

13 Alticri M A A van Schoonhovcn and J D Doll 1977 The ecological role of weeds in insect pest management systems a review illustiated with bean (Phaseohsvulgaris L)cropping systems PANS (Pest Artie News Summ) 23195-205

14 Anmante F 1967 The leaf-cutting ant Atia capiguaraa pest of pastures [in Portuguese] Biol6gico (Sao Paulo) 33113-120 (Rev Appl Entomol A56207)

15Andrade A C and D Puzzi 1951 Preliminary results of experi-merits on the control of Sculptcoris iastantus on sugarcane [in Portuguese] Bioulgico (Sao Paulo) 1744-49 (Rev Appl Entomol A41421422)

16 Appert J 1970 Maliarplia separaelha (rice stem borer) new observations of entomological problems of rice in Madagascar [in French] Agron Trop (Paris) 25329-334

17 Ariadeau Mand Z Harahap 1986 Relevant upland rice breeding objectives Pages 189-197 in Progress in upland rice research Proceedings of the 1985 Jakarta conference International Rice Research Institute 1O Box 933 Manila Philippines

18Asar S and Kisimoto 1975 Seasonal occurrence of the smaller brown planthopper in upland rice [in Japanese] Proc Kanto-Tosan Plant Prot Soc 2228

19 Bakker W 1974 Characterization and ecological aspects of rice yellow mottle virus in Kenya Meded Landbouwhogesh Wage-ningen 152 p

20 Balut F F 1970 White grubs attacking rice Oryza ativa L [in Portuguese] Biol6gico (So Paulo) 36321-322

21 Banerjee S N and L M Pramanik 1967 The lepidopterous stalk borers of rice and their life cycles in the tropics pages 103-124 in The major insect pests ofthe rice plant Proceedings ofa symposium at the International Rice Research Institute September 1964 The Johns Hopkins Press Baltimore Maryland USA

22 Barrion A T and J A Litsinger 1980 Ants a natural enemy of cliaiarac in dryland rice IntRice Res Newsl

5(4)22-23 23 Barrion A T and J A litsinger 1981 The spider fauna of

Philippine rice agroecosystem 1Dryland Philipp Entomnol 5139-166

24 Barrion A T and 1 A Iitsinger 1984 The spider fauna of Philippine rice agroccosystcms II Welland Philipp Eiitomol 611-37

25 Barrion A T1 and J A Litsinger 1984 lihanus (Diptera Tabanidae) eggs an alternative host of rice stei borer (SI3) egg parasite Telenoonis digns (Iliymenoptcra Scelionidae) lit Rice Res Newvsl 9(6)19

26 Barrion A T and J A Litsinger 1985 Identilication of rice leaffolders by wing markings Int Rice Res Newsl 10( 1)24

27 Barrion A T and J A Litsinger 1985 Chctiosspp (Coleoptera Carabidae) a leaffolder ILF)predator lit Rice Res Newsl 10(1)21

28 Bertels A 1970 Insect pests of rice and theircontrol [in Portuguese] Pelotas Brazil Instituto de Expcrimentacao Agropccuararias do Sul Circular 43 24 p

29 Bonzi S M 1982 Ctilo (flisilineus J de Joannis (Lepidoptera Pyralidae) stem borer of irrigated cereals in Upper Volta LioFrench] Agron Trop Riu Rizic Cult Vivrieres Trop 37207-209

30 Bourke T V T L Fenner J N LStihick G Baker EHassan D F OSullivan and C S li 1977 Insect pest survey for the year ending 30th June 1969 Papta New Guinea )epartment of Agrishyculture Stock and Fisheries Port Moresby 57 p

31 Bowden 1 1973 Migration of pests in the tropics Meded Rijksfac Landbouwwet Gent 38785-796

32 Breniere J 1973 The main rice parasites in West Africa and their control Pages 27-42 in Proceedings seminar on plant protection for the rice crop West Africa Rice Development Association Monrovia Liberia

33 Breniere J 1976 The principal insect pests of rice in West Africa and their control West Africa Rice Development Association Monrovia Liberia 52 p

34 Brown ES 1962 The African arinyworn Spoiolptera eXI)Ina (Walker) (Lepidoptera Noctuidae) a review of the literature Cornshymonwealth Institute of Entomology London UK57 p

35 Brown F B 1968 Upland rice in Latin America FAO-International Rice Commission 12th Session of the Working Party on Rice Production and Protection 2-14 Sept Peradeniya Sri lanka Food and Agriculture Organization Rome Italy 6 p

36 Buyckx F J 1 1962 Disease and insect pests of crops inCongo Rwanda and Burundi [iil French] Publications dc LInstilut National pour lestude Agrononiquedu Congo M Weissenbruck S A Brussels Belgium 708 p

37 Cariiio F 0 1981 Role of natural enentics inpopulation suppression and pest management of green rice lealhoppers Ph D thesis University ofthe Philippines at Los Bafitos PhilippinesLaguna 136 -

38 Carpente A J 1983 The development of rice cultivation and extension of vegetable productin in Zanzibar Final Report Vol II1 October 1983 Food and Agricultme Organization Ronte Italy

39 Carroll C P and S1 Risch 1983 Tropical annual cropping systems ant ecology Environ Manage 751-57

4(0 Cates R G 198 1 Host plant predictability and the feeding patterns of monophagous oligophagous and polyphagous insect herbivores Oecologia (Berl) 48319-326

IRPS No 123 January 1987 37

41 Cendcafia S M and F B Caora 1967 Insect pests of rice in the 62 Ene 1 C 1981 I)iiorldhol)ierusihilis Slater (licteroplera Philippines Pages 591-616 in The najor insect pests ofthe rice plant lygacidue Illissinac) a new rice pest front Nigeria lit Rice Res Proceedings of a symposium at the International Rice Research Newsl N3)15 Institute September 1964 The Johns Hopkins Press Baltimore 63 Fugenio T S 1965 rhe effect oi e-vCgetation on rice production Maryland USA 729 p and the possibility of overcoming it by leaf pruning MS thesis

42 Centro Ihternacional de Agricklittira Tropical 1984 Upland rice in University ol the Philippines at Los Baijos ILaguna Ihilippins 69 p Iatin Anerica Pages 93-119 hi An os ervies of uplaind rice research 04 Farrow R A 1975 The Alrican nmigratory locust ii is main Proceedings of the 1982 Boinakamp Ivory Coast upland rice workshop outbreak area of the middle Niger qlinlitatise studies of solitary International Rice Research Institute P0 Blox 933 Manila populations in relation to environmental factors Locusla No II Philippines 198 p

43 Chandra G 1979 Taxonomy ad hn riononlics ofthe imect parasites of 65 Fennal R G 1943 Food crop investigation Windward and rice Icalloppers and planthoppers in the Pihilippines and their Ieeward Islands First report Report onl work dunring the periodiiiportairce iii natural biol)gical cointrol Philipp Fntoltol 4119-139 April 1942-April 1943 Imperial College of Tropical Agriculture

44 Chang T T 1976 The originii colut ion enlIivation dissemination Trinidad 16 p (Rev of Appl Fintornol A3222-25) and diersificition of Asian and African rices -uphytica 25425-44 1 66 Ferreira F 1983 Integrated pest ritaageiret of upland rice [ill

45 Chatierjee P B 1979 Rice leaffolder attacks in India lit Rice Res Portuguese English summary] Pages 323-341 in Upland riceculttre Newsl 4(3)21 factors affecting production E Ferreira T Yamada and F-

46 Cheng Cbien-pan 19(4 Upland rice in Taiwan International Rice Malmolta eds Institutoda Potassae Foslato Institnto Internacional Cnmmission Wiorking Party oit Rice Production ard Protection I()th da Potassa Piracicaha Sao Paulo lraid Meeting 3-10 March 1964 Manila Philippines Food and Agricul- 67 Ferreira E arid F dl Silva Martins 1979 Occurrence of ture Organiaion Rome Italy 5 p Sogatohs orvziola (Muir) inupland rice in Goias Ilrail lit Rice

47 Chcrrett 1 M and 1) 1Peregrine 1976 A review of the status of Rcs Ncwssl 4(4)18 leaf-cutting ants and their control Ann Appl Biol 84124-128 68 Ferreira E and 1F da Silva Martiris 1984 Insect pestof rice ill

48 Chipeta F M 1974 Pests of rice in Malawi Pages 90-95 in Brazil and their control Jill Portugucsej FMIlRAtPA-CNPAF Proceedings Sith eastern Africa cereals research conference 10-15 Goiania Brmiil 67 p March 1974 DRB Mialda ed Ministry of Agriculture lilongwe 69 Ferreira FI F ca Silha Miartins S S Nero and 1 It P de Malawi Carvallio 1982 M tltiplc regression analysis todeterimine the effect of

49 Chu Yau-i 1979 The d atiaging period of the rice seedling fly insect populatiois ott upland rice yield [inIPortuguese Pesqui (Atlterigotiavxigu Stcin) (NItiscide Diptera) oil cori) at Fast Java Agropecu Bras 17671-675 Indonesia Plant Prot Bill 21403413 70 Ferreira F 1F da Silva Martins and F 1PZimmermann 1979

50 Conklin H C 1957 IHaninoo agriculture a report on aln integral Rice varietal resistance to tle lesser corn stalk borer [in Port uguese] systent of shi fti rig cult ivationi in the Pltilippincs Food and Agriculture Pesqui Agropccu Biras 14317-321 Organization Rome Italy 209 p 71 Fleming W F 1968 Biological control of lie hapanese beetle U S

51 Cramer H II 1967 Plant proection and world crop production Dep Agric RLs Sers [ech Bull 1383 78 p Pilaniensclhutz-Nachr 201-524 72 Food and Agriculture Organiation of the United Nations 1982

52 l)ani R C and Majunidar 1978 Prelininary observations on tie Country report surmimary Pages 27-37 in Report on the first incidence of root aphids on differeit rice cultnrcs Sci Cult 4488-89 FAOIUN)IPhail and regional training coiuirse in iniproved ctltural

53 Davidson R I 1979 lffects of root feeding on foliage yield Pages practices for upland rice 5-23 Oct 1981 Northern Region Agrculturd117-120 in T K Ci osb and R B Pottinger eds Proceedings of the Developririt Center Chiang Mai Food and Agricuilture Organiiashy2nd Australian conlerence on grassland invertebrate ecology Govt tion Rome Italy 37 n Printer Wellington New Zealand 73 Franco do Ariaral S 195 1 Residual action of insecticides in turfy

54 l)ean G 1W 1976 Rice insect pests inlaos Int Rice Res Newsl soil--the results of a field experiment [in Portuguese] Biol6gico (Sao 1(2)15 Paulo) 17209-214 (Rev Appl Entomol A41423)

55 I)ean G J W 1978 Insect pests of rice in Laos PANS 24(3)280-289 74 Freeman ) 1970 Report onithe Iban london School of -coroiics 56 )ingle H 1984 Behavior genes and lifc histories conplex adapta- Monographs on Social Anthropology No 41 Athlane Press

tions to uncertain environments Pages 169-194 in A new ecology London UK 317 p novel approaches to interactive systems P W Price C N Slobod- 75 Fuchs T W1 A Harding and 1W Smithlr 1979 Induction and chikoff and W S Gaud cdsl ohn Wiley New York USA termination oldiapause in the sugarcane borer Ann Flitonlol Soc

57 Dobelmann-JP and M Falais 1968 Upland rice cultivation ill tie Am 72271-274 North-western part of Madagascar in 1966-1967 [in French English 76 (hbriel B1) 1975 A review tf the major insect pests of some tpland summary] Trop Agron 2313-21 crops in tlc Philippines Pages 86-111 in Ad Htoc panel of experts on

58 Doniiciano N L 1979 Chemical control of tie lesser corn stalk pest disease arnd weed problems in some rainfed crops Bangkok (lolsonno~paloIgnowslhts) (Zeller 1848) (I-epid(ptera Ihycitid ae) Thailand Food aid Agriculture Organization Roire Italy

on rice in Ponta Gressa Parara 1977-1978 [in Portuguese] Pages 77 Garg D K and N K Shal 1983 White grub outbreak on rainfed 291-303 in Anais 1 Reuniao de tecnicos cmiirizicultura do estado de dryland rice in Uttar Pradeslh Int Rice Res Ncwsl 81(4)16-17 Sao Paulo-Campinas Cati Brazil 78 Garg I) K and IP Tandon 1982 Major insect pests of rice on hilly

59 Dunsmore 1 R 1970 Investigations on tlie varieties pests and tracts of Uttar Pradesh India lilt Rice Res Newsl 7(11)11-12diseases of upland rice (hill padi) in Sarawak Malaysia Int Rice 79 Garrity 1) P 1983 Upland rice research an international bibliograpy Comm Newsl 1929-35 1975-1982 Iternational Rice Research hnstitnte PO Box 933

60 Elias 1R1965 The new decline of rice seedlings is not caused by Manila Philippines 121 p parasites [in Portuguese] Biol6gico (Sao Paulo) 313-6 80 Ghauri M S K 1982 A new subspecies of Iinorh)rone bull

61 Empresa Brasiliera de Pesquisa Agropecuaria 1984 Upland rice itt cornlus Slater (Hemiptera Iygaeidae) front Papua New Guinea on Brazil Pages 121-134 it Av overview of upland rice research rice and carpet grass Ihll lEntoniol Res 72133-137 Proceedings ofthe 1982 Blouak Ivory Coast Upland rice workshop 81 Gonzales F 0 PArregoces R I ihernandef and OT Parada International Rice Research Institute PO Box 933 Manila 1983 Insect and imite pests of rice arid their control in Latin America Philippines [in Spanish] la Federacion Nacional de Arrocerts (Fedearroz)

Bogota Colombia 60 n

38 IRPS No 123 January 1987

82 Grist D H and R J A W Lever 1969 Pests of rice Longmans London UK520 p

83 Hale P R 1979 Two subterranean pests of upland rice in Papua New Guirnca IntRice Res Newsl 4(2)16-17

84 Hale P R and J T Hale 1975 Insect pests of rice in the New Guinea Islands Rice Entomol Newsl 34

85 Harris W V 1969 Termites as pests of crops and trees Common-wealth Institute of Entomology London UK 41 p

86 Hattori 1 1968 A method to distinguish between grass leaf roller (Ctaplha rocis medinalisGuenee) and the rice leaf roller (Susurtia exigua Butler) [in Japanese] Plant Prot (Japan) 23167-170

87 Henry I E and 1A Onsager 1984 Experimental control of the mormon cricket Iahrosshnph by NJsena locustae(Aficrospora Microsporidae) a protozoan parasite of grasshoppers (Ort Acri-didae) Entomophaga 27197-201

88 Hidaka 1 P Vungsilabulr and SKadkao 1974 Studies on ecology and control of the rice gall midge in Thailand Tech Bull 6 TropicalAgricultural Research Center Tokyo Japan 113 p

89 Hincklcy A 1) 1963 Ecology and control of rice planthoppers in Fiji Bull Entomrol Res 53467 481

90 Hirao J 1982 Ecology and chtInical control of the rice leaffolder Jpn Pestic Inf 4114-17

91 Ho DT 1984 Sten borer incifience in rice ecosystems in KenyaEast Africa Pages 307-320 in A overview of upland rice research Proceedings of the 1982 Bouac Ivory Coast upland rice workshop International Rice Research Institute PO Box 933 Manila Philippines

92 Ho )T J 1Njokah and J GKibuka 1983 Studies on rice sten borers in Kenya with emphasis on ifaliarpiaipiraellRag Insect Sci Appl 465-73

93 Ichihara I T Ito A NIatsuo and M Sawada 1971 Chemical control of the rice striped stem borer in upland rice grown in plastic mulch [iii Japanese] Proc Kanto-Tosan Plant Prot Soc 1882

94 IclhiharaIand A Matsuo 1969 Damage caused by rice striped stett borer to upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1686-87

95 Ichhara I atid A MIatsun11970 Conside ration oft lie extraordinary prevalence of rice striped stem borer in upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1778-79

96 Institut de Recherches Agrolnliqes Iropicale et des Cultures Vivrieres 1978 Insect resistance Pages 12-13 in Upland rice 1978 atnual report Bouak Ivory Coast 27 p

97 International Institute for Tropical Agriculture 1983 Aphids itt rice roots I ITA Res Briefs 4(1-3)4-5

98 International Institite fi Tropical Agriculture 1983 Hopperburn in rice caused by brown plathopper IITA Res Briefs 4(l-3)6-7

99 International Rice Research Institute 1975 Annual report for 1974 PO Box 933 Manila Philippines p 216

100 International Rice Restarch Institute 1976 Annual repiort for 1975 PO Box 933 Manila Philippines p 228

101 International Rice Research Institute 1979 Cropping systems program Component technology development and evaluation Page 404 i Annual report for 1978 11O Box 933 Manila Philippines

102 International Rice Research Institute Department of Entomology 1981) Insecticid evaluation 1979 Tables 73 74 PO Box 933Manila Philipes

103 International Rice Research Institute Department of Entonmology 1981 Insecticide evalomtion 1980 Table 81 POBox 933 Manila Philippines

104 International Rice Research Institute 1981 Report of a meeting of Asian rice-based cropping systems entomologists Eleventh cropping systems working group meeting 18-22 May 1981 PO Box 933 Manila Philippines p 132 133 144

105 International Rice Research Institute 1984 Cropping systems program Pages 368-373 in Annual report for 1983 PO Box 933 Manila Philippines

106 International Rice Research Institutc 1984 Upland rice in Asia Pages 45-68 in An overview of upland rice research Proceedings of the 1982 Bouakamp Ivory Coast upland rice workiiop loternational Rice Research Institute 11O Box 933 Manila Philippines

107 Irshad M 1977 Grasshoppers in rice in Pakistan lit Rice Res Newsl 2(3)8-9

108 Ito R and TOno 1964 Upland lice its culture and varieties in Japan lIternational Rice Cormmission loth Meeting of the Working Party on Rice Production and Protection 3-10 March 1964 Manila Philippines Food and Agriculture Organization Rome Italy 12 p

109 Jeffrey M I and V A Dyck 1983 Effect of moonlight on lightshytrap catches of brown planthopper lit Rice Res Newsl 8(6) 13

110 Jennings P R and T Alicia Pineda 1983 Management ofthe rice plnthoppers Sogatodsorivi(coIl in Latin America Presented at the meeting on stable plant resistance FAO and lIndocsian I)inectorate of Food Crops Protection Bali lndonesia 29 Novcniber-2 l)ccemshyber 9 p

IllJimenez Gomez J A 1979 Rice insect pests i lla ios ()riCntales lil Spanish] Pages 99-103 in Rice ciltivation Instituto (olotibiatto Agropecuario Compndio No 29

112 Jongeleen F JJ Soesanto and A Mockidjo 1979 Studies oti pathogens of two species of Scarabacidac froinIndonesia Z Angew Enlomnol 87412-417

113 Jouvenaz 1) PC S lofgren and W A Baiks 1981 Biological control of imported fire antsa review of cirrent knowledge Bull Entomol Soc Am 27203-208

114 Kainoh Y K Shiniiu S MarUani Y Taiaki 1980 lost finding behavior of tie rice bu lplotori+JchinvittjI)allas (liciniptera Corcidac) with special reference to diet pattern ol aggregation and feeding on the rice plant Appl Entomol Zool 15225-233

115 Kakkar K 0 P Sharma and P I Sharma 1977 Note on Ihteronyrchus liodresRedt (Coleopicra I)ynastidae) as asporadic pest of paddy and maie inHimachal Pradesh Corr Agric 156-57

116 Kalode M131 79 Major insect pest problems and their control in upland rice (Unpublishcd mineogriph CWI oJNaaiat lni Chandaratat (198 1)original not seen)

117 Kalshoven L i 98 I The pests of crops in I ldoniesia (translated by P A vatn der Laan) 1 T Ichtiar B3aru-vati Iloeve lakarta Indonesia 701 p

118 Kamal N Q 1981 Suppression of whitebacked planto ner Sogatella fircaifra (Horvath) aLId rice lealfolder (nhal -cis licdb lis (Guenee) populations by natural cnclmie hlI) tiesis Grcgorio Araneta University Foundation Victoncta Park Metto Manila Philippines 179 Ip

119 Karim A N M Reaul 1975 A species ofliealy bug Trioni i70s ceres new oii rice in the Philippines Rice Entoniol Newsl 247

120 Katanyukul Wand C Chandaratat 1981 Insect pestsof uplaid rice and their control Report firstFAOUNlFi Tliailand training course on improved cultural practices for tplad rice Chiang Mai Thailand 5-23 Oct 1981 Food and Agriculture Organiation Rome Italy 22 p

121 Kern F 1955 lhletwioa(I)uninemts) bhhit Biturm(Coleoptera) a new pest of sugarcane inVenezuela [inSpanish with Etnglish sunmary] Agron Trop (Maracay) 527-29 (Rev Appl Eitoniol A4572)

122 Khan K M and S Ghai 1974 White grubs ald their control in India Pesticides (1Bombay) 819-25

123 Kiritani K 1975 Integrated pest management of rice pests Pages 66-70 itt Proceedings platning workshop ott cooperative hIeld research in pest managementP SMotooka ed East-West Center Honololu Hawaii USA

124 Kisimoto R 1967 Ecology of insect vectors forecasting and chemical control Pages 243-255 in The virus diseases of tle rice plant Proceedings of a symposium atthe International Rice Research Institute April 1967 The Johns Hopkins Press altimore Maryshyland USA

125 Kisimoto R and V A Dyck 1976 Climate and rice insects Pages 367-391 in Climate and rice International Rice Research Institute PO Box 933 Manila Philippines

126 Kureha S S Kobayashi SAsari and T Ishihara 1977 Life history and control of the mole cricket Grrllotalpa africana Palisot de Beauvois [in Japanese English translation 7520 1RRI library] Proc Kanto-Tosan Plant Prot Soc 21192-197

127 Kushwaha K S 1981 The black beetle as a pest of rice in Haryana India Int Rice Res Newsl 6(4)14

128 Lawani S M 1982 A review of tile effects of various agronomic practices on cereal stem borers Trop Pest Manage 28266-276

129 Leuck D B 1966 Biology of the lesser cornstalk borer in South Georgia J Econ Entomol 59 797-801

130 lever R J A W 1955 Insect pests of the rice plant in Malaya Int Rice Comm Newsl 1612-21

131 Li Ching-Sing and Shin-Foon Chiu 1951 The study ofthe rice gall midge Iachydiplosis orvza Wood Mason [in Chinese with English summary] 1Taiwan Agric Res 21-13

132 Iitsinger J A 1984 Assesament of the need-based application for rice Proceedings TechnologyTransfer Workshop 15-16 March 1984 International Rice Research Institute 1PO Box 933 Manila Philippines 6 p

133 Litsinger J A 1984 Environmental diversity and insect pest abundance with reference to the Pacific Pages 285-307 in Proceedings sub-regional training course on methods of controlling diseases insects and other pests of plants in the South Pacific 4-20 Oct 1982 Vaini Kingdom of Tonga Nukualofa

134 Litsinger J A RF Apostol and M BObusan 1983 White grub Leucopholis irrorata (Coleoptera Scarabaeidae) pest status popula-tion dynamics and chemical conllol in arice-maize cropping pattern in the Philippines J Econ Entoinol 761133-1138

135 Litsinger J A C de la Cruz F Raymundo A T Barrion M D Lumaban J P Bandong M S Venugopal F Paragna and E Balete 1982 Insect pests and insecticide reponse of dry-seeded bunded rice Pages 179-193 in Report of workshop on cropping systems research in Asia International Rice Research Institute Los Bafos Philippine

136 Litsinger J A M 1) Luinaban J P Bandong A T Barrion W Katanyukul and N Panda 1978 Insect pest management in rainfcd lowland rice agroecosystms of tropical Asia International Rice Research Conference 17-21 April 1978 International Rice Research Institute Los Bafios Philippines

137 Litsinger J A and K Moody 1976 Integrated pest management in multiple cropping systems Pages 293-317 in R I Papendick P A Sanchez and G B Triplett eds Multiple cropping American Society of Agronomy Madison Wisconsin USA

138 Loevinsohn M E 1984 The ecology and control of rice pests in relation to the intensity and synchrony of cultivation Ph D thesis University of London 36) p

139 Loevinsohn M E J A Litsinger N Panda M A Austria R F Apostol C Andrion and J Bandong 1982 A basis for predicting rice insect pest abundance in cropping systems Paper presented at the 1982 International rice research conference on rice and cropping systems 19-23 April 1982 International Rice Research Institute Los Bafios Iaguna Philippines 25 p

140 Lopez A W 1930 Repot of the Entomology Department Pages 145-172 in Annual report of the Philippine Sugar Association 1929-30 Manila

141 Lopez AW 1931 Entomology Department Pages 227-266 in Annual report of the director or research 1930-31 Philippine Sugar Association Manila

142 Lopez A W 1932 Entomology Deparment Pages 252-279 in Annual report of the Philippine Sugar Association 1931-32 Manila

143 Lungibill P and G G Ainslee 1917 The lessercornstalk borer US Dep Agric Agric Res Serv Tech Bull 539 27 p

IRPS No 123 January 1987 39

144 McGuire J U Jr and BS Crandall 1967 Survey of insect pests and plant diseases ofselected food crops of Mexico Central America and Panama U SDepartment of Agriculture International Agriculshytural Development Service Agricultuial Researrh Service October 1967 11 p

145 McNeill Sand T R ESouthwood 1978 The role of nitrogen in the development of insectplant relations Pages 77-9 in lliochemical aspects of plant and animal coevolution J B Ilarbornu ed Academic Press London UK

146 Maebara H 1955 Spatial distribution of lastinstiarlanaeof purplish stem borer (Sesamia iqferens Walker) hibernating in an upland rice field Bull Kagoshima Univ Fac Agric 438-41

147 Mallick E H and N G Hajra 1977 Response of rice plant to defoliation treatments at different developmental stages Indian Agric 2151-56

148 Mansingh A 1971 Physiological classification of dormancies in insects Canl Entomol 103983-10)9

149 Martell C G 1974 Catalogue of plant feeding insects of Mexico [in Spanish] Fitofilo 27(69)1-176

150 Martinez 0 J 1982 Principal rice pests in Ilanos orientalis [in Spanish] P~ages 80-94 in Curso deact nalization de conocimientos en arroz Villaniencis Instituto (olonmbiano Agropecuario

151 Martins J F da Silva F Ferrcira A S Prabhu F 1 P Zimmermann 1980 Use of pesticides for the control of the main subterranean upland rice pests esqui Agropecu Bras 1553-u

152 Martins I F da Silva F Fe eira rod BdamSilveira Pinheiro I Q2 Damage simulation caused by defoliators of upland rice [in Portuguese with English summary] Pesqui Agropecu Bras 171113-1119

153 Martins 1F da Silva N(iyuen van Tan and 1Ida Silveira Pinheiro 1981 Stem borer resistance in upland rice associated with plant morphological characters [in Portuuce Iesqui Agropecu Bras 16187-192

154 Masaki S 1980 Sumnmer diapause Ann Rev Entomiol 251-25 155 Masuzawa T 11 Suwa and F Nakasoji 1983 l)ifkleences of

oviposition preferencc and survival rate of two skipper butterflies Parnara guttata and PIchlpidai iathiais (Iepidoptera HIesperiidac) on rice plant and cogon grass New Entomol 3247-56

156 Maurya D M and C P Vaish 1984 Upland rice in India Pages 69-92 in An overview oftpland rice research Proceedings ofthe 1982 Bouakamp Ivory Coast upland rice workshop International Rice Research Institute IO Box 933 Mnila Philippines

157 Menon M G R R N Katiyai S Kuar and S S Misra 1970 Termites attacking the paddy crop Sci Cult 36294-295

158 Miyahara Y and 1 Nishikubo 1967 Behavior of young nymphs of the red rice root aphid Iholahsipiun ri liabhdontinalisSasaki in relation to movement from the leaf to the roots of upland rice [in Japanese] Pioc Assoc Plant Prot Kyushu 1363-66

159 Mo ljoa Tjien 1952 White grub of upland rice (Holotrichiahelleri Brsk) [in Indonesian with English summary] Contrib Agric Res Stn Bogor No 13134 p

160 Mochida 0 and T Suryana 1979 The occurrence of the brown planthopper and ragged stunt on lowland and upland rice ill Central Java and logyakarta in tile late of January 1979 Entomnolgy Survey Report No 15 Central Research Institute of Agriculture (CRIA) Sukamandi Indonesia I February 1979 9 p

161Monet C 1979 Varietal resistance to upland rice pests [inI French] Pages 35-39 in Annual report 1977 Institut des Savanes Department des Cultures Vivrieres Bouak6 Ivory Coast

162 Monte 0 1942 A caterpillar ofrice-fields[in Portuguese] Biol6gico (Sao Paulo) 8161-163 (Rev Appl Entomol A31414)

163 Munroe I) D 1979 Pest management strategy for hill rice in four resettlement areas in Sarawak Malaysia Pages 89-102 i1 Guidelines for integrated control of rice insect pests Food and Agriculture Organization Plant Production and Protection Paper 14 Rome Italy 115 p

40 IRPS No 123 January 1987

164 Nakasuji F M Ishii and Jen-Zon Ito 1984 Rice skippers in Taiwan and their life histories ZZrC (Taipei) Tech Bull 82 10 p

165 Navas I) 1976 Fall armyworum in rice Pages 99-106 il Ecological animal control by habitat management Proc Tall Timbers Con-ference 28 Feb-I Mar 1974 Fall Timbers Res Stn Misc Publ 6 Tallahassee Horida USA

166 Nickel 1 1 1961 Biological control of rice stem borer A feasibility study Tcch Bull 2 International Rice Rcarch Institute PO Box 933 Manila Philippines IIl p

167 Nigan P M and R A Vernia 1985 Insect pests of uplaild rice in Uttar Pradesh Int Rice Res Newsl 10(6)22

168 Nilakhe S S 1985 -cological observations on spittle bugs with emphasis on their occirence in rice Pcsqtui Agropecu Bras 20407-414

169 Nixon G EJ 195 1 lie association of ats with aphids and coccids Commonwealth Iusti lcte of Entonology Iondon UK 36 p

170 Nurdin F an Ishak Manti 1985 Study oit seedling fly of upland rice Interlational Upland Rice Moitoring Tour Report 26 lebruary-2 MarLI Sukaranti West Sumatra

171 Oka 1 N 1979 Ciltural conttol of the brown planthopper Pages 357-369 in Brown planthopper threat to rice production in Asia International Rice Research Institutc 1p0 Box 933 Manila Philippines

172 Ohni M and A Vilnaii 1975 Rice insect pests of Italy biology of Sipha grcria (Kaltenbach) and its control [in Italian] Riso (Milan) 2459-71

173 Otanes F Q atnd 1 I Sison 194 1 Pests of rice Philipp J Agric 12211-259

174 Painter R I 1951 Insect icsistance in crop plants University Press of Kansits Lawrence Kansas USA 520 p

175 Pantua P C and 1A litsinger 1981 Arthropod abundance in continuous vs biannual rice cropping systeis Int Rice Res Ncwsl 6(2)20-21

176 Pathak M ) and V A l)yck 1975 Sttudics ott insect pests of upland rice Pages 186-191 in Major research in upland rice International Rice Research Institute PO Box 933 Ilanila Philippines

177 Patil B R B M IHasabe and 1) S Ajri 198 I Chemical control of white gritb lIditriihinhorraia Fabr intfesting groundntut paddy atnd sorghum Indian 1Entomol 43232-236

178 Pawar A 1) 1974 (ollection and identification of insect pests of rice rlerminal repirt lInternational Rice Research Institute PO Box 933 Manila Philippines 72 p (tnimco)

179 IerrajU A aid A S Reedy 1965 Observations ott diapausc itt tie steti borer ohrice Rice News Teller (lai-Mar)9-13

180 Pont A C 1972 A review of the Oriental species of theri ona Rondani (I)iptera Muscidac) of econtomic importance Pages 27-102 in Control of sorglium shoot fly M G lomtani and W R Young eds Oxford atd IH1 Ptthl Co New I)clhi India 324 p

181 Prasad S C 1PChand aid M 1PSingh 1983 Incidence of rice gall midge in the upland plateau region of Hihar Oryza 20235-236

182 Rai P S 198 0 Nisaga simpex damage to rice in the hill tracts of South India Int Rice kles News[ 5(I)12

183 Rathorc Y S 1969 Gonocplhahli dcprssunt Fabr as itpredator of (tilo zonwIlhis (Swinhoc) and other moths in field cage at Patnagar Uttar Pradesh Indian 1Entotnol 3127

184 Reed W 1983 Crop losses causcd by insect pists in the developing world Proceedings 10th Internatial Congress of Plant Protection lrighton UK 174-80

185 Reedy M V 1981 Ateactogasterinhens(Walker)a new root-grub pest of tipland paddy and inaize Curr Sci 501076

186 Reveno M A 1968 Catalogue of the insect pests of economic crops in Colombia [in Spaiish] Asociacion Latino Americana de Entomo-logia (AIAE) 129-33

187 Reyes G M 1957 Rice dwarf disease in the Philippines Food and Agriculture Organiiation Plant Prot Bull 617-19

188 Rice S E A A Grigarick and M 0 Way 1982 Effect of leaf and panicle feeding byarmyworm (Lepidoptera Noctuidae) larvae on rice grain yield Environ Entomol 7593-595

189 Risch S J 1) Andow and M A Altieri 1983 Agioccosystem diversity arnd pest control data tentative conclusions and nizw research dirctions Environ Entomol 12625-629

190 Rivera C T 1956 Three organic insecticides in thecontrol of insects affecting upland rice Philipp Agric 39465-472

191 Roffrey 1 1972 Locusta outbreaks in the Philippines Acrida 1177-188

192 Rojas R 1) 1941 The effect of pruning at varying ages upon the yield of Ramairio Phipp Agric 29851-860

193 Roonwal M i 1979 Ternite life and termite control in tropical South Asia Scientilic Pnbl odhpr India 139 p

194 Rosetto C1 F da Silva Martins N C Schmidt and L EAzzini 1978 Damage caused by pasturespittle bugs 4Dvoi)lavopictaandDJ schach) on rice lBragantia Nota 6 (Matcli)25-27

195 Rosztto C1- S S Neto I- Amantc I) Iink 1Garcia-Vieira D M de SoLza N V Bansa t and A M de Olivcira 1971 Rice pests of Brazil [in Portuguese] Pages 149-238 inrFA()-International Rice Commission 2nd meeting of the Rice Committee for the Americas 6-11 l)ecember 1971 Pelotas Rio Grande do Sul Brazil

196 Rothschild G H L 1969 Obseratiois on the armyworm Spodoptera matritia acronrctoiiles Gn (Lep Noctuidae) Bull Entomol Res 59143-160

197 Rothschild G 1- L 1974 Parasites of the armworm Spolopiera mauritiaacroncitoihes in Malaysia Fntornopliaga 19293-299

198 Sachan i N S K Gangwar and 1N Katujar 1980 Field crickets damaging cages in North Eastern IHill Region Indian J Entomol 42526-527

199 Sands 1) P A 1977 The biology and ecology of Lepocorisa (Hemiptera Alydidac) in Papua New Guinea Res Bull 18 Departshyment of Primary Industry Port Moresby Papua New Guinea 104 p

200 Sarawak 1968 titomology division Pages 97-106 in Report of the research branch of the Department of Altriculture 1966 Kuching (Rev AIppl Eintomol A56603)

201 Satpathy JM M S Das and K Naik 1977 Effect of multiple and mixed cropping on the incidence of scine important pestsI Entomol Res 178-85

202 Sauer IHF G 1939 Notes Ott Elasmpnlpualtsligniosellths Zeller (Lep Pyr) a serious pest ofcereals in Sao Paolo [in Portuguese] Arq Inst Biol 10199-206 (Rev Appl Entotnol A28378)

203 Satinders 1L A 1 SKing and C L Vargas S 1983 Crop pests of Central Anerica with t bibliography [in Spanish] Centro Agroshynotnico Tropical de Investigacion y Ensenanza Turrialba Costa Rica (CATIE) Tec Bol 9 90 p

204 Sauphanor B 1985 Some factors of upland rice tolerttce to stem borers in West Africa Inseu Sci Appl 6429434

205 Sen P atnd S Chakravorty 1976 Stentflics in Haritighata Nadia West Bengal India [it Rice Res Newsl 1(12) 15

206 Senapai B and 1M Satpathy 1983 The seedling fly Atherigona orrzae Malloch is a pest of rice in Orissa (India) J Entomol Res 777-79

207 Shhjahan M S Alatn M Ah nad and BN Islam 1983 A new scarabaeid beetle pest of dryland rice in Bangladesh Int Rice Res Ncwsl 8(I)16-17

208 Sharma G L S C Modgol and R C Gautam 1979 Effect of depth of submergence ott incidence of bacterial blight and white grub infestation in transplanted rice Int Rice Res Newsl 4(6)12-13

209 Sharma S 1985 Scarabaeid beetle outbreak itt upland rice Int Rice Res Newsl 1(1(l)24

210 Shiraki T 1917 The paddy borer Schoenobhits incertulas Walker [achaku Agric Exp Stn Formosa 256 p

211 Silva A de Brito and 11P Magalhaes 1981 Insect pests of rice [in Portuguese] Circular Tecnica No 22 EMBRAPA Centro de Pesquisa t ecuaria de Tropico Unido (CPAJU) Belem Brazil 14 p

212 Singh S A 1979 Emergence of rice leaf butterfly as a rice pest in Manipur India int Rice Res Newsl 4(4)16-17

213 SinghS A 1983 Changing trends of insect pestsof rice in Manipur Int Rice Res Newsl 8(f)12

214 Siwi S S and P H van Doesburg 1984 Lenovorisa Latreille in Indonesia (Heteroptera Coreidae Alydinae) Zool Meded (Leiden) 58117-129

215 Soejitno J 1977 Reaction of some upland rice varieties toseedling fly eltherikonaexigua Stein [in Indonesian] Siri Hana I Penyakit 630-35 Lembga I osat Fenclilian Pertanian Bogor Indonesia

216 Socnardi 1967 Insect pests of rice in Indonesia Pages675-683 in The major inse pests of thu rice plant Proceedings of a symposium at the Internatital Rice Rec Wi itt S pternber 1964 The Johns Hopkins Press Baltimore Maryland USA

217 Soto P E A T Petcz and I W Buddenhagen 1976 Survey of insect pests and diseases of rice in different ecological zones in Nigeria Rice Entomol Ncwsl 435-46

218 Soto 11E and Z Ziddiqi 1976 Screening for resistance to African rice insect pests Varietal improsetnent sCmi n r Bouak Ivory Coast 13-17 Sept 1976 West Africa Rice l)evelopment Association Monrovia Iiberia

219 Squire F A 1972 Entomological problems in Bolivia PANS (PestArtic News Sumr) 18249-268

220 Stapely 1 II 1978 The rice leaf roller in the Solomon Islands Int Rice Res Newsl 3(l)13

221 Suenaga H 1937 On aphids attacking the roots of rice Oyo-I)oubts Zasshi 9150-152 [in Japanese] (Rev Appl Entomol A25745)

222 Suenaga H 1952 On the seasonal abundance of three subterranean aphids on upland rice Bull Kyushu Agric Exp Sti 1249-262

223 Sutynso Yas and Dandi Soekaria 1964 Beberapa hana yang ttenperang tana man padi tanah kering [in Indonesian] Rapat kerja Padi Tanah Kering 6 sd 8 Juily 1964 Kaliurang 9 p

224 Tallamy I) W and R F Denno 1983 Alternative life history patterns in risky environments in example from lace bugs Pages 129-147 in Insect life history patterns R F l)cnio and If Dingle eds Spring-Verlag 225 p

225 Tamura 1 1939 Observations on some insect pests of field crops [in Japanese] 1 Plant Prot 26414-420 (Rev Appl EntomolA27-556)

226 Tamura and Suzuki 1964 Influences of the early-transplanting practice in rice culture upon development of the rice stem borer Chilo siql)ressalisWalker Bull Ilokuriku Natil Agric Exp Stn 761-94

227 Tan J P 1924 The rice root aphid Drowhi ihirsoia Philipp Agric 13277-288

228 Tanaka 1 1957 Taxonomy and distribution of some subterranean aphids injurious to upland rice in Japan with description of a new species [in Japanese English summary] Botyu-Kagaku 22168-176

229 Tanaka T 1961 The rice root aphids their ecology and control Spec [full College of Agriculture Utsunomiya University Japan F3 p

230 Tauber M IC A Tauber R Nechols and 1Obrycki 1983 Seasonal activity of parasitoids control by external internal and genetic factors Pages 87-108 itt I)iapause and life cycle strategies in insects V K Brown and I Ilodek eds )r W Junk The Hague Netherlands

231 Taylor W F 1972 Effects of artificial defoliation (simulating pest damage) on varieties of upland rice Exp Agric 879-83

232 feotia T P S and S Natud 1966 Biotoomics of the rice skipper Parnaraniathiis Fabricius (Lepidoptera Hesperiidac) Indian J Entomol 28181-186

233 Teran F 0 1971 Rice pests of Santa Cruz [in Spanish Ministerio de Agricultura Sita Cruz Bolivia Bull 45 8 p

234 Than Itun 1976 Population dynamics of the yellow rice borer Tryporza itcertulas (Walker) and its damage to the rice plant MS thesis University of the Philippines at Ios Bafios Laguna Philippines 92 p

235 Uichanco L B 1911 Pests Pages 55-74 it Rice Philipp Agric 11-200

IRPS No 123 January 1987 41

236 van der Laan P A 195 1 Life history of the rice seedling fly Contrib Gen Agric Res Stn Boor Indonesia 118 15 p

237 van der Laan 11A 1959 Correlation between rainfall in the dry season and the occurrence of the white rice borer (Scirpolhaga innotala WIk) in lava Entomol Fxp Appl 212-20

238 Van Driesche R G 1983 Meaning of percent parasitismin studies of insect parasitism Fnviron Entomol 121 I1- 122

239 Venugopal M S and A litsingcr 1984 Fffect ofcarbofuran on rice growth Prot Ecol 7313-317

240 Vietnam Directorate of Rural Affairs 1962 Annual work progress report on crop improvement program of rice sugarcane vegetable and field crops (lor the period from July 1961 to June 1962) (11) 299 p (Rev Appl Entomol A51526-527)

241 Wan M T K 1972 Observations on rice leaf- and planthoppers in Sarawak (Malaysian Borneo) Malay Agric 148308-335

242 West Africa Rice I)esclopnient Association 1978 Annual research report Vol 2 Statistics Monrovia Liberia

243 West Africa Rice Development Association 1979 Annual research report p47 Monrovia Liberia

244 Williams ) F and CS lofgren 1981 Fli I illy l-468 anew bait toxicant for control of the red imported lire at Fla Entomol 64472477

245 Williams I) 1 1970 The mealybugs OFlomoptera Coccoidea Iscudococcidac) of sugar-cane rice and sorghum Bull Entomol Res 0109-188

246 Williams D) 1 1982 Pulinaria icervi (Signoret) and its allies ott sugar cane and other grasses Btll Entoinol Res 72111-117

247 Wirjosuiardjo S 1976 Study ofchenmical cont rot of insect pests ott upland rice Imiu Pertanian (Jogjakarta) 1323-329

248 Wirjosuhardjo S and N Wirarta 1976 Chemical control of the white grub tIolorithiatlltriin upland rice in M uneng Irobolinggo Indonesia) [in Indonesian] Imut Pertanian (Jogjakarta) 1330-332

249 Yano K T Miyake and V F Eastop 1983 The biology and economic importance of rice aphids (Henuiptera Aphid idae) a review Bull Entomol Res 73539-566

250 Yelp D and P Rhodes 19(4 lhe success of drilled dryland rice iii the Fiji Islands International Rice Commission Working Party on Rice Production and Protection Manila Philippines 3-10 March1964 Food and Agriculture Organization Rome Italy 13 p

251 Young W R 1981 Fifty-five years of research on the sorghum shootfly Insect Sci Appi 23-9

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ISSN 05-3862Other papers in this series TITLES OF NUMBERS 1-45 ARE LISTED ON THE LAST PAGE OF NO46 THOSE OF NUMBERS 46-70 ARE ON THE LAST PAGE OF NO 71-80AND THOSE OF NUMBERS 71-1O0ARE ON

THE LAST PAGE OF NO 101-122

No 101The economics of hybrid rice production in (hina No 113 Boron toxicity in rice No 102 Rice ratooning No 114 Energy analysis rice production systems and rice research No 103 Growth and development of the (Jeep water rice plant No 115 Production risk and optimal fertilizer rates an application of the

No 104 Faridpur acomputer-assisted instruction nmodel forrainfed lowland random coefficient model No 116 Consumer demand for rice grain quality in Thailand Indonesia andrice

No 105 A readingand listening comprehcnsion test inEnglish for non native the Philippines speakers applying for tiraining at IRIRI No 117 Morphological changes in rice panicle development a review of

No 106 Rice grassy stunt virus 2 a new strain of rice grassy stunt in tire literature

Philippines No 118 IRRI-Korca collaborative project for the development of cold-No 107 Physical losses and quality deterioration in rice plstprodnCtion tolerant lines through anthei culture

systemis No 119 Problen soils as potenial areas for adverse soils-tolerant rice No 108 (olptblication ofIRRI materials a survey of translators and varieties in South and Ssuthcasi Asia

publishers No 120 Changes isinall-farm iLtireliing techii)hgy in Thailand and [he No 109 Classification of Philippine rainfall patterns Philippines No 110 Contributions of modern rice varieties to rutrition in Asia No 121 Landforms and modern rice varieties No Ill Changes in rice breeding in 10 Asian countries 1965-84 No 122 Yield stability and modern rice technology No 112 Design parameters affecting the performance of the IRRI-designed

axial-flow pump

Page 19: UPLANDRICE. INSECTl--1i M T -E!R.EOLOU t RTANCE ,AND …delphacid.s3.amazonaws.com › 6958.pdf · The significance of abiotic factors in upland rice insect ecology will be apparent

IRPS No 123 January 1Q87 19

All this appeals to favor insect pest buildup but the notwithstanding the release of stored nutrients in response contrary is true Loevinsohn (138) showed that insect pest to drought that temporarily benefits philoem feeders populations respond exponentially with the proportion of land devoted to rice up to about 75 of the area then the M POR ANCE I I P1A 1) RWI P1St S

rate of response declines In most upland rico areas tie rice crop occupies less than 50i of the area- therefore the Visits to upland ricefields normally reveal 1kw insect pests potential for population buildup is low This relationship Loevinsohn (138) and Locvinsohn ct al (139) compared may not hold for seed pests that can locate small ricefields insect abundance using annual light trap atchcs in upland

Another outcome of the highly diverse flora and phical rainfed lowland and irrigated lowland sites in the Philipshyenvironment of upland rice isthateach upland ricearca has pines Although only one upland site was studied data a unique composition of insect pests showed equal or fewer insects at the upland site than at the

For example in Tanauan Batangas Philippines the rainfcd lowland sites and fewer than at the irrigated sites main upland rice pests are rice leaffolder(RLF) armvworm (Table 3) M Lltiple regression analysis showed that the key Aythntnaseparata WBPH ants Soli psis gentitataand factors explaining insect abundance among a set of cropping white grub Leucopholis irrorata Sixty kilometers away in intensity variables were the number of rice crops grown per Real Quezon the main pests are seedling maggot Athe- year followed by area devoted to rice Table 3 shows that rigona orae flea beetle (Thacitcmnra hasalis thrips the upland rice site had only I rice crop perym rand that rice Stenchaetothrips hiurnmis and rice bug Leptocorisa was planted in only 20Vi of the area Fertiliter and oratorius In a third Philippine upland rice site in Claveria insecticide use had little bea ring on the trend The armters at Misamis Oriental Mindanao the main pests are ants the upland site applied 60 kg N ha higher than at most Solenopsis gemtinata seedling maggot Alieriona orvzae irrigated sites white grubs Holotrichia in)iUaiuoan t and Lucopholiv Insect occurrence at the upland site was omparable to irrorata stem borers Sesamia ifiArelr (hio auriciliusC that at the Cagayan rainfed lowland site parlicularly in 1982stpessallis Scirpophaga incertulas S innoiata and when drought prevented many farmers fron planting(only

4c ona chrrsurapella root aphids Tetraneura niriah- 40(of the rice area eventually was planted that year) M ore dunrinalisand Rhupaosiphunm riahbdminalis lcaffolders favorable weather occurred in 1980 and 1981 which Marasniapataltis and Chaphalocrocis Medinalis army- supported greater rice areas and consequently more insects worm MIthinna separata and rice bug Leltucorisa The most intensive site for insect pests was South Cotabato oratoriusThe great diversity of insect problems is typical Mindanao where farmers plant 23-25 rice crops per year fer upland rice and makes control efforts more difficult with irrigation The Zaragoa Nutva Ecija site also had particularly regarding breeding for insect resistance high pest abundance and at this site the fields were highly

asynchronous Asynchrony was the third most important Low yield potential Iactor in explaining rice insect abundance Most upland rice is grown on low-nutrient and mineral- Rice cropping intensity in upland areas is bound to toxic soils which when combined with eiratic rainfall remain low Only one rainid upland crop is possible per make upland rice a highly riskycrop for the farmerto invest year (Farmers in Claveria Misamis Oriental Mindanao in costly soil amendments and land management Often Philippines tried asecond upland ricecrop i 1984a ycarof upland rice isgrown far from markets making inputs even favorable rains but no harvest was possible b-cause of more expensive Farmers growing a subsistence crop hosi- drought blast and birds) Rice area isaso bound to remain tate to invest in costly inputs to raise yields low because of the high labor and power rcquirelmenlts to

Even having varieties that could double existing yield prepare the land (because of weeds)and because upland rice potential under present management levels would probably wvill not compete with lowland rice as a cash crop Some not prompt many farmers to use purchased insecticide upland rice sites with prolonged rainy seasons howeverare unless tile risk of crop failure were reduced such as it was planted asynchronously such as the Clacra site with irrigation systems in lowland rice The seasonal abundance of Upland rice insect pests

The Tanauan Batangas site is atypical for upland rice attracted toa light trap wasgrapied for crop year 1980-81 in The soils and rainfall pattern are favorable and because of Tanauan FBatangas (Fig I 2) W13P11 was he most its nearness to Manila farmers have cash resources ftm common rice hopper and its population peaked at the sales of vegetables to purchase tertilier They applied an midgrowth stage as it does in lowland rice D-Vac suction average of 60 kg Nha after particle initiation and obtained samples showed an earlier peak than that from a light trap 25-35 tha yields The high fertility isone reason why RLF indicating emigration W13PI usually emigrate after the armyworms and WBPH developed into large numbers vegetative stage Populations in 1980 reached levels that Farmers however used no insecticide on upland rice caused patches of slight yellowing in some fields WBPH although they did on vegetables probably cause hopperburn in Batangas They immigrate

The poor growing conditions fOr rice generally mean year-round as shown by low levels in the light trap BPH poor growth and low fecundity for insect pests fIeding on it however were scarce in the light trap and not collected with

Table 3 Abundance of rice insect pests and factors in rice cropping intensity of Philippine upland rainfed lowland and irrigated lowland locations where light traps were operated daily for atleast I yr -I

0Rainfed Rainfed lowland Irrigated lowlandupland

Pangasinan Cagayan lloilo Santa Maria Nueva Ecija South Cotabato Tanauan Caaringayan Caaringayan Bangag Bangag Bangag Buray

LagunaCale

Batangas Manaoag 1980-81 Santa Motiica 1982-83 Cabanatuan Zaragoza Avancena NaniamanSolana 1981-82 1982-83 Oton

1980-81 1979-80 Oton 1981-82 1981-82 Koronadal Koronadal1980-81 1979-80 1979-80 1983 1983

Insect (kerosene light trapNephotcttixcatch nolyr)virescens 650 1240 500 1800 330 210 3400 1600 160N nigropictus 210 - 360 18400 5500 2900- 140 90 20 - shy 1700 -Reciliadorsalis 710 - 3200 1700180 890 630 140 170 2400 800Nilaparratalugens 130 690 - - 4200 4600300 520 750 180 60 3300 1600 740 310Sogatellafiircifera 1700 31000 13000 8200310 210 3300 610 270 980 1600 6300 -Cyrrorhinus - 12000 6700600 540 3200 290 160 330 120 410Scirpophaga spp 50 410 620 - - 21000 12000150 180 220 270 30 1000Cnaphalocrocismedinalis 70 70 540 610 270 11000 6100200 140 100 6 60 250Chilo suppressalis 0 60 60 1900 300 200- - 30 20 0 6 3 160 -C auricilius - 0 00 - - 20 3 60 20 3 0Sesania inferens 0 - 80 60 3 80 10 51 -Cropping intensity _ -

Rice crops (noyr) 10 10 12 10 10 10 13Rice area (7c) 17 19 20 20 25 2320 85 85 60 70 40 85 85 90Fertilizer (kg Nha) 60 20 85 80 70 7020 0 0 0 30 30Insecticide (no of 40 60 40 300 04 04 0 03 300 13 13 30 50 40 33 33applicationscrop)

__________

IRPS No 123 January 1987 21

Light trapped Insects Drytnd __ 2(notrap-nght) Suctioned Insects (nom )

50 -)o 9 jO40- SOW ffurciWO Light trp

D-Vc suction

20 to 4

20

Egg porasitizahion (M) 75 60

300 SOOampReOftwciffr

45 30shy

15

-insects (notrop-night) 4 Nileporvata Iuen5

Egg parastization()

Light trapped insects (no trop-nght) Suchoned insects (no m 2 )

6 8 Cytorh~nuls hvidipennii 0

4 05 2

50 0 Insects (noltrop-night)40-ui Ipanh pe g

Moohase 1 Seasonal abundance measured by light trap

Rainfall(etod) and D-Vac suction sampler of whitebacked

to ~ hopper Nilaparvata lugens and lea ffolder60)-t Cnaphalocrocis mnedinalis andplthpe g20 L I II I I I parasites and mtrid predator Ortorhjntus livi-Apr May Jun Jul Aug Sep Oct Nov Dec JaIn Feb Mar dipennis set against cropping pattern daily1 9 80o 19 8 1rainfall and moon phase Tanauan Batangas

1980-81

the D- Vac Cyrtorhinus build up on the crop coterminous are collected in light traps during full moon (109)with WBPH buildup Cyrtorhinus disperses during the Because upland rice shares most of the lowland rice pestsrainy season RLF not readily collected in light traps was and those pest species are generally less impotant to uplandprevalent only during the rice crop rice than lowland rice there is a general belief that insect

Nephotettix virescens was more abundant than Nnigro- pests are less important onpicLus in the light trap collections and on the crop GLH upland than lowland rice Arcview of limited data on upland rice shows an averag of population peaked at the reproductive stage as did zigzag 10-21 yield loss from insect pests on a par with that ofleafhopper (ZLH) Crtorhinus tracked the populations of lowland uee determined by the same methodology (132) planthoppers and leafhoppers Both GLH species were (Table 4) McGuire and Crandall (144) made a similarcollected year-round in the light trap indicating immigra- estimate for Central America Cramer (51) howevertion from nearby irrigated rice areas Possibly because of the estimated 35 yieldlow populations loss in South America based on ainsect number did not correlate with similar pest complex - stem borers and seed bugs shymoon phase although reports indicate that more hoppers attacking irrigated rice in North America This estimate did

22 IRPS No 123 January 1987

Light trapped insects D~~nd (no trap-night )

St plht Suctined insects (nom 2

)

D-Voc suction 1520

1o

0 Lih trap 5 ~105 o5

00 Eggparositizotion()5040 NephOtgtha virneens

30 20

I010 A-Insects (notrop -night)

4 Nephatelh nigropccent i

2 o I vV^J - -- ^ VA____ f _

Light trapped insects (no trap-night) Suctioned insects (no rnE

10 -1208 ReCdO doRsol

6 15

4 10 052 iwxALAiamp 0

Light trapped insects (notrap-night) 2Suc oned nsects (nom )

8 CyrtorhmnusI6o nn]10 i0075

4 05

2 0 0

025

2 Seasonal abundance measured by light trap Roinfall (mmd) Moon phase and D-Vac suction sampler of green leafhoppersNephotettix virescens and N ngropicrus and I zigzag leafhopper Recilia dorsalis their egg 60

parasites and micid predator Cyrtorhinus livi-dipennis set against cropping pattern dailyrainfall and moon phase Tanauan Batangas1980-81

40

5 Lf Apr May Jun

1980 Jul Aug Sp Oct Nov I

-aMar

1981

not account for soil pests and was not based on field data in the absence of pests Carbofuran granules are phytotonicAkinsola (I1) cited mainly data from irrigated rice areas in to rice (239) The prevailing hypothesis is that carbofuranAfrica but estimated yield losses of upland rice between 15 prevents soil bacteria from consuming fertilizer The phytoshyand 30 from insects However losses in Thailand over a tonic effect is through greater availability of fertilizer to the4-yr period ranged from I to 13 averaging 5(106) plant Denitrifying bacteria are known to be more active inYield loss data however should be interpreted with aerobic than anaerobic soils therefore it may not be merecaution (184) The Philippine data were all derived from coincidence that the highest recorded yield losses (22-69)trials on farmers fields under farmers normal agronomic occurred in those trials with 12-30 kg basal Nha On themanagement practices except for Bukidnon and Capiz other hand contro with carbofuran granules may not be aswhere researchers used fertilizer Batangas farmers are good as a seed treatment against seed pests and perhapsatypical because ofhigh levels of fertilizer used The trials in yields would have been higher if a systemic seed treatmentBrazil were carried out at an experiment station but insecticide had been used conditions were similar to those of local farmers The 1979 and 1980 trials in Batangas had two full-

Carbofuran granules were used in every trial as a broad- protection treatments one using carbofuran granules andspectrum insecticide applied basally with the seed An the other using bendiocarb WP as a seed treatment Plotsobjection to using insecticides to measure yield losses isthat protected with bendiocarb yielded higher than those proshyinsecticidescandirectlyorindirectlystimulateplantgrowth tected with carbofuran granules The other trials in the

Table 4 Yield losses to insect pests and yield responses to insecticide in upland rice in the Philippines Brazil and Upper Volta

Yield (tha) Yield loss (5) Location Year Cultivar Protected

with Protected

without Check Total Vegetative Reproductive Ripening

Basal N (kgha) Reference

carbofuran carbofuran Protected No with carbofuran

carbofuran

Philippines Pili Camarines Sur Tanauan Batangas

Pangantucan Bukidnon

Dumarao t-apiz

BrazilGoias Minas Ge~ais

Upper Volta6

1975 1974 1974 1975 1975 1976 1977 1978

1979

1980

1979 1980 1979

1977 1978 1979

Bursiging puti Kinanda Dagge Kinanda Dagge Dagge Dagge Dagge UPL Ri5 Dagge UPL Ri5 Dagge UPL RiS UPL Ri5 UPL Ri5 UPL RiS

IAC47 IAC47 IAC47

14 33 11 36 35 31 30 27 37 26 40 30 40 42 46 35

19 21 09

--

-34 24 24 27 --

29 43 31 42 -

47 25

---

10 28 05 27 27 30 26 29 33 28 33 29 43 23 36 11

15 15 06

29 15 55 25 23

3 13 -7 11

-7 18

3 -7 45 22 69

24 28 35

--

21 -11 -20

4 -

-3

23 7

-2 -23 56

-

-

-__0

6 --

-

-23 14

5 0 8 2

22

-

--

0

-

--18 11

3 -10

4 10

7

--7 -

14 -

-1 6

10 -5

8 9 7

-

--

0 0 0 0 0 0 0 0 0 0 0

30 30 30

12 12 12

99 99

100 100 101 101 102 102 102 102 103 103 104 104 104

66 66 66

Farako-Ba

Mean

1977 1978

IRAT10 IRATIO

38 66

--

40 43

-5 35

21

-

10

--

--

--

- 242 243

Z

tJ

-w

24 IRPS No 123 January 1987

Philippines which had two treatments offering protection In the 9 Philippine trials where yield losses were measuredwith and without carbofuran granules relied on further by growth stage 10 of the yield loss (21) occurred at thecontrol with foliar spray and not on seed dressings The vegetative stage and 517 occurred at each of the reproshycarbofuran-protected plots had an 18yield los compared ductive and ripening stages Ibis result is similar to thewith only 10 with foliar sprayS This yield difference is pattern of yield loss in lowland rice measured by successiveprobably due to differences in protection against sown-seed treatments where the crop is unprotected at each growthpests (foliar spray do not control sown-seed pests) rather stage but proticted during the other growth stagesthan phytotonic effects became basal fertilizer wa used in However the insect complexes r sponsible for vegetativeonly 2 of the 10 comparisons stage yield losses in upland rice are entirely diftfrent than in

The recorded yield losses were not related to yield lowland rice The key vegetative stage pests in lowland ricepotential (Fig 3) One might expect higheryielding crops to arc aquatic whereas those in upland rice are sown--seedhave higher percentage of yield losses but this ws not tile (sown in soil) root and seedling pests This dift rence maycase One reason is that insect damage exacerbates plant explain why agricul torists have overlooked tile ilmportanccinjury from drought stress low yielding fields from d ought of upland rice insect pests Many reports of upland rice pestsstress would show disproportionate differences between in Asia ha Ve focused on major lowland species (46 108)insect-protected and -unprotected treatments Shoot and It isalso surprising that tei vegctative stage of upland rice root pests would be more responsible for losses from would record the highest yield losses becatuseseeral studiesdrought stress than sown-scd pests A second reason have shown that upland rice can readily recover from foliageconcerns seed pests which would have a greater impact on reinoval during that stage (117 152 165 188) Upland ricelow tillering low yielding varieties such as )aggc Low poscsses a hitgh obilitV to compensate for ca rly loss oftillering varieties nnot fill in the space created by removal foliage and will even be stinulated by foliage removal toof seeds oweer 1) R i 5 is high yielding because it tillersP1 produce a higher viel than plants ithout any foliageactively which can close the canopy to provide greater weed removed (63 192 231) [ie reason for this apparentcontrol discrepancy hovce is that yield loss at the vegetative

stage does not octUr frol leat arca loss but fron pestsYield loss () removing sown seed and feeding on devcloping tillers and roots -- damage that cannot be readily conpLtnsatCd for

70 - Measuring yield losses in upland rice cnvironrnents is further complicated by a report from Brazil that greater yield losses occurred in response to better insect control60 - =-09 as Insecticide seed treatments resuled in denser plant stands

(n=21) which in turn created a selectively more favorable environshyment for blast (69) This resuIt was parlticurla rlxexacerbated50 when cirbofuran granules were used in seed furrows Aside

ffrom a greater plant stand ca rbofuran iay have auigmented the N lecl and stimulated blast If so ca rbofuran should be40 - replaced with another insecticide or the seeding or fertilizerI rates decreased

Other methods to determine the impact of upland rice30 shy pests wvcrc to interview farners or to develop single speciesbull bull correlations of population levels and yield loss Feneira (66)2 NIM)st (75(1) Golas farmers reporteddid both in Brazil

20- insect losses in ufpland rice and 251 said those lossesS avcragcd 37(1 [his result is consistent with tie yield losses 0 0 measured by field trials in the same statc(Table4) For every

I0 4 of seedlings removC d by leaf-cutting ants a I 7yield loss is predicted Similarly Ferreira (66) found that ai

0 - average of 53 thrips per panicle before panicle initiation meant 3 times more unfillh d grains

-IO0 I 1 1 I I-101 0 2 3 4 5 6 7

Yield (tho) C)N [ROt IM FI IODS 3 Correlation between upland rice yield and yield lols caused by Upland rice insect pests may be controlled by culturalinsects measured in 21 trials in the Philippines Ivory Coast andBrazil genetic biological and chemical methods

IRPS No 123 January 1987 25

Cultural control dry season plowing will have little effect on them Plowing Crop husbandry to reduce upland insect pest populations when the soil is wet from the first monsoon rains will expose can be classified into practices effective in a single field and soil insects that tunnel close to the soil surface to predatory those effective only at the community level Cultural birds chickens dogs and even man (field crickets nole controls should come first in a pest management program crickets and white grubs a re delicacies in man tra ditional They have broad stable effects as pests have little pos- diets) sibility of overcoming them through selection of biotyps The greatest degree i1 inIscct control fron tillage comes (138) On the other hand some cultu ral controls work only from plowing soon after rice ha rvest when most soil insects with high labor or power inputs and connunity-wilt lie close to the soil surface anrd stern borers are in the rice methods require coordination of iany farniters stuble Whitegrubsare mature and tillage exposesthem to

Pantin time The luxury of being able to shift the time of predators Tirnelv control will prtec the crop following pla ntring upland rice is open onl hr la rniers in areas with a rice and lowr the populationIfor succeedinig years Plowingprolonged rainifall pa ttern lI copping intensity or underthe ice stulble helps it decompose killing sten borer rnechanied land preparatiot Gcnerally planlting as soon larvae inside as the rFiinv season begins will lower populatiois of 1110st [requent tillage destroys ait nests bit not termites insect pests Insect populationsarelo aftcra dlvseoll or Tillage and clean culture howes er rcliove food for winter fallow delayed plaiting lets them build tip on termites which then mtay attack a young rice crop Scarab alIternate hosts that gi vigorotsly with the first seasonial beetles also prefer to miposit in eceitly tilled fields Noshyratiris This first flush iof weds cros lItxuttriartlv with iceCult is dibbling faors the survival oftillace Clre Such rineralied N released during the dr season aid made stils i aels tnd insects thtt pttpate or lay eggs- in the soil

available to the plants with the rains It iliso igrtmitcally Zero tilla hio eerl conserves predators ofthe GIH (37) advanttgeous for the rice ciop to tap this trlLItlaI fertility nt it mcttoi lo lessen the impact o0 soil insects and grow vigorotsly to compete with wrus and to tolerate brotdcasting seed or planting in ftltrrows is prefc ired over pest infestation dihlirig in hills Crickets are large enotlgh to eat seedlings in

Larly plantngs terd to escape seedling maggolts Upland a hill atrd tillering normnally ca iriot fill in tlhe missing space rice in Bitangas is planted in Ma and ine itli the Iirst Oftcn dibbll rd is not well covered and iseasily found by rains Fairners are highly mtotiated to plant early tocscape foraging insets ioIdents and birds drlitght and typlotns ricar harct Arn Atrgtst planting Plan uit sii Incrcatsini sced density protects tile cropwould be evcely attacked by It riim orivzo At tti againrst seed and seelling pests especially ants particularly other Philippiie tipland sites Rea l Qteot antd (laelit if the crop will germinate and ermerge quickly M isainis Oriental seedlig mnaggots arnrige ric boe t se hr trftinll itI India rice intcrclopped witi cotton or farnes have a longer growing season and plnit 2 rin or pigeon pci had lower 6IIt and WBIlII poptrlations than mote after the ortset of the rainy season liaaiangas ftrmners rice alone (211) NfailC arid Urplald ir_ 2re coilltori in the plant ca rly becia use ofa shorter raiinfall period arnd the desi re Philippines he chatngC in pest status folii iniercropping is to lliarvet before the typhoor sea son peaks ltighly locatioll-specific (133) cmianarid the net elfc be either

Early plantings cornbind with early-iatiuring varictics nil bcnelficial ortdetriential (189) For example BIatangias provide higher crp tolerartce for white grubs Aestiatior farmers intercrip Itro in their ipland ricefields A liornshyterminates with the first rains but 5-6 moare ineeded for the nrii iotd-Ahis i (I iiiacius) which norlially feeds resting larvae to develop into beetles emerge and lay eggs on taro ill feed or miarbv rice plants Intercropping and for larvae to reach the danriigii g ird-instar stage probably would not afft0 seed pests A crluanion crop (134 In lataigas this normually iseiough tiite for harvest planted ith rice and haing gleater tolerarnce for root

It Brazil howecver delayed planting is a suggested dariage or being toxic nayact isi trapcrop to root-feeditig cultural control for the lesser cornstalk borer and insect pests Acial-fei i ig pests voultId be most affected by pests favored l drought (66 195) Cropping internsity in intrer ropping Shoot aphids leaf beetles sten borers and Brail is low and delayed plraing places the rice crop in the hopplers would be likely ca ntidates to note effcts On the niost stable rainfall period reducing likelihood old rought othlt halnd the corlipalior crop stch~ sitniaie Iiiv be more

In rilniy arca s rinmirsare roi atl to plant other crops benehited than rice from initecropping ( 137) before rice and althNgh it woultl be desirabh to avoid 11echi Weeuling luring the first month after clop infestations by planting rice cuatl lariners lack the labor cstablishent will force pcts such is anyvworris which and time to do so ol a low priority crp prefer grassy cetds ontor tle vorling rice Soil iampsecs will

77age Uplild rice Stils ae light textured and Call be mvnye to the rice crop Cleani citrltirc also lay lore territes plowed duiring the d rv seasor notl only to grin time for to attack I u cetincti inrice ceds i ay he le optinal planting with lie fitst Inilsoon riiains but also to desiccate solition 13) soil insects arid weeds such is ntsedge that foim rhizomes Irtlizr Itneatsing N Iciiliier oin hlwland rice has Htowever becta use most soil insects lie below die plow layer fAtwvored Irigher populains of pliithoppeiv Imlhoppers

26 IRPS No 123 January 1987

and leaffolde rs However because of the need to temper fertilizer use on upland rice o prevent blast fIrtilizer rates will be lower than is comonia r for lowlad rice causing relatixely less hopper and IaIfTfoide r population buildup However inupland rice areas without a history of blast such Is Batangas farmers apply high rates of N and conscqtrently the crop has high infcstation levels of WBPI]and RIF The fhrmrcrs leec insect buildup by splitting the fertili11r in0to two to three applicatiois bit do not applV fertilizer at the vegetati estage because this prodrces a tall crop which illeasily lodge during typhoons Greater crop fertility leads to greater insect survival (better nutrition) larger insects (to cat no re foliage) arid greater fec titd t (to lav nmre eggs) (128 145 171) Not all insect species respondequally hovr Increasing rates (if N l anrid K increased Dialraa s-acchralis and (iaeiurema flea beetle but reduced thrips (66) Zn rcdLuced deadheart density from the lesser cornIstalk borer

Fnldingi Although most Upland rice areas do nIot hawxcaccess to wa ter floodiig the fields is rRrrnriierdcd in areas of Brazil or root bug and white gib control ( 195)

Crop rotatioti Tillage before planting a crop after rc harvest will control stern boreis and other pests renain n the stubble arid will unearth soil pests Planting a non-gratriiaceous crop after rice is recommncrded for termite control (66) Crop rotation is less nttdcd in upland rice culture than in lowlands because atcrop-free period is assured even after harvest of crops following rice

t1ulchilt iryl plastic irli irig designed for weedcoitrol ii upland rice inJapan resulted in ntr highanri liy infestation of (ihilo sulytresvais (93 9-1 95) NIulched Upland rice grew more luxuriantlv and attracted more (Avipositing mothis than u lched rice Also the rice plant and stein borers imatoured 10rC quickly and a partialgeneratiori af vouing ista iNoverwintcred in the stubbleLotili--malitrt varieti Quickly maturing crops reduce the tmbCr of pCst generations that Calt build up Rapid crop establishment olan ca rl v-natu ring vaitvty if)pa 11icLa r will lessen white gib and stem borer damiage White grub larvae intle last iistar are highly destIruCtive so aii early-iiatri ng variety will escape serious root loss Stem borers build 1ip slowlyx in rice to ecormic abundant in a late-Maturing crop anrd caiuse whi tlieads Numibers of somespecies increase exponeitially with each succeeding genera-ti so a xarietyexhat atures I trowh earier thian another will hac less insect damage Early-naturing vaiicties however will not lessen seed pest daiage either ii the soil or oina standing crop

Strwhoi hAIJttam Inrscts tcd irig on thlie aerial portions of rice plats disperse from carly to late plantedfields AS iiiibCirs increase exponentially with eachinsect

gtscnrattm e eiglbrig f rmc~s1(uil upp wh rc generation pests build tIP where neighboring fihrTC e9r1sstagger their plantings II Batangas farmers plant uplandrice within I iro use 120-d s-arieties and plant at the start of the rainy season Ini Misarnis Oriental laricrs stagger their

plantings up to 3 mo apart and use a set of atieties that mature in 4-7 mo Insect pest danage particularly from stem borers is greater in Misarmis Oriental Also seedling maggots damage rice there because rainfall favors frequent planting of maic In Batangas rice escapes seedling maggots bCca usC of distinct drs season and synchron0us early planting with the onset of ains

Therefore combinations of cultural practices ea rlyplanting synchronous planing crop rotation and earlyshymaturing vaneties protect the rice crop against most insect pests

Ilmant resistance Beca tse upland rice has few specialized insect pests and many insect types attack the crop worldwide regional breeding and strong internatiornal cooperation are required

Sown-sced pests arc economically controlled by insectishycide seed treatment and should the relore receive lowxpriori ty inbreeding objectives

Root pests are normally cost ly to control with insecticides but finding resista t sonres mayvbe di fficult Tanaka (229) for exa -ple fi led to find varieties resistant to root aphids Selecting for large root bioniass perhaps should be the strategy for pests sLuch as Whlite grubs that renliove roots Fainter(174) howcvcr cites varietiesof crops tolerant ofor resistant t(i root pests Some sorghun varieties are high in cyanide so rice sorghun might incorporate broadshyspcctruri chemical resistance

Seedling vigor and drought tolerance are high prioritybreeding objectives so tupland rices should have large r root systernsand be able to tolerate ligherlevels of root loss Sonie root loss inolder plants is beneficial if it stimulates new root developnrut and therefore crihances intake of soil nutrients (53) The rice seedling inaggot rtiLotia spp may be

controlled thirough resistant aittics A hi rgc breeding effort on sorghniu has had fair success inideveloping arreties esrstait to Asoccata Rodani (251) A field trial comparing 10 upland ricc varieties showed differences aniong varieties va ring from II to 4(- damaged tilleis (215) Hovver resistarce to 1Iill-0r gtia species may rot cross ox-rti otier

Shiraki (210) reported that upland i-ice stens wcrc harder than lows-land rice stemsperhaps becaIusc (f a higher silicoricontent and thus were more resistant to yellow stern borer hc tortia Iow tillering (f uplaid iice xarneties prodtres a

higher percentage of infested tillers than would occor in higher tillering lowland iice under a siiilar egg density The corollary therefore is to select for high tilleriig upland rices

Stemii bor amcstheh main targes dobrecders inAfrica (571 204 )but nticl oifth e sc reen irig is dotie unde r lo wland co d t osrm s wih A h r~ o l pe i s n acorditions (11 96 218) As with lthrinrnwspecies mayhave to be dealt with independently however initial results show cross resistance between African and Asian species(92)

In Brazil screening Ior insect resistance focttses oil lou I pest problems Varieties tolerant of lasmolwalpus ligi-sellus and resistant to )iatraea saccharalis have been identified (70 153) Current work also emphasies 7iraca limativetntris and Thmialspis (=Iosis) flavouicta (66) These insect pests Ilt onlV citise IIigh yield losses tihey also ire difficult to control chemically ofr by other ilealls

ManV upland rice vaieties aic early maturing and are resistant to pests such as gall midge (I Xl ) Early mIturity is often a highly desirable trait to esuilpe pest buildup but it is not true resista nce bca usc ilthose va reties are plinted late they will be itsdamaged as ain early planted susc ible variety

P~erhaps tile cosiiiopolitall of all upland rice pestsimost are the sced pests (ripenling stagc) which pose a particular problem beta use scuds re ised IsIO for hitnit4ans Ayltoxic substatice introd tcld geticticllyilmay also be toxic to nal iniless it caln he placed old in thfe IdhuffSelectmi fo hairiness or lotng wns kill mIlct sthilrohleiiis dutilig threshing pa rtictfarl maitial threshinfl [hireshers ill complain of itcfimin atid itteascd (fust levels

Attempts itIRRI to fitid tcistatncc to the rice bug plorisa have ilotbeCnl prmitisilg lIIetIattonlids iorc

through the lenilna or palca hillhairs sedstVIor seeds wilh awns that deter bugs promote complaints from threshers Seed bugs liptocorisa and Stcnocoris enter the seed through the opening between tile lemna and palC but varieties with narrow openings are diifficult to mill

Breeding priorities should concentrate ott inotCI gruLtLIps that are most difficult to c nttrol by other meains Steil) horers appear to be the most widespread group that sigiiificantly dalag s rice aind is hard to manage by other Ilnca us

Biological control Very fe stutdic of mitatt~l eneties of rice pests in upland envirollnletits wee in tile wefouiid literature Therefore have not compiled a list of species Ve present a list of natural enemies iOtid in upland ricefields tuic latialln Batangas for some conitno iOldiar pest species ([itble 5)

Muich ofl thc lollowinle itfor ittiol cotlliCs rol Stutdies in the lowlatnds or roi crops otithcr thlin tice

Qu(ailtr o 11ll t11t( Nat trrtf lii hcoeits ight Ile used agaitist ile social insects tilt and termites Sollie predator specice live ill theirf nests n ilnic theirnllt ip-pearanIlCe aMRidbehavior IJtrouMdCing a mote aggressise ant species to displace pest ant codonies is a possibility (169) Patihogens as potential natural encnies have reccived little atteition (I13) Natural ncnuics asa group however hac not been toIl iporta it in Iegulating aintshown be and ternite numbers

The nocturnal behavior aid subteitmien habitat of lole crickets and field crickets protect them from many ntllral enemies There are only limited records of natural cnelics for these groups of pests some pathogens (87) but mainly

IRPS No 123 JattuIny I 271987

parasites - scelionids on eggsand sphecids on ilymphs and adults Sphecid wasps Larra carutmaria (SImitI hI L huzwuensis Rohwer and lsaltiuhta Willi ams speciali in mole crickets pa ralyzing them nd dragging the ninto thcir nests as food lot their voting ()tier sphCcid species Lins aurulenoa (Iabricius) lot maiac (Ashmead ) 1 lhshylssellaus (Smith) and Ml lal)oriosu (Snithf specialie in field clickets

White grUbsO the other hand hae Icti scriuns enuigh to have been extensively studied for cotttroi 1W nati al enemies( 112 140 141) Scoliid parasites C(ajsoMcispp have beetn intrtduced itto tilePhilippites for control of Ielxl ]udisiro la The rcsearchcts w Intttrdced themle during ill era claimed that the white gttionthreak as controlled (142) howevcr it isnormal for pest epidemics to subside and there was no diect evidence to indictte that tfe parasite was responsible A tmore promisitg titctllod might he to identify iso ltcttass prt~f tcartd apply hiactelaln fingal pathogens to tile soil Ihe lalattese beetle in North America has hcci coitrolled hv tite conmnleciall aailable hactcrial prcpa ration itacihlpu pav ofr milksdiscas (71) Spote prtyefott the soil itloted ilrtier reitiain actic fr Veas

Root aphid sardtnctly bugs readilslic tilib to predatots and parlsites once their tetndine ants are controlled The larvac and adtilts of a coccinellid beetle 1Scynms sp prey on llriadlouais in slash and burn upland rice Baits treated with insecticide may be used to control tending ants by killing their ourig (244)

Root-tkcditig blugs false wireoris wireorms and root wccils aic r ofsuch little ifilpoltatice that thenomttll role of liatural Cticniics hias tiot beei asscssed A possibility hoevrl tiglt he tit ItaIIitic niaCIMtodcs lintt1g11en Itese soil-itlhlfbitine mCtitodcs W5uhtfhave to be imiass Produiced

large miltibcr of niatltrll cnemics ircknown ft CltIoriS IC IpblCeii is thtt clttvortnlls ofonlie the crop soott alter land ptel)tritiOti a1tithe bcgit iniig of the raittv

Citsoln hfien tiatllral leniiiy populations aric lo Colltloil WOtuld iii ole mass prodtticing andirclcaslng key spccies

le egg stage is the fpirt (olthe Cedlitgt Ialot life cycle tI hich to f)ctis ftor tittlril elctit cottntrol hplraites o1 pleditors ttsit elleggs highly expoIsYCl I Sgg pailsites or predators stolld probahly e ittass andtbe produced released Littlh i toruialioll is a ailable ott what species to trV o ho to Iniss podLice thiCIi A etiloplhid wasp Icmihlaol sp effctiicly regushy

lates the leatf lcloalapolm2a spioanoiiitr heat andc rice antid should be collsctvcd

leaf beetles whCllose e am] eggs ate laid in the grutindhirO ould be motC difficutlt to control by niatulral enelV Mnll ipuhltiOn But species sUici IS c_gsthispid beetles ith

and larVle oilleascs ate vtilcratble to iparaites and pledators

Natural enemisc of tlirips otl rice are little known

28 IRPS No 123 January 1987

Table S Parasites recorded from upland rice insect pests in Tanauan Batangas Philippines 1977-84

Order Family Species Host stage

I lymttenoptera Braconidae Leaffolders Cnaphalocrocis Marasinia spy

Cardiochilesphilipplinensis Ashmead a Larva

Cotesia angustillasis(Gahan)Cotesia or cypris (Nixon)

Larva Larva

Cotesia or taeniaticornis(Nixon) Larva Cilonusinotakatae Munakata Chelonits SppAlacrocetrts philippinensis (Ashtmead)

Egg to larva Egg to larva Larva

Iclineumonidac Orgilts spp Isclutojppa hireator(Fabricius)

Larva Larva

Itoplectis naraigav(Ashmcad) Larva Tenelhtcha philippiensis (Ashmead) Larva Tenelchha stangli (Ashmead) a

irichomma cnaphalocrosis Uchida Larva Larva

Bethylidac Elasmidac

Goniozms or trianguli r Kieffer lasmtusalhopictus Crawford

Larva Larva

Diptera

Encyrtidac Chalcididac

Tachinidac

Elasmtus spp Copidosomopsisnacohiae (Fidy) Brach nteria evcarinata Gahan Biachyieria lasus (Walker) Ztgohothria ciliata (Wulp)

Larva Egg to larva Pupa Pupa Larva

Argyrophylx nigrotihialis(Blaranov) Larva

l-ymenoptera Braconidac Yellow stern borer Scirpopiaga incertulas

Braco chinensisSzeplegeti Larva CIthelotis spp Chelonus niunakatae Munakata Cotesi flaripes Cameron Stenoracon nicerilli (Bingham) a

Larva Egg to larva Larva Larva

Ichneumonidac Tropobracon schoenobii (Viereck) Amatroinorpiaaccepta metathoracica(Ashmiead)Frihorus sinicus (I lolmgren)

Larva Larva Larva

Ischnojoppaluteator (Fabricius) Larva Isotima nr dammeratiti Rohwer Larva Tenelucha philippinensis (Ashmead) a

Larva Tenehicha stangli (Ashmcad)Trichomma enaphalocrosis Uchida

Larva Lara

Chalcididac Eulophidac Trichogranmmatidae

Xanthopimnpla stemmator (Thunberg) Charops hrachYpterumn Gupta and Malteswary Brachtm eria sppTetrasticttus schoenohii Ferriere Trichorigranm chihunis Ishi

Larva to pupa Larva Pupa Egg Egg

Diptera

Scelionidac Pteromalidae Tachinidae

Tt ichograintnta aponiicuitt (Ashniead) Telenomits rowani (Gahan) a

Trichoinalopsis apattelctena (Crawford) Jeirhaea spp

Egg Fgg Egg pupa Larva

Z4gobothria ciliata (Wulp) Larva

Ilymenoptera lecneumonirlac Gol fringed borer (hilo auricilius

Trichomm enoptalocrosisUchida Larva

Ieioenoptera Braconidae

Aantholimlh steiniator (Thtinberg) a

Pink stet Ioret Sesamia inferens StoInacoil nicrilhi (Bingh am)

Loarva to pupa

larva

Diptera Ilyinenoptera

Armvtnyorms Mythimna separata Spodoptera mauritia acronyetoidesTachinidae Zvgohothriaatropiora( Robinea i -Desvoid y)Eulophidae It-phctrtiscitapadoe (Ashimead)

Larva Larva

(halcididae Braconidac

Brachpin ria spip (oesia spp

Larva larva

IHyntenoptera I)iptera

Rice skipper Ielop ida (halcididae Tachinidae

math ias aidgreen iortedci ciliarNIelan itis eda Brat lhincria sp iii matuqitatt Cameron Irgyrolhirlaxi trotlhiais(marumtv)a1

ismtne Larva Larva

Diptera Ilymenoptera

Tachnidae lelineutionidae

fuown se tilo per Mois frugalisIrgtrophrlax 1igrothiahiv 3 a rat ov) anthopimpa pmnctata (Iabricius)

Larva Larva to pupa

aMost dominant

IRPS No 123 January 1987 29

Normally predatory thrips prey on herbivorous thrips strepsiptera on both Spiders dominate tilenymphaladultNatural enemies of stem bugs --Scolinoplara and Tibraa predators Spittle bugs move to rice from pasture grasses - are in order of importance egg par~isites and predators and biocontrol would be successful only ifcarried out before fungal pathogens and nymphal adult parasites Egg tiley immigrate into ricefields parasites could either be introduced fron other areas or Biological control tactics for hemipterous seed bugs mass produced and released ould follow those outlined for stein bugs

Armyworms are normally lwld in hcck by the activities Coml)arison of environnewtns Natural enemies are of egg and larval parasites When these natural enemies fail perhaps faced with even greaterconstraints than insect pestsusually because of drought armyworns become epidemic to survive ind reproduce in upland environments TheyIt may be worthwhile therefore to release parasites during have adapted ani survived by having wide host ranges the the rainy months following a drought Virus diseases of ability to aestivate or hibernate over unfavorable seasonslarvae would beanotheravnue to exploreforarnyworms and the ability to disperse or by other mechanisms

Parasites have ben traditionallyconsidered for biolegical Comparing the most dominant and therefore the most control attempts against stein borers usually by rekasing adapted species of natural enemies collected in various exotic species( 166) This approach has not met with success environments in the Philippines we can understand how and new avenues should be explored The role of predators different rice environments are arid pathogens is little inderstood and deserves greater Anagqrms olptahiisa mymarid planthopper egg parasiteattention Egg parasitism rates are normally reasonably is cquallyadapted to all Philippine rice environments (Tablehigh but should be supplemented with effective egg 6) It is selective for planthoppers and even parasitizes eggspredators Orthopterans are potential egg predators of the n-iize planthopper Peregrinitsmaidis itt upland areas Metioche vittaticollis (Sta ) Anavipha nifiwnnis (Saus- Alavohis also parasitizes maize planthopper eggs but is sure) and Conocephahs hongipnni (de i-laan) feed on most prevalent on BPH in the irrigated wetlands (onashystem borer eggs etioche and Anaxil)ha specialize in eggs tocerus spp like A optalhilis on BPH and WBPI Iattacks with no hair such as Chilo spp The omnivorous (ono- the eggs of GLH in ell rie environments It is also found cephahsspecializes in eggs cov red with hairs it is a proven parasitizing the eggs ef the white rice licahopper (oimna egg predator of Scirlouphaga spp but has the discouraging spetra Oirosita naias a trichogrammatid egg parasitehabit of eating rice guin (65) attacks BIPH and WBPH in the wetlands and has no other

Grasshoppers have egg (scelionid) and nymphalad ult host A related species 0 aesopi specializes in G1-H in (nemestrinid tachinid and sarcophagid) v irasites Among wetland environments Dryinid strepsipt-an and pipunshypathogens protozoans have been recorded most frequently culid nymphal and adult parasites of planthoppers and Control of locust species by natural enemies could con- leafhoppers also specialize either in hopper species or centrate on their habitual breeding areas environments Among the dryinids Pseuhdgonatoipsflavi-

Leaffolders have rich complexes of natural enemies some feimur prefers the lowlands and BPH while P nuch has no adapted to upland rice (22 27) ranging from egg predators environmental prefirences but is adapted only to WBPH (gryllids coccinellids) to larval parasites (braconids ichncu- The main d ryinid species on GLH aptogonatop)usspp is monids) to larval prcdators (ants and carabids) to larval most dominant in rainfed lowlands and also attacks other pathogeiis (vira and fungal diseases) Efforts to augment leafhoppers within Amnrasca Cicadulinaand Bachaltuathese natural enemies should focus on the egg predators Strepsiptera are more dominant on plarithoppers than (rearing releasing)and viral or fungal pathogens (culturing lealhoppers The Elenchus species in the Philippines attack spraying) all species of planthoppers within the genera Sogawilla

Parasitization of the large conspicuous larvae of Sogatodels NilaparsataHarualia and Opiconsiva Elenshyskippers green horned caterpillars and other polyphagous cusrvasuntaisuiis adapted to the major rice environments Lepidoptera by tachinids and chalcids is normally low ELg but prefers BPH in rainfed arid WBPIH in wetland environshyparasites and predators are perhaps the key to their control ments Ilalictoph(utsmtnroeiand HVpectrus attack G LH

Gall midges are parasitized by pterormalids and platy- and COfimravcctra respectivelygasterids that aestivate within their larval host between Pipunculids are mainly parasitic oii lealhoppers Each seasons Shoot aphids are preyed upon by coccinellids species has a unique environmental preference Tomosshysyrphids and chrysopids varireIa oraetoraprefers Ne7 otetltix nigropictrs and

Planthoppers arid leafhoppers have egg parasites and Pilunwulhtsjavanensisattacks Dehoceplhahls spp which predators as well as nymphialad nlt parasitesand predators are more abundant in upland than irrigated lowland their rich complex of natural enemies includes fungal environments T suhvirescensattacks Nephotettix virescens pathogens The principal egg parasites are myrnarids and and prefers the lowlands Pilpuiulus nttillattls is an trichogramniatids Egg predators are norinally mirid plant upland rice parasite arid attacks only Nephotetix speciesbugs that suck out the yolk Nymphal ad nIt parasites are More than 17 recorded species of leaffolder parasites in dryinids on planthoppers pipunculids on lealhoppers and the Philippines attack species of C(naplwocrocis and

30 IRPS No 123 January 1987

Table 6 Dominant natural enemies of rice pests in upland rainfed lowland and irrigated lowland rice environments of the Philippines

Natural enemy

Host stage Species

Brown planthop-Parasite Egg

MymaridaeAnagrus optabilis(Perkins) Anagrusflaveolus Waterhouse

TrichogrammatidacOligosita naiasGirault

NymphAdult Dryinidac

Pseudogotatopuslaiifemttr Esaki et Hashimoto Elenchidae

Elenchus yasurnatsui Kifune and Hirashima

Environment

Upand Rainfed lowland Irrigated lowland

Ailaparata lugens

x x X x

x

x x

x X X Whitebacked planthoppers Sogatella furcifera and Sogatodes pusanusEgg

MymaridaeAnagrus optabilis(Perkins)

TrichogrammatidacOl~gositanaias Girault

NymphAdult Dryinidae

Pseudogonatopusnudus Perkins Elenchidae

Elenchus yasutatsui Kifune et Hirashima

Parasite Egg

MymaridaeGonatocerus spp

TrichogrammatidaeOligositaaesopi Girault

NymphAdult Pipunculidac

Tomosvaryella subviresccns Loew Tomosvaryella oryzaetora (Koizumi) Pipunclusmutillatus (de Meijere)

DryinidaeIlaplogonatopusspp

Egg Copidosomopsisnacoleiae (Eady)

Larva BraLonidae

Cardiochiles philipphnensisAshnead lchneumonidac

Trichiomnia cnaphalocrosisUchida Temelucha stangli (Ashmead)

Pupa Chalcididae

BrachymeriaexcarinataGahan Brachymerialasus (Walker) Brachymeria nr tachardiaegroup

Parasite Egg

Scelionidae Telenomus rowani (Gahan)

EulophidacTettasticusschoenobhi Ferriere

Larva Braconidae

Stenobraconnicevillei (Bingham) Iclhncurnonidac

Temelucha phiiippinensis(Ashinead)Temelucha stangli (Ashmead)

x x x

x x

X x X

x x Green leafhopper Nephotettix virescens

X x x

x x

X X X X

X

X Rice leaffolders CnaphalocrocisMarasmia

x X x

X

X X

x x x x x X x x x

Yellow stem borer Scirpophaga incertulas

X X x

x

x

x X XAinauromorphaaccepta metathoracica(Ashmead) x

continued on next page

IRPS No 123 January 1987 31

Table 6 cntinued

Natural enemy Environment

Host stage Species Upland Rainfed lowland Irrigated lowland

Larva-pupa Gold fringed borer Chilo attriciiuslchneumonidae

Xanthopimpla stemmator (Thunberg) x aPupa x

Pteromalidae Trichoinalopsis apanteloctena (Crawford) ax

Pink stem borerb Sesamnia inferensParasite

Larva Braconidae

Stenobracon nicevillei (Bingham) x b b Armyworms Mvthinna SpodopteraLarva

Tachinidac Zygobothriaarroptvora (Robineau-Desvoidy) x x

Braconidac Cotesia spp

X Skipper Pelopidas mathias and green horned caterpillar Melanitis leda ismeneF-gg

TrichogramrnatidacTrichogrannaspp x

Scelionidae Undetermined Scelionid

x Larva

Tachinidae Argyroph lax nitrotibialis (Baranov) X x

Pteromalidae Trichomnalopsis apanteloctena (Crawford)

X Predator

CoenagrionidaeAgriocnenis femina femina Brauer x xIschnura senegalensis tambur

Miridae CrtorhinsliidipennisReuter x x

Nabidac Stcnonabis tagalicus (Stal) x

VeliidaeAlicrovelia douglasi atrolineata (Bergroth) x

Coccinellidae Micraspis crocea (Mulsant) X x x

Vespidae Eumenes campaniformis (Fabricius) x

Formicidac Diaecamna spp xOdontoponera transversa (F Smith) x

Carabidae Chaenius spp OphioneaW- aJasciataSchmidt-Goebel

x

Ophionea ishii ishii Habu x x X

Gryllidav Anaxipta longipennis (Saussur-) x xMetioche ittaticollis (Sial) x x x

Tettigoniidae Conocephalus longipennis (de Haan) x x x

aNo rearing made on rainfed lowland bNo rearing made on rainfed lowland or Irrigated lowland

MrasmiaAmong the Laval parasites the braconid Cardio- lowlands and parasitizes oriental maize borer OsIriniachilesphilijpphensis is most adapted to an upland environ- furnacalis (Guenee) larvae The leaffolder pupal parasitesment It also parasitizes Hydelepta indicata a common not only occur equally in all environments but also haveleaffolder of legumes The ichneumonids Trichomma cna- wvide host ranges- parasitizing species of OsIrinia Marucaphalocrosis and Temehcha stangli occur more in the Hydelepta and Honiona - in maize and legumeslowlands T stangli is most adapted to irrigated lowlands The principal egg parasite of yeltow stem borer (YSB)and also parasitizes Chilo and Scirpophaga stem borer Telenomus rowani is widely adapted to all environmentslarvae T cnaphalocrosis is most prevalent in rainfed and has an unusual alternate host -Tabanus eggs (25)

32 IRPS No 123 January 1987

Tetrastictu schoenobii specializes only in YSB and occurs mainly in irrigated lowlands -- further proof that YSB is most adapted to the wetter environments

Larval parasites of YSB are highly environment-specific The braconid Stenohracon nicevillei occurs only in upland rice but also attacks the pink stem borer (PSB) Tieinchwa phiiiplPwnsis an ichneumonid occurs in upland rice and also parasitizes the larvae of (7hiu stqpresaisinatro-morpha accepta melaihoracica like 7 stangi is mnost adapted to the irrigated lowlands

The dark-headed stem borer(l)HSB) (ih pochit3sus was not collected in all rice environments and the gold fringed borer Chilo auricilius nd PSB were collected only in upland rice Xaantiholiplastenrnaloran ichneumonid larval parasite appears to be widely adapted whereas TrihomaslopisapanitI)cteiiaa pteromalid pupal parasite occurs mostly in wetlands and also parasitizes larvae of Chilo suppressais and Pelhuidas mathias

Armyworms Myit hinin and Spodujipera have two main larval parasites - a tachinid Zrgohothriaatropivora in rainfed environments and a braconid Cotesia spp in irrigated lowlands

The butterfly pests Pelopidas and 4elanitishave tricho-grammatid egg parasites found only in irrigated lowland environments

Two larval parasites show environmental preferences --Argrroph av nigrotihialis a tachinid in rainfed habitats and Trichonialopsisapantelkcnela in irrigated ricefields A limgrotibialisalsoattacks the sweet potato hornwormAgrius convolvtdi and T apanteloctena parasitizes DH S1B

Predators as a group show more distinct environmental preferences The aquatic damselflies Coenagrionidae are most abundant in the lowlands Cvriorhinus is also most adapted to the lowlands The nabid Stenonabis tagalicus occurs only in the uplands and isalso prevalent on legumes The coccinellid lady beetles wasps and ants are more prevalent in upland areas AlicrasIpis cit-ea larvae prey ona variety of aphids attacking legumes and maize Etunienes campanonia vespid mud wasp makes nests in trees and is therefore most adapted to the more botanically diverse uplands Soil-dwelling ants cannot tolerate flooding

Upland environments are habitats to arboreal carabid beetles Three species of Chlaeniusbeetles prey on leaffolder larvae and are perhaps more important than parasites (27)

The gryllids and tettigoniids are widely adapted egg predators not only on rice but also on maize and legumes They feed on eggs of most insects that are laid on leaves

The spidercommunity of upland rice environments isrich in species At one site (Batangas Philippines) 31 species have been recorded (23) Of the 176 spider species recorded in Philippine ricefields about one-half (82) occur in upland rice The spiderspecies of upland rice environments overlap more with those of rainfed lowland than with those of irrigated lowland rice (24) The same study showed that of the three environments irrigated rice has (ie greatest spider species diversity foiiowed by rainfed lowland and upland

However of the 10 most prevalent ricefield spiders 9 were abundant in upland rice showing wide adaptation All three spider guilds -- orb-web space-web and huting spiders shywere prevalent in upland ricefields There were however differences in environmental prelerences for some spider species

Of the orb-wcavers Tetragniathamanlihulatawas partishycila rlv abundant 7 japonica was low and Ltucatge dehcorata was absent in upland rice

As in other environments Aiypeiwnaifirtosana was the most dominant space-web spider however its relative numbers were lowerthan for lowland sites Among hunting spiders Ox|opes javanus and Lcosa leatcost fgla were more abundant and Lvcosa lpseudoanmnlata was less abundant in upland compared with lowland rice envishyroniments

Extensive rearing of rice insect pests in upland rainfed lowland and irrigated sites over an 8-yr period in the Philippines provided insight into the effectiveness of parasites as natural enemies (Table 7 8) Most of toe data are from large sample sizes taken over at least one crop which overcomes the pitfalls expressed by Van Driesche (238)

VSB egg parasitization was surprisingly similar across all environments Larval parasitization was highly dynamic among sites and even years within the same site but showed somewhat lowerlevels in uplandsthan in irrigated wetlands This would have been predicted for egg parasites as well since YSB is adapted to the lowlands and so must be its parasites This isevidence that Telenomus rowani is as good at dispersing as its main hosts Trowani clings onto the anal tufts of female YSB and parasitizes eggs as they are laid This phoresy may explain its wide adaptation The more specialized lowland YSB larval parasites therefore appear more effective than their upland counterparts with more alternate hosts

Low collection levels of PSB larvae especially in the wetlands make comparisons between environments difficult Low levels of larval parasifization were recorded in both rainfed environments Except in the Cagayan rainfed wetland site in 1980 low levels ef larval parasitization were evident at irrigated and rainfed sitesalike No differences by environment were evident Higher levels of parasitization occurred in Sarawak during a 1967 outbreak also from tachinids Rothschild (197) indicated that higher parasishytization occurred in upland rice areas

Skipper and green horned caterpillar larval parasitization rates were generally low and showed no environmental effect htighest activity occurred in Cagayan in 1980 (48 parasitization) Although larval parasitization levels of RLF were variable year to year activity appeared greater in the irrigated lowlands than in the uplands Again the highest levels were recorded in Cagayan in 1980 Cagayan has extensive grasslands and fallow around the rice areas and a relatively long rainfall period Rainfall in 1930 was good for crop growth and apparently for increase of larval parasites

IRPS No 123 January 1987 33

Table 7 Parasitization rates of rice planthoppers and green leafhopper in upland rainfed lowland and irrigated lowland environments Philipshypines 1976-840

Environment and site Year Egg parasitization () Nymphaladult parasitization ()

BPH W13P GLH BPII WBPH GLIH

UplandBatangas

Rainfed lowland

1976 1979 1980 1982

15 17 12

2(47)8(39)

4(51)

9(1800)14 (212)

12(304)

8 (101)6 (127)

9(116)

Iloilo

Cagayan

1976 1977 1978 1981 1980 1981 1982

0 (69)

11

44 48 40

-

-

46 55 44

0 (471)

11

52 56 49

0 (200)9 (214)

10 (1259)8(235)6 (900) 3(800)

-7 (462)

11 (270)3 (313)5 (900) 1 (800)

4 (600)10 (725)10 (362)12(870)5 (900) 3 (800)

Pangasinan

Irrigatedlowland

1984 1976 1978 1979 1982

13 (436) 13(379) 14 (457)

6(14)7 (197)

17(198) 11 (486) 18 (275)

3 (81)10 (274)3 (116)

16(871) 19(88)

28(211)9 (764)5 (647)7 (1204) 8(305)

Laguna 1977 11 7 21 1978 31 1979 14 4 4 28 1980 7 8 20 1981 1983

56 7(278)

26 16 (1650)

16 27(1921)

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R I Apostol unpublished 1lo1lo and Pangasinan 136 Cagayan BCanapi unpublished Liuna 175 I C Pantua unpublished 118 234 43 Carino and Shepard unpublished BPH = brown planthopper Nila parvara htgens W131Pi =whitebacked planthopper Sogatelia furcifera GLH = green leafhopper Nephotettrix virescens Figiires in parentheses are sample sizes

Tauher et al (230) reported that internal external and genetic factors influence the seasonal activity of parasitoids

Hopper egg parasites but not nymphaladult parasites were also parlicularly abundant at the Cagayan site possibly the egg parasites have more alternative hopper prey in the weedy areas than nymphaladult paisites

The seasonal dynamics of hopper egg parasites can be seen for upland rice in Batangas(Fig I 2) and for rainfed lowland lice in Pangasinan( 135)and Cagayan( 105) Levels of BPHWBPI-L and GLH egg parasitization were corn-parable in upland and irrigated lowland environments Except for some high rates of paraMsitization early in the season the parasitization rate tended to be steady in all envilonments However hopper nymphaladult parasi-tizution levels were more similar in the rainfed environments and lower than in irrigated rice

Overall parasites do not appear highly effective by themselves against upland or even lowland pests Among natural enemies parasites have attracted the most attention because they can be more readily assesscd

Predators and pathogens lend themselves better to management practices Predators the most important group of natural enemies in riccfields are difficult to quantify On the other hand pathogens appear less im-portant but can be readily cultured and disseminated

The natural enemy commulnity of upland rice is rich in species and differs significantly from that of lowland rice

These beneficial organisms must be conserved by applying insecticides judiciously particularly sprays Seed or soil placement of chemicals minimizes exposure to natural enemies The strategy to derive the greatest benefit from natural enemies isto allow the greatest number of beneficial species to thrive Then perhaps at least one species will be effective against each pest at any tine overcoming the variable abundance of each species within and between years

Programs to introduce exotic species or mass produce indigenous ones are ambitious and expensive because they require trained people on a sustained not ad hoc basis

Chemical control Insecticides offer rapid and efficient control of upland insect pests (190) but-should be used only after other control measures have been considered Insecticides are rarely used on upland rice because of cost (32) Upland rice yields are normally too low to justify the expenditure Also most upland rice is grown as a cash crop Spraying upland rice is more difficult than lowland rice because water is less accessible Government subsidy programs to provide pestishycides to tribal peoples for upland rice production have created problems of toxicity to humans because isolated people have had no experience to be able to handle pesticides safely

However ins-eticide use has beenjustified when a crop is

0Z

Table 8 Parasitization rates of lepidopterous pests in upland rainfed lowland and irrigated lowland environments Philippines 1976-84a

Parasitization (5)

Environment and site Year YSB GFB PSB Earcutting Swarming Skipper Greenborned RLF Brown larva larva caterpillar caterpillar caterpillar caterpillar larva semilooperEgg Larva larva larva larva larva larva

UplandBatangas 1977 - 0 (57) - 0 (63) - 0 (71) - 0 (106) 11(142) 2 (101)1978 43 (4697) 8(451) 5 (38) 3 (40) 6 (61) 2(58) 15 (57) 13 (129) 13 (1286) 4 (30)1979 59(4135) 12(320) 13 (11) 5(28) 9(45) 6(46) 7(23) 8(160) 9(1152) 8(26)

Rainfed lowland Iloilo 1976 - 0 (10) 0(10) 0 (7) 0 (19) 2 13) 15 (180) 5 (76)1978 0 (54) 0 (3) 0 (17) 9 (64) 4 (78) 3 (73) 0 (121) 0 (191)1979 64 (2350) 16 (84) 6 (17) 0(6) 18 (68) 25 (104)

1981 56 (1322) 18(304) 0(5) 12(51) 18(138)Pangasinan 1976 60 (7 7 )b 62(82) 0(111) 5 (39)

1978 58 (53)b 5(51) Cagayan 1979 59(713) 8 (101) 6 (34) 0(7)

1980 0(11) 4 (28) 15 (286)

62 ( 48(177) 61(140)1981 0(5) 12 (2) 13(45) 14 (92)

Irrigatedlowland Laguna 1974 62 8

1974-75 50 41 1975 43 72 1978 96 (623) 21(24) 33 (60)1979 83(2017) 18(110) 8(62) 11 (89) 22(400)1980 78 (1625) 20(75) 0 (16) 4 (72)1983 60

9 (80) 7 (270) 30

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R F Apostol unpublished Iloilo and Pangasnan 136 Cagayan B Canapi unpublished Laguna 175 P CPantua unpublished 118 234 143 Carino and Shepard unpublished YSB = yellow stem borer ScirpophagaincertulasGFB = gold fringed borer Chilo auricilius PSB = pink stem borerSesamia inferens earcutting caterpillar Alythimna separata swarming caterpillar Spcdoptera mauritia acronyctoides skipper Pelopidas mathias reen horned caterpillar Melanitls ledaismene RLF = rice leaffoldersCnaphalocrocismedinalis and Marasmiaspp brown semilooper Mocisfrugalis Figures in parentheses are sample sizes Egg masses

IRPS No 123 January 1987 35

Table 9 Effect of seeding rate and insecticide on plant density and Yield losses from insect pests in upland rice often areyield of Dagge upland rice Batangas Philippines 1976a comparable to those of lowland rainfed or irrigated rice

Seeding rate Insecticide Dosage Plant density Yield (0-Y30o) but usually from different pests The principal (kg aiha) seed treatment (kg aiha) (noIra-row) (tha) groups are soil pests feeding on sown seed and rootsat 14 DE followed by seedling pests These groups have largely gone

50 None - 35 a 20 a unnoticed leading researchers to conclude that stem borers50 Carbofuran F 10 38 ab 23 a and seed bugs are the most important pests (17) Yield loss 10 beCabofuranF100 None - 4551 c 23 a trials that pinpoint major pests need to includc seed

100 Dieldrin WP 10 52 cd 22 a treatments Granules applied in the furrow may not give150 None - 59 d 22 a adequate protection from many seed pests

aAv of 8 fields DE = days after crop emergence In a column means The floristic and pedological diversity of upland rice followed by a common letter are not significantly different at the environments is matched by the diversity of arthropods5level by D)MRT F = flowable Wi = wettable powder both pests and their natural enemies Irrigation has homoshy

genized lowland rice environments Flooded soils tend tothreatened by epidemics of locusts or armyworms New have the same properties and to eliminate soil pests The technology stressing efficiency has minimized insecticide spatial and temporal dominance of cultivated rice has cost benefited specialist pests

Seed treatment is normally inexpensive and can be Upland rice has no specialist arthropod species The economicallyjustified in -nanycases(66) Treating seed with monophagous pests on upland rice prefer the lowlands systemic insecticide (216) but not chlorinated hydrocarbons where rice evolved Rice in the uplands exists only because (247) is most effective in controlling seedling maggots of man and even though lowland rice predates upland riceFoliar sprays have to be repeated to obtain control and specialist species evolved Wheat maize and sorghum do often are washed off by frequent early season rains not have monophagous pests leading us to conclude that Granular insecticides are effective but required dosages are there is no evolutionary advantage for inssects to have too expensive to justify except under heavy infestation narrow host ranges in upland graminaceae(247) When directed against first-instar larvae banding Pests have adapted to the uplands not only by having granules in seed furrows (177) can be effective against white wider plant host ranges but also by having long life cyclesgrubs at low dosages (025 kg aii ha) (134) Seed treatments undergoing dormancy or being dispersive Each pest has a are not promising for white grub control (248) unique life history strategy suited to highly unpredictable

Seed treatment is also effective ainst ants and may be environments In agivenyear someinsect specics an better cheaper than increasing the seeding rate In Batangas exploit upland rice as a food source but no pest is a senious treating 50 kg seed ha was as effective as increasing the one every year on upland rice seeding rate to 100 kgha and using untreated seeds (Table The low yielding potential of upland rice means less 9) In Brazil seed treatments control termites and other soil quantity and lower quality of food for pests to exploit but insects including the lessercornstalk borer(66) Domiciano also less incentive for farmers to undertake control (58) however found seed treatments inconsistent in their The most effective control would be early and synchroshyeffect against high populations of lesser corn stalk borer nous planting of early-maturing upland rice varieties to

Baiting can be an inexpensive way of controlling ants prevent insect pest buildup Breeding for insect resistance seedling maggot flies mole crickets and field crickets Baits should initially focus on stem borers because they are verymade of locally available material and impregnated with difficult to control by other means Biocontrol efforts insecticide can be sparingly distributed taking adantage of should first stress conservation of natural enemies preshythe pests ability to disperse and encounter bait sites dators and pathogens rather than parasites Efficient

The new low-volume hand-held controlled-droplet chemical control includes seed treatment or baiting for seed sprayers may offer an advantage over high-volume knap- seedling and some root pests and using controlled dropletsack sprayes for controlling foliar pests Upland rice sprayers for spot treatments against foliar pests and seed canopies are more open than those in lowland rice and bugs when economic thresholds are reached better droplet penetration should result Controlled droplet applicators would be ideal for seedbugs and defoliators

CONCLUSION

Upland rice is attacked by a wider array of insect pests than is lowland rice mainly beca use of the addition of soil pests Generally population levels of lowland rice pests are lower in upland than lowland rice leading many people to conclude that insect pests are not important on upland rice

36 IRPS No 123 January 1987

REFERENCES CITED

1Abraham C C and K S R Mony 1978 Occurrence ofLeptocorisa acuta Fabr (Coreidae Hemiptera) as a pest of nutmeg trees J Bombay Nat ttistSoc 74553

2 Agyen-Sampong M 1975 HlierogrlphusdiaganensisKrauss (Orthop-tera Acrididae) a new pest of rice in Northern Ghana Ghana J Agric Sci 8249-253

3 Agyen-Samnpong M 1976 Varietal resistance and other inle-ratcd pest control components as factors of rice varietal improvement in West Africa Prcsented at the Second varietal improvement serinar 14-18 Sep 1976 Bouak Ivory Coast West Africa Rice Development Association Monrovia Liberia 18 p

4 Agyen-Sampong M 1977 Outbreak of Ihccrorrdnisorae in rice seedlings near Rokupr Sierra Leone lit Rice Res Newsl 2(6)16-17

5 Agyen-Sampong M 1980 Dirty panicles and rice yield reduction caused by hugs IntRice Res Newsl 5(l)11-12

6 Agyen-Sampong M 1982 The major pest problems of irrigated upland and mangrove swanip rice ecosystems ir the humid tropical Guinea savanna and Salel climate zones Pages 318-328 in Pro-ceedings training course cn concepts techniques and application of irtegrated pest management in rice inWest Africa 10-20 Jan 1982 Fendall Liberia West Africa Rice l)evelopioent Association Monrovia Liberia 505 p

7 Akibo-Betts V I and S A Raymundo 1978 Aphids as rice pests in Sierra Leone lit Rice Res Newsl 3(6)15-16

8 Akinsola F A 1975 Present status of different rice stemborers in parts of Nigeria iicL Entomo Newsl 328

9 Akinsola FA 1979 Ihe biology and ecology of rice stemborers in Nigeria Ph 1) thesis University ol lbadan Nigeria 217 p

10 Akinsola E A 1979 Guidelines of integrated management of insect pests of upland rice in West Africa Presented at the Seminar on integrated management of rice diseases and insect pests at Bobo-Dioulasso Upper Volta 17-22 Sep 1979 West Africa Rice Develop-ment Association Monrovia Liberia

II Akinsola IA 1984 Insect pests of upland rice in Africa Pages 301-305 in An overview of upland rice research Proceedings of the 1982 lIouak Iwsrv Coast upland rice workshop International Rice Research Institute 13O Box 933 Manila Philippines

12 Alam M S V TJohn and Kaung Zan 1985 Insect pests and diseases of rice in Africa Pages 67-82 in Rice improvement in Eastern Central and Southern Africa proceedings of the international rice workshop at Iusaka Zambia 9-19 April 1984

13 Alticri M A A van Schoonhovcn and J D Doll 1977 The ecological role of weeds in insect pest management systems a review illustiated with bean (Phaseohsvulgaris L)cropping systems PANS (Pest Artie News Summ) 23195-205

14 Anmante F 1967 The leaf-cutting ant Atia capiguaraa pest of pastures [in Portuguese] Biol6gico (Sao Paulo) 33113-120 (Rev Appl Entomol A56207)

15Andrade A C and D Puzzi 1951 Preliminary results of experi-merits on the control of Sculptcoris iastantus on sugarcane [in Portuguese] Bioulgico (Sao Paulo) 1744-49 (Rev Appl Entomol A41421422)

16 Appert J 1970 Maliarplia separaelha (rice stem borer) new observations of entomological problems of rice in Madagascar [in French] Agron Trop (Paris) 25329-334

17 Ariadeau Mand Z Harahap 1986 Relevant upland rice breeding objectives Pages 189-197 in Progress in upland rice research Proceedings of the 1985 Jakarta conference International Rice Research Institute 1O Box 933 Manila Philippines

18Asar S and Kisimoto 1975 Seasonal occurrence of the smaller brown planthopper in upland rice [in Japanese] Proc Kanto-Tosan Plant Prot Soc 2228

19 Bakker W 1974 Characterization and ecological aspects of rice yellow mottle virus in Kenya Meded Landbouwhogesh Wage-ningen 152 p

20 Balut F F 1970 White grubs attacking rice Oryza ativa L [in Portuguese] Biol6gico (So Paulo) 36321-322

21 Banerjee S N and L M Pramanik 1967 The lepidopterous stalk borers of rice and their life cycles in the tropics pages 103-124 in The major insect pests ofthe rice plant Proceedings ofa symposium at the International Rice Research Institute September 1964 The Johns Hopkins Press Baltimore Maryland USA

22 Barrion A T and J A Litsinger 1980 Ants a natural enemy of cliaiarac in dryland rice IntRice Res Newsl

5(4)22-23 23 Barrion A T and J A litsinger 1981 The spider fauna of

Philippine rice agroecosystem 1Dryland Philipp Entomnol 5139-166

24 Barrion A T and 1 A Iitsinger 1984 The spider fauna of Philippine rice agroccosystcms II Welland Philipp Eiitomol 611-37

25 Barrion A T1 and J A Litsinger 1984 lihanus (Diptera Tabanidae) eggs an alternative host of rice stei borer (SI3) egg parasite Telenoonis digns (Iliymenoptcra Scelionidae) lit Rice Res Newvsl 9(6)19

26 Barrion A T and J A Litsinger 1985 Identilication of rice leaffolders by wing markings Int Rice Res Newsl 10( 1)24

27 Barrion A T and J A Litsinger 1985 Chctiosspp (Coleoptera Carabidae) a leaffolder ILF)predator lit Rice Res Newsl 10(1)21

28 Bertels A 1970 Insect pests of rice and theircontrol [in Portuguese] Pelotas Brazil Instituto de Expcrimentacao Agropccuararias do Sul Circular 43 24 p

29 Bonzi S M 1982 Ctilo (flisilineus J de Joannis (Lepidoptera Pyralidae) stem borer of irrigated cereals in Upper Volta LioFrench] Agron Trop Riu Rizic Cult Vivrieres Trop 37207-209

30 Bourke T V T L Fenner J N LStihick G Baker EHassan D F OSullivan and C S li 1977 Insect pest survey for the year ending 30th June 1969 Papta New Guinea )epartment of Agrishyculture Stock and Fisheries Port Moresby 57 p

31 Bowden 1 1973 Migration of pests in the tropics Meded Rijksfac Landbouwwet Gent 38785-796

32 Breniere J 1973 The main rice parasites in West Africa and their control Pages 27-42 in Proceedings seminar on plant protection for the rice crop West Africa Rice Development Association Monrovia Liberia

33 Breniere J 1976 The principal insect pests of rice in West Africa and their control West Africa Rice Development Association Monrovia Liberia 52 p

34 Brown ES 1962 The African arinyworn Spoiolptera eXI)Ina (Walker) (Lepidoptera Noctuidae) a review of the literature Cornshymonwealth Institute of Entomology London UK57 p

35 Brown F B 1968 Upland rice in Latin America FAO-International Rice Commission 12th Session of the Working Party on Rice Production and Protection 2-14 Sept Peradeniya Sri lanka Food and Agriculture Organization Rome Italy 6 p

36 Buyckx F J 1 1962 Disease and insect pests of crops inCongo Rwanda and Burundi [iil French] Publications dc LInstilut National pour lestude Agrononiquedu Congo M Weissenbruck S A Brussels Belgium 708 p

37 Cariiio F 0 1981 Role of natural enentics inpopulation suppression and pest management of green rice lealhoppers Ph D thesis University ofthe Philippines at Los Bafitos PhilippinesLaguna 136 -

38 Carpente A J 1983 The development of rice cultivation and extension of vegetable productin in Zanzibar Final Report Vol II1 October 1983 Food and Agricultme Organization Ronte Italy

39 Carroll C P and S1 Risch 1983 Tropical annual cropping systems ant ecology Environ Manage 751-57

4(0 Cates R G 198 1 Host plant predictability and the feeding patterns of monophagous oligophagous and polyphagous insect herbivores Oecologia (Berl) 48319-326

IRPS No 123 January 1987 37

41 Cendcafia S M and F B Caora 1967 Insect pests of rice in the 62 Ene 1 C 1981 I)iiorldhol)ierusihilis Slater (licteroplera Philippines Pages 591-616 in The najor insect pests ofthe rice plant lygacidue Illissinac) a new rice pest front Nigeria lit Rice Res Proceedings of a symposium at the International Rice Research Newsl N3)15 Institute September 1964 The Johns Hopkins Press Baltimore 63 Fugenio T S 1965 rhe effect oi e-vCgetation on rice production Maryland USA 729 p and the possibility of overcoming it by leaf pruning MS thesis

42 Centro Ihternacional de Agricklittira Tropical 1984 Upland rice in University ol the Philippines at Los Baijos ILaguna Ihilippins 69 p Iatin Anerica Pages 93-119 hi An os ervies of uplaind rice research 04 Farrow R A 1975 The Alrican nmigratory locust ii is main Proceedings of the 1982 Boinakamp Ivory Coast upland rice workshop outbreak area of the middle Niger qlinlitatise studies of solitary International Rice Research Institute P0 Blox 933 Manila populations in relation to environmental factors Locusla No II Philippines 198 p

43 Chandra G 1979 Taxonomy ad hn riononlics ofthe imect parasites of 65 Fennal R G 1943 Food crop investigation Windward and rice Icalloppers and planthoppers in the Pihilippines and their Ieeward Islands First report Report onl work dunring the periodiiiportairce iii natural biol)gical cointrol Philipp Fntoltol 4119-139 April 1942-April 1943 Imperial College of Tropical Agriculture

44 Chang T T 1976 The originii colut ion enlIivation dissemination Trinidad 16 p (Rev of Appl Fintornol A3222-25) and diersificition of Asian and African rices -uphytica 25425-44 1 66 Ferreira F 1983 Integrated pest ritaageiret of upland rice [ill

45 Chatierjee P B 1979 Rice leaffolder attacks in India lit Rice Res Portuguese English summary] Pages 323-341 in Upland riceculttre Newsl 4(3)21 factors affecting production E Ferreira T Yamada and F-

46 Cheng Cbien-pan 19(4 Upland rice in Taiwan International Rice Malmolta eds Institutoda Potassae Foslato Institnto Internacional Cnmmission Wiorking Party oit Rice Production ard Protection I()th da Potassa Piracicaha Sao Paulo lraid Meeting 3-10 March 1964 Manila Philippines Food and Agricul- 67 Ferreira E arid F dl Silva Martins 1979 Occurrence of ture Organiaion Rome Italy 5 p Sogatohs orvziola (Muir) inupland rice in Goias Ilrail lit Rice

47 Chcrrett 1 M and 1) 1Peregrine 1976 A review of the status of Rcs Ncwssl 4(4)18 leaf-cutting ants and their control Ann Appl Biol 84124-128 68 Ferreira E and 1F da Silva Martiris 1984 Insect pestof rice ill

48 Chipeta F M 1974 Pests of rice in Malawi Pages 90-95 in Brazil and their control Jill Portugucsej FMIlRAtPA-CNPAF Proceedings Sith eastern Africa cereals research conference 10-15 Goiania Brmiil 67 p March 1974 DRB Mialda ed Ministry of Agriculture lilongwe 69 Ferreira FI F ca Silha Miartins S S Nero and 1 It P de Malawi Carvallio 1982 M tltiplc regression analysis todeterimine the effect of

49 Chu Yau-i 1979 The d atiaging period of the rice seedling fly insect populatiois ott upland rice yield [inIPortuguese Pesqui (Atlterigotiavxigu Stcin) (NItiscide Diptera) oil cori) at Fast Java Agropecu Bras 17671-675 Indonesia Plant Prot Bill 21403413 70 Ferreira F 1F da Silva Martins and F 1PZimmermann 1979

50 Conklin H C 1957 IHaninoo agriculture a report on aln integral Rice varietal resistance to tle lesser corn stalk borer [in Port uguese] systent of shi fti rig cult ivationi in the Pltilippincs Food and Agriculture Pesqui Agropccu Biras 14317-321 Organization Rome Italy 209 p 71 Fleming W F 1968 Biological control of lie hapanese beetle U S

51 Cramer H II 1967 Plant proection and world crop production Dep Agric RLs Sers [ech Bull 1383 78 p Pilaniensclhutz-Nachr 201-524 72 Food and Agriculture Organiation of the United Nations 1982

52 l)ani R C and Majunidar 1978 Prelininary observations on tie Country report surmimary Pages 27-37 in Report on the first incidence of root aphids on differeit rice cultnrcs Sci Cult 4488-89 FAOIUN)IPhail and regional training coiuirse in iniproved ctltural

53 Davidson R I 1979 lffects of root feeding on foliage yield Pages practices for upland rice 5-23 Oct 1981 Northern Region Agrculturd117-120 in T K Ci osb and R B Pottinger eds Proceedings of the Developririt Center Chiang Mai Food and Agricuilture Organiiashy2nd Australian conlerence on grassland invertebrate ecology Govt tion Rome Italy 37 n Printer Wellington New Zealand 73 Franco do Ariaral S 195 1 Residual action of insecticides in turfy

54 l)ean G 1W 1976 Rice insect pests inlaos Int Rice Res Newsl soil--the results of a field experiment [in Portuguese] Biol6gico (Sao 1(2)15 Paulo) 17209-214 (Rev Appl Entomol A41423)

55 I)ean G J W 1978 Insect pests of rice in Laos PANS 24(3)280-289 74 Freeman ) 1970 Report onithe Iban london School of -coroiics 56 )ingle H 1984 Behavior genes and lifc histories conplex adapta- Monographs on Social Anthropology No 41 Athlane Press

tions to uncertain environments Pages 169-194 in A new ecology London UK 317 p novel approaches to interactive systems P W Price C N Slobod- 75 Fuchs T W1 A Harding and 1W Smithlr 1979 Induction and chikoff and W S Gaud cdsl ohn Wiley New York USA termination oldiapause in the sugarcane borer Ann Flitonlol Soc

57 Dobelmann-JP and M Falais 1968 Upland rice cultivation ill tie Am 72271-274 North-western part of Madagascar in 1966-1967 [in French English 76 (hbriel B1) 1975 A review tf the major insect pests of some tpland summary] Trop Agron 2313-21 crops in tlc Philippines Pages 86-111 in Ad Htoc panel of experts on

58 Doniiciano N L 1979 Chemical control of tie lesser corn stalk pest disease arnd weed problems in some rainfed crops Bangkok (lolsonno~paloIgnowslhts) (Zeller 1848) (I-epid(ptera Ihycitid ae) Thailand Food aid Agriculture Organization Roire Italy

on rice in Ponta Gressa Parara 1977-1978 [in Portuguese] Pages 77 Garg D K and N K Shal 1983 White grub outbreak on rainfed 291-303 in Anais 1 Reuniao de tecnicos cmiirizicultura do estado de dryland rice in Uttar Pradeslh Int Rice Res Ncwsl 81(4)16-17 Sao Paulo-Campinas Cati Brazil 78 Garg I) K and IP Tandon 1982 Major insect pests of rice on hilly

59 Dunsmore 1 R 1970 Investigations on tlie varieties pests and tracts of Uttar Pradesh India lilt Rice Res Newsl 7(11)11-12diseases of upland rice (hill padi) in Sarawak Malaysia Int Rice 79 Garrity 1) P 1983 Upland rice research an international bibliograpy Comm Newsl 1929-35 1975-1982 Iternational Rice Research hnstitnte PO Box 933

60 Elias 1R1965 The new decline of rice seedlings is not caused by Manila Philippines 121 p parasites [in Portuguese] Biol6gico (Sao Paulo) 313-6 80 Ghauri M S K 1982 A new subspecies of Iinorh)rone bull

61 Empresa Brasiliera de Pesquisa Agropecuaria 1984 Upland rice itt cornlus Slater (Hemiptera Iygaeidae) front Papua New Guinea on Brazil Pages 121-134 it Av overview of upland rice research rice and carpet grass Ihll lEntoniol Res 72133-137 Proceedings ofthe 1982 Blouak Ivory Coast Upland rice workshop 81 Gonzales F 0 PArregoces R I ihernandef and OT Parada International Rice Research Institute PO Box 933 Manila 1983 Insect and imite pests of rice arid their control in Latin America Philippines [in Spanish] la Federacion Nacional de Arrocerts (Fedearroz)

Bogota Colombia 60 n

38 IRPS No 123 January 1987

82 Grist D H and R J A W Lever 1969 Pests of rice Longmans London UK520 p

83 Hale P R 1979 Two subterranean pests of upland rice in Papua New Guirnca IntRice Res Newsl 4(2)16-17

84 Hale P R and J T Hale 1975 Insect pests of rice in the New Guinea Islands Rice Entomol Newsl 34

85 Harris W V 1969 Termites as pests of crops and trees Common-wealth Institute of Entomology London UK 41 p

86 Hattori 1 1968 A method to distinguish between grass leaf roller (Ctaplha rocis medinalisGuenee) and the rice leaf roller (Susurtia exigua Butler) [in Japanese] Plant Prot (Japan) 23167-170

87 Henry I E and 1A Onsager 1984 Experimental control of the mormon cricket Iahrosshnph by NJsena locustae(Aficrospora Microsporidae) a protozoan parasite of grasshoppers (Ort Acri-didae) Entomophaga 27197-201

88 Hidaka 1 P Vungsilabulr and SKadkao 1974 Studies on ecology and control of the rice gall midge in Thailand Tech Bull 6 TropicalAgricultural Research Center Tokyo Japan 113 p

89 Hincklcy A 1) 1963 Ecology and control of rice planthoppers in Fiji Bull Entomrol Res 53467 481

90 Hirao J 1982 Ecology and chtInical control of the rice leaffolder Jpn Pestic Inf 4114-17

91 Ho DT 1984 Sten borer incifience in rice ecosystems in KenyaEast Africa Pages 307-320 in A overview of upland rice research Proceedings of the 1982 Bouac Ivory Coast upland rice workshop International Rice Research Institute PO Box 933 Manila Philippines

92 Ho )T J 1Njokah and J GKibuka 1983 Studies on rice sten borers in Kenya with emphasis on ifaliarpiaipiraellRag Insect Sci Appl 465-73

93 Ichihara I T Ito A NIatsuo and M Sawada 1971 Chemical control of the rice striped stem borer in upland rice grown in plastic mulch [iii Japanese] Proc Kanto-Tosan Plant Prot Soc 1882

94 IclhiharaIand A Matsuo 1969 Damage caused by rice striped stett borer to upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1686-87

95 Ichhara I atid A MIatsun11970 Conside ration oft lie extraordinary prevalence of rice striped stem borer in upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1778-79

96 Institut de Recherches Agrolnliqes Iropicale et des Cultures Vivrieres 1978 Insect resistance Pages 12-13 in Upland rice 1978 atnual report Bouak Ivory Coast 27 p

97 International Institute for Tropical Agriculture 1983 Aphids itt rice roots I ITA Res Briefs 4(1-3)4-5

98 International Institite fi Tropical Agriculture 1983 Hopperburn in rice caused by brown plathopper IITA Res Briefs 4(l-3)6-7

99 International Rice Research Institute 1975 Annual report for 1974 PO Box 933 Manila Philippines p 216

100 International Rice Restarch Institute 1976 Annual repiort for 1975 PO Box 933 Manila Philippines p 228

101 International Rice Research Institute 1979 Cropping systems program Component technology development and evaluation Page 404 i Annual report for 1978 11O Box 933 Manila Philippines

102 International Rice Research Institute Department of Entomology 1981) Insecticid evaluation 1979 Tables 73 74 PO Box 933Manila Philipes

103 International Rice Research Institute Department of Entonmology 1981 Insecticide evalomtion 1980 Table 81 POBox 933 Manila Philippines

104 International Rice Research Institute 1981 Report of a meeting of Asian rice-based cropping systems entomologists Eleventh cropping systems working group meeting 18-22 May 1981 PO Box 933 Manila Philippines p 132 133 144

105 International Rice Research Institute 1984 Cropping systems program Pages 368-373 in Annual report for 1983 PO Box 933 Manila Philippines

106 International Rice Research Institutc 1984 Upland rice in Asia Pages 45-68 in An overview of upland rice research Proceedings of the 1982 Bouakamp Ivory Coast upland rice workiiop loternational Rice Research Institute 11O Box 933 Manila Philippines

107 Irshad M 1977 Grasshoppers in rice in Pakistan lit Rice Res Newsl 2(3)8-9

108 Ito R and TOno 1964 Upland lice its culture and varieties in Japan lIternational Rice Cormmission loth Meeting of the Working Party on Rice Production and Protection 3-10 March 1964 Manila Philippines Food and Agriculture Organization Rome Italy 12 p

109 Jeffrey M I and V A Dyck 1983 Effect of moonlight on lightshytrap catches of brown planthopper lit Rice Res Newsl 8(6) 13

110 Jennings P R and T Alicia Pineda 1983 Management ofthe rice plnthoppers Sogatodsorivi(coIl in Latin America Presented at the meeting on stable plant resistance FAO and lIndocsian I)inectorate of Food Crops Protection Bali lndonesia 29 Novcniber-2 l)ccemshyber 9 p

IllJimenez Gomez J A 1979 Rice insect pests i lla ios ()riCntales lil Spanish] Pages 99-103 in Rice ciltivation Instituto (olotibiatto Agropecuario Compndio No 29

112 Jongeleen F JJ Soesanto and A Mockidjo 1979 Studies oti pathogens of two species of Scarabacidac froinIndonesia Z Angew Enlomnol 87412-417

113 Jouvenaz 1) PC S lofgren and W A Baiks 1981 Biological control of imported fire antsa review of cirrent knowledge Bull Entomol Soc Am 27203-208

114 Kainoh Y K Shiniiu S MarUani Y Taiaki 1980 lost finding behavior of tie rice bu lplotori+JchinvittjI)allas (liciniptera Corcidac) with special reference to diet pattern ol aggregation and feeding on the rice plant Appl Entomol Zool 15225-233

115 Kakkar K 0 P Sharma and P I Sharma 1977 Note on Ihteronyrchus liodresRedt (Coleopicra I)ynastidae) as asporadic pest of paddy and maie inHimachal Pradesh Corr Agric 156-57

116 Kalode M131 79 Major insect pest problems and their control in upland rice (Unpublishcd mineogriph CWI oJNaaiat lni Chandaratat (198 1)original not seen)

117 Kalshoven L i 98 I The pests of crops in I ldoniesia (translated by P A vatn der Laan) 1 T Ichtiar B3aru-vati Iloeve lakarta Indonesia 701 p

118 Kamal N Q 1981 Suppression of whitebacked planto ner Sogatella fircaifra (Horvath) aLId rice lealfolder (nhal -cis licdb lis (Guenee) populations by natural cnclmie hlI) tiesis Grcgorio Araneta University Foundation Victoncta Park Metto Manila Philippines 179 Ip

119 Karim A N M Reaul 1975 A species ofliealy bug Trioni i70s ceres new oii rice in the Philippines Rice Entoniol Newsl 247

120 Katanyukul Wand C Chandaratat 1981 Insect pestsof uplaid rice and their control Report firstFAOUNlFi Tliailand training course on improved cultural practices for tplad rice Chiang Mai Thailand 5-23 Oct 1981 Food and Agriculture Organiation Rome Italy 22 p

121 Kern F 1955 lhletwioa(I)uninemts) bhhit Biturm(Coleoptera) a new pest of sugarcane inVenezuela [inSpanish with Etnglish sunmary] Agron Trop (Maracay) 527-29 (Rev Appl Eitoniol A4572)

122 Khan K M and S Ghai 1974 White grubs ald their control in India Pesticides (1Bombay) 819-25

123 Kiritani K 1975 Integrated pest management of rice pests Pages 66-70 itt Proceedings platning workshop ott cooperative hIeld research in pest managementP SMotooka ed East-West Center Honololu Hawaii USA

124 Kisimoto R 1967 Ecology of insect vectors forecasting and chemical control Pages 243-255 in The virus diseases of tle rice plant Proceedings of a symposium atthe International Rice Research Institute April 1967 The Johns Hopkins Press altimore Maryshyland USA

125 Kisimoto R and V A Dyck 1976 Climate and rice insects Pages 367-391 in Climate and rice International Rice Research Institute PO Box 933 Manila Philippines

126 Kureha S S Kobayashi SAsari and T Ishihara 1977 Life history and control of the mole cricket Grrllotalpa africana Palisot de Beauvois [in Japanese English translation 7520 1RRI library] Proc Kanto-Tosan Plant Prot Soc 21192-197

127 Kushwaha K S 1981 The black beetle as a pest of rice in Haryana India Int Rice Res Newsl 6(4)14

128 Lawani S M 1982 A review of tile effects of various agronomic practices on cereal stem borers Trop Pest Manage 28266-276

129 Leuck D B 1966 Biology of the lesser cornstalk borer in South Georgia J Econ Entomol 59 797-801

130 lever R J A W 1955 Insect pests of the rice plant in Malaya Int Rice Comm Newsl 1612-21

131 Li Ching-Sing and Shin-Foon Chiu 1951 The study ofthe rice gall midge Iachydiplosis orvza Wood Mason [in Chinese with English summary] 1Taiwan Agric Res 21-13

132 Iitsinger J A 1984 Assesament of the need-based application for rice Proceedings TechnologyTransfer Workshop 15-16 March 1984 International Rice Research Institute 1PO Box 933 Manila Philippines 6 p

133 Litsinger J A 1984 Environmental diversity and insect pest abundance with reference to the Pacific Pages 285-307 in Proceedings sub-regional training course on methods of controlling diseases insects and other pests of plants in the South Pacific 4-20 Oct 1982 Vaini Kingdom of Tonga Nukualofa

134 Litsinger J A RF Apostol and M BObusan 1983 White grub Leucopholis irrorata (Coleoptera Scarabaeidae) pest status popula-tion dynamics and chemical conllol in arice-maize cropping pattern in the Philippines J Econ Entoinol 761133-1138

135 Litsinger J A C de la Cruz F Raymundo A T Barrion M D Lumaban J P Bandong M S Venugopal F Paragna and E Balete 1982 Insect pests and insecticide reponse of dry-seeded bunded rice Pages 179-193 in Report of workshop on cropping systems research in Asia International Rice Research Institute Los Bafos Philippine

136 Litsinger J A M 1) Luinaban J P Bandong A T Barrion W Katanyukul and N Panda 1978 Insect pest management in rainfcd lowland rice agroecosystms of tropical Asia International Rice Research Conference 17-21 April 1978 International Rice Research Institute Los Bafios Philippines

137 Litsinger J A and K Moody 1976 Integrated pest management in multiple cropping systems Pages 293-317 in R I Papendick P A Sanchez and G B Triplett eds Multiple cropping American Society of Agronomy Madison Wisconsin USA

138 Loevinsohn M E 1984 The ecology and control of rice pests in relation to the intensity and synchrony of cultivation Ph D thesis University of London 36) p

139 Loevinsohn M E J A Litsinger N Panda M A Austria R F Apostol C Andrion and J Bandong 1982 A basis for predicting rice insect pest abundance in cropping systems Paper presented at the 1982 International rice research conference on rice and cropping systems 19-23 April 1982 International Rice Research Institute Los Bafios Iaguna Philippines 25 p

140 Lopez A W 1930 Repot of the Entomology Department Pages 145-172 in Annual report of the Philippine Sugar Association 1929-30 Manila

141 Lopez AW 1931 Entomology Department Pages 227-266 in Annual report of the director or research 1930-31 Philippine Sugar Association Manila

142 Lopez A W 1932 Entomology Deparment Pages 252-279 in Annual report of the Philippine Sugar Association 1931-32 Manila

143 Lungibill P and G G Ainslee 1917 The lessercornstalk borer US Dep Agric Agric Res Serv Tech Bull 539 27 p

IRPS No 123 January 1987 39

144 McGuire J U Jr and BS Crandall 1967 Survey of insect pests and plant diseases ofselected food crops of Mexico Central America and Panama U SDepartment of Agriculture International Agriculshytural Development Service Agricultuial Researrh Service October 1967 11 p

145 McNeill Sand T R ESouthwood 1978 The role of nitrogen in the development of insectplant relations Pages 77-9 in lliochemical aspects of plant and animal coevolution J B Ilarbornu ed Academic Press London UK

146 Maebara H 1955 Spatial distribution of lastinstiarlanaeof purplish stem borer (Sesamia iqferens Walker) hibernating in an upland rice field Bull Kagoshima Univ Fac Agric 438-41

147 Mallick E H and N G Hajra 1977 Response of rice plant to defoliation treatments at different developmental stages Indian Agric 2151-56

148 Mansingh A 1971 Physiological classification of dormancies in insects Canl Entomol 103983-10)9

149 Martell C G 1974 Catalogue of plant feeding insects of Mexico [in Spanish] Fitofilo 27(69)1-176

150 Martinez 0 J 1982 Principal rice pests in Ilanos orientalis [in Spanish] P~ages 80-94 in Curso deact nalization de conocimientos en arroz Villaniencis Instituto (olonmbiano Agropecuario

151 Martins J F da Silva F Ferrcira A S Prabhu F 1 P Zimmermann 1980 Use of pesticides for the control of the main subterranean upland rice pests esqui Agropecu Bras 1553-u

152 Martins I F da Silva F Fe eira rod BdamSilveira Pinheiro I Q2 Damage simulation caused by defoliators of upland rice [in Portuguese with English summary] Pesqui Agropecu Bras 171113-1119

153 Martins 1F da Silva N(iyuen van Tan and 1Ida Silveira Pinheiro 1981 Stem borer resistance in upland rice associated with plant morphological characters [in Portuuce Iesqui Agropecu Bras 16187-192

154 Masaki S 1980 Sumnmer diapause Ann Rev Entomiol 251-25 155 Masuzawa T 11 Suwa and F Nakasoji 1983 l)ifkleences of

oviposition preferencc and survival rate of two skipper butterflies Parnara guttata and PIchlpidai iathiais (Iepidoptera HIesperiidac) on rice plant and cogon grass New Entomol 3247-56

156 Maurya D M and C P Vaish 1984 Upland rice in India Pages 69-92 in An overview oftpland rice research Proceedings ofthe 1982 Bouakamp Ivory Coast upland rice workshop International Rice Research Institute IO Box 933 Mnila Philippines

157 Menon M G R R N Katiyai S Kuar and S S Misra 1970 Termites attacking the paddy crop Sci Cult 36294-295

158 Miyahara Y and 1 Nishikubo 1967 Behavior of young nymphs of the red rice root aphid Iholahsipiun ri liabhdontinalisSasaki in relation to movement from the leaf to the roots of upland rice [in Japanese] Pioc Assoc Plant Prot Kyushu 1363-66

159 Mo ljoa Tjien 1952 White grub of upland rice (Holotrichiahelleri Brsk) [in Indonesian with English summary] Contrib Agric Res Stn Bogor No 13134 p

160 Mochida 0 and T Suryana 1979 The occurrence of the brown planthopper and ragged stunt on lowland and upland rice ill Central Java and logyakarta in tile late of January 1979 Entomnolgy Survey Report No 15 Central Research Institute of Agriculture (CRIA) Sukamandi Indonesia I February 1979 9 p

161Monet C 1979 Varietal resistance to upland rice pests [inI French] Pages 35-39 in Annual report 1977 Institut des Savanes Department des Cultures Vivrieres Bouak6 Ivory Coast

162 Monte 0 1942 A caterpillar ofrice-fields[in Portuguese] Biol6gico (Sao Paulo) 8161-163 (Rev Appl Entomol A31414)

163 Munroe I) D 1979 Pest management strategy for hill rice in four resettlement areas in Sarawak Malaysia Pages 89-102 i1 Guidelines for integrated control of rice insect pests Food and Agriculture Organization Plant Production and Protection Paper 14 Rome Italy 115 p

40 IRPS No 123 January 1987

164 Nakasuji F M Ishii and Jen-Zon Ito 1984 Rice skippers in Taiwan and their life histories ZZrC (Taipei) Tech Bull 82 10 p

165 Navas I) 1976 Fall armyworum in rice Pages 99-106 il Ecological animal control by habitat management Proc Tall Timbers Con-ference 28 Feb-I Mar 1974 Fall Timbers Res Stn Misc Publ 6 Tallahassee Horida USA

166 Nickel 1 1 1961 Biological control of rice stem borer A feasibility study Tcch Bull 2 International Rice Rcarch Institute PO Box 933 Manila Philippines IIl p

167 Nigan P M and R A Vernia 1985 Insect pests of uplaild rice in Uttar Pradesh Int Rice Res Newsl 10(6)22

168 Nilakhe S S 1985 -cological observations on spittle bugs with emphasis on their occirence in rice Pcsqtui Agropecu Bras 20407-414

169 Nixon G EJ 195 1 lie association of ats with aphids and coccids Commonwealth Iusti lcte of Entonology Iondon UK 36 p

170 Nurdin F an Ishak Manti 1985 Study oit seedling fly of upland rice Interlational Upland Rice Moitoring Tour Report 26 lebruary-2 MarLI Sukaranti West Sumatra

171 Oka 1 N 1979 Ciltural conttol of the brown planthopper Pages 357-369 in Brown planthopper threat to rice production in Asia International Rice Research Institutc 1p0 Box 933 Manila Philippines

172 Ohni M and A Vilnaii 1975 Rice insect pests of Italy biology of Sipha grcria (Kaltenbach) and its control [in Italian] Riso (Milan) 2459-71

173 Otanes F Q atnd 1 I Sison 194 1 Pests of rice Philipp J Agric 12211-259

174 Painter R I 1951 Insect icsistance in crop plants University Press of Kansits Lawrence Kansas USA 520 p

175 Pantua P C and 1A litsinger 1981 Arthropod abundance in continuous vs biannual rice cropping systeis Int Rice Res Ncwsl 6(2)20-21

176 Pathak M ) and V A l)yck 1975 Sttudics ott insect pests of upland rice Pages 186-191 in Major research in upland rice International Rice Research Institute PO Box 933 Ilanila Philippines

177 Patil B R B M IHasabe and 1) S Ajri 198 I Chemical control of white gritb lIditriihinhorraia Fabr intfesting groundntut paddy atnd sorghum Indian 1Entomol 43232-236

178 Pawar A 1) 1974 (ollection and identification of insect pests of rice rlerminal repirt lInternational Rice Research Institute PO Box 933 Manila Philippines 72 p (tnimco)

179 IerrajU A aid A S Reedy 1965 Observations ott diapausc itt tie steti borer ohrice Rice News Teller (lai-Mar)9-13

180 Pont A C 1972 A review of the Oriental species of theri ona Rondani (I)iptera Muscidac) of econtomic importance Pages 27-102 in Control of sorglium shoot fly M G lomtani and W R Young eds Oxford atd IH1 Ptthl Co New I)clhi India 324 p

181 Prasad S C 1PChand aid M 1PSingh 1983 Incidence of rice gall midge in the upland plateau region of Hihar Oryza 20235-236

182 Rai P S 198 0 Nisaga simpex damage to rice in the hill tracts of South India Int Rice kles News[ 5(I)12

183 Rathorc Y S 1969 Gonocplhahli dcprssunt Fabr as itpredator of (tilo zonwIlhis (Swinhoc) and other moths in field cage at Patnagar Uttar Pradesh Indian 1Entotnol 3127

184 Reed W 1983 Crop losses causcd by insect pists in the developing world Proceedings 10th Internatial Congress of Plant Protection lrighton UK 174-80

185 Reedy M V 1981 Ateactogasterinhens(Walker)a new root-grub pest of tipland paddy and inaize Curr Sci 501076

186 Reveno M A 1968 Catalogue of the insect pests of economic crops in Colombia [in Spaiish] Asociacion Latino Americana de Entomo-logia (AIAE) 129-33

187 Reyes G M 1957 Rice dwarf disease in the Philippines Food and Agriculture Organiiation Plant Prot Bull 617-19

188 Rice S E A A Grigarick and M 0 Way 1982 Effect of leaf and panicle feeding byarmyworm (Lepidoptera Noctuidae) larvae on rice grain yield Environ Entomol 7593-595

189 Risch S J 1) Andow and M A Altieri 1983 Agioccosystem diversity arnd pest control data tentative conclusions and nizw research dirctions Environ Entomol 12625-629

190 Rivera C T 1956 Three organic insecticides in thecontrol of insects affecting upland rice Philipp Agric 39465-472

191 Roffrey 1 1972 Locusta outbreaks in the Philippines Acrida 1177-188

192 Rojas R 1) 1941 The effect of pruning at varying ages upon the yield of Ramairio Phipp Agric 29851-860

193 Roonwal M i 1979 Ternite life and termite control in tropical South Asia Scientilic Pnbl odhpr India 139 p

194 Rosetto C1 F da Silva Martins N C Schmidt and L EAzzini 1978 Damage caused by pasturespittle bugs 4Dvoi)lavopictaandDJ schach) on rice lBragantia Nota 6 (Matcli)25-27

195 Rosztto C1- S S Neto I- Amantc I) Iink 1Garcia-Vieira D M de SoLza N V Bansa t and A M de Olivcira 1971 Rice pests of Brazil [in Portuguese] Pages 149-238 inrFA()-International Rice Commission 2nd meeting of the Rice Committee for the Americas 6-11 l)ecember 1971 Pelotas Rio Grande do Sul Brazil

196 Rothschild G H L 1969 Obseratiois on the armyworm Spodoptera matritia acronrctoiiles Gn (Lep Noctuidae) Bull Entomol Res 59143-160

197 Rothschild G 1- L 1974 Parasites of the armworm Spolopiera mauritiaacroncitoihes in Malaysia Fntornopliaga 19293-299

198 Sachan i N S K Gangwar and 1N Katujar 1980 Field crickets damaging cages in North Eastern IHill Region Indian J Entomol 42526-527

199 Sands 1) P A 1977 The biology and ecology of Lepocorisa (Hemiptera Alydidac) in Papua New Guinea Res Bull 18 Departshyment of Primary Industry Port Moresby Papua New Guinea 104 p

200 Sarawak 1968 titomology division Pages 97-106 in Report of the research branch of the Department of Altriculture 1966 Kuching (Rev AIppl Eintomol A56603)

201 Satpathy JM M S Das and K Naik 1977 Effect of multiple and mixed cropping on the incidence of scine important pestsI Entomol Res 178-85

202 Sauer IHF G 1939 Notes Ott Elasmpnlpualtsligniosellths Zeller (Lep Pyr) a serious pest ofcereals in Sao Paolo [in Portuguese] Arq Inst Biol 10199-206 (Rev Appl Entotnol A28378)

203 Satinders 1L A 1 SKing and C L Vargas S 1983 Crop pests of Central Anerica with t bibliography [in Spanish] Centro Agroshynotnico Tropical de Investigacion y Ensenanza Turrialba Costa Rica (CATIE) Tec Bol 9 90 p

204 Sauphanor B 1985 Some factors of upland rice tolerttce to stem borers in West Africa Inseu Sci Appl 6429434

205 Sen P atnd S Chakravorty 1976 Stentflics in Haritighata Nadia West Bengal India [it Rice Res Newsl 1(12) 15

206 Senapai B and 1M Satpathy 1983 The seedling fly Atherigona orrzae Malloch is a pest of rice in Orissa (India) J Entomol Res 777-79

207 Shhjahan M S Alatn M Ah nad and BN Islam 1983 A new scarabaeid beetle pest of dryland rice in Bangladesh Int Rice Res Ncwsl 8(I)16-17

208 Sharma G L S C Modgol and R C Gautam 1979 Effect of depth of submergence ott incidence of bacterial blight and white grub infestation in transplanted rice Int Rice Res Newsl 4(6)12-13

209 Sharma S 1985 Scarabaeid beetle outbreak itt upland rice Int Rice Res Newsl 1(1(l)24

210 Shiraki T 1917 The paddy borer Schoenobhits incertulas Walker [achaku Agric Exp Stn Formosa 256 p

211 Silva A de Brito and 11P Magalhaes 1981 Insect pests of rice [in Portuguese] Circular Tecnica No 22 EMBRAPA Centro de Pesquisa t ecuaria de Tropico Unido (CPAJU) Belem Brazil 14 p

212 Singh S A 1979 Emergence of rice leaf butterfly as a rice pest in Manipur India int Rice Res Newsl 4(4)16-17

213 SinghS A 1983 Changing trends of insect pestsof rice in Manipur Int Rice Res Newsl 8(f)12

214 Siwi S S and P H van Doesburg 1984 Lenovorisa Latreille in Indonesia (Heteroptera Coreidae Alydinae) Zool Meded (Leiden) 58117-129

215 Soejitno J 1977 Reaction of some upland rice varieties toseedling fly eltherikonaexigua Stein [in Indonesian] Siri Hana I Penyakit 630-35 Lembga I osat Fenclilian Pertanian Bogor Indonesia

216 Socnardi 1967 Insect pests of rice in Indonesia Pages675-683 in The major inse pests of thu rice plant Proceedings of a symposium at the Internatital Rice Rec Wi itt S pternber 1964 The Johns Hopkins Press Baltimore Maryland USA

217 Soto P E A T Petcz and I W Buddenhagen 1976 Survey of insect pests and diseases of rice in different ecological zones in Nigeria Rice Entomol Ncwsl 435-46

218 Soto 11E and Z Ziddiqi 1976 Screening for resistance to African rice insect pests Varietal improsetnent sCmi n r Bouak Ivory Coast 13-17 Sept 1976 West Africa Rice l)evelopment Association Monrovia Iiberia

219 Squire F A 1972 Entomological problems in Bolivia PANS (PestArtic News Sumr) 18249-268

220 Stapely 1 II 1978 The rice leaf roller in the Solomon Islands Int Rice Res Newsl 3(l)13

221 Suenaga H 1937 On aphids attacking the roots of rice Oyo-I)oubts Zasshi 9150-152 [in Japanese] (Rev Appl Entomol A25745)

222 Suenaga H 1952 On the seasonal abundance of three subterranean aphids on upland rice Bull Kyushu Agric Exp Sti 1249-262

223 Sutynso Yas and Dandi Soekaria 1964 Beberapa hana yang ttenperang tana man padi tanah kering [in Indonesian] Rapat kerja Padi Tanah Kering 6 sd 8 Juily 1964 Kaliurang 9 p

224 Tallamy I) W and R F Denno 1983 Alternative life history patterns in risky environments in example from lace bugs Pages 129-147 in Insect life history patterns R F l)cnio and If Dingle eds Spring-Verlag 225 p

225 Tamura 1 1939 Observations on some insect pests of field crops [in Japanese] 1 Plant Prot 26414-420 (Rev Appl EntomolA27-556)

226 Tamura and Suzuki 1964 Influences of the early-transplanting practice in rice culture upon development of the rice stem borer Chilo siql)ressalisWalker Bull Ilokuriku Natil Agric Exp Stn 761-94

227 Tan J P 1924 The rice root aphid Drowhi ihirsoia Philipp Agric 13277-288

228 Tanaka 1 1957 Taxonomy and distribution of some subterranean aphids injurious to upland rice in Japan with description of a new species [in Japanese English summary] Botyu-Kagaku 22168-176

229 Tanaka T 1961 The rice root aphids their ecology and control Spec [full College of Agriculture Utsunomiya University Japan F3 p

230 Tauber M IC A Tauber R Nechols and 1Obrycki 1983 Seasonal activity of parasitoids control by external internal and genetic factors Pages 87-108 itt I)iapause and life cycle strategies in insects V K Brown and I Ilodek eds )r W Junk The Hague Netherlands

231 Taylor W F 1972 Effects of artificial defoliation (simulating pest damage) on varieties of upland rice Exp Agric 879-83

232 feotia T P S and S Natud 1966 Biotoomics of the rice skipper Parnaraniathiis Fabricius (Lepidoptera Hesperiidac) Indian J Entomol 28181-186

233 Teran F 0 1971 Rice pests of Santa Cruz [in Spanish Ministerio de Agricultura Sita Cruz Bolivia Bull 45 8 p

234 Than Itun 1976 Population dynamics of the yellow rice borer Tryporza itcertulas (Walker) and its damage to the rice plant MS thesis University of the Philippines at Ios Bafios Laguna Philippines 92 p

235 Uichanco L B 1911 Pests Pages 55-74 it Rice Philipp Agric 11-200

IRPS No 123 January 1987 41

236 van der Laan P A 195 1 Life history of the rice seedling fly Contrib Gen Agric Res Stn Boor Indonesia 118 15 p

237 van der Laan 11A 1959 Correlation between rainfall in the dry season and the occurrence of the white rice borer (Scirpolhaga innotala WIk) in lava Entomol Fxp Appl 212-20

238 Van Driesche R G 1983 Meaning of percent parasitismin studies of insect parasitism Fnviron Entomol 121 I1- 122

239 Venugopal M S and A litsingcr 1984 Fffect ofcarbofuran on rice growth Prot Ecol 7313-317

240 Vietnam Directorate of Rural Affairs 1962 Annual work progress report on crop improvement program of rice sugarcane vegetable and field crops (lor the period from July 1961 to June 1962) (11) 299 p (Rev Appl Entomol A51526-527)

241 Wan M T K 1972 Observations on rice leaf- and planthoppers in Sarawak (Malaysian Borneo) Malay Agric 148308-335

242 West Africa Rice I)esclopnient Association 1978 Annual research report Vol 2 Statistics Monrovia Liberia

243 West Africa Rice Development Association 1979 Annual research report p47 Monrovia Liberia

244 Williams ) F and CS lofgren 1981 Fli I illy l-468 anew bait toxicant for control of the red imported lire at Fla Entomol 64472477

245 Williams I) 1 1970 The mealybugs OFlomoptera Coccoidea Iscudococcidac) of sugar-cane rice and sorghum Bull Entomol Res 0109-188

246 Williams D) 1 1982 Pulinaria icervi (Signoret) and its allies ott sugar cane and other grasses Btll Entoinol Res 72111-117

247 Wirjosuiardjo S 1976 Study ofchenmical cont rot of insect pests ott upland rice Imiu Pertanian (Jogjakarta) 1323-329

248 Wirjosuhardjo S and N Wirarta 1976 Chemical control of the white grub tIolorithiatlltriin upland rice in M uneng Irobolinggo Indonesia) [in Indonesian] Imut Pertanian (Jogjakarta) 1330-332

249 Yano K T Miyake and V F Eastop 1983 The biology and economic importance of rice aphids (Henuiptera Aphid idae) a review Bull Entomol Res 73539-566

250 Yelp D and P Rhodes 19(4 lhe success of drilled dryland rice iii the Fiji Islands International Rice Commission Working Party on Rice Production and Protection Manila Philippines 3-10 March1964 Food and Agriculture Organization Rome Italy 13 p

251 Young W R 1981 Fifty-five years of research on the sorghum shootfly Insect Sci Appi 23-9

IRR Irch Institute Co EN CAS DE NON REMISE RENYOVER A

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ISSN 05-3862Other papers in this series TITLES OF NUMBERS 1-45 ARE LISTED ON THE LAST PAGE OF NO46 THOSE OF NUMBERS 46-70 ARE ON THE LAST PAGE OF NO 71-80AND THOSE OF NUMBERS 71-1O0ARE ON

THE LAST PAGE OF NO 101-122

No 101The economics of hybrid rice production in (hina No 113 Boron toxicity in rice No 102 Rice ratooning No 114 Energy analysis rice production systems and rice research No 103 Growth and development of the (Jeep water rice plant No 115 Production risk and optimal fertilizer rates an application of the

No 104 Faridpur acomputer-assisted instruction nmodel forrainfed lowland random coefficient model No 116 Consumer demand for rice grain quality in Thailand Indonesia andrice

No 105 A readingand listening comprehcnsion test inEnglish for non native the Philippines speakers applying for tiraining at IRIRI No 117 Morphological changes in rice panicle development a review of

No 106 Rice grassy stunt virus 2 a new strain of rice grassy stunt in tire literature

Philippines No 118 IRRI-Korca collaborative project for the development of cold-No 107 Physical losses and quality deterioration in rice plstprodnCtion tolerant lines through anthei culture

systemis No 119 Problen soils as potenial areas for adverse soils-tolerant rice No 108 (olptblication ofIRRI materials a survey of translators and varieties in South and Ssuthcasi Asia

publishers No 120 Changes isinall-farm iLtireliing techii)hgy in Thailand and [he No 109 Classification of Philippine rainfall patterns Philippines No 110 Contributions of modern rice varieties to rutrition in Asia No 121 Landforms and modern rice varieties No Ill Changes in rice breeding in 10 Asian countries 1965-84 No 122 Yield stability and modern rice technology No 112 Design parameters affecting the performance of the IRRI-designed

axial-flow pump

Page 20: UPLANDRICE. INSECTl--1i M T -E!R.EOLOU t RTANCE ,AND …delphacid.s3.amazonaws.com › 6958.pdf · The significance of abiotic factors in upland rice insect ecology will be apparent

Table 3 Abundance of rice insect pests and factors in rice cropping intensity of Philippine upland rainfed lowland and irrigated lowland locations where light traps were operated daily for atleast I yr -I

0Rainfed Rainfed lowland Irrigated lowlandupland

Pangasinan Cagayan lloilo Santa Maria Nueva Ecija South Cotabato Tanauan Caaringayan Caaringayan Bangag Bangag Bangag Buray

LagunaCale

Batangas Manaoag 1980-81 Santa Motiica 1982-83 Cabanatuan Zaragoza Avancena NaniamanSolana 1981-82 1982-83 Oton

1980-81 1979-80 Oton 1981-82 1981-82 Koronadal Koronadal1980-81 1979-80 1979-80 1983 1983

Insect (kerosene light trapNephotcttixcatch nolyr)virescens 650 1240 500 1800 330 210 3400 1600 160N nigropictus 210 - 360 18400 5500 2900- 140 90 20 - shy 1700 -Reciliadorsalis 710 - 3200 1700180 890 630 140 170 2400 800Nilaparratalugens 130 690 - - 4200 4600300 520 750 180 60 3300 1600 740 310Sogatellafiircifera 1700 31000 13000 8200310 210 3300 610 270 980 1600 6300 -Cyrrorhinus - 12000 6700600 540 3200 290 160 330 120 410Scirpophaga spp 50 410 620 - - 21000 12000150 180 220 270 30 1000Cnaphalocrocismedinalis 70 70 540 610 270 11000 6100200 140 100 6 60 250Chilo suppressalis 0 60 60 1900 300 200- - 30 20 0 6 3 160 -C auricilius - 0 00 - - 20 3 60 20 3 0Sesania inferens 0 - 80 60 3 80 10 51 -Cropping intensity _ -

Rice crops (noyr) 10 10 12 10 10 10 13Rice area (7c) 17 19 20 20 25 2320 85 85 60 70 40 85 85 90Fertilizer (kg Nha) 60 20 85 80 70 7020 0 0 0 30 30Insecticide (no of 40 60 40 300 04 04 0 03 300 13 13 30 50 40 33 33applicationscrop)

__________

IRPS No 123 January 1987 21

Light trapped Insects Drytnd __ 2(notrap-nght) Suctioned Insects (nom )

50 -)o 9 jO40- SOW ffurciWO Light trp

D-Vc suction

20 to 4

20

Egg porasitizahion (M) 75 60

300 SOOampReOftwciffr

45 30shy

15

-insects (notrop-night) 4 Nileporvata Iuen5

Egg parastization()

Light trapped insects (no trop-nght) Suchoned insects (no m 2 )

6 8 Cytorh~nuls hvidipennii 0

4 05 2

50 0 Insects (noltrop-night)40-ui Ipanh pe g

Moohase 1 Seasonal abundance measured by light trap

Rainfall(etod) and D-Vac suction sampler of whitebacked

to ~ hopper Nilaparvata lugens and lea ffolder60)-t Cnaphalocrocis mnedinalis andplthpe g20 L I II I I I parasites and mtrid predator Ortorhjntus livi-Apr May Jun Jul Aug Sep Oct Nov Dec JaIn Feb Mar dipennis set against cropping pattern daily1 9 80o 19 8 1rainfall and moon phase Tanauan Batangas

1980-81

the D- Vac Cyrtorhinus build up on the crop coterminous are collected in light traps during full moon (109)with WBPH buildup Cyrtorhinus disperses during the Because upland rice shares most of the lowland rice pestsrainy season RLF not readily collected in light traps was and those pest species are generally less impotant to uplandprevalent only during the rice crop rice than lowland rice there is a general belief that insect

Nephotettix virescens was more abundant than Nnigro- pests are less important onpicLus in the light trap collections and on the crop GLH upland than lowland rice Arcview of limited data on upland rice shows an averag of population peaked at the reproductive stage as did zigzag 10-21 yield loss from insect pests on a par with that ofleafhopper (ZLH) Crtorhinus tracked the populations of lowland uee determined by the same methodology (132) planthoppers and leafhoppers Both GLH species were (Table 4) McGuire and Crandall (144) made a similarcollected year-round in the light trap indicating immigra- estimate for Central America Cramer (51) howevertion from nearby irrigated rice areas Possibly because of the estimated 35 yieldlow populations loss in South America based on ainsect number did not correlate with similar pest complex - stem borers and seed bugs shymoon phase although reports indicate that more hoppers attacking irrigated rice in North America This estimate did

22 IRPS No 123 January 1987

Light trapped insects D~~nd (no trap-night )

St plht Suctined insects (nom 2

)

D-Voc suction 1520

1o

0 Lih trap 5 ~105 o5

00 Eggparositizotion()5040 NephOtgtha virneens

30 20

I010 A-Insects (notrop -night)

4 Nephatelh nigropccent i

2 o I vV^J - -- ^ VA____ f _

Light trapped insects (no trap-night) Suctioned insects (no rnE

10 -1208 ReCdO doRsol

6 15

4 10 052 iwxALAiamp 0

Light trapped insects (notrap-night) 2Suc oned nsects (nom )

8 CyrtorhmnusI6o nn]10 i0075

4 05

2 0 0

025

2 Seasonal abundance measured by light trap Roinfall (mmd) Moon phase and D-Vac suction sampler of green leafhoppersNephotettix virescens and N ngropicrus and I zigzag leafhopper Recilia dorsalis their egg 60

parasites and micid predator Cyrtorhinus livi-dipennis set against cropping pattern dailyrainfall and moon phase Tanauan Batangas1980-81

40

5 Lf Apr May Jun

1980 Jul Aug Sp Oct Nov I

-aMar

1981

not account for soil pests and was not based on field data in the absence of pests Carbofuran granules are phytotonicAkinsola (I1) cited mainly data from irrigated rice areas in to rice (239) The prevailing hypothesis is that carbofuranAfrica but estimated yield losses of upland rice between 15 prevents soil bacteria from consuming fertilizer The phytoshyand 30 from insects However losses in Thailand over a tonic effect is through greater availability of fertilizer to the4-yr period ranged from I to 13 averaging 5(106) plant Denitrifying bacteria are known to be more active inYield loss data however should be interpreted with aerobic than anaerobic soils therefore it may not be merecaution (184) The Philippine data were all derived from coincidence that the highest recorded yield losses (22-69)trials on farmers fields under farmers normal agronomic occurred in those trials with 12-30 kg basal Nha On themanagement practices except for Bukidnon and Capiz other hand contro with carbofuran granules may not be aswhere researchers used fertilizer Batangas farmers are good as a seed treatment against seed pests and perhapsatypical because ofhigh levels of fertilizer used The trials in yields would have been higher if a systemic seed treatmentBrazil were carried out at an experiment station but insecticide had been used conditions were similar to those of local farmers The 1979 and 1980 trials in Batangas had two full-

Carbofuran granules were used in every trial as a broad- protection treatments one using carbofuran granules andspectrum insecticide applied basally with the seed An the other using bendiocarb WP as a seed treatment Plotsobjection to using insecticides to measure yield losses isthat protected with bendiocarb yielded higher than those proshyinsecticidescandirectlyorindirectlystimulateplantgrowth tected with carbofuran granules The other trials in the

Table 4 Yield losses to insect pests and yield responses to insecticide in upland rice in the Philippines Brazil and Upper Volta

Yield (tha) Yield loss (5) Location Year Cultivar Protected

with Protected

without Check Total Vegetative Reproductive Ripening

Basal N (kgha) Reference

carbofuran carbofuran Protected No with carbofuran

carbofuran

Philippines Pili Camarines Sur Tanauan Batangas

Pangantucan Bukidnon

Dumarao t-apiz

BrazilGoias Minas Ge~ais

Upper Volta6

1975 1974 1974 1975 1975 1976 1977 1978

1979

1980

1979 1980 1979

1977 1978 1979

Bursiging puti Kinanda Dagge Kinanda Dagge Dagge Dagge Dagge UPL Ri5 Dagge UPL Ri5 Dagge UPL RiS UPL Ri5 UPL Ri5 UPL RiS

IAC47 IAC47 IAC47

14 33 11 36 35 31 30 27 37 26 40 30 40 42 46 35

19 21 09

--

-34 24 24 27 --

29 43 31 42 -

47 25

---

10 28 05 27 27 30 26 29 33 28 33 29 43 23 36 11

15 15 06

29 15 55 25 23

3 13 -7 11

-7 18

3 -7 45 22 69

24 28 35

--

21 -11 -20

4 -

-3

23 7

-2 -23 56

-

-

-__0

6 --

-

-23 14

5 0 8 2

22

-

--

0

-

--18 11

3 -10

4 10

7

--7 -

14 -

-1 6

10 -5

8 9 7

-

--

0 0 0 0 0 0 0 0 0 0 0

30 30 30

12 12 12

99 99

100 100 101 101 102 102 102 102 103 103 104 104 104

66 66 66

Farako-Ba

Mean

1977 1978

IRAT10 IRATIO

38 66

--

40 43

-5 35

21

-

10

--

--

--

- 242 243

Z

tJ

-w

24 IRPS No 123 January 1987

Philippines which had two treatments offering protection In the 9 Philippine trials where yield losses were measuredwith and without carbofuran granules relied on further by growth stage 10 of the yield loss (21) occurred at thecontrol with foliar spray and not on seed dressings The vegetative stage and 517 occurred at each of the reproshycarbofuran-protected plots had an 18yield los compared ductive and ripening stages Ibis result is similar to thewith only 10 with foliar sprayS This yield difference is pattern of yield loss in lowland rice measured by successiveprobably due to differences in protection against sown-seed treatments where the crop is unprotected at each growthpests (foliar spray do not control sown-seed pests) rather stage but proticted during the other growth stagesthan phytotonic effects became basal fertilizer wa used in However the insect complexes r sponsible for vegetativeonly 2 of the 10 comparisons stage yield losses in upland rice are entirely diftfrent than in

The recorded yield losses were not related to yield lowland rice The key vegetative stage pests in lowland ricepotential (Fig 3) One might expect higheryielding crops to arc aquatic whereas those in upland rice are sown--seedhave higher percentage of yield losses but this ws not tile (sown in soil) root and seedling pests This dift rence maycase One reason is that insect damage exacerbates plant explain why agricul torists have overlooked tile ilmportanccinjury from drought stress low yielding fields from d ought of upland rice insect pests Many reports of upland rice pestsstress would show disproportionate differences between in Asia ha Ve focused on major lowland species (46 108)insect-protected and -unprotected treatments Shoot and It isalso surprising that tei vegctative stage of upland rice root pests would be more responsible for losses from would record the highest yield losses becatuseseeral studiesdrought stress than sown-scd pests A second reason have shown that upland rice can readily recover from foliageconcerns seed pests which would have a greater impact on reinoval during that stage (117 152 165 188) Upland ricelow tillering low yielding varieties such as )aggc Low poscsses a hitgh obilitV to compensate for ca rly loss oftillering varieties nnot fill in the space created by removal foliage and will even be stinulated by foliage removal toof seeds oweer 1) R i 5 is high yielding because it tillersP1 produce a higher viel than plants ithout any foliageactively which can close the canopy to provide greater weed removed (63 192 231) [ie reason for this apparentcontrol discrepancy hovce is that yield loss at the vegetative

stage does not octUr frol leat arca loss but fron pestsYield loss () removing sown seed and feeding on devcloping tillers and roots -- damage that cannot be readily conpLtnsatCd for

70 - Measuring yield losses in upland rice cnvironrnents is further complicated by a report from Brazil that greater yield losses occurred in response to better insect control60 - =-09 as Insecticide seed treatments resuled in denser plant stands

(n=21) which in turn created a selectively more favorable environshyment for blast (69) This resuIt was parlticurla rlxexacerbated50 when cirbofuran granules were used in seed furrows Aside

ffrom a greater plant stand ca rbofuran iay have auigmented the N lecl and stimulated blast If so ca rbofuran should be40 - replaced with another insecticide or the seeding or fertilizerI rates decreased

Other methods to determine the impact of upland rice30 shy pests wvcrc to interview farners or to develop single speciesbull bull correlations of population levels and yield loss Feneira (66)2 NIM)st (75(1) Golas farmers reporteddid both in Brazil

20- insect losses in ufpland rice and 251 said those lossesS avcragcd 37(1 [his result is consistent with tie yield losses 0 0 measured by field trials in the same statc(Table4) For every

I0 4 of seedlings removC d by leaf-cutting ants a I 7yield loss is predicted Similarly Ferreira (66) found that ai

0 - average of 53 thrips per panicle before panicle initiation meant 3 times more unfillh d grains

-IO0 I 1 1 I I-101 0 2 3 4 5 6 7

Yield (tho) C)N [ROt IM FI IODS 3 Correlation between upland rice yield and yield lols caused by Upland rice insect pests may be controlled by culturalinsects measured in 21 trials in the Philippines Ivory Coast andBrazil genetic biological and chemical methods

IRPS No 123 January 1987 25

Cultural control dry season plowing will have little effect on them Plowing Crop husbandry to reduce upland insect pest populations when the soil is wet from the first monsoon rains will expose can be classified into practices effective in a single field and soil insects that tunnel close to the soil surface to predatory those effective only at the community level Cultural birds chickens dogs and even man (field crickets nole controls should come first in a pest management program crickets and white grubs a re delicacies in man tra ditional They have broad stable effects as pests have little pos- diets) sibility of overcoming them through selection of biotyps The greatest degree i1 inIscct control fron tillage comes (138) On the other hand some cultu ral controls work only from plowing soon after rice ha rvest when most soil insects with high labor or power inputs and connunity-wilt lie close to the soil surface anrd stern borers are in the rice methods require coordination of iany farniters stuble Whitegrubsare mature and tillage exposesthem to

Pantin time The luxury of being able to shift the time of predators Tirnelv control will prtec the crop following pla ntring upland rice is open onl hr la rniers in areas with a rice and lowr the populationIfor succeedinig years Plowingprolonged rainifall pa ttern lI copping intensity or underthe ice stulble helps it decompose killing sten borer rnechanied land preparatiot Gcnerally planlting as soon larvae inside as the rFiinv season begins will lower populatiois of 1110st [requent tillage destroys ait nests bit not termites insect pests Insect populationsarelo aftcra dlvseoll or Tillage and clean culture howes er rcliove food for winter fallow delayed plaiting lets them build tip on termites which then mtay attack a young rice crop Scarab alIternate hosts that gi vigorotsly with the first seasonial beetles also prefer to miposit in eceitly tilled fields Noshyratiris This first flush iof weds cros lItxuttriartlv with iceCult is dibbling faors the survival oftillace Clre Such rineralied N released during the dr season aid made stils i aels tnd insects thtt pttpate or lay eggs- in the soil

available to the plants with the rains It iliso igrtmitcally Zero tilla hio eerl conserves predators ofthe GIH (37) advanttgeous for the rice ciop to tap this trlLItlaI fertility nt it mcttoi lo lessen the impact o0 soil insects and grow vigorotsly to compete with wrus and to tolerate brotdcasting seed or planting in ftltrrows is prefc ired over pest infestation dihlirig in hills Crickets are large enotlgh to eat seedlings in

Larly plantngs terd to escape seedling maggolts Upland a hill atrd tillering normnally ca iriot fill in tlhe missing space rice in Bitangas is planted in Ma and ine itli the Iirst Oftcn dibbll rd is not well covered and iseasily found by rains Fairners are highly mtotiated to plant early tocscape foraging insets ioIdents and birds drlitght and typlotns ricar harct Arn Atrgtst planting Plan uit sii Incrcatsini sced density protects tile cropwould be evcely attacked by It riim orivzo At tti againrst seed and seelling pests especially ants particularly other Philippiie tipland sites Rea l Qteot antd (laelit if the crop will germinate and ermerge quickly M isainis Oriental seedlig mnaggots arnrige ric boe t se hr trftinll itI India rice intcrclopped witi cotton or farnes have a longer growing season and plnit 2 rin or pigeon pci had lower 6IIt and WBIlII poptrlations than mote after the ortset of the rainy season liaaiangas ftrmners rice alone (211) NfailC arid Urplald ir_ 2re coilltori in the plant ca rly becia use ofa shorter raiinfall period arnd the desi re Philippines he chatngC in pest status folii iniercropping is to lliarvet before the typhoor sea son peaks ltighly locatioll-specific (133) cmianarid the net elfc be either

Early plantings cornbind with early-iatiuring varictics nil bcnelficial ortdetriential (189) For example BIatangias provide higher crp tolerartce for white grubs Aestiatior farmers intercrip Itro in their ipland ricefields A liornshyterminates with the first rains but 5-6 moare ineeded for the nrii iotd-Ahis i (I iiiacius) which norlially feeds resting larvae to develop into beetles emerge and lay eggs on taro ill feed or miarbv rice plants Intercropping and for larvae to reach the danriigii g ird-instar stage probably would not afft0 seed pests A crluanion crop (134 In lataigas this normually iseiough tiite for harvest planted ith rice and haing gleater tolerarnce for root

It Brazil howecver delayed planting is a suggested dariage or being toxic nayact isi trapcrop to root-feeditig cultural control for the lesser cornstalk borer and insect pests Acial-fei i ig pests voultId be most affected by pests favored l drought (66 195) Cropping internsity in intrer ropping Shoot aphids leaf beetles sten borers and Brail is low and delayed plraing places the rice crop in the hopplers would be likely ca ntidates to note effcts On the niost stable rainfall period reducing likelihood old rought othlt halnd the corlipalior crop stch~ sitniaie Iiiv be more

In rilniy arca s rinmirsare roi atl to plant other crops benehited than rice from initecropping ( 137) before rice and althNgh it woultl be desirabh to avoid 11echi Weeuling luring the first month after clop infestations by planting rice cuatl lariners lack the labor cstablishent will force pcts such is anyvworris which and time to do so ol a low priority crp prefer grassy cetds ontor tle vorling rice Soil iampsecs will

77age Uplild rice Stils ae light textured and Call be mvnye to the rice crop Cleani citrltirc also lay lore territes plowed duiring the d rv seasor notl only to grin time for to attack I u cetincti inrice ceds i ay he le optinal planting with lie fitst Inilsoon riiains but also to desiccate solition 13) soil insects arid weeds such is ntsedge that foim rhizomes Irtlizr Itneatsing N Iciiliier oin hlwland rice has Htowever becta use most soil insects lie below die plow layer fAtwvored Irigher populains of pliithoppeiv Imlhoppers

26 IRPS No 123 January 1987

and leaffolde rs However because of the need to temper fertilizer use on upland rice o prevent blast fIrtilizer rates will be lower than is comonia r for lowlad rice causing relatixely less hopper and IaIfTfoide r population buildup However inupland rice areas without a history of blast such Is Batangas farmers apply high rates of N and conscqtrently the crop has high infcstation levels of WBPI]and RIF The fhrmrcrs leec insect buildup by splitting the fertili11r in0to two to three applicatiois bit do not applV fertilizer at the vegetati estage because this prodrces a tall crop which illeasily lodge during typhoons Greater crop fertility leads to greater insect survival (better nutrition) larger insects (to cat no re foliage) arid greater fec titd t (to lav nmre eggs) (128 145 171) Not all insect species respondequally hovr Increasing rates (if N l anrid K increased Dialraa s-acchralis and (iaeiurema flea beetle but reduced thrips (66) Zn rcdLuced deadheart density from the lesser cornIstalk borer

Fnldingi Although most Upland rice areas do nIot hawxcaccess to wa ter floodiig the fields is rRrrnriierdcd in areas of Brazil or root bug and white gib control ( 195)

Crop rotatioti Tillage before planting a crop after rc harvest will control stern boreis and other pests renain n the stubble arid will unearth soil pests Planting a non-gratriiaceous crop after rice is recommncrded for termite control (66) Crop rotation is less nttdcd in upland rice culture than in lowlands because atcrop-free period is assured even after harvest of crops following rice

t1ulchilt iryl plastic irli irig designed for weedcoitrol ii upland rice inJapan resulted in ntr highanri liy infestation of (ihilo sulytresvais (93 9-1 95) NIulched Upland rice grew more luxuriantlv and attracted more (Avipositing mothis than u lched rice Also the rice plant and stein borers imatoured 10rC quickly and a partialgeneratiori af vouing ista iNoverwintcred in the stubbleLotili--malitrt varieti Quickly maturing crops reduce the tmbCr of pCst generations that Calt build up Rapid crop establishment olan ca rl v-natu ring vaitvty if)pa 11icLa r will lessen white gib and stem borer damiage White grub larvae intle last iistar are highly destIruCtive so aii early-iiatri ng variety will escape serious root loss Stem borers build 1ip slowlyx in rice to ecormic abundant in a late-Maturing crop anrd caiuse whi tlieads Numibers of somespecies increase exponeitially with each succeeding genera-ti so a xarietyexhat atures I trowh earier thian another will hac less insect damage Early-naturing vaiicties however will not lessen seed pest daiage either ii the soil or oina standing crop

Strwhoi hAIJttam Inrscts tcd irig on thlie aerial portions of rice plats disperse from carly to late plantedfields AS iiiibCirs increase exponentially with eachinsect

gtscnrattm e eiglbrig f rmc~s1(uil upp wh rc generation pests build tIP where neighboring fihrTC e9r1sstagger their plantings II Batangas farmers plant uplandrice within I iro use 120-d s-arieties and plant at the start of the rainy season Ini Misarnis Oriental laricrs stagger their

plantings up to 3 mo apart and use a set of atieties that mature in 4-7 mo Insect pest danage particularly from stem borers is greater in Misarmis Oriental Also seedling maggots damage rice there because rainfall favors frequent planting of maic In Batangas rice escapes seedling maggots bCca usC of distinct drs season and synchron0us early planting with the onset of ains

Therefore combinations of cultural practices ea rlyplanting synchronous planing crop rotation and earlyshymaturing vaneties protect the rice crop against most insect pests

Ilmant resistance Beca tse upland rice has few specialized insect pests and many insect types attack the crop worldwide regional breeding and strong internatiornal cooperation are required

Sown-sced pests arc economically controlled by insectishycide seed treatment and should the relore receive lowxpriori ty inbreeding objectives

Root pests are normally cost ly to control with insecticides but finding resista t sonres mayvbe di fficult Tanaka (229) for exa -ple fi led to find varieties resistant to root aphids Selecting for large root bioniass perhaps should be the strategy for pests sLuch as Whlite grubs that renliove roots Fainter(174) howcvcr cites varietiesof crops tolerant ofor resistant t(i root pests Some sorghun varieties are high in cyanide so rice sorghun might incorporate broadshyspcctruri chemical resistance

Seedling vigor and drought tolerance are high prioritybreeding objectives so tupland rices should have large r root systernsand be able to tolerate ligherlevels of root loss Sonie root loss inolder plants is beneficial if it stimulates new root developnrut and therefore crihances intake of soil nutrients (53) The rice seedling inaggot rtiLotia spp may be

controlled thirough resistant aittics A hi rgc breeding effort on sorghniu has had fair success inideveloping arreties esrstait to Asoccata Rodani (251) A field trial comparing 10 upland ricc varieties showed differences aniong varieties va ring from II to 4(- damaged tilleis (215) Hovver resistarce to 1Iill-0r gtia species may rot cross ox-rti otier

Shiraki (210) reported that upland i-ice stens wcrc harder than lows-land rice stemsperhaps becaIusc (f a higher silicoricontent and thus were more resistant to yellow stern borer hc tortia Iow tillering (f uplaid iice xarneties prodtres a

higher percentage of infested tillers than would occor in higher tillering lowland iice under a siiilar egg density The corollary therefore is to select for high tilleriig upland rices

Stemii bor amcstheh main targes dobrecders inAfrica (571 204 )but nticl oifth e sc reen irig is dotie unde r lo wland co d t osrm s wih A h r~ o l pe i s n acorditions (11 96 218) As with lthrinrnwspecies mayhave to be dealt with independently however initial results show cross resistance between African and Asian species(92)

In Brazil screening Ior insect resistance focttses oil lou I pest problems Varieties tolerant of lasmolwalpus ligi-sellus and resistant to )iatraea saccharalis have been identified (70 153) Current work also emphasies 7iraca limativetntris and Thmialspis (=Iosis) flavouicta (66) These insect pests Ilt onlV citise IIigh yield losses tihey also ire difficult to control chemically ofr by other ilealls

ManV upland rice vaieties aic early maturing and are resistant to pests such as gall midge (I Xl ) Early mIturity is often a highly desirable trait to esuilpe pest buildup but it is not true resista nce bca usc ilthose va reties are plinted late they will be itsdamaged as ain early planted susc ible variety

P~erhaps tile cosiiiopolitall of all upland rice pestsimost are the sced pests (ripenling stagc) which pose a particular problem beta use scuds re ised IsIO for hitnit4ans Ayltoxic substatice introd tcld geticticllyilmay also be toxic to nal iniless it caln he placed old in thfe IdhuffSelectmi fo hairiness or lotng wns kill mIlct sthilrohleiiis dutilig threshing pa rtictfarl maitial threshinfl [hireshers ill complain of itcfimin atid itteascd (fust levels

Attempts itIRRI to fitid tcistatncc to the rice bug plorisa have ilotbeCnl prmitisilg lIIetIattonlids iorc

through the lenilna or palca hillhairs sedstVIor seeds wilh awns that deter bugs promote complaints from threshers Seed bugs liptocorisa and Stcnocoris enter the seed through the opening between tile lemna and palC but varieties with narrow openings are diifficult to mill

Breeding priorities should concentrate ott inotCI gruLtLIps that are most difficult to c nttrol by other meains Steil) horers appear to be the most widespread group that sigiiificantly dalag s rice aind is hard to manage by other Ilnca us

Biological control Very fe stutdic of mitatt~l eneties of rice pests in upland envirollnletits wee in tile wefouiid literature Therefore have not compiled a list of species Ve present a list of natural enemies iOtid in upland ricefields tuic latialln Batangas for some conitno iOldiar pest species ([itble 5)

Muich ofl thc lollowinle itfor ittiol cotlliCs rol Stutdies in the lowlatnds or roi crops otithcr thlin tice

Qu(ailtr o 11ll t11t( Nat trrtf lii hcoeits ight Ile used agaitist ile social insects tilt and termites Sollie predator specice live ill theirf nests n ilnic theirnllt ip-pearanIlCe aMRidbehavior IJtrouMdCing a mote aggressise ant species to displace pest ant codonies is a possibility (169) Patihogens as potential natural encnies have reccived little atteition (I13) Natural ncnuics asa group however hac not been toIl iporta it in Iegulating aintshown be and ternite numbers

The nocturnal behavior aid subteitmien habitat of lole crickets and field crickets protect them from many ntllral enemies There are only limited records of natural cnelics for these groups of pests some pathogens (87) but mainly

IRPS No 123 JattuIny I 271987

parasites - scelionids on eggsand sphecids on ilymphs and adults Sphecid wasps Larra carutmaria (SImitI hI L huzwuensis Rohwer and lsaltiuhta Willi ams speciali in mole crickets pa ralyzing them nd dragging the ninto thcir nests as food lot their voting ()tier sphCcid species Lins aurulenoa (Iabricius) lot maiac (Ashmead ) 1 lhshylssellaus (Smith) and Ml lal)oriosu (Snithf specialie in field clickets

White grUbsO the other hand hae Icti scriuns enuigh to have been extensively studied for cotttroi 1W nati al enemies( 112 140 141) Scoliid parasites C(ajsoMcispp have beetn intrtduced itto tilePhilippites for control of Ielxl ]udisiro la The rcsearchcts w Intttrdced themle during ill era claimed that the white gttionthreak as controlled (142) howevcr it isnormal for pest epidemics to subside and there was no diect evidence to indictte that tfe parasite was responsible A tmore promisitg titctllod might he to identify iso ltcttass prt~f tcartd apply hiactelaln fingal pathogens to tile soil Ihe lalattese beetle in North America has hcci coitrolled hv tite conmnleciall aailable hactcrial prcpa ration itacihlpu pav ofr milksdiscas (71) Spote prtyefott the soil itloted ilrtier reitiain actic fr Veas

Root aphid sardtnctly bugs readilslic tilib to predatots and parlsites once their tetndine ants are controlled The larvac and adtilts of a coccinellid beetle 1Scynms sp prey on llriadlouais in slash and burn upland rice Baits treated with insecticide may be used to control tending ants by killing their ourig (244)

Root-tkcditig blugs false wireoris wireorms and root wccils aic r ofsuch little ifilpoltatice that thenomttll role of liatural Cticniics hias tiot beei asscssed A possibility hoevrl tiglt he tit ItaIIitic niaCIMtodcs lintt1g11en Itese soil-itlhlfbitine mCtitodcs W5uhtfhave to be imiass Produiced

large miltibcr of niatltrll cnemics ircknown ft CltIoriS IC IpblCeii is thtt clttvortnlls ofonlie the crop soott alter land ptel)tritiOti a1tithe bcgit iniig of the raittv

Citsoln hfien tiatllral leniiiy populations aric lo Colltloil WOtuld iii ole mass prodtticing andirclcaslng key spccies

le egg stage is the fpirt (olthe Cedlitgt Ialot life cycle tI hich to f)ctis ftor tittlril elctit cottntrol hplraites o1 pleditors ttsit elleggs highly expoIsYCl I Sgg pailsites or predators stolld probahly e ittass andtbe produced released Littlh i toruialioll is a ailable ott what species to trV o ho to Iniss podLice thiCIi A etiloplhid wasp Icmihlaol sp effctiicly regushy

lates the leatf lcloalapolm2a spioanoiiitr heat andc rice antid should be collsctvcd

leaf beetles whCllose e am] eggs ate laid in the grutindhirO ould be motC difficutlt to control by niatulral enelV Mnll ipuhltiOn But species sUici IS c_gsthispid beetles ith

and larVle oilleascs ate vtilcratble to iparaites and pledators

Natural enemisc of tlirips otl rice are little known

28 IRPS No 123 January 1987

Table S Parasites recorded from upland rice insect pests in Tanauan Batangas Philippines 1977-84

Order Family Species Host stage

I lymttenoptera Braconidae Leaffolders Cnaphalocrocis Marasinia spy

Cardiochilesphilipplinensis Ashmead a Larva

Cotesia angustillasis(Gahan)Cotesia or cypris (Nixon)

Larva Larva

Cotesia or taeniaticornis(Nixon) Larva Cilonusinotakatae Munakata Chelonits SppAlacrocetrts philippinensis (Ashtmead)

Egg to larva Egg to larva Larva

Iclineumonidac Orgilts spp Isclutojppa hireator(Fabricius)

Larva Larva

Itoplectis naraigav(Ashmcad) Larva Tenelhtcha philippiensis (Ashmead) Larva Tenelchha stangli (Ashmead) a

irichomma cnaphalocrosis Uchida Larva Larva

Bethylidac Elasmidac

Goniozms or trianguli r Kieffer lasmtusalhopictus Crawford

Larva Larva

Diptera

Encyrtidac Chalcididac

Tachinidac

Elasmtus spp Copidosomopsisnacohiae (Fidy) Brach nteria evcarinata Gahan Biachyieria lasus (Walker) Ztgohothria ciliata (Wulp)

Larva Egg to larva Pupa Pupa Larva

Argyrophylx nigrotihialis(Blaranov) Larva

l-ymenoptera Braconidac Yellow stern borer Scirpopiaga incertulas

Braco chinensisSzeplegeti Larva CIthelotis spp Chelonus niunakatae Munakata Cotesi flaripes Cameron Stenoracon nicerilli (Bingham) a

Larva Egg to larva Larva Larva

Ichneumonidac Tropobracon schoenobii (Viereck) Amatroinorpiaaccepta metathoracica(Ashmiead)Frihorus sinicus (I lolmgren)

Larva Larva Larva

Ischnojoppaluteator (Fabricius) Larva Isotima nr dammeratiti Rohwer Larva Tenelucha philippinensis (Ashmead) a

Larva Tenehicha stangli (Ashmcad)Trichomma enaphalocrosis Uchida

Larva Lara

Chalcididac Eulophidac Trichogranmmatidae

Xanthopimnpla stemmator (Thunberg) Charops hrachYpterumn Gupta and Malteswary Brachtm eria sppTetrasticttus schoenohii Ferriere Trichorigranm chihunis Ishi

Larva to pupa Larva Pupa Egg Egg

Diptera

Scelionidac Pteromalidae Tachinidae

Tt ichograintnta aponiicuitt (Ashniead) Telenomits rowani (Gahan) a

Trichoinalopsis apattelctena (Crawford) Jeirhaea spp

Egg Fgg Egg pupa Larva

Z4gobothria ciliata (Wulp) Larva

Ilymenoptera lecneumonirlac Gol fringed borer (hilo auricilius

Trichomm enoptalocrosisUchida Larva

Ieioenoptera Braconidae

Aantholimlh steiniator (Thtinberg) a

Pink stet Ioret Sesamia inferens StoInacoil nicrilhi (Bingh am)

Loarva to pupa

larva

Diptera Ilyinenoptera

Armvtnyorms Mythimna separata Spodoptera mauritia acronyetoidesTachinidae Zvgohothriaatropiora( Robinea i -Desvoid y)Eulophidae It-phctrtiscitapadoe (Ashimead)

Larva Larva

(halcididae Braconidac

Brachpin ria spip (oesia spp

Larva larva

IHyntenoptera I)iptera

Rice skipper Ielop ida (halcididae Tachinidae

math ias aidgreen iortedci ciliarNIelan itis eda Brat lhincria sp iii matuqitatt Cameron Irgyrolhirlaxi trotlhiais(marumtv)a1

ismtne Larva Larva

Diptera Ilymenoptera

Tachnidae lelineutionidae

fuown se tilo per Mois frugalisIrgtrophrlax 1igrothiahiv 3 a rat ov) anthopimpa pmnctata (Iabricius)

Larva Larva to pupa

aMost dominant

IRPS No 123 January 1987 29

Normally predatory thrips prey on herbivorous thrips strepsiptera on both Spiders dominate tilenymphaladultNatural enemies of stem bugs --Scolinoplara and Tibraa predators Spittle bugs move to rice from pasture grasses - are in order of importance egg par~isites and predators and biocontrol would be successful only ifcarried out before fungal pathogens and nymphal adult parasites Egg tiley immigrate into ricefields parasites could either be introduced fron other areas or Biological control tactics for hemipterous seed bugs mass produced and released ould follow those outlined for stein bugs

Armyworms are normally lwld in hcck by the activities Coml)arison of environnewtns Natural enemies are of egg and larval parasites When these natural enemies fail perhaps faced with even greaterconstraints than insect pestsusually because of drought armyworns become epidemic to survive ind reproduce in upland environments TheyIt may be worthwhile therefore to release parasites during have adapted ani survived by having wide host ranges the the rainy months following a drought Virus diseases of ability to aestivate or hibernate over unfavorable seasonslarvae would beanotheravnue to exploreforarnyworms and the ability to disperse or by other mechanisms

Parasites have ben traditionallyconsidered for biolegical Comparing the most dominant and therefore the most control attempts against stein borers usually by rekasing adapted species of natural enemies collected in various exotic species( 166) This approach has not met with success environments in the Philippines we can understand how and new avenues should be explored The role of predators different rice environments are arid pathogens is little inderstood and deserves greater Anagqrms olptahiisa mymarid planthopper egg parasiteattention Egg parasitism rates are normally reasonably is cquallyadapted to all Philippine rice environments (Tablehigh but should be supplemented with effective egg 6) It is selective for planthoppers and even parasitizes eggspredators Orthopterans are potential egg predators of the n-iize planthopper Peregrinitsmaidis itt upland areas Metioche vittaticollis (Sta ) Anavipha nifiwnnis (Saus- Alavohis also parasitizes maize planthopper eggs but is sure) and Conocephahs hongipnni (de i-laan) feed on most prevalent on BPH in the irrigated wetlands (onashystem borer eggs etioche and Anaxil)ha specialize in eggs tocerus spp like A optalhilis on BPH and WBPI Iattacks with no hair such as Chilo spp The omnivorous (ono- the eggs of GLH in ell rie environments It is also found cephahsspecializes in eggs cov red with hairs it is a proven parasitizing the eggs ef the white rice licahopper (oimna egg predator of Scirlouphaga spp but has the discouraging spetra Oirosita naias a trichogrammatid egg parasitehabit of eating rice guin (65) attacks BIPH and WBPH in the wetlands and has no other

Grasshoppers have egg (scelionid) and nymphalad ult host A related species 0 aesopi specializes in G1-H in (nemestrinid tachinid and sarcophagid) v irasites Among wetland environments Dryinid strepsipt-an and pipunshypathogens protozoans have been recorded most frequently culid nymphal and adult parasites of planthoppers and Control of locust species by natural enemies could con- leafhoppers also specialize either in hopper species or centrate on their habitual breeding areas environments Among the dryinids Pseuhdgonatoipsflavi-

Leaffolders have rich complexes of natural enemies some feimur prefers the lowlands and BPH while P nuch has no adapted to upland rice (22 27) ranging from egg predators environmental prefirences but is adapted only to WBPH (gryllids coccinellids) to larval parasites (braconids ichncu- The main d ryinid species on GLH aptogonatop)usspp is monids) to larval prcdators (ants and carabids) to larval most dominant in rainfed lowlands and also attacks other pathogeiis (vira and fungal diseases) Efforts to augment leafhoppers within Amnrasca Cicadulinaand Bachaltuathese natural enemies should focus on the egg predators Strepsiptera are more dominant on plarithoppers than (rearing releasing)and viral or fungal pathogens (culturing lealhoppers The Elenchus species in the Philippines attack spraying) all species of planthoppers within the genera Sogawilla

Parasitization of the large conspicuous larvae of Sogatodels NilaparsataHarualia and Opiconsiva Elenshyskippers green horned caterpillars and other polyphagous cusrvasuntaisuiis adapted to the major rice environments Lepidoptera by tachinids and chalcids is normally low ELg but prefers BPH in rainfed arid WBPIH in wetland environshyparasites and predators are perhaps the key to their control ments Ilalictoph(utsmtnroeiand HVpectrus attack G LH

Gall midges are parasitized by pterormalids and platy- and COfimravcctra respectivelygasterids that aestivate within their larval host between Pipunculids are mainly parasitic oii lealhoppers Each seasons Shoot aphids are preyed upon by coccinellids species has a unique environmental preference Tomosshysyrphids and chrysopids varireIa oraetoraprefers Ne7 otetltix nigropictrs and

Planthoppers arid leafhoppers have egg parasites and Pilunwulhtsjavanensisattacks Dehoceplhahls spp which predators as well as nymphialad nlt parasitesand predators are more abundant in upland than irrigated lowland their rich complex of natural enemies includes fungal environments T suhvirescensattacks Nephotettix virescens pathogens The principal egg parasites are myrnarids and and prefers the lowlands Pilpuiulus nttillattls is an trichogramniatids Egg predators are norinally mirid plant upland rice parasite arid attacks only Nephotetix speciesbugs that suck out the yolk Nymphal ad nIt parasites are More than 17 recorded species of leaffolder parasites in dryinids on planthoppers pipunculids on lealhoppers and the Philippines attack species of C(naplwocrocis and

30 IRPS No 123 January 1987

Table 6 Dominant natural enemies of rice pests in upland rainfed lowland and irrigated lowland rice environments of the Philippines

Natural enemy

Host stage Species

Brown planthop-Parasite Egg

MymaridaeAnagrus optabilis(Perkins) Anagrusflaveolus Waterhouse

TrichogrammatidacOligosita naiasGirault

NymphAdult Dryinidac

Pseudogotatopuslaiifemttr Esaki et Hashimoto Elenchidae

Elenchus yasurnatsui Kifune and Hirashima

Environment

Upand Rainfed lowland Irrigated lowland

Ailaparata lugens

x x X x

x

x x

x X X Whitebacked planthoppers Sogatella furcifera and Sogatodes pusanusEgg

MymaridaeAnagrus optabilis(Perkins)

TrichogrammatidacOl~gositanaias Girault

NymphAdult Dryinidae

Pseudogonatopusnudus Perkins Elenchidae

Elenchus yasutatsui Kifune et Hirashima

Parasite Egg

MymaridaeGonatocerus spp

TrichogrammatidaeOligositaaesopi Girault

NymphAdult Pipunculidac

Tomosvaryella subviresccns Loew Tomosvaryella oryzaetora (Koizumi) Pipunclusmutillatus (de Meijere)

DryinidaeIlaplogonatopusspp

Egg Copidosomopsisnacoleiae (Eady)

Larva BraLonidae

Cardiochiles philipphnensisAshnead lchneumonidac

Trichiomnia cnaphalocrosisUchida Temelucha stangli (Ashmead)

Pupa Chalcididae

BrachymeriaexcarinataGahan Brachymerialasus (Walker) Brachymeria nr tachardiaegroup

Parasite Egg

Scelionidae Telenomus rowani (Gahan)

EulophidacTettasticusschoenobhi Ferriere

Larva Braconidae

Stenobraconnicevillei (Bingham) Iclhncurnonidac

Temelucha phiiippinensis(Ashinead)Temelucha stangli (Ashmead)

x x x

x x

X x X

x x Green leafhopper Nephotettix virescens

X x x

x x

X X X X

X

X Rice leaffolders CnaphalocrocisMarasmia

x X x

X

X X

x x x x x X x x x

Yellow stem borer Scirpophaga incertulas

X X x

x

x

x X XAinauromorphaaccepta metathoracica(Ashmead) x

continued on next page

IRPS No 123 January 1987 31

Table 6 cntinued

Natural enemy Environment

Host stage Species Upland Rainfed lowland Irrigated lowland

Larva-pupa Gold fringed borer Chilo attriciiuslchneumonidae

Xanthopimpla stemmator (Thunberg) x aPupa x

Pteromalidae Trichoinalopsis apanteloctena (Crawford) ax

Pink stem borerb Sesamnia inferensParasite

Larva Braconidae

Stenobracon nicevillei (Bingham) x b b Armyworms Mvthinna SpodopteraLarva

Tachinidac Zygobothriaarroptvora (Robineau-Desvoidy) x x

Braconidac Cotesia spp

X Skipper Pelopidas mathias and green horned caterpillar Melanitis leda ismeneF-gg

TrichogramrnatidacTrichogrannaspp x

Scelionidae Undetermined Scelionid

x Larva

Tachinidae Argyroph lax nitrotibialis (Baranov) X x

Pteromalidae Trichomnalopsis apanteloctena (Crawford)

X Predator

CoenagrionidaeAgriocnenis femina femina Brauer x xIschnura senegalensis tambur

Miridae CrtorhinsliidipennisReuter x x

Nabidac Stcnonabis tagalicus (Stal) x

VeliidaeAlicrovelia douglasi atrolineata (Bergroth) x

Coccinellidae Micraspis crocea (Mulsant) X x x

Vespidae Eumenes campaniformis (Fabricius) x

Formicidac Diaecamna spp xOdontoponera transversa (F Smith) x

Carabidae Chaenius spp OphioneaW- aJasciataSchmidt-Goebel

x

Ophionea ishii ishii Habu x x X

Gryllidav Anaxipta longipennis (Saussur-) x xMetioche ittaticollis (Sial) x x x

Tettigoniidae Conocephalus longipennis (de Haan) x x x

aNo rearing made on rainfed lowland bNo rearing made on rainfed lowland or Irrigated lowland

MrasmiaAmong the Laval parasites the braconid Cardio- lowlands and parasitizes oriental maize borer OsIriniachilesphilijpphensis is most adapted to an upland environ- furnacalis (Guenee) larvae The leaffolder pupal parasitesment It also parasitizes Hydelepta indicata a common not only occur equally in all environments but also haveleaffolder of legumes The ichneumonids Trichomma cna- wvide host ranges- parasitizing species of OsIrinia Marucaphalocrosis and Temehcha stangli occur more in the Hydelepta and Honiona - in maize and legumeslowlands T stangli is most adapted to irrigated lowlands The principal egg parasite of yeltow stem borer (YSB)and also parasitizes Chilo and Scirpophaga stem borer Telenomus rowani is widely adapted to all environmentslarvae T cnaphalocrosis is most prevalent in rainfed and has an unusual alternate host -Tabanus eggs (25)

32 IRPS No 123 January 1987

Tetrastictu schoenobii specializes only in YSB and occurs mainly in irrigated lowlands -- further proof that YSB is most adapted to the wetter environments

Larval parasites of YSB are highly environment-specific The braconid Stenohracon nicevillei occurs only in upland rice but also attacks the pink stem borer (PSB) Tieinchwa phiiiplPwnsis an ichneumonid occurs in upland rice and also parasitizes the larvae of (7hiu stqpresaisinatro-morpha accepta melaihoracica like 7 stangi is mnost adapted to the irrigated lowlands

The dark-headed stem borer(l)HSB) (ih pochit3sus was not collected in all rice environments and the gold fringed borer Chilo auricilius nd PSB were collected only in upland rice Xaantiholiplastenrnaloran ichneumonid larval parasite appears to be widely adapted whereas TrihomaslopisapanitI)cteiiaa pteromalid pupal parasite occurs mostly in wetlands and also parasitizes larvae of Chilo suppressais and Pelhuidas mathias

Armyworms Myit hinin and Spodujipera have two main larval parasites - a tachinid Zrgohothriaatropivora in rainfed environments and a braconid Cotesia spp in irrigated lowlands

The butterfly pests Pelopidas and 4elanitishave tricho-grammatid egg parasites found only in irrigated lowland environments

Two larval parasites show environmental preferences --Argrroph av nigrotihialis a tachinid in rainfed habitats and Trichonialopsisapantelkcnela in irrigated ricefields A limgrotibialisalsoattacks the sweet potato hornwormAgrius convolvtdi and T apanteloctena parasitizes DH S1B

Predators as a group show more distinct environmental preferences The aquatic damselflies Coenagrionidae are most abundant in the lowlands Cvriorhinus is also most adapted to the lowlands The nabid Stenonabis tagalicus occurs only in the uplands and isalso prevalent on legumes The coccinellid lady beetles wasps and ants are more prevalent in upland areas AlicrasIpis cit-ea larvae prey ona variety of aphids attacking legumes and maize Etunienes campanonia vespid mud wasp makes nests in trees and is therefore most adapted to the more botanically diverse uplands Soil-dwelling ants cannot tolerate flooding

Upland environments are habitats to arboreal carabid beetles Three species of Chlaeniusbeetles prey on leaffolder larvae and are perhaps more important than parasites (27)

The gryllids and tettigoniids are widely adapted egg predators not only on rice but also on maize and legumes They feed on eggs of most insects that are laid on leaves

The spidercommunity of upland rice environments isrich in species At one site (Batangas Philippines) 31 species have been recorded (23) Of the 176 spider species recorded in Philippine ricefields about one-half (82) occur in upland rice The spiderspecies of upland rice environments overlap more with those of rainfed lowland than with those of irrigated lowland rice (24) The same study showed that of the three environments irrigated rice has (ie greatest spider species diversity foiiowed by rainfed lowland and upland

However of the 10 most prevalent ricefield spiders 9 were abundant in upland rice showing wide adaptation All three spider guilds -- orb-web space-web and huting spiders shywere prevalent in upland ricefields There were however differences in environmental prelerences for some spider species

Of the orb-wcavers Tetragniathamanlihulatawas partishycila rlv abundant 7 japonica was low and Ltucatge dehcorata was absent in upland rice

As in other environments Aiypeiwnaifirtosana was the most dominant space-web spider however its relative numbers were lowerthan for lowland sites Among hunting spiders Ox|opes javanus and Lcosa leatcost fgla were more abundant and Lvcosa lpseudoanmnlata was less abundant in upland compared with lowland rice envishyroniments

Extensive rearing of rice insect pests in upland rainfed lowland and irrigated sites over an 8-yr period in the Philippines provided insight into the effectiveness of parasites as natural enemies (Table 7 8) Most of toe data are from large sample sizes taken over at least one crop which overcomes the pitfalls expressed by Van Driesche (238)

VSB egg parasitization was surprisingly similar across all environments Larval parasitization was highly dynamic among sites and even years within the same site but showed somewhat lowerlevels in uplandsthan in irrigated wetlands This would have been predicted for egg parasites as well since YSB is adapted to the lowlands and so must be its parasites This isevidence that Telenomus rowani is as good at dispersing as its main hosts Trowani clings onto the anal tufts of female YSB and parasitizes eggs as they are laid This phoresy may explain its wide adaptation The more specialized lowland YSB larval parasites therefore appear more effective than their upland counterparts with more alternate hosts

Low collection levels of PSB larvae especially in the wetlands make comparisons between environments difficult Low levels of larval parasifization were recorded in both rainfed environments Except in the Cagayan rainfed wetland site in 1980 low levels ef larval parasitization were evident at irrigated and rainfed sitesalike No differences by environment were evident Higher levels of parasitization occurred in Sarawak during a 1967 outbreak also from tachinids Rothschild (197) indicated that higher parasishytization occurred in upland rice areas

Skipper and green horned caterpillar larval parasitization rates were generally low and showed no environmental effect htighest activity occurred in Cagayan in 1980 (48 parasitization) Although larval parasitization levels of RLF were variable year to year activity appeared greater in the irrigated lowlands than in the uplands Again the highest levels were recorded in Cagayan in 1980 Cagayan has extensive grasslands and fallow around the rice areas and a relatively long rainfall period Rainfall in 1930 was good for crop growth and apparently for increase of larval parasites

IRPS No 123 January 1987 33

Table 7 Parasitization rates of rice planthoppers and green leafhopper in upland rainfed lowland and irrigated lowland environments Philipshypines 1976-840

Environment and site Year Egg parasitization () Nymphaladult parasitization ()

BPH W13P GLH BPII WBPH GLIH

UplandBatangas

Rainfed lowland

1976 1979 1980 1982

15 17 12

2(47)8(39)

4(51)

9(1800)14 (212)

12(304)

8 (101)6 (127)

9(116)

Iloilo

Cagayan

1976 1977 1978 1981 1980 1981 1982

0 (69)

11

44 48 40

-

-

46 55 44

0 (471)

11

52 56 49

0 (200)9 (214)

10 (1259)8(235)6 (900) 3(800)

-7 (462)

11 (270)3 (313)5 (900) 1 (800)

4 (600)10 (725)10 (362)12(870)5 (900) 3 (800)

Pangasinan

Irrigatedlowland

1984 1976 1978 1979 1982

13 (436) 13(379) 14 (457)

6(14)7 (197)

17(198) 11 (486) 18 (275)

3 (81)10 (274)3 (116)

16(871) 19(88)

28(211)9 (764)5 (647)7 (1204) 8(305)

Laguna 1977 11 7 21 1978 31 1979 14 4 4 28 1980 7 8 20 1981 1983

56 7(278)

26 16 (1650)

16 27(1921)

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R I Apostol unpublished 1lo1lo and Pangasinan 136 Cagayan BCanapi unpublished Liuna 175 I C Pantua unpublished 118 234 43 Carino and Shepard unpublished BPH = brown planthopper Nila parvara htgens W131Pi =whitebacked planthopper Sogatelia furcifera GLH = green leafhopper Nephotettrix virescens Figiires in parentheses are sample sizes

Tauher et al (230) reported that internal external and genetic factors influence the seasonal activity of parasitoids

Hopper egg parasites but not nymphaladult parasites were also parlicularly abundant at the Cagayan site possibly the egg parasites have more alternative hopper prey in the weedy areas than nymphaladult paisites

The seasonal dynamics of hopper egg parasites can be seen for upland rice in Batangas(Fig I 2) and for rainfed lowland lice in Pangasinan( 135)and Cagayan( 105) Levels of BPHWBPI-L and GLH egg parasitization were corn-parable in upland and irrigated lowland environments Except for some high rates of paraMsitization early in the season the parasitization rate tended to be steady in all envilonments However hopper nymphaladult parasi-tizution levels were more similar in the rainfed environments and lower than in irrigated rice

Overall parasites do not appear highly effective by themselves against upland or even lowland pests Among natural enemies parasites have attracted the most attention because they can be more readily assesscd

Predators and pathogens lend themselves better to management practices Predators the most important group of natural enemies in riccfields are difficult to quantify On the other hand pathogens appear less im-portant but can be readily cultured and disseminated

The natural enemy commulnity of upland rice is rich in species and differs significantly from that of lowland rice

These beneficial organisms must be conserved by applying insecticides judiciously particularly sprays Seed or soil placement of chemicals minimizes exposure to natural enemies The strategy to derive the greatest benefit from natural enemies isto allow the greatest number of beneficial species to thrive Then perhaps at least one species will be effective against each pest at any tine overcoming the variable abundance of each species within and between years

Programs to introduce exotic species or mass produce indigenous ones are ambitious and expensive because they require trained people on a sustained not ad hoc basis

Chemical control Insecticides offer rapid and efficient control of upland insect pests (190) but-should be used only after other control measures have been considered Insecticides are rarely used on upland rice because of cost (32) Upland rice yields are normally too low to justify the expenditure Also most upland rice is grown as a cash crop Spraying upland rice is more difficult than lowland rice because water is less accessible Government subsidy programs to provide pestishycides to tribal peoples for upland rice production have created problems of toxicity to humans because isolated people have had no experience to be able to handle pesticides safely

However ins-eticide use has beenjustified when a crop is

0Z

Table 8 Parasitization rates of lepidopterous pests in upland rainfed lowland and irrigated lowland environments Philippines 1976-84a

Parasitization (5)

Environment and site Year YSB GFB PSB Earcutting Swarming Skipper Greenborned RLF Brown larva larva caterpillar caterpillar caterpillar caterpillar larva semilooperEgg Larva larva larva larva larva larva

UplandBatangas 1977 - 0 (57) - 0 (63) - 0 (71) - 0 (106) 11(142) 2 (101)1978 43 (4697) 8(451) 5 (38) 3 (40) 6 (61) 2(58) 15 (57) 13 (129) 13 (1286) 4 (30)1979 59(4135) 12(320) 13 (11) 5(28) 9(45) 6(46) 7(23) 8(160) 9(1152) 8(26)

Rainfed lowland Iloilo 1976 - 0 (10) 0(10) 0 (7) 0 (19) 2 13) 15 (180) 5 (76)1978 0 (54) 0 (3) 0 (17) 9 (64) 4 (78) 3 (73) 0 (121) 0 (191)1979 64 (2350) 16 (84) 6 (17) 0(6) 18 (68) 25 (104)

1981 56 (1322) 18(304) 0(5) 12(51) 18(138)Pangasinan 1976 60 (7 7 )b 62(82) 0(111) 5 (39)

1978 58 (53)b 5(51) Cagayan 1979 59(713) 8 (101) 6 (34) 0(7)

1980 0(11) 4 (28) 15 (286)

62 ( 48(177) 61(140)1981 0(5) 12 (2) 13(45) 14 (92)

Irrigatedlowland Laguna 1974 62 8

1974-75 50 41 1975 43 72 1978 96 (623) 21(24) 33 (60)1979 83(2017) 18(110) 8(62) 11 (89) 22(400)1980 78 (1625) 20(75) 0 (16) 4 (72)1983 60

9 (80) 7 (270) 30

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R F Apostol unpublished Iloilo and Pangasnan 136 Cagayan B Canapi unpublished Laguna 175 P CPantua unpublished 118 234 143 Carino and Shepard unpublished YSB = yellow stem borer ScirpophagaincertulasGFB = gold fringed borer Chilo auricilius PSB = pink stem borerSesamia inferens earcutting caterpillar Alythimna separata swarming caterpillar Spcdoptera mauritia acronyctoides skipper Pelopidas mathias reen horned caterpillar Melanitls ledaismene RLF = rice leaffoldersCnaphalocrocismedinalis and Marasmiaspp brown semilooper Mocisfrugalis Figures in parentheses are sample sizes Egg masses

IRPS No 123 January 1987 35

Table 9 Effect of seeding rate and insecticide on plant density and Yield losses from insect pests in upland rice often areyield of Dagge upland rice Batangas Philippines 1976a comparable to those of lowland rainfed or irrigated rice

Seeding rate Insecticide Dosage Plant density Yield (0-Y30o) but usually from different pests The principal (kg aiha) seed treatment (kg aiha) (noIra-row) (tha) groups are soil pests feeding on sown seed and rootsat 14 DE followed by seedling pests These groups have largely gone

50 None - 35 a 20 a unnoticed leading researchers to conclude that stem borers50 Carbofuran F 10 38 ab 23 a and seed bugs are the most important pests (17) Yield loss 10 beCabofuranF100 None - 4551 c 23 a trials that pinpoint major pests need to includc seed

100 Dieldrin WP 10 52 cd 22 a treatments Granules applied in the furrow may not give150 None - 59 d 22 a adequate protection from many seed pests

aAv of 8 fields DE = days after crop emergence In a column means The floristic and pedological diversity of upland rice followed by a common letter are not significantly different at the environments is matched by the diversity of arthropods5level by D)MRT F = flowable Wi = wettable powder both pests and their natural enemies Irrigation has homoshy

genized lowland rice environments Flooded soils tend tothreatened by epidemics of locusts or armyworms New have the same properties and to eliminate soil pests The technology stressing efficiency has minimized insecticide spatial and temporal dominance of cultivated rice has cost benefited specialist pests

Seed treatment is normally inexpensive and can be Upland rice has no specialist arthropod species The economicallyjustified in -nanycases(66) Treating seed with monophagous pests on upland rice prefer the lowlands systemic insecticide (216) but not chlorinated hydrocarbons where rice evolved Rice in the uplands exists only because (247) is most effective in controlling seedling maggots of man and even though lowland rice predates upland riceFoliar sprays have to be repeated to obtain control and specialist species evolved Wheat maize and sorghum do often are washed off by frequent early season rains not have monophagous pests leading us to conclude that Granular insecticides are effective but required dosages are there is no evolutionary advantage for inssects to have too expensive to justify except under heavy infestation narrow host ranges in upland graminaceae(247) When directed against first-instar larvae banding Pests have adapted to the uplands not only by having granules in seed furrows (177) can be effective against white wider plant host ranges but also by having long life cyclesgrubs at low dosages (025 kg aii ha) (134) Seed treatments undergoing dormancy or being dispersive Each pest has a are not promising for white grub control (248) unique life history strategy suited to highly unpredictable

Seed treatment is also effective ainst ants and may be environments In agivenyear someinsect specics an better cheaper than increasing the seeding rate In Batangas exploit upland rice as a food source but no pest is a senious treating 50 kg seed ha was as effective as increasing the one every year on upland rice seeding rate to 100 kgha and using untreated seeds (Table The low yielding potential of upland rice means less 9) In Brazil seed treatments control termites and other soil quantity and lower quality of food for pests to exploit but insects including the lessercornstalk borer(66) Domiciano also less incentive for farmers to undertake control (58) however found seed treatments inconsistent in their The most effective control would be early and synchroshyeffect against high populations of lesser corn stalk borer nous planting of early-maturing upland rice varieties to

Baiting can be an inexpensive way of controlling ants prevent insect pest buildup Breeding for insect resistance seedling maggot flies mole crickets and field crickets Baits should initially focus on stem borers because they are verymade of locally available material and impregnated with difficult to control by other means Biocontrol efforts insecticide can be sparingly distributed taking adantage of should first stress conservation of natural enemies preshythe pests ability to disperse and encounter bait sites dators and pathogens rather than parasites Efficient

The new low-volume hand-held controlled-droplet chemical control includes seed treatment or baiting for seed sprayers may offer an advantage over high-volume knap- seedling and some root pests and using controlled dropletsack sprayes for controlling foliar pests Upland rice sprayers for spot treatments against foliar pests and seed canopies are more open than those in lowland rice and bugs when economic thresholds are reached better droplet penetration should result Controlled droplet applicators would be ideal for seedbugs and defoliators

CONCLUSION

Upland rice is attacked by a wider array of insect pests than is lowland rice mainly beca use of the addition of soil pests Generally population levels of lowland rice pests are lower in upland than lowland rice leading many people to conclude that insect pests are not important on upland rice

36 IRPS No 123 January 1987

REFERENCES CITED

1Abraham C C and K S R Mony 1978 Occurrence ofLeptocorisa acuta Fabr (Coreidae Hemiptera) as a pest of nutmeg trees J Bombay Nat ttistSoc 74553

2 Agyen-Sampong M 1975 HlierogrlphusdiaganensisKrauss (Orthop-tera Acrididae) a new pest of rice in Northern Ghana Ghana J Agric Sci 8249-253

3 Agyen-Samnpong M 1976 Varietal resistance and other inle-ratcd pest control components as factors of rice varietal improvement in West Africa Prcsented at the Second varietal improvement serinar 14-18 Sep 1976 Bouak Ivory Coast West Africa Rice Development Association Monrovia Liberia 18 p

4 Agyen-Sampong M 1977 Outbreak of Ihccrorrdnisorae in rice seedlings near Rokupr Sierra Leone lit Rice Res Newsl 2(6)16-17

5 Agyen-Sampong M 1980 Dirty panicles and rice yield reduction caused by hugs IntRice Res Newsl 5(l)11-12

6 Agyen-Sampong M 1982 The major pest problems of irrigated upland and mangrove swanip rice ecosystems ir the humid tropical Guinea savanna and Salel climate zones Pages 318-328 in Pro-ceedings training course cn concepts techniques and application of irtegrated pest management in rice inWest Africa 10-20 Jan 1982 Fendall Liberia West Africa Rice l)evelopioent Association Monrovia Liberia 505 p

7 Akibo-Betts V I and S A Raymundo 1978 Aphids as rice pests in Sierra Leone lit Rice Res Newsl 3(6)15-16

8 Akinsola F A 1975 Present status of different rice stemborers in parts of Nigeria iicL Entomo Newsl 328

9 Akinsola FA 1979 Ihe biology and ecology of rice stemborers in Nigeria Ph 1) thesis University ol lbadan Nigeria 217 p

10 Akinsola E A 1979 Guidelines of integrated management of insect pests of upland rice in West Africa Presented at the Seminar on integrated management of rice diseases and insect pests at Bobo-Dioulasso Upper Volta 17-22 Sep 1979 West Africa Rice Develop-ment Association Monrovia Liberia

II Akinsola IA 1984 Insect pests of upland rice in Africa Pages 301-305 in An overview of upland rice research Proceedings of the 1982 lIouak Iwsrv Coast upland rice workshop International Rice Research Institute 13O Box 933 Manila Philippines

12 Alam M S V TJohn and Kaung Zan 1985 Insect pests and diseases of rice in Africa Pages 67-82 in Rice improvement in Eastern Central and Southern Africa proceedings of the international rice workshop at Iusaka Zambia 9-19 April 1984

13 Alticri M A A van Schoonhovcn and J D Doll 1977 The ecological role of weeds in insect pest management systems a review illustiated with bean (Phaseohsvulgaris L)cropping systems PANS (Pest Artie News Summ) 23195-205

14 Anmante F 1967 The leaf-cutting ant Atia capiguaraa pest of pastures [in Portuguese] Biol6gico (Sao Paulo) 33113-120 (Rev Appl Entomol A56207)

15Andrade A C and D Puzzi 1951 Preliminary results of experi-merits on the control of Sculptcoris iastantus on sugarcane [in Portuguese] Bioulgico (Sao Paulo) 1744-49 (Rev Appl Entomol A41421422)

16 Appert J 1970 Maliarplia separaelha (rice stem borer) new observations of entomological problems of rice in Madagascar [in French] Agron Trop (Paris) 25329-334

17 Ariadeau Mand Z Harahap 1986 Relevant upland rice breeding objectives Pages 189-197 in Progress in upland rice research Proceedings of the 1985 Jakarta conference International Rice Research Institute 1O Box 933 Manila Philippines

18Asar S and Kisimoto 1975 Seasonal occurrence of the smaller brown planthopper in upland rice [in Japanese] Proc Kanto-Tosan Plant Prot Soc 2228

19 Bakker W 1974 Characterization and ecological aspects of rice yellow mottle virus in Kenya Meded Landbouwhogesh Wage-ningen 152 p

20 Balut F F 1970 White grubs attacking rice Oryza ativa L [in Portuguese] Biol6gico (So Paulo) 36321-322

21 Banerjee S N and L M Pramanik 1967 The lepidopterous stalk borers of rice and their life cycles in the tropics pages 103-124 in The major insect pests ofthe rice plant Proceedings ofa symposium at the International Rice Research Institute September 1964 The Johns Hopkins Press Baltimore Maryland USA

22 Barrion A T and J A Litsinger 1980 Ants a natural enemy of cliaiarac in dryland rice IntRice Res Newsl

5(4)22-23 23 Barrion A T and J A litsinger 1981 The spider fauna of

Philippine rice agroecosystem 1Dryland Philipp Entomnol 5139-166

24 Barrion A T and 1 A Iitsinger 1984 The spider fauna of Philippine rice agroccosystcms II Welland Philipp Eiitomol 611-37

25 Barrion A T1 and J A Litsinger 1984 lihanus (Diptera Tabanidae) eggs an alternative host of rice stei borer (SI3) egg parasite Telenoonis digns (Iliymenoptcra Scelionidae) lit Rice Res Newvsl 9(6)19

26 Barrion A T and J A Litsinger 1985 Identilication of rice leaffolders by wing markings Int Rice Res Newsl 10( 1)24

27 Barrion A T and J A Litsinger 1985 Chctiosspp (Coleoptera Carabidae) a leaffolder ILF)predator lit Rice Res Newsl 10(1)21

28 Bertels A 1970 Insect pests of rice and theircontrol [in Portuguese] Pelotas Brazil Instituto de Expcrimentacao Agropccuararias do Sul Circular 43 24 p

29 Bonzi S M 1982 Ctilo (flisilineus J de Joannis (Lepidoptera Pyralidae) stem borer of irrigated cereals in Upper Volta LioFrench] Agron Trop Riu Rizic Cult Vivrieres Trop 37207-209

30 Bourke T V T L Fenner J N LStihick G Baker EHassan D F OSullivan and C S li 1977 Insect pest survey for the year ending 30th June 1969 Papta New Guinea )epartment of Agrishyculture Stock and Fisheries Port Moresby 57 p

31 Bowden 1 1973 Migration of pests in the tropics Meded Rijksfac Landbouwwet Gent 38785-796

32 Breniere J 1973 The main rice parasites in West Africa and their control Pages 27-42 in Proceedings seminar on plant protection for the rice crop West Africa Rice Development Association Monrovia Liberia

33 Breniere J 1976 The principal insect pests of rice in West Africa and their control West Africa Rice Development Association Monrovia Liberia 52 p

34 Brown ES 1962 The African arinyworn Spoiolptera eXI)Ina (Walker) (Lepidoptera Noctuidae) a review of the literature Cornshymonwealth Institute of Entomology London UK57 p

35 Brown F B 1968 Upland rice in Latin America FAO-International Rice Commission 12th Session of the Working Party on Rice Production and Protection 2-14 Sept Peradeniya Sri lanka Food and Agriculture Organization Rome Italy 6 p

36 Buyckx F J 1 1962 Disease and insect pests of crops inCongo Rwanda and Burundi [iil French] Publications dc LInstilut National pour lestude Agrononiquedu Congo M Weissenbruck S A Brussels Belgium 708 p

37 Cariiio F 0 1981 Role of natural enentics inpopulation suppression and pest management of green rice lealhoppers Ph D thesis University ofthe Philippines at Los Bafitos PhilippinesLaguna 136 -

38 Carpente A J 1983 The development of rice cultivation and extension of vegetable productin in Zanzibar Final Report Vol II1 October 1983 Food and Agricultme Organization Ronte Italy

39 Carroll C P and S1 Risch 1983 Tropical annual cropping systems ant ecology Environ Manage 751-57

4(0 Cates R G 198 1 Host plant predictability and the feeding patterns of monophagous oligophagous and polyphagous insect herbivores Oecologia (Berl) 48319-326

IRPS No 123 January 1987 37

41 Cendcafia S M and F B Caora 1967 Insect pests of rice in the 62 Ene 1 C 1981 I)iiorldhol)ierusihilis Slater (licteroplera Philippines Pages 591-616 in The najor insect pests ofthe rice plant lygacidue Illissinac) a new rice pest front Nigeria lit Rice Res Proceedings of a symposium at the International Rice Research Newsl N3)15 Institute September 1964 The Johns Hopkins Press Baltimore 63 Fugenio T S 1965 rhe effect oi e-vCgetation on rice production Maryland USA 729 p and the possibility of overcoming it by leaf pruning MS thesis

42 Centro Ihternacional de Agricklittira Tropical 1984 Upland rice in University ol the Philippines at Los Baijos ILaguna Ihilippins 69 p Iatin Anerica Pages 93-119 hi An os ervies of uplaind rice research 04 Farrow R A 1975 The Alrican nmigratory locust ii is main Proceedings of the 1982 Boinakamp Ivory Coast upland rice workshop outbreak area of the middle Niger qlinlitatise studies of solitary International Rice Research Institute P0 Blox 933 Manila populations in relation to environmental factors Locusla No II Philippines 198 p

43 Chandra G 1979 Taxonomy ad hn riononlics ofthe imect parasites of 65 Fennal R G 1943 Food crop investigation Windward and rice Icalloppers and planthoppers in the Pihilippines and their Ieeward Islands First report Report onl work dunring the periodiiiportairce iii natural biol)gical cointrol Philipp Fntoltol 4119-139 April 1942-April 1943 Imperial College of Tropical Agriculture

44 Chang T T 1976 The originii colut ion enlIivation dissemination Trinidad 16 p (Rev of Appl Fintornol A3222-25) and diersificition of Asian and African rices -uphytica 25425-44 1 66 Ferreira F 1983 Integrated pest ritaageiret of upland rice [ill

45 Chatierjee P B 1979 Rice leaffolder attacks in India lit Rice Res Portuguese English summary] Pages 323-341 in Upland riceculttre Newsl 4(3)21 factors affecting production E Ferreira T Yamada and F-

46 Cheng Cbien-pan 19(4 Upland rice in Taiwan International Rice Malmolta eds Institutoda Potassae Foslato Institnto Internacional Cnmmission Wiorking Party oit Rice Production ard Protection I()th da Potassa Piracicaha Sao Paulo lraid Meeting 3-10 March 1964 Manila Philippines Food and Agricul- 67 Ferreira E arid F dl Silva Martins 1979 Occurrence of ture Organiaion Rome Italy 5 p Sogatohs orvziola (Muir) inupland rice in Goias Ilrail lit Rice

47 Chcrrett 1 M and 1) 1Peregrine 1976 A review of the status of Rcs Ncwssl 4(4)18 leaf-cutting ants and their control Ann Appl Biol 84124-128 68 Ferreira E and 1F da Silva Martiris 1984 Insect pestof rice ill

48 Chipeta F M 1974 Pests of rice in Malawi Pages 90-95 in Brazil and their control Jill Portugucsej FMIlRAtPA-CNPAF Proceedings Sith eastern Africa cereals research conference 10-15 Goiania Brmiil 67 p March 1974 DRB Mialda ed Ministry of Agriculture lilongwe 69 Ferreira FI F ca Silha Miartins S S Nero and 1 It P de Malawi Carvallio 1982 M tltiplc regression analysis todeterimine the effect of

49 Chu Yau-i 1979 The d atiaging period of the rice seedling fly insect populatiois ott upland rice yield [inIPortuguese Pesqui (Atlterigotiavxigu Stcin) (NItiscide Diptera) oil cori) at Fast Java Agropecu Bras 17671-675 Indonesia Plant Prot Bill 21403413 70 Ferreira F 1F da Silva Martins and F 1PZimmermann 1979

50 Conklin H C 1957 IHaninoo agriculture a report on aln integral Rice varietal resistance to tle lesser corn stalk borer [in Port uguese] systent of shi fti rig cult ivationi in the Pltilippincs Food and Agriculture Pesqui Agropccu Biras 14317-321 Organization Rome Italy 209 p 71 Fleming W F 1968 Biological control of lie hapanese beetle U S

51 Cramer H II 1967 Plant proection and world crop production Dep Agric RLs Sers [ech Bull 1383 78 p Pilaniensclhutz-Nachr 201-524 72 Food and Agriculture Organiation of the United Nations 1982

52 l)ani R C and Majunidar 1978 Prelininary observations on tie Country report surmimary Pages 27-37 in Report on the first incidence of root aphids on differeit rice cultnrcs Sci Cult 4488-89 FAOIUN)IPhail and regional training coiuirse in iniproved ctltural

53 Davidson R I 1979 lffects of root feeding on foliage yield Pages practices for upland rice 5-23 Oct 1981 Northern Region Agrculturd117-120 in T K Ci osb and R B Pottinger eds Proceedings of the Developririt Center Chiang Mai Food and Agricuilture Organiiashy2nd Australian conlerence on grassland invertebrate ecology Govt tion Rome Italy 37 n Printer Wellington New Zealand 73 Franco do Ariaral S 195 1 Residual action of insecticides in turfy

54 l)ean G 1W 1976 Rice insect pests inlaos Int Rice Res Newsl soil--the results of a field experiment [in Portuguese] Biol6gico (Sao 1(2)15 Paulo) 17209-214 (Rev Appl Entomol A41423)

55 I)ean G J W 1978 Insect pests of rice in Laos PANS 24(3)280-289 74 Freeman ) 1970 Report onithe Iban london School of -coroiics 56 )ingle H 1984 Behavior genes and lifc histories conplex adapta- Monographs on Social Anthropology No 41 Athlane Press

tions to uncertain environments Pages 169-194 in A new ecology London UK 317 p novel approaches to interactive systems P W Price C N Slobod- 75 Fuchs T W1 A Harding and 1W Smithlr 1979 Induction and chikoff and W S Gaud cdsl ohn Wiley New York USA termination oldiapause in the sugarcane borer Ann Flitonlol Soc

57 Dobelmann-JP and M Falais 1968 Upland rice cultivation ill tie Am 72271-274 North-western part of Madagascar in 1966-1967 [in French English 76 (hbriel B1) 1975 A review tf the major insect pests of some tpland summary] Trop Agron 2313-21 crops in tlc Philippines Pages 86-111 in Ad Htoc panel of experts on

58 Doniiciano N L 1979 Chemical control of tie lesser corn stalk pest disease arnd weed problems in some rainfed crops Bangkok (lolsonno~paloIgnowslhts) (Zeller 1848) (I-epid(ptera Ihycitid ae) Thailand Food aid Agriculture Organization Roire Italy

on rice in Ponta Gressa Parara 1977-1978 [in Portuguese] Pages 77 Garg D K and N K Shal 1983 White grub outbreak on rainfed 291-303 in Anais 1 Reuniao de tecnicos cmiirizicultura do estado de dryland rice in Uttar Pradeslh Int Rice Res Ncwsl 81(4)16-17 Sao Paulo-Campinas Cati Brazil 78 Garg I) K and IP Tandon 1982 Major insect pests of rice on hilly

59 Dunsmore 1 R 1970 Investigations on tlie varieties pests and tracts of Uttar Pradesh India lilt Rice Res Newsl 7(11)11-12diseases of upland rice (hill padi) in Sarawak Malaysia Int Rice 79 Garrity 1) P 1983 Upland rice research an international bibliograpy Comm Newsl 1929-35 1975-1982 Iternational Rice Research hnstitnte PO Box 933

60 Elias 1R1965 The new decline of rice seedlings is not caused by Manila Philippines 121 p parasites [in Portuguese] Biol6gico (Sao Paulo) 313-6 80 Ghauri M S K 1982 A new subspecies of Iinorh)rone bull

61 Empresa Brasiliera de Pesquisa Agropecuaria 1984 Upland rice itt cornlus Slater (Hemiptera Iygaeidae) front Papua New Guinea on Brazil Pages 121-134 it Av overview of upland rice research rice and carpet grass Ihll lEntoniol Res 72133-137 Proceedings ofthe 1982 Blouak Ivory Coast Upland rice workshop 81 Gonzales F 0 PArregoces R I ihernandef and OT Parada International Rice Research Institute PO Box 933 Manila 1983 Insect and imite pests of rice arid their control in Latin America Philippines [in Spanish] la Federacion Nacional de Arrocerts (Fedearroz)

Bogota Colombia 60 n

38 IRPS No 123 January 1987

82 Grist D H and R J A W Lever 1969 Pests of rice Longmans London UK520 p

83 Hale P R 1979 Two subterranean pests of upland rice in Papua New Guirnca IntRice Res Newsl 4(2)16-17

84 Hale P R and J T Hale 1975 Insect pests of rice in the New Guinea Islands Rice Entomol Newsl 34

85 Harris W V 1969 Termites as pests of crops and trees Common-wealth Institute of Entomology London UK 41 p

86 Hattori 1 1968 A method to distinguish between grass leaf roller (Ctaplha rocis medinalisGuenee) and the rice leaf roller (Susurtia exigua Butler) [in Japanese] Plant Prot (Japan) 23167-170

87 Henry I E and 1A Onsager 1984 Experimental control of the mormon cricket Iahrosshnph by NJsena locustae(Aficrospora Microsporidae) a protozoan parasite of grasshoppers (Ort Acri-didae) Entomophaga 27197-201

88 Hidaka 1 P Vungsilabulr and SKadkao 1974 Studies on ecology and control of the rice gall midge in Thailand Tech Bull 6 TropicalAgricultural Research Center Tokyo Japan 113 p

89 Hincklcy A 1) 1963 Ecology and control of rice planthoppers in Fiji Bull Entomrol Res 53467 481

90 Hirao J 1982 Ecology and chtInical control of the rice leaffolder Jpn Pestic Inf 4114-17

91 Ho DT 1984 Sten borer incifience in rice ecosystems in KenyaEast Africa Pages 307-320 in A overview of upland rice research Proceedings of the 1982 Bouac Ivory Coast upland rice workshop International Rice Research Institute PO Box 933 Manila Philippines

92 Ho )T J 1Njokah and J GKibuka 1983 Studies on rice sten borers in Kenya with emphasis on ifaliarpiaipiraellRag Insect Sci Appl 465-73

93 Ichihara I T Ito A NIatsuo and M Sawada 1971 Chemical control of the rice striped stem borer in upland rice grown in plastic mulch [iii Japanese] Proc Kanto-Tosan Plant Prot Soc 1882

94 IclhiharaIand A Matsuo 1969 Damage caused by rice striped stett borer to upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1686-87

95 Ichhara I atid A MIatsun11970 Conside ration oft lie extraordinary prevalence of rice striped stem borer in upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1778-79

96 Institut de Recherches Agrolnliqes Iropicale et des Cultures Vivrieres 1978 Insect resistance Pages 12-13 in Upland rice 1978 atnual report Bouak Ivory Coast 27 p

97 International Institute for Tropical Agriculture 1983 Aphids itt rice roots I ITA Res Briefs 4(1-3)4-5

98 International Institite fi Tropical Agriculture 1983 Hopperburn in rice caused by brown plathopper IITA Res Briefs 4(l-3)6-7

99 International Rice Research Institute 1975 Annual report for 1974 PO Box 933 Manila Philippines p 216

100 International Rice Restarch Institute 1976 Annual repiort for 1975 PO Box 933 Manila Philippines p 228

101 International Rice Research Institute 1979 Cropping systems program Component technology development and evaluation Page 404 i Annual report for 1978 11O Box 933 Manila Philippines

102 International Rice Research Institute Department of Entomology 1981) Insecticid evaluation 1979 Tables 73 74 PO Box 933Manila Philipes

103 International Rice Research Institute Department of Entonmology 1981 Insecticide evalomtion 1980 Table 81 POBox 933 Manila Philippines

104 International Rice Research Institute 1981 Report of a meeting of Asian rice-based cropping systems entomologists Eleventh cropping systems working group meeting 18-22 May 1981 PO Box 933 Manila Philippines p 132 133 144

105 International Rice Research Institute 1984 Cropping systems program Pages 368-373 in Annual report for 1983 PO Box 933 Manila Philippines

106 International Rice Research Institutc 1984 Upland rice in Asia Pages 45-68 in An overview of upland rice research Proceedings of the 1982 Bouakamp Ivory Coast upland rice workiiop loternational Rice Research Institute 11O Box 933 Manila Philippines

107 Irshad M 1977 Grasshoppers in rice in Pakistan lit Rice Res Newsl 2(3)8-9

108 Ito R and TOno 1964 Upland lice its culture and varieties in Japan lIternational Rice Cormmission loth Meeting of the Working Party on Rice Production and Protection 3-10 March 1964 Manila Philippines Food and Agriculture Organization Rome Italy 12 p

109 Jeffrey M I and V A Dyck 1983 Effect of moonlight on lightshytrap catches of brown planthopper lit Rice Res Newsl 8(6) 13

110 Jennings P R and T Alicia Pineda 1983 Management ofthe rice plnthoppers Sogatodsorivi(coIl in Latin America Presented at the meeting on stable plant resistance FAO and lIndocsian I)inectorate of Food Crops Protection Bali lndonesia 29 Novcniber-2 l)ccemshyber 9 p

IllJimenez Gomez J A 1979 Rice insect pests i lla ios ()riCntales lil Spanish] Pages 99-103 in Rice ciltivation Instituto (olotibiatto Agropecuario Compndio No 29

112 Jongeleen F JJ Soesanto and A Mockidjo 1979 Studies oti pathogens of two species of Scarabacidac froinIndonesia Z Angew Enlomnol 87412-417

113 Jouvenaz 1) PC S lofgren and W A Baiks 1981 Biological control of imported fire antsa review of cirrent knowledge Bull Entomol Soc Am 27203-208

114 Kainoh Y K Shiniiu S MarUani Y Taiaki 1980 lost finding behavior of tie rice bu lplotori+JchinvittjI)allas (liciniptera Corcidac) with special reference to diet pattern ol aggregation and feeding on the rice plant Appl Entomol Zool 15225-233

115 Kakkar K 0 P Sharma and P I Sharma 1977 Note on Ihteronyrchus liodresRedt (Coleopicra I)ynastidae) as asporadic pest of paddy and maie inHimachal Pradesh Corr Agric 156-57

116 Kalode M131 79 Major insect pest problems and their control in upland rice (Unpublishcd mineogriph CWI oJNaaiat lni Chandaratat (198 1)original not seen)

117 Kalshoven L i 98 I The pests of crops in I ldoniesia (translated by P A vatn der Laan) 1 T Ichtiar B3aru-vati Iloeve lakarta Indonesia 701 p

118 Kamal N Q 1981 Suppression of whitebacked planto ner Sogatella fircaifra (Horvath) aLId rice lealfolder (nhal -cis licdb lis (Guenee) populations by natural cnclmie hlI) tiesis Grcgorio Araneta University Foundation Victoncta Park Metto Manila Philippines 179 Ip

119 Karim A N M Reaul 1975 A species ofliealy bug Trioni i70s ceres new oii rice in the Philippines Rice Entoniol Newsl 247

120 Katanyukul Wand C Chandaratat 1981 Insect pestsof uplaid rice and their control Report firstFAOUNlFi Tliailand training course on improved cultural practices for tplad rice Chiang Mai Thailand 5-23 Oct 1981 Food and Agriculture Organiation Rome Italy 22 p

121 Kern F 1955 lhletwioa(I)uninemts) bhhit Biturm(Coleoptera) a new pest of sugarcane inVenezuela [inSpanish with Etnglish sunmary] Agron Trop (Maracay) 527-29 (Rev Appl Eitoniol A4572)

122 Khan K M and S Ghai 1974 White grubs ald their control in India Pesticides (1Bombay) 819-25

123 Kiritani K 1975 Integrated pest management of rice pests Pages 66-70 itt Proceedings platning workshop ott cooperative hIeld research in pest managementP SMotooka ed East-West Center Honololu Hawaii USA

124 Kisimoto R 1967 Ecology of insect vectors forecasting and chemical control Pages 243-255 in The virus diseases of tle rice plant Proceedings of a symposium atthe International Rice Research Institute April 1967 The Johns Hopkins Press altimore Maryshyland USA

125 Kisimoto R and V A Dyck 1976 Climate and rice insects Pages 367-391 in Climate and rice International Rice Research Institute PO Box 933 Manila Philippines

126 Kureha S S Kobayashi SAsari and T Ishihara 1977 Life history and control of the mole cricket Grrllotalpa africana Palisot de Beauvois [in Japanese English translation 7520 1RRI library] Proc Kanto-Tosan Plant Prot Soc 21192-197

127 Kushwaha K S 1981 The black beetle as a pest of rice in Haryana India Int Rice Res Newsl 6(4)14

128 Lawani S M 1982 A review of tile effects of various agronomic practices on cereal stem borers Trop Pest Manage 28266-276

129 Leuck D B 1966 Biology of the lesser cornstalk borer in South Georgia J Econ Entomol 59 797-801

130 lever R J A W 1955 Insect pests of the rice plant in Malaya Int Rice Comm Newsl 1612-21

131 Li Ching-Sing and Shin-Foon Chiu 1951 The study ofthe rice gall midge Iachydiplosis orvza Wood Mason [in Chinese with English summary] 1Taiwan Agric Res 21-13

132 Iitsinger J A 1984 Assesament of the need-based application for rice Proceedings TechnologyTransfer Workshop 15-16 March 1984 International Rice Research Institute 1PO Box 933 Manila Philippines 6 p

133 Litsinger J A 1984 Environmental diversity and insect pest abundance with reference to the Pacific Pages 285-307 in Proceedings sub-regional training course on methods of controlling diseases insects and other pests of plants in the South Pacific 4-20 Oct 1982 Vaini Kingdom of Tonga Nukualofa

134 Litsinger J A RF Apostol and M BObusan 1983 White grub Leucopholis irrorata (Coleoptera Scarabaeidae) pest status popula-tion dynamics and chemical conllol in arice-maize cropping pattern in the Philippines J Econ Entoinol 761133-1138

135 Litsinger J A C de la Cruz F Raymundo A T Barrion M D Lumaban J P Bandong M S Venugopal F Paragna and E Balete 1982 Insect pests and insecticide reponse of dry-seeded bunded rice Pages 179-193 in Report of workshop on cropping systems research in Asia International Rice Research Institute Los Bafos Philippine

136 Litsinger J A M 1) Luinaban J P Bandong A T Barrion W Katanyukul and N Panda 1978 Insect pest management in rainfcd lowland rice agroecosystms of tropical Asia International Rice Research Conference 17-21 April 1978 International Rice Research Institute Los Bafios Philippines

137 Litsinger J A and K Moody 1976 Integrated pest management in multiple cropping systems Pages 293-317 in R I Papendick P A Sanchez and G B Triplett eds Multiple cropping American Society of Agronomy Madison Wisconsin USA

138 Loevinsohn M E 1984 The ecology and control of rice pests in relation to the intensity and synchrony of cultivation Ph D thesis University of London 36) p

139 Loevinsohn M E J A Litsinger N Panda M A Austria R F Apostol C Andrion and J Bandong 1982 A basis for predicting rice insect pest abundance in cropping systems Paper presented at the 1982 International rice research conference on rice and cropping systems 19-23 April 1982 International Rice Research Institute Los Bafios Iaguna Philippines 25 p

140 Lopez A W 1930 Repot of the Entomology Department Pages 145-172 in Annual report of the Philippine Sugar Association 1929-30 Manila

141 Lopez AW 1931 Entomology Department Pages 227-266 in Annual report of the director or research 1930-31 Philippine Sugar Association Manila

142 Lopez A W 1932 Entomology Deparment Pages 252-279 in Annual report of the Philippine Sugar Association 1931-32 Manila

143 Lungibill P and G G Ainslee 1917 The lessercornstalk borer US Dep Agric Agric Res Serv Tech Bull 539 27 p

IRPS No 123 January 1987 39

144 McGuire J U Jr and BS Crandall 1967 Survey of insect pests and plant diseases ofselected food crops of Mexico Central America and Panama U SDepartment of Agriculture International Agriculshytural Development Service Agricultuial Researrh Service October 1967 11 p

145 McNeill Sand T R ESouthwood 1978 The role of nitrogen in the development of insectplant relations Pages 77-9 in lliochemical aspects of plant and animal coevolution J B Ilarbornu ed Academic Press London UK

146 Maebara H 1955 Spatial distribution of lastinstiarlanaeof purplish stem borer (Sesamia iqferens Walker) hibernating in an upland rice field Bull Kagoshima Univ Fac Agric 438-41

147 Mallick E H and N G Hajra 1977 Response of rice plant to defoliation treatments at different developmental stages Indian Agric 2151-56

148 Mansingh A 1971 Physiological classification of dormancies in insects Canl Entomol 103983-10)9

149 Martell C G 1974 Catalogue of plant feeding insects of Mexico [in Spanish] Fitofilo 27(69)1-176

150 Martinez 0 J 1982 Principal rice pests in Ilanos orientalis [in Spanish] P~ages 80-94 in Curso deact nalization de conocimientos en arroz Villaniencis Instituto (olonmbiano Agropecuario

151 Martins J F da Silva F Ferrcira A S Prabhu F 1 P Zimmermann 1980 Use of pesticides for the control of the main subterranean upland rice pests esqui Agropecu Bras 1553-u

152 Martins I F da Silva F Fe eira rod BdamSilveira Pinheiro I Q2 Damage simulation caused by defoliators of upland rice [in Portuguese with English summary] Pesqui Agropecu Bras 171113-1119

153 Martins 1F da Silva N(iyuen van Tan and 1Ida Silveira Pinheiro 1981 Stem borer resistance in upland rice associated with plant morphological characters [in Portuuce Iesqui Agropecu Bras 16187-192

154 Masaki S 1980 Sumnmer diapause Ann Rev Entomiol 251-25 155 Masuzawa T 11 Suwa and F Nakasoji 1983 l)ifkleences of

oviposition preferencc and survival rate of two skipper butterflies Parnara guttata and PIchlpidai iathiais (Iepidoptera HIesperiidac) on rice plant and cogon grass New Entomol 3247-56

156 Maurya D M and C P Vaish 1984 Upland rice in India Pages 69-92 in An overview oftpland rice research Proceedings ofthe 1982 Bouakamp Ivory Coast upland rice workshop International Rice Research Institute IO Box 933 Mnila Philippines

157 Menon M G R R N Katiyai S Kuar and S S Misra 1970 Termites attacking the paddy crop Sci Cult 36294-295

158 Miyahara Y and 1 Nishikubo 1967 Behavior of young nymphs of the red rice root aphid Iholahsipiun ri liabhdontinalisSasaki in relation to movement from the leaf to the roots of upland rice [in Japanese] Pioc Assoc Plant Prot Kyushu 1363-66

159 Mo ljoa Tjien 1952 White grub of upland rice (Holotrichiahelleri Brsk) [in Indonesian with English summary] Contrib Agric Res Stn Bogor No 13134 p

160 Mochida 0 and T Suryana 1979 The occurrence of the brown planthopper and ragged stunt on lowland and upland rice ill Central Java and logyakarta in tile late of January 1979 Entomnolgy Survey Report No 15 Central Research Institute of Agriculture (CRIA) Sukamandi Indonesia I February 1979 9 p

161Monet C 1979 Varietal resistance to upland rice pests [inI French] Pages 35-39 in Annual report 1977 Institut des Savanes Department des Cultures Vivrieres Bouak6 Ivory Coast

162 Monte 0 1942 A caterpillar ofrice-fields[in Portuguese] Biol6gico (Sao Paulo) 8161-163 (Rev Appl Entomol A31414)

163 Munroe I) D 1979 Pest management strategy for hill rice in four resettlement areas in Sarawak Malaysia Pages 89-102 i1 Guidelines for integrated control of rice insect pests Food and Agriculture Organization Plant Production and Protection Paper 14 Rome Italy 115 p

40 IRPS No 123 January 1987

164 Nakasuji F M Ishii and Jen-Zon Ito 1984 Rice skippers in Taiwan and their life histories ZZrC (Taipei) Tech Bull 82 10 p

165 Navas I) 1976 Fall armyworum in rice Pages 99-106 il Ecological animal control by habitat management Proc Tall Timbers Con-ference 28 Feb-I Mar 1974 Fall Timbers Res Stn Misc Publ 6 Tallahassee Horida USA

166 Nickel 1 1 1961 Biological control of rice stem borer A feasibility study Tcch Bull 2 International Rice Rcarch Institute PO Box 933 Manila Philippines IIl p

167 Nigan P M and R A Vernia 1985 Insect pests of uplaild rice in Uttar Pradesh Int Rice Res Newsl 10(6)22

168 Nilakhe S S 1985 -cological observations on spittle bugs with emphasis on their occirence in rice Pcsqtui Agropecu Bras 20407-414

169 Nixon G EJ 195 1 lie association of ats with aphids and coccids Commonwealth Iusti lcte of Entonology Iondon UK 36 p

170 Nurdin F an Ishak Manti 1985 Study oit seedling fly of upland rice Interlational Upland Rice Moitoring Tour Report 26 lebruary-2 MarLI Sukaranti West Sumatra

171 Oka 1 N 1979 Ciltural conttol of the brown planthopper Pages 357-369 in Brown planthopper threat to rice production in Asia International Rice Research Institutc 1p0 Box 933 Manila Philippines

172 Ohni M and A Vilnaii 1975 Rice insect pests of Italy biology of Sipha grcria (Kaltenbach) and its control [in Italian] Riso (Milan) 2459-71

173 Otanes F Q atnd 1 I Sison 194 1 Pests of rice Philipp J Agric 12211-259

174 Painter R I 1951 Insect icsistance in crop plants University Press of Kansits Lawrence Kansas USA 520 p

175 Pantua P C and 1A litsinger 1981 Arthropod abundance in continuous vs biannual rice cropping systeis Int Rice Res Ncwsl 6(2)20-21

176 Pathak M ) and V A l)yck 1975 Sttudics ott insect pests of upland rice Pages 186-191 in Major research in upland rice International Rice Research Institute PO Box 933 Ilanila Philippines

177 Patil B R B M IHasabe and 1) S Ajri 198 I Chemical control of white gritb lIditriihinhorraia Fabr intfesting groundntut paddy atnd sorghum Indian 1Entomol 43232-236

178 Pawar A 1) 1974 (ollection and identification of insect pests of rice rlerminal repirt lInternational Rice Research Institute PO Box 933 Manila Philippines 72 p (tnimco)

179 IerrajU A aid A S Reedy 1965 Observations ott diapausc itt tie steti borer ohrice Rice News Teller (lai-Mar)9-13

180 Pont A C 1972 A review of the Oriental species of theri ona Rondani (I)iptera Muscidac) of econtomic importance Pages 27-102 in Control of sorglium shoot fly M G lomtani and W R Young eds Oxford atd IH1 Ptthl Co New I)clhi India 324 p

181 Prasad S C 1PChand aid M 1PSingh 1983 Incidence of rice gall midge in the upland plateau region of Hihar Oryza 20235-236

182 Rai P S 198 0 Nisaga simpex damage to rice in the hill tracts of South India Int Rice kles News[ 5(I)12

183 Rathorc Y S 1969 Gonocplhahli dcprssunt Fabr as itpredator of (tilo zonwIlhis (Swinhoc) and other moths in field cage at Patnagar Uttar Pradesh Indian 1Entotnol 3127

184 Reed W 1983 Crop losses causcd by insect pists in the developing world Proceedings 10th Internatial Congress of Plant Protection lrighton UK 174-80

185 Reedy M V 1981 Ateactogasterinhens(Walker)a new root-grub pest of tipland paddy and inaize Curr Sci 501076

186 Reveno M A 1968 Catalogue of the insect pests of economic crops in Colombia [in Spaiish] Asociacion Latino Americana de Entomo-logia (AIAE) 129-33

187 Reyes G M 1957 Rice dwarf disease in the Philippines Food and Agriculture Organiiation Plant Prot Bull 617-19

188 Rice S E A A Grigarick and M 0 Way 1982 Effect of leaf and panicle feeding byarmyworm (Lepidoptera Noctuidae) larvae on rice grain yield Environ Entomol 7593-595

189 Risch S J 1) Andow and M A Altieri 1983 Agioccosystem diversity arnd pest control data tentative conclusions and nizw research dirctions Environ Entomol 12625-629

190 Rivera C T 1956 Three organic insecticides in thecontrol of insects affecting upland rice Philipp Agric 39465-472

191 Roffrey 1 1972 Locusta outbreaks in the Philippines Acrida 1177-188

192 Rojas R 1) 1941 The effect of pruning at varying ages upon the yield of Ramairio Phipp Agric 29851-860

193 Roonwal M i 1979 Ternite life and termite control in tropical South Asia Scientilic Pnbl odhpr India 139 p

194 Rosetto C1 F da Silva Martins N C Schmidt and L EAzzini 1978 Damage caused by pasturespittle bugs 4Dvoi)lavopictaandDJ schach) on rice lBragantia Nota 6 (Matcli)25-27

195 Rosztto C1- S S Neto I- Amantc I) Iink 1Garcia-Vieira D M de SoLza N V Bansa t and A M de Olivcira 1971 Rice pests of Brazil [in Portuguese] Pages 149-238 inrFA()-International Rice Commission 2nd meeting of the Rice Committee for the Americas 6-11 l)ecember 1971 Pelotas Rio Grande do Sul Brazil

196 Rothschild G H L 1969 Obseratiois on the armyworm Spodoptera matritia acronrctoiiles Gn (Lep Noctuidae) Bull Entomol Res 59143-160

197 Rothschild G 1- L 1974 Parasites of the armworm Spolopiera mauritiaacroncitoihes in Malaysia Fntornopliaga 19293-299

198 Sachan i N S K Gangwar and 1N Katujar 1980 Field crickets damaging cages in North Eastern IHill Region Indian J Entomol 42526-527

199 Sands 1) P A 1977 The biology and ecology of Lepocorisa (Hemiptera Alydidac) in Papua New Guinea Res Bull 18 Departshyment of Primary Industry Port Moresby Papua New Guinea 104 p

200 Sarawak 1968 titomology division Pages 97-106 in Report of the research branch of the Department of Altriculture 1966 Kuching (Rev AIppl Eintomol A56603)

201 Satpathy JM M S Das and K Naik 1977 Effect of multiple and mixed cropping on the incidence of scine important pestsI Entomol Res 178-85

202 Sauer IHF G 1939 Notes Ott Elasmpnlpualtsligniosellths Zeller (Lep Pyr) a serious pest ofcereals in Sao Paolo [in Portuguese] Arq Inst Biol 10199-206 (Rev Appl Entotnol A28378)

203 Satinders 1L A 1 SKing and C L Vargas S 1983 Crop pests of Central Anerica with t bibliography [in Spanish] Centro Agroshynotnico Tropical de Investigacion y Ensenanza Turrialba Costa Rica (CATIE) Tec Bol 9 90 p

204 Sauphanor B 1985 Some factors of upland rice tolerttce to stem borers in West Africa Inseu Sci Appl 6429434

205 Sen P atnd S Chakravorty 1976 Stentflics in Haritighata Nadia West Bengal India [it Rice Res Newsl 1(12) 15

206 Senapai B and 1M Satpathy 1983 The seedling fly Atherigona orrzae Malloch is a pest of rice in Orissa (India) J Entomol Res 777-79

207 Shhjahan M S Alatn M Ah nad and BN Islam 1983 A new scarabaeid beetle pest of dryland rice in Bangladesh Int Rice Res Ncwsl 8(I)16-17

208 Sharma G L S C Modgol and R C Gautam 1979 Effect of depth of submergence ott incidence of bacterial blight and white grub infestation in transplanted rice Int Rice Res Newsl 4(6)12-13

209 Sharma S 1985 Scarabaeid beetle outbreak itt upland rice Int Rice Res Newsl 1(1(l)24

210 Shiraki T 1917 The paddy borer Schoenobhits incertulas Walker [achaku Agric Exp Stn Formosa 256 p

211 Silva A de Brito and 11P Magalhaes 1981 Insect pests of rice [in Portuguese] Circular Tecnica No 22 EMBRAPA Centro de Pesquisa t ecuaria de Tropico Unido (CPAJU) Belem Brazil 14 p

212 Singh S A 1979 Emergence of rice leaf butterfly as a rice pest in Manipur India int Rice Res Newsl 4(4)16-17

213 SinghS A 1983 Changing trends of insect pestsof rice in Manipur Int Rice Res Newsl 8(f)12

214 Siwi S S and P H van Doesburg 1984 Lenovorisa Latreille in Indonesia (Heteroptera Coreidae Alydinae) Zool Meded (Leiden) 58117-129

215 Soejitno J 1977 Reaction of some upland rice varieties toseedling fly eltherikonaexigua Stein [in Indonesian] Siri Hana I Penyakit 630-35 Lembga I osat Fenclilian Pertanian Bogor Indonesia

216 Socnardi 1967 Insect pests of rice in Indonesia Pages675-683 in The major inse pests of thu rice plant Proceedings of a symposium at the Internatital Rice Rec Wi itt S pternber 1964 The Johns Hopkins Press Baltimore Maryland USA

217 Soto P E A T Petcz and I W Buddenhagen 1976 Survey of insect pests and diseases of rice in different ecological zones in Nigeria Rice Entomol Ncwsl 435-46

218 Soto 11E and Z Ziddiqi 1976 Screening for resistance to African rice insect pests Varietal improsetnent sCmi n r Bouak Ivory Coast 13-17 Sept 1976 West Africa Rice l)evelopment Association Monrovia Iiberia

219 Squire F A 1972 Entomological problems in Bolivia PANS (PestArtic News Sumr) 18249-268

220 Stapely 1 II 1978 The rice leaf roller in the Solomon Islands Int Rice Res Newsl 3(l)13

221 Suenaga H 1937 On aphids attacking the roots of rice Oyo-I)oubts Zasshi 9150-152 [in Japanese] (Rev Appl Entomol A25745)

222 Suenaga H 1952 On the seasonal abundance of three subterranean aphids on upland rice Bull Kyushu Agric Exp Sti 1249-262

223 Sutynso Yas and Dandi Soekaria 1964 Beberapa hana yang ttenperang tana man padi tanah kering [in Indonesian] Rapat kerja Padi Tanah Kering 6 sd 8 Juily 1964 Kaliurang 9 p

224 Tallamy I) W and R F Denno 1983 Alternative life history patterns in risky environments in example from lace bugs Pages 129-147 in Insect life history patterns R F l)cnio and If Dingle eds Spring-Verlag 225 p

225 Tamura 1 1939 Observations on some insect pests of field crops [in Japanese] 1 Plant Prot 26414-420 (Rev Appl EntomolA27-556)

226 Tamura and Suzuki 1964 Influences of the early-transplanting practice in rice culture upon development of the rice stem borer Chilo siql)ressalisWalker Bull Ilokuriku Natil Agric Exp Stn 761-94

227 Tan J P 1924 The rice root aphid Drowhi ihirsoia Philipp Agric 13277-288

228 Tanaka 1 1957 Taxonomy and distribution of some subterranean aphids injurious to upland rice in Japan with description of a new species [in Japanese English summary] Botyu-Kagaku 22168-176

229 Tanaka T 1961 The rice root aphids their ecology and control Spec [full College of Agriculture Utsunomiya University Japan F3 p

230 Tauber M IC A Tauber R Nechols and 1Obrycki 1983 Seasonal activity of parasitoids control by external internal and genetic factors Pages 87-108 itt I)iapause and life cycle strategies in insects V K Brown and I Ilodek eds )r W Junk The Hague Netherlands

231 Taylor W F 1972 Effects of artificial defoliation (simulating pest damage) on varieties of upland rice Exp Agric 879-83

232 feotia T P S and S Natud 1966 Biotoomics of the rice skipper Parnaraniathiis Fabricius (Lepidoptera Hesperiidac) Indian J Entomol 28181-186

233 Teran F 0 1971 Rice pests of Santa Cruz [in Spanish Ministerio de Agricultura Sita Cruz Bolivia Bull 45 8 p

234 Than Itun 1976 Population dynamics of the yellow rice borer Tryporza itcertulas (Walker) and its damage to the rice plant MS thesis University of the Philippines at Ios Bafios Laguna Philippines 92 p

235 Uichanco L B 1911 Pests Pages 55-74 it Rice Philipp Agric 11-200

IRPS No 123 January 1987 41

236 van der Laan P A 195 1 Life history of the rice seedling fly Contrib Gen Agric Res Stn Boor Indonesia 118 15 p

237 van der Laan 11A 1959 Correlation between rainfall in the dry season and the occurrence of the white rice borer (Scirpolhaga innotala WIk) in lava Entomol Fxp Appl 212-20

238 Van Driesche R G 1983 Meaning of percent parasitismin studies of insect parasitism Fnviron Entomol 121 I1- 122

239 Venugopal M S and A litsingcr 1984 Fffect ofcarbofuran on rice growth Prot Ecol 7313-317

240 Vietnam Directorate of Rural Affairs 1962 Annual work progress report on crop improvement program of rice sugarcane vegetable and field crops (lor the period from July 1961 to June 1962) (11) 299 p (Rev Appl Entomol A51526-527)

241 Wan M T K 1972 Observations on rice leaf- and planthoppers in Sarawak (Malaysian Borneo) Malay Agric 148308-335

242 West Africa Rice I)esclopnient Association 1978 Annual research report Vol 2 Statistics Monrovia Liberia

243 West Africa Rice Development Association 1979 Annual research report p47 Monrovia Liberia

244 Williams ) F and CS lofgren 1981 Fli I illy l-468 anew bait toxicant for control of the red imported lire at Fla Entomol 64472477

245 Williams I) 1 1970 The mealybugs OFlomoptera Coccoidea Iscudococcidac) of sugar-cane rice and sorghum Bull Entomol Res 0109-188

246 Williams D) 1 1982 Pulinaria icervi (Signoret) and its allies ott sugar cane and other grasses Btll Entoinol Res 72111-117

247 Wirjosuiardjo S 1976 Study ofchenmical cont rot of insect pests ott upland rice Imiu Pertanian (Jogjakarta) 1323-329

248 Wirjosuhardjo S and N Wirarta 1976 Chemical control of the white grub tIolorithiatlltriin upland rice in M uneng Irobolinggo Indonesia) [in Indonesian] Imut Pertanian (Jogjakarta) 1330-332

249 Yano K T Miyake and V F Eastop 1983 The biology and economic importance of rice aphids (Henuiptera Aphid idae) a review Bull Entomol Res 73539-566

250 Yelp D and P Rhodes 19(4 lhe success of drilled dryland rice iii the Fiji Islands International Rice Commission Working Party on Rice Production and Protection Manila Philippines 3-10 March1964 Food and Agriculture Organization Rome Italy 13 p

251 Young W R 1981 Fifty-five years of research on the sorghum shootfly Insect Sci Appi 23-9

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axial-flow pump

Page 21: UPLANDRICE. INSECTl--1i M T -E!R.EOLOU t RTANCE ,AND …delphacid.s3.amazonaws.com › 6958.pdf · The significance of abiotic factors in upland rice insect ecology will be apparent

__________

IRPS No 123 January 1987 21

Light trapped Insects Drytnd __ 2(notrap-nght) Suctioned Insects (nom )

50 -)o 9 jO40- SOW ffurciWO Light trp

D-Vc suction

20 to 4

20

Egg porasitizahion (M) 75 60

300 SOOampReOftwciffr

45 30shy

15

-insects (notrop-night) 4 Nileporvata Iuen5

Egg parastization()

Light trapped insects (no trop-nght) Suchoned insects (no m 2 )

6 8 Cytorh~nuls hvidipennii 0

4 05 2

50 0 Insects (noltrop-night)40-ui Ipanh pe g

Moohase 1 Seasonal abundance measured by light trap

Rainfall(etod) and D-Vac suction sampler of whitebacked

to ~ hopper Nilaparvata lugens and lea ffolder60)-t Cnaphalocrocis mnedinalis andplthpe g20 L I II I I I parasites and mtrid predator Ortorhjntus livi-Apr May Jun Jul Aug Sep Oct Nov Dec JaIn Feb Mar dipennis set against cropping pattern daily1 9 80o 19 8 1rainfall and moon phase Tanauan Batangas

1980-81

the D- Vac Cyrtorhinus build up on the crop coterminous are collected in light traps during full moon (109)with WBPH buildup Cyrtorhinus disperses during the Because upland rice shares most of the lowland rice pestsrainy season RLF not readily collected in light traps was and those pest species are generally less impotant to uplandprevalent only during the rice crop rice than lowland rice there is a general belief that insect

Nephotettix virescens was more abundant than Nnigro- pests are less important onpicLus in the light trap collections and on the crop GLH upland than lowland rice Arcview of limited data on upland rice shows an averag of population peaked at the reproductive stage as did zigzag 10-21 yield loss from insect pests on a par with that ofleafhopper (ZLH) Crtorhinus tracked the populations of lowland uee determined by the same methodology (132) planthoppers and leafhoppers Both GLH species were (Table 4) McGuire and Crandall (144) made a similarcollected year-round in the light trap indicating immigra- estimate for Central America Cramer (51) howevertion from nearby irrigated rice areas Possibly because of the estimated 35 yieldlow populations loss in South America based on ainsect number did not correlate with similar pest complex - stem borers and seed bugs shymoon phase although reports indicate that more hoppers attacking irrigated rice in North America This estimate did

22 IRPS No 123 January 1987

Light trapped insects D~~nd (no trap-night )

St plht Suctined insects (nom 2

)

D-Voc suction 1520

1o

0 Lih trap 5 ~105 o5

00 Eggparositizotion()5040 NephOtgtha virneens

30 20

I010 A-Insects (notrop -night)

4 Nephatelh nigropccent i

2 o I vV^J - -- ^ VA____ f _

Light trapped insects (no trap-night) Suctioned insects (no rnE

10 -1208 ReCdO doRsol

6 15

4 10 052 iwxALAiamp 0

Light trapped insects (notrap-night) 2Suc oned nsects (nom )

8 CyrtorhmnusI6o nn]10 i0075

4 05

2 0 0

025

2 Seasonal abundance measured by light trap Roinfall (mmd) Moon phase and D-Vac suction sampler of green leafhoppersNephotettix virescens and N ngropicrus and I zigzag leafhopper Recilia dorsalis their egg 60

parasites and micid predator Cyrtorhinus livi-dipennis set against cropping pattern dailyrainfall and moon phase Tanauan Batangas1980-81

40

5 Lf Apr May Jun

1980 Jul Aug Sp Oct Nov I

-aMar

1981

not account for soil pests and was not based on field data in the absence of pests Carbofuran granules are phytotonicAkinsola (I1) cited mainly data from irrigated rice areas in to rice (239) The prevailing hypothesis is that carbofuranAfrica but estimated yield losses of upland rice between 15 prevents soil bacteria from consuming fertilizer The phytoshyand 30 from insects However losses in Thailand over a tonic effect is through greater availability of fertilizer to the4-yr period ranged from I to 13 averaging 5(106) plant Denitrifying bacteria are known to be more active inYield loss data however should be interpreted with aerobic than anaerobic soils therefore it may not be merecaution (184) The Philippine data were all derived from coincidence that the highest recorded yield losses (22-69)trials on farmers fields under farmers normal agronomic occurred in those trials with 12-30 kg basal Nha On themanagement practices except for Bukidnon and Capiz other hand contro with carbofuran granules may not be aswhere researchers used fertilizer Batangas farmers are good as a seed treatment against seed pests and perhapsatypical because ofhigh levels of fertilizer used The trials in yields would have been higher if a systemic seed treatmentBrazil were carried out at an experiment station but insecticide had been used conditions were similar to those of local farmers The 1979 and 1980 trials in Batangas had two full-

Carbofuran granules were used in every trial as a broad- protection treatments one using carbofuran granules andspectrum insecticide applied basally with the seed An the other using bendiocarb WP as a seed treatment Plotsobjection to using insecticides to measure yield losses isthat protected with bendiocarb yielded higher than those proshyinsecticidescandirectlyorindirectlystimulateplantgrowth tected with carbofuran granules The other trials in the

Table 4 Yield losses to insect pests and yield responses to insecticide in upland rice in the Philippines Brazil and Upper Volta

Yield (tha) Yield loss (5) Location Year Cultivar Protected

with Protected

without Check Total Vegetative Reproductive Ripening

Basal N (kgha) Reference

carbofuran carbofuran Protected No with carbofuran

carbofuran

Philippines Pili Camarines Sur Tanauan Batangas

Pangantucan Bukidnon

Dumarao t-apiz

BrazilGoias Minas Ge~ais

Upper Volta6

1975 1974 1974 1975 1975 1976 1977 1978

1979

1980

1979 1980 1979

1977 1978 1979

Bursiging puti Kinanda Dagge Kinanda Dagge Dagge Dagge Dagge UPL Ri5 Dagge UPL Ri5 Dagge UPL RiS UPL Ri5 UPL Ri5 UPL RiS

IAC47 IAC47 IAC47

14 33 11 36 35 31 30 27 37 26 40 30 40 42 46 35

19 21 09

--

-34 24 24 27 --

29 43 31 42 -

47 25

---

10 28 05 27 27 30 26 29 33 28 33 29 43 23 36 11

15 15 06

29 15 55 25 23

3 13 -7 11

-7 18

3 -7 45 22 69

24 28 35

--

21 -11 -20

4 -

-3

23 7

-2 -23 56

-

-

-__0

6 --

-

-23 14

5 0 8 2

22

-

--

0

-

--18 11

3 -10

4 10

7

--7 -

14 -

-1 6

10 -5

8 9 7

-

--

0 0 0 0 0 0 0 0 0 0 0

30 30 30

12 12 12

99 99

100 100 101 101 102 102 102 102 103 103 104 104 104

66 66 66

Farako-Ba

Mean

1977 1978

IRAT10 IRATIO

38 66

--

40 43

-5 35

21

-

10

--

--

--

- 242 243

Z

tJ

-w

24 IRPS No 123 January 1987

Philippines which had two treatments offering protection In the 9 Philippine trials where yield losses were measuredwith and without carbofuran granules relied on further by growth stage 10 of the yield loss (21) occurred at thecontrol with foliar spray and not on seed dressings The vegetative stage and 517 occurred at each of the reproshycarbofuran-protected plots had an 18yield los compared ductive and ripening stages Ibis result is similar to thewith only 10 with foliar sprayS This yield difference is pattern of yield loss in lowland rice measured by successiveprobably due to differences in protection against sown-seed treatments where the crop is unprotected at each growthpests (foliar spray do not control sown-seed pests) rather stage but proticted during the other growth stagesthan phytotonic effects became basal fertilizer wa used in However the insect complexes r sponsible for vegetativeonly 2 of the 10 comparisons stage yield losses in upland rice are entirely diftfrent than in

The recorded yield losses were not related to yield lowland rice The key vegetative stage pests in lowland ricepotential (Fig 3) One might expect higheryielding crops to arc aquatic whereas those in upland rice are sown--seedhave higher percentage of yield losses but this ws not tile (sown in soil) root and seedling pests This dift rence maycase One reason is that insect damage exacerbates plant explain why agricul torists have overlooked tile ilmportanccinjury from drought stress low yielding fields from d ought of upland rice insect pests Many reports of upland rice pestsstress would show disproportionate differences between in Asia ha Ve focused on major lowland species (46 108)insect-protected and -unprotected treatments Shoot and It isalso surprising that tei vegctative stage of upland rice root pests would be more responsible for losses from would record the highest yield losses becatuseseeral studiesdrought stress than sown-scd pests A second reason have shown that upland rice can readily recover from foliageconcerns seed pests which would have a greater impact on reinoval during that stage (117 152 165 188) Upland ricelow tillering low yielding varieties such as )aggc Low poscsses a hitgh obilitV to compensate for ca rly loss oftillering varieties nnot fill in the space created by removal foliage and will even be stinulated by foliage removal toof seeds oweer 1) R i 5 is high yielding because it tillersP1 produce a higher viel than plants ithout any foliageactively which can close the canopy to provide greater weed removed (63 192 231) [ie reason for this apparentcontrol discrepancy hovce is that yield loss at the vegetative

stage does not octUr frol leat arca loss but fron pestsYield loss () removing sown seed and feeding on devcloping tillers and roots -- damage that cannot be readily conpLtnsatCd for

70 - Measuring yield losses in upland rice cnvironrnents is further complicated by a report from Brazil that greater yield losses occurred in response to better insect control60 - =-09 as Insecticide seed treatments resuled in denser plant stands

(n=21) which in turn created a selectively more favorable environshyment for blast (69) This resuIt was parlticurla rlxexacerbated50 when cirbofuran granules were used in seed furrows Aside

ffrom a greater plant stand ca rbofuran iay have auigmented the N lecl and stimulated blast If so ca rbofuran should be40 - replaced with another insecticide or the seeding or fertilizerI rates decreased

Other methods to determine the impact of upland rice30 shy pests wvcrc to interview farners or to develop single speciesbull bull correlations of population levels and yield loss Feneira (66)2 NIM)st (75(1) Golas farmers reporteddid both in Brazil

20- insect losses in ufpland rice and 251 said those lossesS avcragcd 37(1 [his result is consistent with tie yield losses 0 0 measured by field trials in the same statc(Table4) For every

I0 4 of seedlings removC d by leaf-cutting ants a I 7yield loss is predicted Similarly Ferreira (66) found that ai

0 - average of 53 thrips per panicle before panicle initiation meant 3 times more unfillh d grains

-IO0 I 1 1 I I-101 0 2 3 4 5 6 7

Yield (tho) C)N [ROt IM FI IODS 3 Correlation between upland rice yield and yield lols caused by Upland rice insect pests may be controlled by culturalinsects measured in 21 trials in the Philippines Ivory Coast andBrazil genetic biological and chemical methods

IRPS No 123 January 1987 25

Cultural control dry season plowing will have little effect on them Plowing Crop husbandry to reduce upland insect pest populations when the soil is wet from the first monsoon rains will expose can be classified into practices effective in a single field and soil insects that tunnel close to the soil surface to predatory those effective only at the community level Cultural birds chickens dogs and even man (field crickets nole controls should come first in a pest management program crickets and white grubs a re delicacies in man tra ditional They have broad stable effects as pests have little pos- diets) sibility of overcoming them through selection of biotyps The greatest degree i1 inIscct control fron tillage comes (138) On the other hand some cultu ral controls work only from plowing soon after rice ha rvest when most soil insects with high labor or power inputs and connunity-wilt lie close to the soil surface anrd stern borers are in the rice methods require coordination of iany farniters stuble Whitegrubsare mature and tillage exposesthem to

Pantin time The luxury of being able to shift the time of predators Tirnelv control will prtec the crop following pla ntring upland rice is open onl hr la rniers in areas with a rice and lowr the populationIfor succeedinig years Plowingprolonged rainifall pa ttern lI copping intensity or underthe ice stulble helps it decompose killing sten borer rnechanied land preparatiot Gcnerally planlting as soon larvae inside as the rFiinv season begins will lower populatiois of 1110st [requent tillage destroys ait nests bit not termites insect pests Insect populationsarelo aftcra dlvseoll or Tillage and clean culture howes er rcliove food for winter fallow delayed plaiting lets them build tip on termites which then mtay attack a young rice crop Scarab alIternate hosts that gi vigorotsly with the first seasonial beetles also prefer to miposit in eceitly tilled fields Noshyratiris This first flush iof weds cros lItxuttriartlv with iceCult is dibbling faors the survival oftillace Clre Such rineralied N released during the dr season aid made stils i aels tnd insects thtt pttpate or lay eggs- in the soil

available to the plants with the rains It iliso igrtmitcally Zero tilla hio eerl conserves predators ofthe GIH (37) advanttgeous for the rice ciop to tap this trlLItlaI fertility nt it mcttoi lo lessen the impact o0 soil insects and grow vigorotsly to compete with wrus and to tolerate brotdcasting seed or planting in ftltrrows is prefc ired over pest infestation dihlirig in hills Crickets are large enotlgh to eat seedlings in

Larly plantngs terd to escape seedling maggolts Upland a hill atrd tillering normnally ca iriot fill in tlhe missing space rice in Bitangas is planted in Ma and ine itli the Iirst Oftcn dibbll rd is not well covered and iseasily found by rains Fairners are highly mtotiated to plant early tocscape foraging insets ioIdents and birds drlitght and typlotns ricar harct Arn Atrgtst planting Plan uit sii Incrcatsini sced density protects tile cropwould be evcely attacked by It riim orivzo At tti againrst seed and seelling pests especially ants particularly other Philippiie tipland sites Rea l Qteot antd (laelit if the crop will germinate and ermerge quickly M isainis Oriental seedlig mnaggots arnrige ric boe t se hr trftinll itI India rice intcrclopped witi cotton or farnes have a longer growing season and plnit 2 rin or pigeon pci had lower 6IIt and WBIlII poptrlations than mote after the ortset of the rainy season liaaiangas ftrmners rice alone (211) NfailC arid Urplald ir_ 2re coilltori in the plant ca rly becia use ofa shorter raiinfall period arnd the desi re Philippines he chatngC in pest status folii iniercropping is to lliarvet before the typhoor sea son peaks ltighly locatioll-specific (133) cmianarid the net elfc be either

Early plantings cornbind with early-iatiuring varictics nil bcnelficial ortdetriential (189) For example BIatangias provide higher crp tolerartce for white grubs Aestiatior farmers intercrip Itro in their ipland ricefields A liornshyterminates with the first rains but 5-6 moare ineeded for the nrii iotd-Ahis i (I iiiacius) which norlially feeds resting larvae to develop into beetles emerge and lay eggs on taro ill feed or miarbv rice plants Intercropping and for larvae to reach the danriigii g ird-instar stage probably would not afft0 seed pests A crluanion crop (134 In lataigas this normually iseiough tiite for harvest planted ith rice and haing gleater tolerarnce for root

It Brazil howecver delayed planting is a suggested dariage or being toxic nayact isi trapcrop to root-feeditig cultural control for the lesser cornstalk borer and insect pests Acial-fei i ig pests voultId be most affected by pests favored l drought (66 195) Cropping internsity in intrer ropping Shoot aphids leaf beetles sten borers and Brail is low and delayed plraing places the rice crop in the hopplers would be likely ca ntidates to note effcts On the niost stable rainfall period reducing likelihood old rought othlt halnd the corlipalior crop stch~ sitniaie Iiiv be more

In rilniy arca s rinmirsare roi atl to plant other crops benehited than rice from initecropping ( 137) before rice and althNgh it woultl be desirabh to avoid 11echi Weeuling luring the first month after clop infestations by planting rice cuatl lariners lack the labor cstablishent will force pcts such is anyvworris which and time to do so ol a low priority crp prefer grassy cetds ontor tle vorling rice Soil iampsecs will

77age Uplild rice Stils ae light textured and Call be mvnye to the rice crop Cleani citrltirc also lay lore territes plowed duiring the d rv seasor notl only to grin time for to attack I u cetincti inrice ceds i ay he le optinal planting with lie fitst Inilsoon riiains but also to desiccate solition 13) soil insects arid weeds such is ntsedge that foim rhizomes Irtlizr Itneatsing N Iciiliier oin hlwland rice has Htowever becta use most soil insects lie below die plow layer fAtwvored Irigher populains of pliithoppeiv Imlhoppers

26 IRPS No 123 January 1987

and leaffolde rs However because of the need to temper fertilizer use on upland rice o prevent blast fIrtilizer rates will be lower than is comonia r for lowlad rice causing relatixely less hopper and IaIfTfoide r population buildup However inupland rice areas without a history of blast such Is Batangas farmers apply high rates of N and conscqtrently the crop has high infcstation levels of WBPI]and RIF The fhrmrcrs leec insect buildup by splitting the fertili11r in0to two to three applicatiois bit do not applV fertilizer at the vegetati estage because this prodrces a tall crop which illeasily lodge during typhoons Greater crop fertility leads to greater insect survival (better nutrition) larger insects (to cat no re foliage) arid greater fec titd t (to lav nmre eggs) (128 145 171) Not all insect species respondequally hovr Increasing rates (if N l anrid K increased Dialraa s-acchralis and (iaeiurema flea beetle but reduced thrips (66) Zn rcdLuced deadheart density from the lesser cornIstalk borer

Fnldingi Although most Upland rice areas do nIot hawxcaccess to wa ter floodiig the fields is rRrrnriierdcd in areas of Brazil or root bug and white gib control ( 195)

Crop rotatioti Tillage before planting a crop after rc harvest will control stern boreis and other pests renain n the stubble arid will unearth soil pests Planting a non-gratriiaceous crop after rice is recommncrded for termite control (66) Crop rotation is less nttdcd in upland rice culture than in lowlands because atcrop-free period is assured even after harvest of crops following rice

t1ulchilt iryl plastic irli irig designed for weedcoitrol ii upland rice inJapan resulted in ntr highanri liy infestation of (ihilo sulytresvais (93 9-1 95) NIulched Upland rice grew more luxuriantlv and attracted more (Avipositing mothis than u lched rice Also the rice plant and stein borers imatoured 10rC quickly and a partialgeneratiori af vouing ista iNoverwintcred in the stubbleLotili--malitrt varieti Quickly maturing crops reduce the tmbCr of pCst generations that Calt build up Rapid crop establishment olan ca rl v-natu ring vaitvty if)pa 11icLa r will lessen white gib and stem borer damiage White grub larvae intle last iistar are highly destIruCtive so aii early-iiatri ng variety will escape serious root loss Stem borers build 1ip slowlyx in rice to ecormic abundant in a late-Maturing crop anrd caiuse whi tlieads Numibers of somespecies increase exponeitially with each succeeding genera-ti so a xarietyexhat atures I trowh earier thian another will hac less insect damage Early-naturing vaiicties however will not lessen seed pest daiage either ii the soil or oina standing crop

Strwhoi hAIJttam Inrscts tcd irig on thlie aerial portions of rice plats disperse from carly to late plantedfields AS iiiibCirs increase exponentially with eachinsect

gtscnrattm e eiglbrig f rmc~s1(uil upp wh rc generation pests build tIP where neighboring fihrTC e9r1sstagger their plantings II Batangas farmers plant uplandrice within I iro use 120-d s-arieties and plant at the start of the rainy season Ini Misarnis Oriental laricrs stagger their

plantings up to 3 mo apart and use a set of atieties that mature in 4-7 mo Insect pest danage particularly from stem borers is greater in Misarmis Oriental Also seedling maggots damage rice there because rainfall favors frequent planting of maic In Batangas rice escapes seedling maggots bCca usC of distinct drs season and synchron0us early planting with the onset of ains

Therefore combinations of cultural practices ea rlyplanting synchronous planing crop rotation and earlyshymaturing vaneties protect the rice crop against most insect pests

Ilmant resistance Beca tse upland rice has few specialized insect pests and many insect types attack the crop worldwide regional breeding and strong internatiornal cooperation are required

Sown-sced pests arc economically controlled by insectishycide seed treatment and should the relore receive lowxpriori ty inbreeding objectives

Root pests are normally cost ly to control with insecticides but finding resista t sonres mayvbe di fficult Tanaka (229) for exa -ple fi led to find varieties resistant to root aphids Selecting for large root bioniass perhaps should be the strategy for pests sLuch as Whlite grubs that renliove roots Fainter(174) howcvcr cites varietiesof crops tolerant ofor resistant t(i root pests Some sorghun varieties are high in cyanide so rice sorghun might incorporate broadshyspcctruri chemical resistance

Seedling vigor and drought tolerance are high prioritybreeding objectives so tupland rices should have large r root systernsand be able to tolerate ligherlevels of root loss Sonie root loss inolder plants is beneficial if it stimulates new root developnrut and therefore crihances intake of soil nutrients (53) The rice seedling inaggot rtiLotia spp may be

controlled thirough resistant aittics A hi rgc breeding effort on sorghniu has had fair success inideveloping arreties esrstait to Asoccata Rodani (251) A field trial comparing 10 upland ricc varieties showed differences aniong varieties va ring from II to 4(- damaged tilleis (215) Hovver resistarce to 1Iill-0r gtia species may rot cross ox-rti otier

Shiraki (210) reported that upland i-ice stens wcrc harder than lows-land rice stemsperhaps becaIusc (f a higher silicoricontent and thus were more resistant to yellow stern borer hc tortia Iow tillering (f uplaid iice xarneties prodtres a

higher percentage of infested tillers than would occor in higher tillering lowland iice under a siiilar egg density The corollary therefore is to select for high tilleriig upland rices

Stemii bor amcstheh main targes dobrecders inAfrica (571 204 )but nticl oifth e sc reen irig is dotie unde r lo wland co d t osrm s wih A h r~ o l pe i s n acorditions (11 96 218) As with lthrinrnwspecies mayhave to be dealt with independently however initial results show cross resistance between African and Asian species(92)

In Brazil screening Ior insect resistance focttses oil lou I pest problems Varieties tolerant of lasmolwalpus ligi-sellus and resistant to )iatraea saccharalis have been identified (70 153) Current work also emphasies 7iraca limativetntris and Thmialspis (=Iosis) flavouicta (66) These insect pests Ilt onlV citise IIigh yield losses tihey also ire difficult to control chemically ofr by other ilealls

ManV upland rice vaieties aic early maturing and are resistant to pests such as gall midge (I Xl ) Early mIturity is often a highly desirable trait to esuilpe pest buildup but it is not true resista nce bca usc ilthose va reties are plinted late they will be itsdamaged as ain early planted susc ible variety

P~erhaps tile cosiiiopolitall of all upland rice pestsimost are the sced pests (ripenling stagc) which pose a particular problem beta use scuds re ised IsIO for hitnit4ans Ayltoxic substatice introd tcld geticticllyilmay also be toxic to nal iniless it caln he placed old in thfe IdhuffSelectmi fo hairiness or lotng wns kill mIlct sthilrohleiiis dutilig threshing pa rtictfarl maitial threshinfl [hireshers ill complain of itcfimin atid itteascd (fust levels

Attempts itIRRI to fitid tcistatncc to the rice bug plorisa have ilotbeCnl prmitisilg lIIetIattonlids iorc

through the lenilna or palca hillhairs sedstVIor seeds wilh awns that deter bugs promote complaints from threshers Seed bugs liptocorisa and Stcnocoris enter the seed through the opening between tile lemna and palC but varieties with narrow openings are diifficult to mill

Breeding priorities should concentrate ott inotCI gruLtLIps that are most difficult to c nttrol by other meains Steil) horers appear to be the most widespread group that sigiiificantly dalag s rice aind is hard to manage by other Ilnca us

Biological control Very fe stutdic of mitatt~l eneties of rice pests in upland envirollnletits wee in tile wefouiid literature Therefore have not compiled a list of species Ve present a list of natural enemies iOtid in upland ricefields tuic latialln Batangas for some conitno iOldiar pest species ([itble 5)

Muich ofl thc lollowinle itfor ittiol cotlliCs rol Stutdies in the lowlatnds or roi crops otithcr thlin tice

Qu(ailtr o 11ll t11t( Nat trrtf lii hcoeits ight Ile used agaitist ile social insects tilt and termites Sollie predator specice live ill theirf nests n ilnic theirnllt ip-pearanIlCe aMRidbehavior IJtrouMdCing a mote aggressise ant species to displace pest ant codonies is a possibility (169) Patihogens as potential natural encnies have reccived little atteition (I13) Natural ncnuics asa group however hac not been toIl iporta it in Iegulating aintshown be and ternite numbers

The nocturnal behavior aid subteitmien habitat of lole crickets and field crickets protect them from many ntllral enemies There are only limited records of natural cnelics for these groups of pests some pathogens (87) but mainly

IRPS No 123 JattuIny I 271987

parasites - scelionids on eggsand sphecids on ilymphs and adults Sphecid wasps Larra carutmaria (SImitI hI L huzwuensis Rohwer and lsaltiuhta Willi ams speciali in mole crickets pa ralyzing them nd dragging the ninto thcir nests as food lot their voting ()tier sphCcid species Lins aurulenoa (Iabricius) lot maiac (Ashmead ) 1 lhshylssellaus (Smith) and Ml lal)oriosu (Snithf specialie in field clickets

White grUbsO the other hand hae Icti scriuns enuigh to have been extensively studied for cotttroi 1W nati al enemies( 112 140 141) Scoliid parasites C(ajsoMcispp have beetn intrtduced itto tilePhilippites for control of Ielxl ]udisiro la The rcsearchcts w Intttrdced themle during ill era claimed that the white gttionthreak as controlled (142) howevcr it isnormal for pest epidemics to subside and there was no diect evidence to indictte that tfe parasite was responsible A tmore promisitg titctllod might he to identify iso ltcttass prt~f tcartd apply hiactelaln fingal pathogens to tile soil Ihe lalattese beetle in North America has hcci coitrolled hv tite conmnleciall aailable hactcrial prcpa ration itacihlpu pav ofr milksdiscas (71) Spote prtyefott the soil itloted ilrtier reitiain actic fr Veas

Root aphid sardtnctly bugs readilslic tilib to predatots and parlsites once their tetndine ants are controlled The larvac and adtilts of a coccinellid beetle 1Scynms sp prey on llriadlouais in slash and burn upland rice Baits treated with insecticide may be used to control tending ants by killing their ourig (244)

Root-tkcditig blugs false wireoris wireorms and root wccils aic r ofsuch little ifilpoltatice that thenomttll role of liatural Cticniics hias tiot beei asscssed A possibility hoevrl tiglt he tit ItaIIitic niaCIMtodcs lintt1g11en Itese soil-itlhlfbitine mCtitodcs W5uhtfhave to be imiass Produiced

large miltibcr of niatltrll cnemics ircknown ft CltIoriS IC IpblCeii is thtt clttvortnlls ofonlie the crop soott alter land ptel)tritiOti a1tithe bcgit iniig of the raittv

Citsoln hfien tiatllral leniiiy populations aric lo Colltloil WOtuld iii ole mass prodtticing andirclcaslng key spccies

le egg stage is the fpirt (olthe Cedlitgt Ialot life cycle tI hich to f)ctis ftor tittlril elctit cottntrol hplraites o1 pleditors ttsit elleggs highly expoIsYCl I Sgg pailsites or predators stolld probahly e ittass andtbe produced released Littlh i toruialioll is a ailable ott what species to trV o ho to Iniss podLice thiCIi A etiloplhid wasp Icmihlaol sp effctiicly regushy

lates the leatf lcloalapolm2a spioanoiiitr heat andc rice antid should be collsctvcd

leaf beetles whCllose e am] eggs ate laid in the grutindhirO ould be motC difficutlt to control by niatulral enelV Mnll ipuhltiOn But species sUici IS c_gsthispid beetles ith

and larVle oilleascs ate vtilcratble to iparaites and pledators

Natural enemisc of tlirips otl rice are little known

28 IRPS No 123 January 1987

Table S Parasites recorded from upland rice insect pests in Tanauan Batangas Philippines 1977-84

Order Family Species Host stage

I lymttenoptera Braconidae Leaffolders Cnaphalocrocis Marasinia spy

Cardiochilesphilipplinensis Ashmead a Larva

Cotesia angustillasis(Gahan)Cotesia or cypris (Nixon)

Larva Larva

Cotesia or taeniaticornis(Nixon) Larva Cilonusinotakatae Munakata Chelonits SppAlacrocetrts philippinensis (Ashtmead)

Egg to larva Egg to larva Larva

Iclineumonidac Orgilts spp Isclutojppa hireator(Fabricius)

Larva Larva

Itoplectis naraigav(Ashmcad) Larva Tenelhtcha philippiensis (Ashmead) Larva Tenelchha stangli (Ashmead) a

irichomma cnaphalocrosis Uchida Larva Larva

Bethylidac Elasmidac

Goniozms or trianguli r Kieffer lasmtusalhopictus Crawford

Larva Larva

Diptera

Encyrtidac Chalcididac

Tachinidac

Elasmtus spp Copidosomopsisnacohiae (Fidy) Brach nteria evcarinata Gahan Biachyieria lasus (Walker) Ztgohothria ciliata (Wulp)

Larva Egg to larva Pupa Pupa Larva

Argyrophylx nigrotihialis(Blaranov) Larva

l-ymenoptera Braconidac Yellow stern borer Scirpopiaga incertulas

Braco chinensisSzeplegeti Larva CIthelotis spp Chelonus niunakatae Munakata Cotesi flaripes Cameron Stenoracon nicerilli (Bingham) a

Larva Egg to larva Larva Larva

Ichneumonidac Tropobracon schoenobii (Viereck) Amatroinorpiaaccepta metathoracica(Ashmiead)Frihorus sinicus (I lolmgren)

Larva Larva Larva

Ischnojoppaluteator (Fabricius) Larva Isotima nr dammeratiti Rohwer Larva Tenelucha philippinensis (Ashmead) a

Larva Tenehicha stangli (Ashmcad)Trichomma enaphalocrosis Uchida

Larva Lara

Chalcididac Eulophidac Trichogranmmatidae

Xanthopimnpla stemmator (Thunberg) Charops hrachYpterumn Gupta and Malteswary Brachtm eria sppTetrasticttus schoenohii Ferriere Trichorigranm chihunis Ishi

Larva to pupa Larva Pupa Egg Egg

Diptera

Scelionidac Pteromalidae Tachinidae

Tt ichograintnta aponiicuitt (Ashniead) Telenomits rowani (Gahan) a

Trichoinalopsis apattelctena (Crawford) Jeirhaea spp

Egg Fgg Egg pupa Larva

Z4gobothria ciliata (Wulp) Larva

Ilymenoptera lecneumonirlac Gol fringed borer (hilo auricilius

Trichomm enoptalocrosisUchida Larva

Ieioenoptera Braconidae

Aantholimlh steiniator (Thtinberg) a

Pink stet Ioret Sesamia inferens StoInacoil nicrilhi (Bingh am)

Loarva to pupa

larva

Diptera Ilyinenoptera

Armvtnyorms Mythimna separata Spodoptera mauritia acronyetoidesTachinidae Zvgohothriaatropiora( Robinea i -Desvoid y)Eulophidae It-phctrtiscitapadoe (Ashimead)

Larva Larva

(halcididae Braconidac

Brachpin ria spip (oesia spp

Larva larva

IHyntenoptera I)iptera

Rice skipper Ielop ida (halcididae Tachinidae

math ias aidgreen iortedci ciliarNIelan itis eda Brat lhincria sp iii matuqitatt Cameron Irgyrolhirlaxi trotlhiais(marumtv)a1

ismtne Larva Larva

Diptera Ilymenoptera

Tachnidae lelineutionidae

fuown se tilo per Mois frugalisIrgtrophrlax 1igrothiahiv 3 a rat ov) anthopimpa pmnctata (Iabricius)

Larva Larva to pupa

aMost dominant

IRPS No 123 January 1987 29

Normally predatory thrips prey on herbivorous thrips strepsiptera on both Spiders dominate tilenymphaladultNatural enemies of stem bugs --Scolinoplara and Tibraa predators Spittle bugs move to rice from pasture grasses - are in order of importance egg par~isites and predators and biocontrol would be successful only ifcarried out before fungal pathogens and nymphal adult parasites Egg tiley immigrate into ricefields parasites could either be introduced fron other areas or Biological control tactics for hemipterous seed bugs mass produced and released ould follow those outlined for stein bugs

Armyworms are normally lwld in hcck by the activities Coml)arison of environnewtns Natural enemies are of egg and larval parasites When these natural enemies fail perhaps faced with even greaterconstraints than insect pestsusually because of drought armyworns become epidemic to survive ind reproduce in upland environments TheyIt may be worthwhile therefore to release parasites during have adapted ani survived by having wide host ranges the the rainy months following a drought Virus diseases of ability to aestivate or hibernate over unfavorable seasonslarvae would beanotheravnue to exploreforarnyworms and the ability to disperse or by other mechanisms

Parasites have ben traditionallyconsidered for biolegical Comparing the most dominant and therefore the most control attempts against stein borers usually by rekasing adapted species of natural enemies collected in various exotic species( 166) This approach has not met with success environments in the Philippines we can understand how and new avenues should be explored The role of predators different rice environments are arid pathogens is little inderstood and deserves greater Anagqrms olptahiisa mymarid planthopper egg parasiteattention Egg parasitism rates are normally reasonably is cquallyadapted to all Philippine rice environments (Tablehigh but should be supplemented with effective egg 6) It is selective for planthoppers and even parasitizes eggspredators Orthopterans are potential egg predators of the n-iize planthopper Peregrinitsmaidis itt upland areas Metioche vittaticollis (Sta ) Anavipha nifiwnnis (Saus- Alavohis also parasitizes maize planthopper eggs but is sure) and Conocephahs hongipnni (de i-laan) feed on most prevalent on BPH in the irrigated wetlands (onashystem borer eggs etioche and Anaxil)ha specialize in eggs tocerus spp like A optalhilis on BPH and WBPI Iattacks with no hair such as Chilo spp The omnivorous (ono- the eggs of GLH in ell rie environments It is also found cephahsspecializes in eggs cov red with hairs it is a proven parasitizing the eggs ef the white rice licahopper (oimna egg predator of Scirlouphaga spp but has the discouraging spetra Oirosita naias a trichogrammatid egg parasitehabit of eating rice guin (65) attacks BIPH and WBPH in the wetlands and has no other

Grasshoppers have egg (scelionid) and nymphalad ult host A related species 0 aesopi specializes in G1-H in (nemestrinid tachinid and sarcophagid) v irasites Among wetland environments Dryinid strepsipt-an and pipunshypathogens protozoans have been recorded most frequently culid nymphal and adult parasites of planthoppers and Control of locust species by natural enemies could con- leafhoppers also specialize either in hopper species or centrate on their habitual breeding areas environments Among the dryinids Pseuhdgonatoipsflavi-

Leaffolders have rich complexes of natural enemies some feimur prefers the lowlands and BPH while P nuch has no adapted to upland rice (22 27) ranging from egg predators environmental prefirences but is adapted only to WBPH (gryllids coccinellids) to larval parasites (braconids ichncu- The main d ryinid species on GLH aptogonatop)usspp is monids) to larval prcdators (ants and carabids) to larval most dominant in rainfed lowlands and also attacks other pathogeiis (vira and fungal diseases) Efforts to augment leafhoppers within Amnrasca Cicadulinaand Bachaltuathese natural enemies should focus on the egg predators Strepsiptera are more dominant on plarithoppers than (rearing releasing)and viral or fungal pathogens (culturing lealhoppers The Elenchus species in the Philippines attack spraying) all species of planthoppers within the genera Sogawilla

Parasitization of the large conspicuous larvae of Sogatodels NilaparsataHarualia and Opiconsiva Elenshyskippers green horned caterpillars and other polyphagous cusrvasuntaisuiis adapted to the major rice environments Lepidoptera by tachinids and chalcids is normally low ELg but prefers BPH in rainfed arid WBPIH in wetland environshyparasites and predators are perhaps the key to their control ments Ilalictoph(utsmtnroeiand HVpectrus attack G LH

Gall midges are parasitized by pterormalids and platy- and COfimravcctra respectivelygasterids that aestivate within their larval host between Pipunculids are mainly parasitic oii lealhoppers Each seasons Shoot aphids are preyed upon by coccinellids species has a unique environmental preference Tomosshysyrphids and chrysopids varireIa oraetoraprefers Ne7 otetltix nigropictrs and

Planthoppers arid leafhoppers have egg parasites and Pilunwulhtsjavanensisattacks Dehoceplhahls spp which predators as well as nymphialad nlt parasitesand predators are more abundant in upland than irrigated lowland their rich complex of natural enemies includes fungal environments T suhvirescensattacks Nephotettix virescens pathogens The principal egg parasites are myrnarids and and prefers the lowlands Pilpuiulus nttillattls is an trichogramniatids Egg predators are norinally mirid plant upland rice parasite arid attacks only Nephotetix speciesbugs that suck out the yolk Nymphal ad nIt parasites are More than 17 recorded species of leaffolder parasites in dryinids on planthoppers pipunculids on lealhoppers and the Philippines attack species of C(naplwocrocis and

30 IRPS No 123 January 1987

Table 6 Dominant natural enemies of rice pests in upland rainfed lowland and irrigated lowland rice environments of the Philippines

Natural enemy

Host stage Species

Brown planthop-Parasite Egg

MymaridaeAnagrus optabilis(Perkins) Anagrusflaveolus Waterhouse

TrichogrammatidacOligosita naiasGirault

NymphAdult Dryinidac

Pseudogotatopuslaiifemttr Esaki et Hashimoto Elenchidae

Elenchus yasurnatsui Kifune and Hirashima

Environment

Upand Rainfed lowland Irrigated lowland

Ailaparata lugens

x x X x

x

x x

x X X Whitebacked planthoppers Sogatella furcifera and Sogatodes pusanusEgg

MymaridaeAnagrus optabilis(Perkins)

TrichogrammatidacOl~gositanaias Girault

NymphAdult Dryinidae

Pseudogonatopusnudus Perkins Elenchidae

Elenchus yasutatsui Kifune et Hirashima

Parasite Egg

MymaridaeGonatocerus spp

TrichogrammatidaeOligositaaesopi Girault

NymphAdult Pipunculidac

Tomosvaryella subviresccns Loew Tomosvaryella oryzaetora (Koizumi) Pipunclusmutillatus (de Meijere)

DryinidaeIlaplogonatopusspp

Egg Copidosomopsisnacoleiae (Eady)

Larva BraLonidae

Cardiochiles philipphnensisAshnead lchneumonidac

Trichiomnia cnaphalocrosisUchida Temelucha stangli (Ashmead)

Pupa Chalcididae

BrachymeriaexcarinataGahan Brachymerialasus (Walker) Brachymeria nr tachardiaegroup

Parasite Egg

Scelionidae Telenomus rowani (Gahan)

EulophidacTettasticusschoenobhi Ferriere

Larva Braconidae

Stenobraconnicevillei (Bingham) Iclhncurnonidac

Temelucha phiiippinensis(Ashinead)Temelucha stangli (Ashmead)

x x x

x x

X x X

x x Green leafhopper Nephotettix virescens

X x x

x x

X X X X

X

X Rice leaffolders CnaphalocrocisMarasmia

x X x

X

X X

x x x x x X x x x

Yellow stem borer Scirpophaga incertulas

X X x

x

x

x X XAinauromorphaaccepta metathoracica(Ashmead) x

continued on next page

IRPS No 123 January 1987 31

Table 6 cntinued

Natural enemy Environment

Host stage Species Upland Rainfed lowland Irrigated lowland

Larva-pupa Gold fringed borer Chilo attriciiuslchneumonidae

Xanthopimpla stemmator (Thunberg) x aPupa x

Pteromalidae Trichoinalopsis apanteloctena (Crawford) ax

Pink stem borerb Sesamnia inferensParasite

Larva Braconidae

Stenobracon nicevillei (Bingham) x b b Armyworms Mvthinna SpodopteraLarva

Tachinidac Zygobothriaarroptvora (Robineau-Desvoidy) x x

Braconidac Cotesia spp

X Skipper Pelopidas mathias and green horned caterpillar Melanitis leda ismeneF-gg

TrichogramrnatidacTrichogrannaspp x

Scelionidae Undetermined Scelionid

x Larva

Tachinidae Argyroph lax nitrotibialis (Baranov) X x

Pteromalidae Trichomnalopsis apanteloctena (Crawford)

X Predator

CoenagrionidaeAgriocnenis femina femina Brauer x xIschnura senegalensis tambur

Miridae CrtorhinsliidipennisReuter x x

Nabidac Stcnonabis tagalicus (Stal) x

VeliidaeAlicrovelia douglasi atrolineata (Bergroth) x

Coccinellidae Micraspis crocea (Mulsant) X x x

Vespidae Eumenes campaniformis (Fabricius) x

Formicidac Diaecamna spp xOdontoponera transversa (F Smith) x

Carabidae Chaenius spp OphioneaW- aJasciataSchmidt-Goebel

x

Ophionea ishii ishii Habu x x X

Gryllidav Anaxipta longipennis (Saussur-) x xMetioche ittaticollis (Sial) x x x

Tettigoniidae Conocephalus longipennis (de Haan) x x x

aNo rearing made on rainfed lowland bNo rearing made on rainfed lowland or Irrigated lowland

MrasmiaAmong the Laval parasites the braconid Cardio- lowlands and parasitizes oriental maize borer OsIriniachilesphilijpphensis is most adapted to an upland environ- furnacalis (Guenee) larvae The leaffolder pupal parasitesment It also parasitizes Hydelepta indicata a common not only occur equally in all environments but also haveleaffolder of legumes The ichneumonids Trichomma cna- wvide host ranges- parasitizing species of OsIrinia Marucaphalocrosis and Temehcha stangli occur more in the Hydelepta and Honiona - in maize and legumeslowlands T stangli is most adapted to irrigated lowlands The principal egg parasite of yeltow stem borer (YSB)and also parasitizes Chilo and Scirpophaga stem borer Telenomus rowani is widely adapted to all environmentslarvae T cnaphalocrosis is most prevalent in rainfed and has an unusual alternate host -Tabanus eggs (25)

32 IRPS No 123 January 1987

Tetrastictu schoenobii specializes only in YSB and occurs mainly in irrigated lowlands -- further proof that YSB is most adapted to the wetter environments

Larval parasites of YSB are highly environment-specific The braconid Stenohracon nicevillei occurs only in upland rice but also attacks the pink stem borer (PSB) Tieinchwa phiiiplPwnsis an ichneumonid occurs in upland rice and also parasitizes the larvae of (7hiu stqpresaisinatro-morpha accepta melaihoracica like 7 stangi is mnost adapted to the irrigated lowlands

The dark-headed stem borer(l)HSB) (ih pochit3sus was not collected in all rice environments and the gold fringed borer Chilo auricilius nd PSB were collected only in upland rice Xaantiholiplastenrnaloran ichneumonid larval parasite appears to be widely adapted whereas TrihomaslopisapanitI)cteiiaa pteromalid pupal parasite occurs mostly in wetlands and also parasitizes larvae of Chilo suppressais and Pelhuidas mathias

Armyworms Myit hinin and Spodujipera have two main larval parasites - a tachinid Zrgohothriaatropivora in rainfed environments and a braconid Cotesia spp in irrigated lowlands

The butterfly pests Pelopidas and 4elanitishave tricho-grammatid egg parasites found only in irrigated lowland environments

Two larval parasites show environmental preferences --Argrroph av nigrotihialis a tachinid in rainfed habitats and Trichonialopsisapantelkcnela in irrigated ricefields A limgrotibialisalsoattacks the sweet potato hornwormAgrius convolvtdi and T apanteloctena parasitizes DH S1B

Predators as a group show more distinct environmental preferences The aquatic damselflies Coenagrionidae are most abundant in the lowlands Cvriorhinus is also most adapted to the lowlands The nabid Stenonabis tagalicus occurs only in the uplands and isalso prevalent on legumes The coccinellid lady beetles wasps and ants are more prevalent in upland areas AlicrasIpis cit-ea larvae prey ona variety of aphids attacking legumes and maize Etunienes campanonia vespid mud wasp makes nests in trees and is therefore most adapted to the more botanically diverse uplands Soil-dwelling ants cannot tolerate flooding

Upland environments are habitats to arboreal carabid beetles Three species of Chlaeniusbeetles prey on leaffolder larvae and are perhaps more important than parasites (27)

The gryllids and tettigoniids are widely adapted egg predators not only on rice but also on maize and legumes They feed on eggs of most insects that are laid on leaves

The spidercommunity of upland rice environments isrich in species At one site (Batangas Philippines) 31 species have been recorded (23) Of the 176 spider species recorded in Philippine ricefields about one-half (82) occur in upland rice The spiderspecies of upland rice environments overlap more with those of rainfed lowland than with those of irrigated lowland rice (24) The same study showed that of the three environments irrigated rice has (ie greatest spider species diversity foiiowed by rainfed lowland and upland

However of the 10 most prevalent ricefield spiders 9 were abundant in upland rice showing wide adaptation All three spider guilds -- orb-web space-web and huting spiders shywere prevalent in upland ricefields There were however differences in environmental prelerences for some spider species

Of the orb-wcavers Tetragniathamanlihulatawas partishycila rlv abundant 7 japonica was low and Ltucatge dehcorata was absent in upland rice

As in other environments Aiypeiwnaifirtosana was the most dominant space-web spider however its relative numbers were lowerthan for lowland sites Among hunting spiders Ox|opes javanus and Lcosa leatcost fgla were more abundant and Lvcosa lpseudoanmnlata was less abundant in upland compared with lowland rice envishyroniments

Extensive rearing of rice insect pests in upland rainfed lowland and irrigated sites over an 8-yr period in the Philippines provided insight into the effectiveness of parasites as natural enemies (Table 7 8) Most of toe data are from large sample sizes taken over at least one crop which overcomes the pitfalls expressed by Van Driesche (238)

VSB egg parasitization was surprisingly similar across all environments Larval parasitization was highly dynamic among sites and even years within the same site but showed somewhat lowerlevels in uplandsthan in irrigated wetlands This would have been predicted for egg parasites as well since YSB is adapted to the lowlands and so must be its parasites This isevidence that Telenomus rowani is as good at dispersing as its main hosts Trowani clings onto the anal tufts of female YSB and parasitizes eggs as they are laid This phoresy may explain its wide adaptation The more specialized lowland YSB larval parasites therefore appear more effective than their upland counterparts with more alternate hosts

Low collection levels of PSB larvae especially in the wetlands make comparisons between environments difficult Low levels of larval parasifization were recorded in both rainfed environments Except in the Cagayan rainfed wetland site in 1980 low levels ef larval parasitization were evident at irrigated and rainfed sitesalike No differences by environment were evident Higher levels of parasitization occurred in Sarawak during a 1967 outbreak also from tachinids Rothschild (197) indicated that higher parasishytization occurred in upland rice areas

Skipper and green horned caterpillar larval parasitization rates were generally low and showed no environmental effect htighest activity occurred in Cagayan in 1980 (48 parasitization) Although larval parasitization levels of RLF were variable year to year activity appeared greater in the irrigated lowlands than in the uplands Again the highest levels were recorded in Cagayan in 1980 Cagayan has extensive grasslands and fallow around the rice areas and a relatively long rainfall period Rainfall in 1930 was good for crop growth and apparently for increase of larval parasites

IRPS No 123 January 1987 33

Table 7 Parasitization rates of rice planthoppers and green leafhopper in upland rainfed lowland and irrigated lowland environments Philipshypines 1976-840

Environment and site Year Egg parasitization () Nymphaladult parasitization ()

BPH W13P GLH BPII WBPH GLIH

UplandBatangas

Rainfed lowland

1976 1979 1980 1982

15 17 12

2(47)8(39)

4(51)

9(1800)14 (212)

12(304)

8 (101)6 (127)

9(116)

Iloilo

Cagayan

1976 1977 1978 1981 1980 1981 1982

0 (69)

11

44 48 40

-

-

46 55 44

0 (471)

11

52 56 49

0 (200)9 (214)

10 (1259)8(235)6 (900) 3(800)

-7 (462)

11 (270)3 (313)5 (900) 1 (800)

4 (600)10 (725)10 (362)12(870)5 (900) 3 (800)

Pangasinan

Irrigatedlowland

1984 1976 1978 1979 1982

13 (436) 13(379) 14 (457)

6(14)7 (197)

17(198) 11 (486) 18 (275)

3 (81)10 (274)3 (116)

16(871) 19(88)

28(211)9 (764)5 (647)7 (1204) 8(305)

Laguna 1977 11 7 21 1978 31 1979 14 4 4 28 1980 7 8 20 1981 1983

56 7(278)

26 16 (1650)

16 27(1921)

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R I Apostol unpublished 1lo1lo and Pangasinan 136 Cagayan BCanapi unpublished Liuna 175 I C Pantua unpublished 118 234 43 Carino and Shepard unpublished BPH = brown planthopper Nila parvara htgens W131Pi =whitebacked planthopper Sogatelia furcifera GLH = green leafhopper Nephotettrix virescens Figiires in parentheses are sample sizes

Tauher et al (230) reported that internal external and genetic factors influence the seasonal activity of parasitoids

Hopper egg parasites but not nymphaladult parasites were also parlicularly abundant at the Cagayan site possibly the egg parasites have more alternative hopper prey in the weedy areas than nymphaladult paisites

The seasonal dynamics of hopper egg parasites can be seen for upland rice in Batangas(Fig I 2) and for rainfed lowland lice in Pangasinan( 135)and Cagayan( 105) Levels of BPHWBPI-L and GLH egg parasitization were corn-parable in upland and irrigated lowland environments Except for some high rates of paraMsitization early in the season the parasitization rate tended to be steady in all envilonments However hopper nymphaladult parasi-tizution levels were more similar in the rainfed environments and lower than in irrigated rice

Overall parasites do not appear highly effective by themselves against upland or even lowland pests Among natural enemies parasites have attracted the most attention because they can be more readily assesscd

Predators and pathogens lend themselves better to management practices Predators the most important group of natural enemies in riccfields are difficult to quantify On the other hand pathogens appear less im-portant but can be readily cultured and disseminated

The natural enemy commulnity of upland rice is rich in species and differs significantly from that of lowland rice

These beneficial organisms must be conserved by applying insecticides judiciously particularly sprays Seed or soil placement of chemicals minimizes exposure to natural enemies The strategy to derive the greatest benefit from natural enemies isto allow the greatest number of beneficial species to thrive Then perhaps at least one species will be effective against each pest at any tine overcoming the variable abundance of each species within and between years

Programs to introduce exotic species or mass produce indigenous ones are ambitious and expensive because they require trained people on a sustained not ad hoc basis

Chemical control Insecticides offer rapid and efficient control of upland insect pests (190) but-should be used only after other control measures have been considered Insecticides are rarely used on upland rice because of cost (32) Upland rice yields are normally too low to justify the expenditure Also most upland rice is grown as a cash crop Spraying upland rice is more difficult than lowland rice because water is less accessible Government subsidy programs to provide pestishycides to tribal peoples for upland rice production have created problems of toxicity to humans because isolated people have had no experience to be able to handle pesticides safely

However ins-eticide use has beenjustified when a crop is

0Z

Table 8 Parasitization rates of lepidopterous pests in upland rainfed lowland and irrigated lowland environments Philippines 1976-84a

Parasitization (5)

Environment and site Year YSB GFB PSB Earcutting Swarming Skipper Greenborned RLF Brown larva larva caterpillar caterpillar caterpillar caterpillar larva semilooperEgg Larva larva larva larva larva larva

UplandBatangas 1977 - 0 (57) - 0 (63) - 0 (71) - 0 (106) 11(142) 2 (101)1978 43 (4697) 8(451) 5 (38) 3 (40) 6 (61) 2(58) 15 (57) 13 (129) 13 (1286) 4 (30)1979 59(4135) 12(320) 13 (11) 5(28) 9(45) 6(46) 7(23) 8(160) 9(1152) 8(26)

Rainfed lowland Iloilo 1976 - 0 (10) 0(10) 0 (7) 0 (19) 2 13) 15 (180) 5 (76)1978 0 (54) 0 (3) 0 (17) 9 (64) 4 (78) 3 (73) 0 (121) 0 (191)1979 64 (2350) 16 (84) 6 (17) 0(6) 18 (68) 25 (104)

1981 56 (1322) 18(304) 0(5) 12(51) 18(138)Pangasinan 1976 60 (7 7 )b 62(82) 0(111) 5 (39)

1978 58 (53)b 5(51) Cagayan 1979 59(713) 8 (101) 6 (34) 0(7)

1980 0(11) 4 (28) 15 (286)

62 ( 48(177) 61(140)1981 0(5) 12 (2) 13(45) 14 (92)

Irrigatedlowland Laguna 1974 62 8

1974-75 50 41 1975 43 72 1978 96 (623) 21(24) 33 (60)1979 83(2017) 18(110) 8(62) 11 (89) 22(400)1980 78 (1625) 20(75) 0 (16) 4 (72)1983 60

9 (80) 7 (270) 30

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R F Apostol unpublished Iloilo and Pangasnan 136 Cagayan B Canapi unpublished Laguna 175 P CPantua unpublished 118 234 143 Carino and Shepard unpublished YSB = yellow stem borer ScirpophagaincertulasGFB = gold fringed borer Chilo auricilius PSB = pink stem borerSesamia inferens earcutting caterpillar Alythimna separata swarming caterpillar Spcdoptera mauritia acronyctoides skipper Pelopidas mathias reen horned caterpillar Melanitls ledaismene RLF = rice leaffoldersCnaphalocrocismedinalis and Marasmiaspp brown semilooper Mocisfrugalis Figures in parentheses are sample sizes Egg masses

IRPS No 123 January 1987 35

Table 9 Effect of seeding rate and insecticide on plant density and Yield losses from insect pests in upland rice often areyield of Dagge upland rice Batangas Philippines 1976a comparable to those of lowland rainfed or irrigated rice

Seeding rate Insecticide Dosage Plant density Yield (0-Y30o) but usually from different pests The principal (kg aiha) seed treatment (kg aiha) (noIra-row) (tha) groups are soil pests feeding on sown seed and rootsat 14 DE followed by seedling pests These groups have largely gone

50 None - 35 a 20 a unnoticed leading researchers to conclude that stem borers50 Carbofuran F 10 38 ab 23 a and seed bugs are the most important pests (17) Yield loss 10 beCabofuranF100 None - 4551 c 23 a trials that pinpoint major pests need to includc seed

100 Dieldrin WP 10 52 cd 22 a treatments Granules applied in the furrow may not give150 None - 59 d 22 a adequate protection from many seed pests

aAv of 8 fields DE = days after crop emergence In a column means The floristic and pedological diversity of upland rice followed by a common letter are not significantly different at the environments is matched by the diversity of arthropods5level by D)MRT F = flowable Wi = wettable powder both pests and their natural enemies Irrigation has homoshy

genized lowland rice environments Flooded soils tend tothreatened by epidemics of locusts or armyworms New have the same properties and to eliminate soil pests The technology stressing efficiency has minimized insecticide spatial and temporal dominance of cultivated rice has cost benefited specialist pests

Seed treatment is normally inexpensive and can be Upland rice has no specialist arthropod species The economicallyjustified in -nanycases(66) Treating seed with monophagous pests on upland rice prefer the lowlands systemic insecticide (216) but not chlorinated hydrocarbons where rice evolved Rice in the uplands exists only because (247) is most effective in controlling seedling maggots of man and even though lowland rice predates upland riceFoliar sprays have to be repeated to obtain control and specialist species evolved Wheat maize and sorghum do often are washed off by frequent early season rains not have monophagous pests leading us to conclude that Granular insecticides are effective but required dosages are there is no evolutionary advantage for inssects to have too expensive to justify except under heavy infestation narrow host ranges in upland graminaceae(247) When directed against first-instar larvae banding Pests have adapted to the uplands not only by having granules in seed furrows (177) can be effective against white wider plant host ranges but also by having long life cyclesgrubs at low dosages (025 kg aii ha) (134) Seed treatments undergoing dormancy or being dispersive Each pest has a are not promising for white grub control (248) unique life history strategy suited to highly unpredictable

Seed treatment is also effective ainst ants and may be environments In agivenyear someinsect specics an better cheaper than increasing the seeding rate In Batangas exploit upland rice as a food source but no pest is a senious treating 50 kg seed ha was as effective as increasing the one every year on upland rice seeding rate to 100 kgha and using untreated seeds (Table The low yielding potential of upland rice means less 9) In Brazil seed treatments control termites and other soil quantity and lower quality of food for pests to exploit but insects including the lessercornstalk borer(66) Domiciano also less incentive for farmers to undertake control (58) however found seed treatments inconsistent in their The most effective control would be early and synchroshyeffect against high populations of lesser corn stalk borer nous planting of early-maturing upland rice varieties to

Baiting can be an inexpensive way of controlling ants prevent insect pest buildup Breeding for insect resistance seedling maggot flies mole crickets and field crickets Baits should initially focus on stem borers because they are verymade of locally available material and impregnated with difficult to control by other means Biocontrol efforts insecticide can be sparingly distributed taking adantage of should first stress conservation of natural enemies preshythe pests ability to disperse and encounter bait sites dators and pathogens rather than parasites Efficient

The new low-volume hand-held controlled-droplet chemical control includes seed treatment or baiting for seed sprayers may offer an advantage over high-volume knap- seedling and some root pests and using controlled dropletsack sprayes for controlling foliar pests Upland rice sprayers for spot treatments against foliar pests and seed canopies are more open than those in lowland rice and bugs when economic thresholds are reached better droplet penetration should result Controlled droplet applicators would be ideal for seedbugs and defoliators

CONCLUSION

Upland rice is attacked by a wider array of insect pests than is lowland rice mainly beca use of the addition of soil pests Generally population levels of lowland rice pests are lower in upland than lowland rice leading many people to conclude that insect pests are not important on upland rice

36 IRPS No 123 January 1987

REFERENCES CITED

1Abraham C C and K S R Mony 1978 Occurrence ofLeptocorisa acuta Fabr (Coreidae Hemiptera) as a pest of nutmeg trees J Bombay Nat ttistSoc 74553

2 Agyen-Sampong M 1975 HlierogrlphusdiaganensisKrauss (Orthop-tera Acrididae) a new pest of rice in Northern Ghana Ghana J Agric Sci 8249-253

3 Agyen-Samnpong M 1976 Varietal resistance and other inle-ratcd pest control components as factors of rice varietal improvement in West Africa Prcsented at the Second varietal improvement serinar 14-18 Sep 1976 Bouak Ivory Coast West Africa Rice Development Association Monrovia Liberia 18 p

4 Agyen-Sampong M 1977 Outbreak of Ihccrorrdnisorae in rice seedlings near Rokupr Sierra Leone lit Rice Res Newsl 2(6)16-17

5 Agyen-Sampong M 1980 Dirty panicles and rice yield reduction caused by hugs IntRice Res Newsl 5(l)11-12

6 Agyen-Sampong M 1982 The major pest problems of irrigated upland and mangrove swanip rice ecosystems ir the humid tropical Guinea savanna and Salel climate zones Pages 318-328 in Pro-ceedings training course cn concepts techniques and application of irtegrated pest management in rice inWest Africa 10-20 Jan 1982 Fendall Liberia West Africa Rice l)evelopioent Association Monrovia Liberia 505 p

7 Akibo-Betts V I and S A Raymundo 1978 Aphids as rice pests in Sierra Leone lit Rice Res Newsl 3(6)15-16

8 Akinsola F A 1975 Present status of different rice stemborers in parts of Nigeria iicL Entomo Newsl 328

9 Akinsola FA 1979 Ihe biology and ecology of rice stemborers in Nigeria Ph 1) thesis University ol lbadan Nigeria 217 p

10 Akinsola E A 1979 Guidelines of integrated management of insect pests of upland rice in West Africa Presented at the Seminar on integrated management of rice diseases and insect pests at Bobo-Dioulasso Upper Volta 17-22 Sep 1979 West Africa Rice Develop-ment Association Monrovia Liberia

II Akinsola IA 1984 Insect pests of upland rice in Africa Pages 301-305 in An overview of upland rice research Proceedings of the 1982 lIouak Iwsrv Coast upland rice workshop International Rice Research Institute 13O Box 933 Manila Philippines

12 Alam M S V TJohn and Kaung Zan 1985 Insect pests and diseases of rice in Africa Pages 67-82 in Rice improvement in Eastern Central and Southern Africa proceedings of the international rice workshop at Iusaka Zambia 9-19 April 1984

13 Alticri M A A van Schoonhovcn and J D Doll 1977 The ecological role of weeds in insect pest management systems a review illustiated with bean (Phaseohsvulgaris L)cropping systems PANS (Pest Artie News Summ) 23195-205

14 Anmante F 1967 The leaf-cutting ant Atia capiguaraa pest of pastures [in Portuguese] Biol6gico (Sao Paulo) 33113-120 (Rev Appl Entomol A56207)

15Andrade A C and D Puzzi 1951 Preliminary results of experi-merits on the control of Sculptcoris iastantus on sugarcane [in Portuguese] Bioulgico (Sao Paulo) 1744-49 (Rev Appl Entomol A41421422)

16 Appert J 1970 Maliarplia separaelha (rice stem borer) new observations of entomological problems of rice in Madagascar [in French] Agron Trop (Paris) 25329-334

17 Ariadeau Mand Z Harahap 1986 Relevant upland rice breeding objectives Pages 189-197 in Progress in upland rice research Proceedings of the 1985 Jakarta conference International Rice Research Institute 1O Box 933 Manila Philippines

18Asar S and Kisimoto 1975 Seasonal occurrence of the smaller brown planthopper in upland rice [in Japanese] Proc Kanto-Tosan Plant Prot Soc 2228

19 Bakker W 1974 Characterization and ecological aspects of rice yellow mottle virus in Kenya Meded Landbouwhogesh Wage-ningen 152 p

20 Balut F F 1970 White grubs attacking rice Oryza ativa L [in Portuguese] Biol6gico (So Paulo) 36321-322

21 Banerjee S N and L M Pramanik 1967 The lepidopterous stalk borers of rice and their life cycles in the tropics pages 103-124 in The major insect pests ofthe rice plant Proceedings ofa symposium at the International Rice Research Institute September 1964 The Johns Hopkins Press Baltimore Maryland USA

22 Barrion A T and J A Litsinger 1980 Ants a natural enemy of cliaiarac in dryland rice IntRice Res Newsl

5(4)22-23 23 Barrion A T and J A litsinger 1981 The spider fauna of

Philippine rice agroecosystem 1Dryland Philipp Entomnol 5139-166

24 Barrion A T and 1 A Iitsinger 1984 The spider fauna of Philippine rice agroccosystcms II Welland Philipp Eiitomol 611-37

25 Barrion A T1 and J A Litsinger 1984 lihanus (Diptera Tabanidae) eggs an alternative host of rice stei borer (SI3) egg parasite Telenoonis digns (Iliymenoptcra Scelionidae) lit Rice Res Newvsl 9(6)19

26 Barrion A T and J A Litsinger 1985 Identilication of rice leaffolders by wing markings Int Rice Res Newsl 10( 1)24

27 Barrion A T and J A Litsinger 1985 Chctiosspp (Coleoptera Carabidae) a leaffolder ILF)predator lit Rice Res Newsl 10(1)21

28 Bertels A 1970 Insect pests of rice and theircontrol [in Portuguese] Pelotas Brazil Instituto de Expcrimentacao Agropccuararias do Sul Circular 43 24 p

29 Bonzi S M 1982 Ctilo (flisilineus J de Joannis (Lepidoptera Pyralidae) stem borer of irrigated cereals in Upper Volta LioFrench] Agron Trop Riu Rizic Cult Vivrieres Trop 37207-209

30 Bourke T V T L Fenner J N LStihick G Baker EHassan D F OSullivan and C S li 1977 Insect pest survey for the year ending 30th June 1969 Papta New Guinea )epartment of Agrishyculture Stock and Fisheries Port Moresby 57 p

31 Bowden 1 1973 Migration of pests in the tropics Meded Rijksfac Landbouwwet Gent 38785-796

32 Breniere J 1973 The main rice parasites in West Africa and their control Pages 27-42 in Proceedings seminar on plant protection for the rice crop West Africa Rice Development Association Monrovia Liberia

33 Breniere J 1976 The principal insect pests of rice in West Africa and their control West Africa Rice Development Association Monrovia Liberia 52 p

34 Brown ES 1962 The African arinyworn Spoiolptera eXI)Ina (Walker) (Lepidoptera Noctuidae) a review of the literature Cornshymonwealth Institute of Entomology London UK57 p

35 Brown F B 1968 Upland rice in Latin America FAO-International Rice Commission 12th Session of the Working Party on Rice Production and Protection 2-14 Sept Peradeniya Sri lanka Food and Agriculture Organization Rome Italy 6 p

36 Buyckx F J 1 1962 Disease and insect pests of crops inCongo Rwanda and Burundi [iil French] Publications dc LInstilut National pour lestude Agrononiquedu Congo M Weissenbruck S A Brussels Belgium 708 p

37 Cariiio F 0 1981 Role of natural enentics inpopulation suppression and pest management of green rice lealhoppers Ph D thesis University ofthe Philippines at Los Bafitos PhilippinesLaguna 136 -

38 Carpente A J 1983 The development of rice cultivation and extension of vegetable productin in Zanzibar Final Report Vol II1 October 1983 Food and Agricultme Organization Ronte Italy

39 Carroll C P and S1 Risch 1983 Tropical annual cropping systems ant ecology Environ Manage 751-57

4(0 Cates R G 198 1 Host plant predictability and the feeding patterns of monophagous oligophagous and polyphagous insect herbivores Oecologia (Berl) 48319-326

IRPS No 123 January 1987 37

41 Cendcafia S M and F B Caora 1967 Insect pests of rice in the 62 Ene 1 C 1981 I)iiorldhol)ierusihilis Slater (licteroplera Philippines Pages 591-616 in The najor insect pests ofthe rice plant lygacidue Illissinac) a new rice pest front Nigeria lit Rice Res Proceedings of a symposium at the International Rice Research Newsl N3)15 Institute September 1964 The Johns Hopkins Press Baltimore 63 Fugenio T S 1965 rhe effect oi e-vCgetation on rice production Maryland USA 729 p and the possibility of overcoming it by leaf pruning MS thesis

42 Centro Ihternacional de Agricklittira Tropical 1984 Upland rice in University ol the Philippines at Los Baijos ILaguna Ihilippins 69 p Iatin Anerica Pages 93-119 hi An os ervies of uplaind rice research 04 Farrow R A 1975 The Alrican nmigratory locust ii is main Proceedings of the 1982 Boinakamp Ivory Coast upland rice workshop outbreak area of the middle Niger qlinlitatise studies of solitary International Rice Research Institute P0 Blox 933 Manila populations in relation to environmental factors Locusla No II Philippines 198 p

43 Chandra G 1979 Taxonomy ad hn riononlics ofthe imect parasites of 65 Fennal R G 1943 Food crop investigation Windward and rice Icalloppers and planthoppers in the Pihilippines and their Ieeward Islands First report Report onl work dunring the periodiiiportairce iii natural biol)gical cointrol Philipp Fntoltol 4119-139 April 1942-April 1943 Imperial College of Tropical Agriculture

44 Chang T T 1976 The originii colut ion enlIivation dissemination Trinidad 16 p (Rev of Appl Fintornol A3222-25) and diersificition of Asian and African rices -uphytica 25425-44 1 66 Ferreira F 1983 Integrated pest ritaageiret of upland rice [ill

45 Chatierjee P B 1979 Rice leaffolder attacks in India lit Rice Res Portuguese English summary] Pages 323-341 in Upland riceculttre Newsl 4(3)21 factors affecting production E Ferreira T Yamada and F-

46 Cheng Cbien-pan 19(4 Upland rice in Taiwan International Rice Malmolta eds Institutoda Potassae Foslato Institnto Internacional Cnmmission Wiorking Party oit Rice Production ard Protection I()th da Potassa Piracicaha Sao Paulo lraid Meeting 3-10 March 1964 Manila Philippines Food and Agricul- 67 Ferreira E arid F dl Silva Martins 1979 Occurrence of ture Organiaion Rome Italy 5 p Sogatohs orvziola (Muir) inupland rice in Goias Ilrail lit Rice

47 Chcrrett 1 M and 1) 1Peregrine 1976 A review of the status of Rcs Ncwssl 4(4)18 leaf-cutting ants and their control Ann Appl Biol 84124-128 68 Ferreira E and 1F da Silva Martiris 1984 Insect pestof rice ill

48 Chipeta F M 1974 Pests of rice in Malawi Pages 90-95 in Brazil and their control Jill Portugucsej FMIlRAtPA-CNPAF Proceedings Sith eastern Africa cereals research conference 10-15 Goiania Brmiil 67 p March 1974 DRB Mialda ed Ministry of Agriculture lilongwe 69 Ferreira FI F ca Silha Miartins S S Nero and 1 It P de Malawi Carvallio 1982 M tltiplc regression analysis todeterimine the effect of

49 Chu Yau-i 1979 The d atiaging period of the rice seedling fly insect populatiois ott upland rice yield [inIPortuguese Pesqui (Atlterigotiavxigu Stcin) (NItiscide Diptera) oil cori) at Fast Java Agropecu Bras 17671-675 Indonesia Plant Prot Bill 21403413 70 Ferreira F 1F da Silva Martins and F 1PZimmermann 1979

50 Conklin H C 1957 IHaninoo agriculture a report on aln integral Rice varietal resistance to tle lesser corn stalk borer [in Port uguese] systent of shi fti rig cult ivationi in the Pltilippincs Food and Agriculture Pesqui Agropccu Biras 14317-321 Organization Rome Italy 209 p 71 Fleming W F 1968 Biological control of lie hapanese beetle U S

51 Cramer H II 1967 Plant proection and world crop production Dep Agric RLs Sers [ech Bull 1383 78 p Pilaniensclhutz-Nachr 201-524 72 Food and Agriculture Organiation of the United Nations 1982

52 l)ani R C and Majunidar 1978 Prelininary observations on tie Country report surmimary Pages 27-37 in Report on the first incidence of root aphids on differeit rice cultnrcs Sci Cult 4488-89 FAOIUN)IPhail and regional training coiuirse in iniproved ctltural

53 Davidson R I 1979 lffects of root feeding on foliage yield Pages practices for upland rice 5-23 Oct 1981 Northern Region Agrculturd117-120 in T K Ci osb and R B Pottinger eds Proceedings of the Developririt Center Chiang Mai Food and Agricuilture Organiiashy2nd Australian conlerence on grassland invertebrate ecology Govt tion Rome Italy 37 n Printer Wellington New Zealand 73 Franco do Ariaral S 195 1 Residual action of insecticides in turfy

54 l)ean G 1W 1976 Rice insect pests inlaos Int Rice Res Newsl soil--the results of a field experiment [in Portuguese] Biol6gico (Sao 1(2)15 Paulo) 17209-214 (Rev Appl Entomol A41423)

55 I)ean G J W 1978 Insect pests of rice in Laos PANS 24(3)280-289 74 Freeman ) 1970 Report onithe Iban london School of -coroiics 56 )ingle H 1984 Behavior genes and lifc histories conplex adapta- Monographs on Social Anthropology No 41 Athlane Press

tions to uncertain environments Pages 169-194 in A new ecology London UK 317 p novel approaches to interactive systems P W Price C N Slobod- 75 Fuchs T W1 A Harding and 1W Smithlr 1979 Induction and chikoff and W S Gaud cdsl ohn Wiley New York USA termination oldiapause in the sugarcane borer Ann Flitonlol Soc

57 Dobelmann-JP and M Falais 1968 Upland rice cultivation ill tie Am 72271-274 North-western part of Madagascar in 1966-1967 [in French English 76 (hbriel B1) 1975 A review tf the major insect pests of some tpland summary] Trop Agron 2313-21 crops in tlc Philippines Pages 86-111 in Ad Htoc panel of experts on

58 Doniiciano N L 1979 Chemical control of tie lesser corn stalk pest disease arnd weed problems in some rainfed crops Bangkok (lolsonno~paloIgnowslhts) (Zeller 1848) (I-epid(ptera Ihycitid ae) Thailand Food aid Agriculture Organization Roire Italy

on rice in Ponta Gressa Parara 1977-1978 [in Portuguese] Pages 77 Garg D K and N K Shal 1983 White grub outbreak on rainfed 291-303 in Anais 1 Reuniao de tecnicos cmiirizicultura do estado de dryland rice in Uttar Pradeslh Int Rice Res Ncwsl 81(4)16-17 Sao Paulo-Campinas Cati Brazil 78 Garg I) K and IP Tandon 1982 Major insect pests of rice on hilly

59 Dunsmore 1 R 1970 Investigations on tlie varieties pests and tracts of Uttar Pradesh India lilt Rice Res Newsl 7(11)11-12diseases of upland rice (hill padi) in Sarawak Malaysia Int Rice 79 Garrity 1) P 1983 Upland rice research an international bibliograpy Comm Newsl 1929-35 1975-1982 Iternational Rice Research hnstitnte PO Box 933

60 Elias 1R1965 The new decline of rice seedlings is not caused by Manila Philippines 121 p parasites [in Portuguese] Biol6gico (Sao Paulo) 313-6 80 Ghauri M S K 1982 A new subspecies of Iinorh)rone bull

61 Empresa Brasiliera de Pesquisa Agropecuaria 1984 Upland rice itt cornlus Slater (Hemiptera Iygaeidae) front Papua New Guinea on Brazil Pages 121-134 it Av overview of upland rice research rice and carpet grass Ihll lEntoniol Res 72133-137 Proceedings ofthe 1982 Blouak Ivory Coast Upland rice workshop 81 Gonzales F 0 PArregoces R I ihernandef and OT Parada International Rice Research Institute PO Box 933 Manila 1983 Insect and imite pests of rice arid their control in Latin America Philippines [in Spanish] la Federacion Nacional de Arrocerts (Fedearroz)

Bogota Colombia 60 n

38 IRPS No 123 January 1987

82 Grist D H and R J A W Lever 1969 Pests of rice Longmans London UK520 p

83 Hale P R 1979 Two subterranean pests of upland rice in Papua New Guirnca IntRice Res Newsl 4(2)16-17

84 Hale P R and J T Hale 1975 Insect pests of rice in the New Guinea Islands Rice Entomol Newsl 34

85 Harris W V 1969 Termites as pests of crops and trees Common-wealth Institute of Entomology London UK 41 p

86 Hattori 1 1968 A method to distinguish between grass leaf roller (Ctaplha rocis medinalisGuenee) and the rice leaf roller (Susurtia exigua Butler) [in Japanese] Plant Prot (Japan) 23167-170

87 Henry I E and 1A Onsager 1984 Experimental control of the mormon cricket Iahrosshnph by NJsena locustae(Aficrospora Microsporidae) a protozoan parasite of grasshoppers (Ort Acri-didae) Entomophaga 27197-201

88 Hidaka 1 P Vungsilabulr and SKadkao 1974 Studies on ecology and control of the rice gall midge in Thailand Tech Bull 6 TropicalAgricultural Research Center Tokyo Japan 113 p

89 Hincklcy A 1) 1963 Ecology and control of rice planthoppers in Fiji Bull Entomrol Res 53467 481

90 Hirao J 1982 Ecology and chtInical control of the rice leaffolder Jpn Pestic Inf 4114-17

91 Ho DT 1984 Sten borer incifience in rice ecosystems in KenyaEast Africa Pages 307-320 in A overview of upland rice research Proceedings of the 1982 Bouac Ivory Coast upland rice workshop International Rice Research Institute PO Box 933 Manila Philippines

92 Ho )T J 1Njokah and J GKibuka 1983 Studies on rice sten borers in Kenya with emphasis on ifaliarpiaipiraellRag Insect Sci Appl 465-73

93 Ichihara I T Ito A NIatsuo and M Sawada 1971 Chemical control of the rice striped stem borer in upland rice grown in plastic mulch [iii Japanese] Proc Kanto-Tosan Plant Prot Soc 1882

94 IclhiharaIand A Matsuo 1969 Damage caused by rice striped stett borer to upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1686-87

95 Ichhara I atid A MIatsun11970 Conside ration oft lie extraordinary prevalence of rice striped stem borer in upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1778-79

96 Institut de Recherches Agrolnliqes Iropicale et des Cultures Vivrieres 1978 Insect resistance Pages 12-13 in Upland rice 1978 atnual report Bouak Ivory Coast 27 p

97 International Institute for Tropical Agriculture 1983 Aphids itt rice roots I ITA Res Briefs 4(1-3)4-5

98 International Institite fi Tropical Agriculture 1983 Hopperburn in rice caused by brown plathopper IITA Res Briefs 4(l-3)6-7

99 International Rice Research Institute 1975 Annual report for 1974 PO Box 933 Manila Philippines p 216

100 International Rice Restarch Institute 1976 Annual repiort for 1975 PO Box 933 Manila Philippines p 228

101 International Rice Research Institute 1979 Cropping systems program Component technology development and evaluation Page 404 i Annual report for 1978 11O Box 933 Manila Philippines

102 International Rice Research Institute Department of Entomology 1981) Insecticid evaluation 1979 Tables 73 74 PO Box 933Manila Philipes

103 International Rice Research Institute Department of Entonmology 1981 Insecticide evalomtion 1980 Table 81 POBox 933 Manila Philippines

104 International Rice Research Institute 1981 Report of a meeting of Asian rice-based cropping systems entomologists Eleventh cropping systems working group meeting 18-22 May 1981 PO Box 933 Manila Philippines p 132 133 144

105 International Rice Research Institute 1984 Cropping systems program Pages 368-373 in Annual report for 1983 PO Box 933 Manila Philippines

106 International Rice Research Institutc 1984 Upland rice in Asia Pages 45-68 in An overview of upland rice research Proceedings of the 1982 Bouakamp Ivory Coast upland rice workiiop loternational Rice Research Institute 11O Box 933 Manila Philippines

107 Irshad M 1977 Grasshoppers in rice in Pakistan lit Rice Res Newsl 2(3)8-9

108 Ito R and TOno 1964 Upland lice its culture and varieties in Japan lIternational Rice Cormmission loth Meeting of the Working Party on Rice Production and Protection 3-10 March 1964 Manila Philippines Food and Agriculture Organization Rome Italy 12 p

109 Jeffrey M I and V A Dyck 1983 Effect of moonlight on lightshytrap catches of brown planthopper lit Rice Res Newsl 8(6) 13

110 Jennings P R and T Alicia Pineda 1983 Management ofthe rice plnthoppers Sogatodsorivi(coIl in Latin America Presented at the meeting on stable plant resistance FAO and lIndocsian I)inectorate of Food Crops Protection Bali lndonesia 29 Novcniber-2 l)ccemshyber 9 p

IllJimenez Gomez J A 1979 Rice insect pests i lla ios ()riCntales lil Spanish] Pages 99-103 in Rice ciltivation Instituto (olotibiatto Agropecuario Compndio No 29

112 Jongeleen F JJ Soesanto and A Mockidjo 1979 Studies oti pathogens of two species of Scarabacidac froinIndonesia Z Angew Enlomnol 87412-417

113 Jouvenaz 1) PC S lofgren and W A Baiks 1981 Biological control of imported fire antsa review of cirrent knowledge Bull Entomol Soc Am 27203-208

114 Kainoh Y K Shiniiu S MarUani Y Taiaki 1980 lost finding behavior of tie rice bu lplotori+JchinvittjI)allas (liciniptera Corcidac) with special reference to diet pattern ol aggregation and feeding on the rice plant Appl Entomol Zool 15225-233

115 Kakkar K 0 P Sharma and P I Sharma 1977 Note on Ihteronyrchus liodresRedt (Coleopicra I)ynastidae) as asporadic pest of paddy and maie inHimachal Pradesh Corr Agric 156-57

116 Kalode M131 79 Major insect pest problems and their control in upland rice (Unpublishcd mineogriph CWI oJNaaiat lni Chandaratat (198 1)original not seen)

117 Kalshoven L i 98 I The pests of crops in I ldoniesia (translated by P A vatn der Laan) 1 T Ichtiar B3aru-vati Iloeve lakarta Indonesia 701 p

118 Kamal N Q 1981 Suppression of whitebacked planto ner Sogatella fircaifra (Horvath) aLId rice lealfolder (nhal -cis licdb lis (Guenee) populations by natural cnclmie hlI) tiesis Grcgorio Araneta University Foundation Victoncta Park Metto Manila Philippines 179 Ip

119 Karim A N M Reaul 1975 A species ofliealy bug Trioni i70s ceres new oii rice in the Philippines Rice Entoniol Newsl 247

120 Katanyukul Wand C Chandaratat 1981 Insect pestsof uplaid rice and their control Report firstFAOUNlFi Tliailand training course on improved cultural practices for tplad rice Chiang Mai Thailand 5-23 Oct 1981 Food and Agriculture Organiation Rome Italy 22 p

121 Kern F 1955 lhletwioa(I)uninemts) bhhit Biturm(Coleoptera) a new pest of sugarcane inVenezuela [inSpanish with Etnglish sunmary] Agron Trop (Maracay) 527-29 (Rev Appl Eitoniol A4572)

122 Khan K M and S Ghai 1974 White grubs ald their control in India Pesticides (1Bombay) 819-25

123 Kiritani K 1975 Integrated pest management of rice pests Pages 66-70 itt Proceedings platning workshop ott cooperative hIeld research in pest managementP SMotooka ed East-West Center Honololu Hawaii USA

124 Kisimoto R 1967 Ecology of insect vectors forecasting and chemical control Pages 243-255 in The virus diseases of tle rice plant Proceedings of a symposium atthe International Rice Research Institute April 1967 The Johns Hopkins Press altimore Maryshyland USA

125 Kisimoto R and V A Dyck 1976 Climate and rice insects Pages 367-391 in Climate and rice International Rice Research Institute PO Box 933 Manila Philippines

126 Kureha S S Kobayashi SAsari and T Ishihara 1977 Life history and control of the mole cricket Grrllotalpa africana Palisot de Beauvois [in Japanese English translation 7520 1RRI library] Proc Kanto-Tosan Plant Prot Soc 21192-197

127 Kushwaha K S 1981 The black beetle as a pest of rice in Haryana India Int Rice Res Newsl 6(4)14

128 Lawani S M 1982 A review of tile effects of various agronomic practices on cereal stem borers Trop Pest Manage 28266-276

129 Leuck D B 1966 Biology of the lesser cornstalk borer in South Georgia J Econ Entomol 59 797-801

130 lever R J A W 1955 Insect pests of the rice plant in Malaya Int Rice Comm Newsl 1612-21

131 Li Ching-Sing and Shin-Foon Chiu 1951 The study ofthe rice gall midge Iachydiplosis orvza Wood Mason [in Chinese with English summary] 1Taiwan Agric Res 21-13

132 Iitsinger J A 1984 Assesament of the need-based application for rice Proceedings TechnologyTransfer Workshop 15-16 March 1984 International Rice Research Institute 1PO Box 933 Manila Philippines 6 p

133 Litsinger J A 1984 Environmental diversity and insect pest abundance with reference to the Pacific Pages 285-307 in Proceedings sub-regional training course on methods of controlling diseases insects and other pests of plants in the South Pacific 4-20 Oct 1982 Vaini Kingdom of Tonga Nukualofa

134 Litsinger J A RF Apostol and M BObusan 1983 White grub Leucopholis irrorata (Coleoptera Scarabaeidae) pest status popula-tion dynamics and chemical conllol in arice-maize cropping pattern in the Philippines J Econ Entoinol 761133-1138

135 Litsinger J A C de la Cruz F Raymundo A T Barrion M D Lumaban J P Bandong M S Venugopal F Paragna and E Balete 1982 Insect pests and insecticide reponse of dry-seeded bunded rice Pages 179-193 in Report of workshop on cropping systems research in Asia International Rice Research Institute Los Bafos Philippine

136 Litsinger J A M 1) Luinaban J P Bandong A T Barrion W Katanyukul and N Panda 1978 Insect pest management in rainfcd lowland rice agroecosystms of tropical Asia International Rice Research Conference 17-21 April 1978 International Rice Research Institute Los Bafios Philippines

137 Litsinger J A and K Moody 1976 Integrated pest management in multiple cropping systems Pages 293-317 in R I Papendick P A Sanchez and G B Triplett eds Multiple cropping American Society of Agronomy Madison Wisconsin USA

138 Loevinsohn M E 1984 The ecology and control of rice pests in relation to the intensity and synchrony of cultivation Ph D thesis University of London 36) p

139 Loevinsohn M E J A Litsinger N Panda M A Austria R F Apostol C Andrion and J Bandong 1982 A basis for predicting rice insect pest abundance in cropping systems Paper presented at the 1982 International rice research conference on rice and cropping systems 19-23 April 1982 International Rice Research Institute Los Bafios Iaguna Philippines 25 p

140 Lopez A W 1930 Repot of the Entomology Department Pages 145-172 in Annual report of the Philippine Sugar Association 1929-30 Manila

141 Lopez AW 1931 Entomology Department Pages 227-266 in Annual report of the director or research 1930-31 Philippine Sugar Association Manila

142 Lopez A W 1932 Entomology Deparment Pages 252-279 in Annual report of the Philippine Sugar Association 1931-32 Manila

143 Lungibill P and G G Ainslee 1917 The lessercornstalk borer US Dep Agric Agric Res Serv Tech Bull 539 27 p

IRPS No 123 January 1987 39

144 McGuire J U Jr and BS Crandall 1967 Survey of insect pests and plant diseases ofselected food crops of Mexico Central America and Panama U SDepartment of Agriculture International Agriculshytural Development Service Agricultuial Researrh Service October 1967 11 p

145 McNeill Sand T R ESouthwood 1978 The role of nitrogen in the development of insectplant relations Pages 77-9 in lliochemical aspects of plant and animal coevolution J B Ilarbornu ed Academic Press London UK

146 Maebara H 1955 Spatial distribution of lastinstiarlanaeof purplish stem borer (Sesamia iqferens Walker) hibernating in an upland rice field Bull Kagoshima Univ Fac Agric 438-41

147 Mallick E H and N G Hajra 1977 Response of rice plant to defoliation treatments at different developmental stages Indian Agric 2151-56

148 Mansingh A 1971 Physiological classification of dormancies in insects Canl Entomol 103983-10)9

149 Martell C G 1974 Catalogue of plant feeding insects of Mexico [in Spanish] Fitofilo 27(69)1-176

150 Martinez 0 J 1982 Principal rice pests in Ilanos orientalis [in Spanish] P~ages 80-94 in Curso deact nalization de conocimientos en arroz Villaniencis Instituto (olonmbiano Agropecuario

151 Martins J F da Silva F Ferrcira A S Prabhu F 1 P Zimmermann 1980 Use of pesticides for the control of the main subterranean upland rice pests esqui Agropecu Bras 1553-u

152 Martins I F da Silva F Fe eira rod BdamSilveira Pinheiro I Q2 Damage simulation caused by defoliators of upland rice [in Portuguese with English summary] Pesqui Agropecu Bras 171113-1119

153 Martins 1F da Silva N(iyuen van Tan and 1Ida Silveira Pinheiro 1981 Stem borer resistance in upland rice associated with plant morphological characters [in Portuuce Iesqui Agropecu Bras 16187-192

154 Masaki S 1980 Sumnmer diapause Ann Rev Entomiol 251-25 155 Masuzawa T 11 Suwa and F Nakasoji 1983 l)ifkleences of

oviposition preferencc and survival rate of two skipper butterflies Parnara guttata and PIchlpidai iathiais (Iepidoptera HIesperiidac) on rice plant and cogon grass New Entomol 3247-56

156 Maurya D M and C P Vaish 1984 Upland rice in India Pages 69-92 in An overview oftpland rice research Proceedings ofthe 1982 Bouakamp Ivory Coast upland rice workshop International Rice Research Institute IO Box 933 Mnila Philippines

157 Menon M G R R N Katiyai S Kuar and S S Misra 1970 Termites attacking the paddy crop Sci Cult 36294-295

158 Miyahara Y and 1 Nishikubo 1967 Behavior of young nymphs of the red rice root aphid Iholahsipiun ri liabhdontinalisSasaki in relation to movement from the leaf to the roots of upland rice [in Japanese] Pioc Assoc Plant Prot Kyushu 1363-66

159 Mo ljoa Tjien 1952 White grub of upland rice (Holotrichiahelleri Brsk) [in Indonesian with English summary] Contrib Agric Res Stn Bogor No 13134 p

160 Mochida 0 and T Suryana 1979 The occurrence of the brown planthopper and ragged stunt on lowland and upland rice ill Central Java and logyakarta in tile late of January 1979 Entomnolgy Survey Report No 15 Central Research Institute of Agriculture (CRIA) Sukamandi Indonesia I February 1979 9 p

161Monet C 1979 Varietal resistance to upland rice pests [inI French] Pages 35-39 in Annual report 1977 Institut des Savanes Department des Cultures Vivrieres Bouak6 Ivory Coast

162 Monte 0 1942 A caterpillar ofrice-fields[in Portuguese] Biol6gico (Sao Paulo) 8161-163 (Rev Appl Entomol A31414)

163 Munroe I) D 1979 Pest management strategy for hill rice in four resettlement areas in Sarawak Malaysia Pages 89-102 i1 Guidelines for integrated control of rice insect pests Food and Agriculture Organization Plant Production and Protection Paper 14 Rome Italy 115 p

40 IRPS No 123 January 1987

164 Nakasuji F M Ishii and Jen-Zon Ito 1984 Rice skippers in Taiwan and their life histories ZZrC (Taipei) Tech Bull 82 10 p

165 Navas I) 1976 Fall armyworum in rice Pages 99-106 il Ecological animal control by habitat management Proc Tall Timbers Con-ference 28 Feb-I Mar 1974 Fall Timbers Res Stn Misc Publ 6 Tallahassee Horida USA

166 Nickel 1 1 1961 Biological control of rice stem borer A feasibility study Tcch Bull 2 International Rice Rcarch Institute PO Box 933 Manila Philippines IIl p

167 Nigan P M and R A Vernia 1985 Insect pests of uplaild rice in Uttar Pradesh Int Rice Res Newsl 10(6)22

168 Nilakhe S S 1985 -cological observations on spittle bugs with emphasis on their occirence in rice Pcsqtui Agropecu Bras 20407-414

169 Nixon G EJ 195 1 lie association of ats with aphids and coccids Commonwealth Iusti lcte of Entonology Iondon UK 36 p

170 Nurdin F an Ishak Manti 1985 Study oit seedling fly of upland rice Interlational Upland Rice Moitoring Tour Report 26 lebruary-2 MarLI Sukaranti West Sumatra

171 Oka 1 N 1979 Ciltural conttol of the brown planthopper Pages 357-369 in Brown planthopper threat to rice production in Asia International Rice Research Institutc 1p0 Box 933 Manila Philippines

172 Ohni M and A Vilnaii 1975 Rice insect pests of Italy biology of Sipha grcria (Kaltenbach) and its control [in Italian] Riso (Milan) 2459-71

173 Otanes F Q atnd 1 I Sison 194 1 Pests of rice Philipp J Agric 12211-259

174 Painter R I 1951 Insect icsistance in crop plants University Press of Kansits Lawrence Kansas USA 520 p

175 Pantua P C and 1A litsinger 1981 Arthropod abundance in continuous vs biannual rice cropping systeis Int Rice Res Ncwsl 6(2)20-21

176 Pathak M ) and V A l)yck 1975 Sttudics ott insect pests of upland rice Pages 186-191 in Major research in upland rice International Rice Research Institute PO Box 933 Ilanila Philippines

177 Patil B R B M IHasabe and 1) S Ajri 198 I Chemical control of white gritb lIditriihinhorraia Fabr intfesting groundntut paddy atnd sorghum Indian 1Entomol 43232-236

178 Pawar A 1) 1974 (ollection and identification of insect pests of rice rlerminal repirt lInternational Rice Research Institute PO Box 933 Manila Philippines 72 p (tnimco)

179 IerrajU A aid A S Reedy 1965 Observations ott diapausc itt tie steti borer ohrice Rice News Teller (lai-Mar)9-13

180 Pont A C 1972 A review of the Oriental species of theri ona Rondani (I)iptera Muscidac) of econtomic importance Pages 27-102 in Control of sorglium shoot fly M G lomtani and W R Young eds Oxford atd IH1 Ptthl Co New I)clhi India 324 p

181 Prasad S C 1PChand aid M 1PSingh 1983 Incidence of rice gall midge in the upland plateau region of Hihar Oryza 20235-236

182 Rai P S 198 0 Nisaga simpex damage to rice in the hill tracts of South India Int Rice kles News[ 5(I)12

183 Rathorc Y S 1969 Gonocplhahli dcprssunt Fabr as itpredator of (tilo zonwIlhis (Swinhoc) and other moths in field cage at Patnagar Uttar Pradesh Indian 1Entotnol 3127

184 Reed W 1983 Crop losses causcd by insect pists in the developing world Proceedings 10th Internatial Congress of Plant Protection lrighton UK 174-80

185 Reedy M V 1981 Ateactogasterinhens(Walker)a new root-grub pest of tipland paddy and inaize Curr Sci 501076

186 Reveno M A 1968 Catalogue of the insect pests of economic crops in Colombia [in Spaiish] Asociacion Latino Americana de Entomo-logia (AIAE) 129-33

187 Reyes G M 1957 Rice dwarf disease in the Philippines Food and Agriculture Organiiation Plant Prot Bull 617-19

188 Rice S E A A Grigarick and M 0 Way 1982 Effect of leaf and panicle feeding byarmyworm (Lepidoptera Noctuidae) larvae on rice grain yield Environ Entomol 7593-595

189 Risch S J 1) Andow and M A Altieri 1983 Agioccosystem diversity arnd pest control data tentative conclusions and nizw research dirctions Environ Entomol 12625-629

190 Rivera C T 1956 Three organic insecticides in thecontrol of insects affecting upland rice Philipp Agric 39465-472

191 Roffrey 1 1972 Locusta outbreaks in the Philippines Acrida 1177-188

192 Rojas R 1) 1941 The effect of pruning at varying ages upon the yield of Ramairio Phipp Agric 29851-860

193 Roonwal M i 1979 Ternite life and termite control in tropical South Asia Scientilic Pnbl odhpr India 139 p

194 Rosetto C1 F da Silva Martins N C Schmidt and L EAzzini 1978 Damage caused by pasturespittle bugs 4Dvoi)lavopictaandDJ schach) on rice lBragantia Nota 6 (Matcli)25-27

195 Rosztto C1- S S Neto I- Amantc I) Iink 1Garcia-Vieira D M de SoLza N V Bansa t and A M de Olivcira 1971 Rice pests of Brazil [in Portuguese] Pages 149-238 inrFA()-International Rice Commission 2nd meeting of the Rice Committee for the Americas 6-11 l)ecember 1971 Pelotas Rio Grande do Sul Brazil

196 Rothschild G H L 1969 Obseratiois on the armyworm Spodoptera matritia acronrctoiiles Gn (Lep Noctuidae) Bull Entomol Res 59143-160

197 Rothschild G 1- L 1974 Parasites of the armworm Spolopiera mauritiaacroncitoihes in Malaysia Fntornopliaga 19293-299

198 Sachan i N S K Gangwar and 1N Katujar 1980 Field crickets damaging cages in North Eastern IHill Region Indian J Entomol 42526-527

199 Sands 1) P A 1977 The biology and ecology of Lepocorisa (Hemiptera Alydidac) in Papua New Guinea Res Bull 18 Departshyment of Primary Industry Port Moresby Papua New Guinea 104 p

200 Sarawak 1968 titomology division Pages 97-106 in Report of the research branch of the Department of Altriculture 1966 Kuching (Rev AIppl Eintomol A56603)

201 Satpathy JM M S Das and K Naik 1977 Effect of multiple and mixed cropping on the incidence of scine important pestsI Entomol Res 178-85

202 Sauer IHF G 1939 Notes Ott Elasmpnlpualtsligniosellths Zeller (Lep Pyr) a serious pest ofcereals in Sao Paolo [in Portuguese] Arq Inst Biol 10199-206 (Rev Appl Entotnol A28378)

203 Satinders 1L A 1 SKing and C L Vargas S 1983 Crop pests of Central Anerica with t bibliography [in Spanish] Centro Agroshynotnico Tropical de Investigacion y Ensenanza Turrialba Costa Rica (CATIE) Tec Bol 9 90 p

204 Sauphanor B 1985 Some factors of upland rice tolerttce to stem borers in West Africa Inseu Sci Appl 6429434

205 Sen P atnd S Chakravorty 1976 Stentflics in Haritighata Nadia West Bengal India [it Rice Res Newsl 1(12) 15

206 Senapai B and 1M Satpathy 1983 The seedling fly Atherigona orrzae Malloch is a pest of rice in Orissa (India) J Entomol Res 777-79

207 Shhjahan M S Alatn M Ah nad and BN Islam 1983 A new scarabaeid beetle pest of dryland rice in Bangladesh Int Rice Res Ncwsl 8(I)16-17

208 Sharma G L S C Modgol and R C Gautam 1979 Effect of depth of submergence ott incidence of bacterial blight and white grub infestation in transplanted rice Int Rice Res Newsl 4(6)12-13

209 Sharma S 1985 Scarabaeid beetle outbreak itt upland rice Int Rice Res Newsl 1(1(l)24

210 Shiraki T 1917 The paddy borer Schoenobhits incertulas Walker [achaku Agric Exp Stn Formosa 256 p

211 Silva A de Brito and 11P Magalhaes 1981 Insect pests of rice [in Portuguese] Circular Tecnica No 22 EMBRAPA Centro de Pesquisa t ecuaria de Tropico Unido (CPAJU) Belem Brazil 14 p

212 Singh S A 1979 Emergence of rice leaf butterfly as a rice pest in Manipur India int Rice Res Newsl 4(4)16-17

213 SinghS A 1983 Changing trends of insect pestsof rice in Manipur Int Rice Res Newsl 8(f)12

214 Siwi S S and P H van Doesburg 1984 Lenovorisa Latreille in Indonesia (Heteroptera Coreidae Alydinae) Zool Meded (Leiden) 58117-129

215 Soejitno J 1977 Reaction of some upland rice varieties toseedling fly eltherikonaexigua Stein [in Indonesian] Siri Hana I Penyakit 630-35 Lembga I osat Fenclilian Pertanian Bogor Indonesia

216 Socnardi 1967 Insect pests of rice in Indonesia Pages675-683 in The major inse pests of thu rice plant Proceedings of a symposium at the Internatital Rice Rec Wi itt S pternber 1964 The Johns Hopkins Press Baltimore Maryland USA

217 Soto P E A T Petcz and I W Buddenhagen 1976 Survey of insect pests and diseases of rice in different ecological zones in Nigeria Rice Entomol Ncwsl 435-46

218 Soto 11E and Z Ziddiqi 1976 Screening for resistance to African rice insect pests Varietal improsetnent sCmi n r Bouak Ivory Coast 13-17 Sept 1976 West Africa Rice l)evelopment Association Monrovia Iiberia

219 Squire F A 1972 Entomological problems in Bolivia PANS (PestArtic News Sumr) 18249-268

220 Stapely 1 II 1978 The rice leaf roller in the Solomon Islands Int Rice Res Newsl 3(l)13

221 Suenaga H 1937 On aphids attacking the roots of rice Oyo-I)oubts Zasshi 9150-152 [in Japanese] (Rev Appl Entomol A25745)

222 Suenaga H 1952 On the seasonal abundance of three subterranean aphids on upland rice Bull Kyushu Agric Exp Sti 1249-262

223 Sutynso Yas and Dandi Soekaria 1964 Beberapa hana yang ttenperang tana man padi tanah kering [in Indonesian] Rapat kerja Padi Tanah Kering 6 sd 8 Juily 1964 Kaliurang 9 p

224 Tallamy I) W and R F Denno 1983 Alternative life history patterns in risky environments in example from lace bugs Pages 129-147 in Insect life history patterns R F l)cnio and If Dingle eds Spring-Verlag 225 p

225 Tamura 1 1939 Observations on some insect pests of field crops [in Japanese] 1 Plant Prot 26414-420 (Rev Appl EntomolA27-556)

226 Tamura and Suzuki 1964 Influences of the early-transplanting practice in rice culture upon development of the rice stem borer Chilo siql)ressalisWalker Bull Ilokuriku Natil Agric Exp Stn 761-94

227 Tan J P 1924 The rice root aphid Drowhi ihirsoia Philipp Agric 13277-288

228 Tanaka 1 1957 Taxonomy and distribution of some subterranean aphids injurious to upland rice in Japan with description of a new species [in Japanese English summary] Botyu-Kagaku 22168-176

229 Tanaka T 1961 The rice root aphids their ecology and control Spec [full College of Agriculture Utsunomiya University Japan F3 p

230 Tauber M IC A Tauber R Nechols and 1Obrycki 1983 Seasonal activity of parasitoids control by external internal and genetic factors Pages 87-108 itt I)iapause and life cycle strategies in insects V K Brown and I Ilodek eds )r W Junk The Hague Netherlands

231 Taylor W F 1972 Effects of artificial defoliation (simulating pest damage) on varieties of upland rice Exp Agric 879-83

232 feotia T P S and S Natud 1966 Biotoomics of the rice skipper Parnaraniathiis Fabricius (Lepidoptera Hesperiidac) Indian J Entomol 28181-186

233 Teran F 0 1971 Rice pests of Santa Cruz [in Spanish Ministerio de Agricultura Sita Cruz Bolivia Bull 45 8 p

234 Than Itun 1976 Population dynamics of the yellow rice borer Tryporza itcertulas (Walker) and its damage to the rice plant MS thesis University of the Philippines at Ios Bafios Laguna Philippines 92 p

235 Uichanco L B 1911 Pests Pages 55-74 it Rice Philipp Agric 11-200

IRPS No 123 January 1987 41

236 van der Laan P A 195 1 Life history of the rice seedling fly Contrib Gen Agric Res Stn Boor Indonesia 118 15 p

237 van der Laan 11A 1959 Correlation between rainfall in the dry season and the occurrence of the white rice borer (Scirpolhaga innotala WIk) in lava Entomol Fxp Appl 212-20

238 Van Driesche R G 1983 Meaning of percent parasitismin studies of insect parasitism Fnviron Entomol 121 I1- 122

239 Venugopal M S and A litsingcr 1984 Fffect ofcarbofuran on rice growth Prot Ecol 7313-317

240 Vietnam Directorate of Rural Affairs 1962 Annual work progress report on crop improvement program of rice sugarcane vegetable and field crops (lor the period from July 1961 to June 1962) (11) 299 p (Rev Appl Entomol A51526-527)

241 Wan M T K 1972 Observations on rice leaf- and planthoppers in Sarawak (Malaysian Borneo) Malay Agric 148308-335

242 West Africa Rice I)esclopnient Association 1978 Annual research report Vol 2 Statistics Monrovia Liberia

243 West Africa Rice Development Association 1979 Annual research report p47 Monrovia Liberia

244 Williams ) F and CS lofgren 1981 Fli I illy l-468 anew bait toxicant for control of the red imported lire at Fla Entomol 64472477

245 Williams I) 1 1970 The mealybugs OFlomoptera Coccoidea Iscudococcidac) of sugar-cane rice and sorghum Bull Entomol Res 0109-188

246 Williams D) 1 1982 Pulinaria icervi (Signoret) and its allies ott sugar cane and other grasses Btll Entoinol Res 72111-117

247 Wirjosuiardjo S 1976 Study ofchenmical cont rot of insect pests ott upland rice Imiu Pertanian (Jogjakarta) 1323-329

248 Wirjosuhardjo S and N Wirarta 1976 Chemical control of the white grub tIolorithiatlltriin upland rice in M uneng Irobolinggo Indonesia) [in Indonesian] Imut Pertanian (Jogjakarta) 1330-332

249 Yano K T Miyake and V F Eastop 1983 The biology and economic importance of rice aphids (Henuiptera Aphid idae) a review Bull Entomol Res 73539-566

250 Yelp D and P Rhodes 19(4 lhe success of drilled dryland rice iii the Fiji Islands International Rice Commission Working Party on Rice Production and Protection Manila Philippines 3-10 March1964 Food and Agriculture Organization Rome Italy 13 p

251 Young W R 1981 Fifty-five years of research on the sorghum shootfly Insect Sci Appi 23-9

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No 101The economics of hybrid rice production in (hina No 113 Boron toxicity in rice No 102 Rice ratooning No 114 Energy analysis rice production systems and rice research No 103 Growth and development of the (Jeep water rice plant No 115 Production risk and optimal fertilizer rates an application of the

No 104 Faridpur acomputer-assisted instruction nmodel forrainfed lowland random coefficient model No 116 Consumer demand for rice grain quality in Thailand Indonesia andrice

No 105 A readingand listening comprehcnsion test inEnglish for non native the Philippines speakers applying for tiraining at IRIRI No 117 Morphological changes in rice panicle development a review of

No 106 Rice grassy stunt virus 2 a new strain of rice grassy stunt in tire literature

Philippines No 118 IRRI-Korca collaborative project for the development of cold-No 107 Physical losses and quality deterioration in rice plstprodnCtion tolerant lines through anthei culture

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publishers No 120 Changes isinall-farm iLtireliing techii)hgy in Thailand and [he No 109 Classification of Philippine rainfall patterns Philippines No 110 Contributions of modern rice varieties to rutrition in Asia No 121 Landforms and modern rice varieties No Ill Changes in rice breeding in 10 Asian countries 1965-84 No 122 Yield stability and modern rice technology No 112 Design parameters affecting the performance of the IRRI-designed

axial-flow pump

Page 22: UPLANDRICE. INSECTl--1i M T -E!R.EOLOU t RTANCE ,AND …delphacid.s3.amazonaws.com › 6958.pdf · The significance of abiotic factors in upland rice insect ecology will be apparent

22 IRPS No 123 January 1987

Light trapped insects D~~nd (no trap-night )

St plht Suctined insects (nom 2

)

D-Voc suction 1520

1o

0 Lih trap 5 ~105 o5

00 Eggparositizotion()5040 NephOtgtha virneens

30 20

I010 A-Insects (notrop -night)

4 Nephatelh nigropccent i

2 o I vV^J - -- ^ VA____ f _

Light trapped insects (no trap-night) Suctioned insects (no rnE

10 -1208 ReCdO doRsol

6 15

4 10 052 iwxALAiamp 0

Light trapped insects (notrap-night) 2Suc oned nsects (nom )

8 CyrtorhmnusI6o nn]10 i0075

4 05

2 0 0

025

2 Seasonal abundance measured by light trap Roinfall (mmd) Moon phase and D-Vac suction sampler of green leafhoppersNephotettix virescens and N ngropicrus and I zigzag leafhopper Recilia dorsalis their egg 60

parasites and micid predator Cyrtorhinus livi-dipennis set against cropping pattern dailyrainfall and moon phase Tanauan Batangas1980-81

40

5 Lf Apr May Jun

1980 Jul Aug Sp Oct Nov I

-aMar

1981

not account for soil pests and was not based on field data in the absence of pests Carbofuran granules are phytotonicAkinsola (I1) cited mainly data from irrigated rice areas in to rice (239) The prevailing hypothesis is that carbofuranAfrica but estimated yield losses of upland rice between 15 prevents soil bacteria from consuming fertilizer The phytoshyand 30 from insects However losses in Thailand over a tonic effect is through greater availability of fertilizer to the4-yr period ranged from I to 13 averaging 5(106) plant Denitrifying bacteria are known to be more active inYield loss data however should be interpreted with aerobic than anaerobic soils therefore it may not be merecaution (184) The Philippine data were all derived from coincidence that the highest recorded yield losses (22-69)trials on farmers fields under farmers normal agronomic occurred in those trials with 12-30 kg basal Nha On themanagement practices except for Bukidnon and Capiz other hand contro with carbofuran granules may not be aswhere researchers used fertilizer Batangas farmers are good as a seed treatment against seed pests and perhapsatypical because ofhigh levels of fertilizer used The trials in yields would have been higher if a systemic seed treatmentBrazil were carried out at an experiment station but insecticide had been used conditions were similar to those of local farmers The 1979 and 1980 trials in Batangas had two full-

Carbofuran granules were used in every trial as a broad- protection treatments one using carbofuran granules andspectrum insecticide applied basally with the seed An the other using bendiocarb WP as a seed treatment Plotsobjection to using insecticides to measure yield losses isthat protected with bendiocarb yielded higher than those proshyinsecticidescandirectlyorindirectlystimulateplantgrowth tected with carbofuran granules The other trials in the

Table 4 Yield losses to insect pests and yield responses to insecticide in upland rice in the Philippines Brazil and Upper Volta

Yield (tha) Yield loss (5) Location Year Cultivar Protected

with Protected

without Check Total Vegetative Reproductive Ripening

Basal N (kgha) Reference

carbofuran carbofuran Protected No with carbofuran

carbofuran

Philippines Pili Camarines Sur Tanauan Batangas

Pangantucan Bukidnon

Dumarao t-apiz

BrazilGoias Minas Ge~ais

Upper Volta6

1975 1974 1974 1975 1975 1976 1977 1978

1979

1980

1979 1980 1979

1977 1978 1979

Bursiging puti Kinanda Dagge Kinanda Dagge Dagge Dagge Dagge UPL Ri5 Dagge UPL Ri5 Dagge UPL RiS UPL Ri5 UPL Ri5 UPL RiS

IAC47 IAC47 IAC47

14 33 11 36 35 31 30 27 37 26 40 30 40 42 46 35

19 21 09

--

-34 24 24 27 --

29 43 31 42 -

47 25

---

10 28 05 27 27 30 26 29 33 28 33 29 43 23 36 11

15 15 06

29 15 55 25 23

3 13 -7 11

-7 18

3 -7 45 22 69

24 28 35

--

21 -11 -20

4 -

-3

23 7

-2 -23 56

-

-

-__0

6 --

-

-23 14

5 0 8 2

22

-

--

0

-

--18 11

3 -10

4 10

7

--7 -

14 -

-1 6

10 -5

8 9 7

-

--

0 0 0 0 0 0 0 0 0 0 0

30 30 30

12 12 12

99 99

100 100 101 101 102 102 102 102 103 103 104 104 104

66 66 66

Farako-Ba

Mean

1977 1978

IRAT10 IRATIO

38 66

--

40 43

-5 35

21

-

10

--

--

--

- 242 243

Z

tJ

-w

24 IRPS No 123 January 1987

Philippines which had two treatments offering protection In the 9 Philippine trials where yield losses were measuredwith and without carbofuran granules relied on further by growth stage 10 of the yield loss (21) occurred at thecontrol with foliar spray and not on seed dressings The vegetative stage and 517 occurred at each of the reproshycarbofuran-protected plots had an 18yield los compared ductive and ripening stages Ibis result is similar to thewith only 10 with foliar sprayS This yield difference is pattern of yield loss in lowland rice measured by successiveprobably due to differences in protection against sown-seed treatments where the crop is unprotected at each growthpests (foliar spray do not control sown-seed pests) rather stage but proticted during the other growth stagesthan phytotonic effects became basal fertilizer wa used in However the insect complexes r sponsible for vegetativeonly 2 of the 10 comparisons stage yield losses in upland rice are entirely diftfrent than in

The recorded yield losses were not related to yield lowland rice The key vegetative stage pests in lowland ricepotential (Fig 3) One might expect higheryielding crops to arc aquatic whereas those in upland rice are sown--seedhave higher percentage of yield losses but this ws not tile (sown in soil) root and seedling pests This dift rence maycase One reason is that insect damage exacerbates plant explain why agricul torists have overlooked tile ilmportanccinjury from drought stress low yielding fields from d ought of upland rice insect pests Many reports of upland rice pestsstress would show disproportionate differences between in Asia ha Ve focused on major lowland species (46 108)insect-protected and -unprotected treatments Shoot and It isalso surprising that tei vegctative stage of upland rice root pests would be more responsible for losses from would record the highest yield losses becatuseseeral studiesdrought stress than sown-scd pests A second reason have shown that upland rice can readily recover from foliageconcerns seed pests which would have a greater impact on reinoval during that stage (117 152 165 188) Upland ricelow tillering low yielding varieties such as )aggc Low poscsses a hitgh obilitV to compensate for ca rly loss oftillering varieties nnot fill in the space created by removal foliage and will even be stinulated by foliage removal toof seeds oweer 1) R i 5 is high yielding because it tillersP1 produce a higher viel than plants ithout any foliageactively which can close the canopy to provide greater weed removed (63 192 231) [ie reason for this apparentcontrol discrepancy hovce is that yield loss at the vegetative

stage does not octUr frol leat arca loss but fron pestsYield loss () removing sown seed and feeding on devcloping tillers and roots -- damage that cannot be readily conpLtnsatCd for

70 - Measuring yield losses in upland rice cnvironrnents is further complicated by a report from Brazil that greater yield losses occurred in response to better insect control60 - =-09 as Insecticide seed treatments resuled in denser plant stands

(n=21) which in turn created a selectively more favorable environshyment for blast (69) This resuIt was parlticurla rlxexacerbated50 when cirbofuran granules were used in seed furrows Aside

ffrom a greater plant stand ca rbofuran iay have auigmented the N lecl and stimulated blast If so ca rbofuran should be40 - replaced with another insecticide or the seeding or fertilizerI rates decreased

Other methods to determine the impact of upland rice30 shy pests wvcrc to interview farners or to develop single speciesbull bull correlations of population levels and yield loss Feneira (66)2 NIM)st (75(1) Golas farmers reporteddid both in Brazil

20- insect losses in ufpland rice and 251 said those lossesS avcragcd 37(1 [his result is consistent with tie yield losses 0 0 measured by field trials in the same statc(Table4) For every

I0 4 of seedlings removC d by leaf-cutting ants a I 7yield loss is predicted Similarly Ferreira (66) found that ai

0 - average of 53 thrips per panicle before panicle initiation meant 3 times more unfillh d grains

-IO0 I 1 1 I I-101 0 2 3 4 5 6 7

Yield (tho) C)N [ROt IM FI IODS 3 Correlation between upland rice yield and yield lols caused by Upland rice insect pests may be controlled by culturalinsects measured in 21 trials in the Philippines Ivory Coast andBrazil genetic biological and chemical methods

IRPS No 123 January 1987 25

Cultural control dry season plowing will have little effect on them Plowing Crop husbandry to reduce upland insect pest populations when the soil is wet from the first monsoon rains will expose can be classified into practices effective in a single field and soil insects that tunnel close to the soil surface to predatory those effective only at the community level Cultural birds chickens dogs and even man (field crickets nole controls should come first in a pest management program crickets and white grubs a re delicacies in man tra ditional They have broad stable effects as pests have little pos- diets) sibility of overcoming them through selection of biotyps The greatest degree i1 inIscct control fron tillage comes (138) On the other hand some cultu ral controls work only from plowing soon after rice ha rvest when most soil insects with high labor or power inputs and connunity-wilt lie close to the soil surface anrd stern borers are in the rice methods require coordination of iany farniters stuble Whitegrubsare mature and tillage exposesthem to

Pantin time The luxury of being able to shift the time of predators Tirnelv control will prtec the crop following pla ntring upland rice is open onl hr la rniers in areas with a rice and lowr the populationIfor succeedinig years Plowingprolonged rainifall pa ttern lI copping intensity or underthe ice stulble helps it decompose killing sten borer rnechanied land preparatiot Gcnerally planlting as soon larvae inside as the rFiinv season begins will lower populatiois of 1110st [requent tillage destroys ait nests bit not termites insect pests Insect populationsarelo aftcra dlvseoll or Tillage and clean culture howes er rcliove food for winter fallow delayed plaiting lets them build tip on termites which then mtay attack a young rice crop Scarab alIternate hosts that gi vigorotsly with the first seasonial beetles also prefer to miposit in eceitly tilled fields Noshyratiris This first flush iof weds cros lItxuttriartlv with iceCult is dibbling faors the survival oftillace Clre Such rineralied N released during the dr season aid made stils i aels tnd insects thtt pttpate or lay eggs- in the soil

available to the plants with the rains It iliso igrtmitcally Zero tilla hio eerl conserves predators ofthe GIH (37) advanttgeous for the rice ciop to tap this trlLItlaI fertility nt it mcttoi lo lessen the impact o0 soil insects and grow vigorotsly to compete with wrus and to tolerate brotdcasting seed or planting in ftltrrows is prefc ired over pest infestation dihlirig in hills Crickets are large enotlgh to eat seedlings in

Larly plantngs terd to escape seedling maggolts Upland a hill atrd tillering normnally ca iriot fill in tlhe missing space rice in Bitangas is planted in Ma and ine itli the Iirst Oftcn dibbll rd is not well covered and iseasily found by rains Fairners are highly mtotiated to plant early tocscape foraging insets ioIdents and birds drlitght and typlotns ricar harct Arn Atrgtst planting Plan uit sii Incrcatsini sced density protects tile cropwould be evcely attacked by It riim orivzo At tti againrst seed and seelling pests especially ants particularly other Philippiie tipland sites Rea l Qteot antd (laelit if the crop will germinate and ermerge quickly M isainis Oriental seedlig mnaggots arnrige ric boe t se hr trftinll itI India rice intcrclopped witi cotton or farnes have a longer growing season and plnit 2 rin or pigeon pci had lower 6IIt and WBIlII poptrlations than mote after the ortset of the rainy season liaaiangas ftrmners rice alone (211) NfailC arid Urplald ir_ 2re coilltori in the plant ca rly becia use ofa shorter raiinfall period arnd the desi re Philippines he chatngC in pest status folii iniercropping is to lliarvet before the typhoor sea son peaks ltighly locatioll-specific (133) cmianarid the net elfc be either

Early plantings cornbind with early-iatiuring varictics nil bcnelficial ortdetriential (189) For example BIatangias provide higher crp tolerartce for white grubs Aestiatior farmers intercrip Itro in their ipland ricefields A liornshyterminates with the first rains but 5-6 moare ineeded for the nrii iotd-Ahis i (I iiiacius) which norlially feeds resting larvae to develop into beetles emerge and lay eggs on taro ill feed or miarbv rice plants Intercropping and for larvae to reach the danriigii g ird-instar stage probably would not afft0 seed pests A crluanion crop (134 In lataigas this normually iseiough tiite for harvest planted ith rice and haing gleater tolerarnce for root

It Brazil howecver delayed planting is a suggested dariage or being toxic nayact isi trapcrop to root-feeditig cultural control for the lesser cornstalk borer and insect pests Acial-fei i ig pests voultId be most affected by pests favored l drought (66 195) Cropping internsity in intrer ropping Shoot aphids leaf beetles sten borers and Brail is low and delayed plraing places the rice crop in the hopplers would be likely ca ntidates to note effcts On the niost stable rainfall period reducing likelihood old rought othlt halnd the corlipalior crop stch~ sitniaie Iiiv be more

In rilniy arca s rinmirsare roi atl to plant other crops benehited than rice from initecropping ( 137) before rice and althNgh it woultl be desirabh to avoid 11echi Weeuling luring the first month after clop infestations by planting rice cuatl lariners lack the labor cstablishent will force pcts such is anyvworris which and time to do so ol a low priority crp prefer grassy cetds ontor tle vorling rice Soil iampsecs will

77age Uplild rice Stils ae light textured and Call be mvnye to the rice crop Cleani citrltirc also lay lore territes plowed duiring the d rv seasor notl only to grin time for to attack I u cetincti inrice ceds i ay he le optinal planting with lie fitst Inilsoon riiains but also to desiccate solition 13) soil insects arid weeds such is ntsedge that foim rhizomes Irtlizr Itneatsing N Iciiliier oin hlwland rice has Htowever becta use most soil insects lie below die plow layer fAtwvored Irigher populains of pliithoppeiv Imlhoppers

26 IRPS No 123 January 1987

and leaffolde rs However because of the need to temper fertilizer use on upland rice o prevent blast fIrtilizer rates will be lower than is comonia r for lowlad rice causing relatixely less hopper and IaIfTfoide r population buildup However inupland rice areas without a history of blast such Is Batangas farmers apply high rates of N and conscqtrently the crop has high infcstation levels of WBPI]and RIF The fhrmrcrs leec insect buildup by splitting the fertili11r in0to two to three applicatiois bit do not applV fertilizer at the vegetati estage because this prodrces a tall crop which illeasily lodge during typhoons Greater crop fertility leads to greater insect survival (better nutrition) larger insects (to cat no re foliage) arid greater fec titd t (to lav nmre eggs) (128 145 171) Not all insect species respondequally hovr Increasing rates (if N l anrid K increased Dialraa s-acchralis and (iaeiurema flea beetle but reduced thrips (66) Zn rcdLuced deadheart density from the lesser cornIstalk borer

Fnldingi Although most Upland rice areas do nIot hawxcaccess to wa ter floodiig the fields is rRrrnriierdcd in areas of Brazil or root bug and white gib control ( 195)

Crop rotatioti Tillage before planting a crop after rc harvest will control stern boreis and other pests renain n the stubble arid will unearth soil pests Planting a non-gratriiaceous crop after rice is recommncrded for termite control (66) Crop rotation is less nttdcd in upland rice culture than in lowlands because atcrop-free period is assured even after harvest of crops following rice

t1ulchilt iryl plastic irli irig designed for weedcoitrol ii upland rice inJapan resulted in ntr highanri liy infestation of (ihilo sulytresvais (93 9-1 95) NIulched Upland rice grew more luxuriantlv and attracted more (Avipositing mothis than u lched rice Also the rice plant and stein borers imatoured 10rC quickly and a partialgeneratiori af vouing ista iNoverwintcred in the stubbleLotili--malitrt varieti Quickly maturing crops reduce the tmbCr of pCst generations that Calt build up Rapid crop establishment olan ca rl v-natu ring vaitvty if)pa 11icLa r will lessen white gib and stem borer damiage White grub larvae intle last iistar are highly destIruCtive so aii early-iiatri ng variety will escape serious root loss Stem borers build 1ip slowlyx in rice to ecormic abundant in a late-Maturing crop anrd caiuse whi tlieads Numibers of somespecies increase exponeitially with each succeeding genera-ti so a xarietyexhat atures I trowh earier thian another will hac less insect damage Early-naturing vaiicties however will not lessen seed pest daiage either ii the soil or oina standing crop

Strwhoi hAIJttam Inrscts tcd irig on thlie aerial portions of rice plats disperse from carly to late plantedfields AS iiiibCirs increase exponentially with eachinsect

gtscnrattm e eiglbrig f rmc~s1(uil upp wh rc generation pests build tIP where neighboring fihrTC e9r1sstagger their plantings II Batangas farmers plant uplandrice within I iro use 120-d s-arieties and plant at the start of the rainy season Ini Misarnis Oriental laricrs stagger their

plantings up to 3 mo apart and use a set of atieties that mature in 4-7 mo Insect pest danage particularly from stem borers is greater in Misarmis Oriental Also seedling maggots damage rice there because rainfall favors frequent planting of maic In Batangas rice escapes seedling maggots bCca usC of distinct drs season and synchron0us early planting with the onset of ains

Therefore combinations of cultural practices ea rlyplanting synchronous planing crop rotation and earlyshymaturing vaneties protect the rice crop against most insect pests

Ilmant resistance Beca tse upland rice has few specialized insect pests and many insect types attack the crop worldwide regional breeding and strong internatiornal cooperation are required

Sown-sced pests arc economically controlled by insectishycide seed treatment and should the relore receive lowxpriori ty inbreeding objectives

Root pests are normally cost ly to control with insecticides but finding resista t sonres mayvbe di fficult Tanaka (229) for exa -ple fi led to find varieties resistant to root aphids Selecting for large root bioniass perhaps should be the strategy for pests sLuch as Whlite grubs that renliove roots Fainter(174) howcvcr cites varietiesof crops tolerant ofor resistant t(i root pests Some sorghun varieties are high in cyanide so rice sorghun might incorporate broadshyspcctruri chemical resistance

Seedling vigor and drought tolerance are high prioritybreeding objectives so tupland rices should have large r root systernsand be able to tolerate ligherlevels of root loss Sonie root loss inolder plants is beneficial if it stimulates new root developnrut and therefore crihances intake of soil nutrients (53) The rice seedling inaggot rtiLotia spp may be

controlled thirough resistant aittics A hi rgc breeding effort on sorghniu has had fair success inideveloping arreties esrstait to Asoccata Rodani (251) A field trial comparing 10 upland ricc varieties showed differences aniong varieties va ring from II to 4(- damaged tilleis (215) Hovver resistarce to 1Iill-0r gtia species may rot cross ox-rti otier

Shiraki (210) reported that upland i-ice stens wcrc harder than lows-land rice stemsperhaps becaIusc (f a higher silicoricontent and thus were more resistant to yellow stern borer hc tortia Iow tillering (f uplaid iice xarneties prodtres a

higher percentage of infested tillers than would occor in higher tillering lowland iice under a siiilar egg density The corollary therefore is to select for high tilleriig upland rices

Stemii bor amcstheh main targes dobrecders inAfrica (571 204 )but nticl oifth e sc reen irig is dotie unde r lo wland co d t osrm s wih A h r~ o l pe i s n acorditions (11 96 218) As with lthrinrnwspecies mayhave to be dealt with independently however initial results show cross resistance between African and Asian species(92)

In Brazil screening Ior insect resistance focttses oil lou I pest problems Varieties tolerant of lasmolwalpus ligi-sellus and resistant to )iatraea saccharalis have been identified (70 153) Current work also emphasies 7iraca limativetntris and Thmialspis (=Iosis) flavouicta (66) These insect pests Ilt onlV citise IIigh yield losses tihey also ire difficult to control chemically ofr by other ilealls

ManV upland rice vaieties aic early maturing and are resistant to pests such as gall midge (I Xl ) Early mIturity is often a highly desirable trait to esuilpe pest buildup but it is not true resista nce bca usc ilthose va reties are plinted late they will be itsdamaged as ain early planted susc ible variety

P~erhaps tile cosiiiopolitall of all upland rice pestsimost are the sced pests (ripenling stagc) which pose a particular problem beta use scuds re ised IsIO for hitnit4ans Ayltoxic substatice introd tcld geticticllyilmay also be toxic to nal iniless it caln he placed old in thfe IdhuffSelectmi fo hairiness or lotng wns kill mIlct sthilrohleiiis dutilig threshing pa rtictfarl maitial threshinfl [hireshers ill complain of itcfimin atid itteascd (fust levels

Attempts itIRRI to fitid tcistatncc to the rice bug plorisa have ilotbeCnl prmitisilg lIIetIattonlids iorc

through the lenilna or palca hillhairs sedstVIor seeds wilh awns that deter bugs promote complaints from threshers Seed bugs liptocorisa and Stcnocoris enter the seed through the opening between tile lemna and palC but varieties with narrow openings are diifficult to mill

Breeding priorities should concentrate ott inotCI gruLtLIps that are most difficult to c nttrol by other meains Steil) horers appear to be the most widespread group that sigiiificantly dalag s rice aind is hard to manage by other Ilnca us

Biological control Very fe stutdic of mitatt~l eneties of rice pests in upland envirollnletits wee in tile wefouiid literature Therefore have not compiled a list of species Ve present a list of natural enemies iOtid in upland ricefields tuic latialln Batangas for some conitno iOldiar pest species ([itble 5)

Muich ofl thc lollowinle itfor ittiol cotlliCs rol Stutdies in the lowlatnds or roi crops otithcr thlin tice

Qu(ailtr o 11ll t11t( Nat trrtf lii hcoeits ight Ile used agaitist ile social insects tilt and termites Sollie predator specice live ill theirf nests n ilnic theirnllt ip-pearanIlCe aMRidbehavior IJtrouMdCing a mote aggressise ant species to displace pest ant codonies is a possibility (169) Patihogens as potential natural encnies have reccived little atteition (I13) Natural ncnuics asa group however hac not been toIl iporta it in Iegulating aintshown be and ternite numbers

The nocturnal behavior aid subteitmien habitat of lole crickets and field crickets protect them from many ntllral enemies There are only limited records of natural cnelics for these groups of pests some pathogens (87) but mainly

IRPS No 123 JattuIny I 271987

parasites - scelionids on eggsand sphecids on ilymphs and adults Sphecid wasps Larra carutmaria (SImitI hI L huzwuensis Rohwer and lsaltiuhta Willi ams speciali in mole crickets pa ralyzing them nd dragging the ninto thcir nests as food lot their voting ()tier sphCcid species Lins aurulenoa (Iabricius) lot maiac (Ashmead ) 1 lhshylssellaus (Smith) and Ml lal)oriosu (Snithf specialie in field clickets

White grUbsO the other hand hae Icti scriuns enuigh to have been extensively studied for cotttroi 1W nati al enemies( 112 140 141) Scoliid parasites C(ajsoMcispp have beetn intrtduced itto tilePhilippites for control of Ielxl ]udisiro la The rcsearchcts w Intttrdced themle during ill era claimed that the white gttionthreak as controlled (142) howevcr it isnormal for pest epidemics to subside and there was no diect evidence to indictte that tfe parasite was responsible A tmore promisitg titctllod might he to identify iso ltcttass prt~f tcartd apply hiactelaln fingal pathogens to tile soil Ihe lalattese beetle in North America has hcci coitrolled hv tite conmnleciall aailable hactcrial prcpa ration itacihlpu pav ofr milksdiscas (71) Spote prtyefott the soil itloted ilrtier reitiain actic fr Veas

Root aphid sardtnctly bugs readilslic tilib to predatots and parlsites once their tetndine ants are controlled The larvac and adtilts of a coccinellid beetle 1Scynms sp prey on llriadlouais in slash and burn upland rice Baits treated with insecticide may be used to control tending ants by killing their ourig (244)

Root-tkcditig blugs false wireoris wireorms and root wccils aic r ofsuch little ifilpoltatice that thenomttll role of liatural Cticniics hias tiot beei asscssed A possibility hoevrl tiglt he tit ItaIIitic niaCIMtodcs lintt1g11en Itese soil-itlhlfbitine mCtitodcs W5uhtfhave to be imiass Produiced

large miltibcr of niatltrll cnemics ircknown ft CltIoriS IC IpblCeii is thtt clttvortnlls ofonlie the crop soott alter land ptel)tritiOti a1tithe bcgit iniig of the raittv

Citsoln hfien tiatllral leniiiy populations aric lo Colltloil WOtuld iii ole mass prodtticing andirclcaslng key spccies

le egg stage is the fpirt (olthe Cedlitgt Ialot life cycle tI hich to f)ctis ftor tittlril elctit cottntrol hplraites o1 pleditors ttsit elleggs highly expoIsYCl I Sgg pailsites or predators stolld probahly e ittass andtbe produced released Littlh i toruialioll is a ailable ott what species to trV o ho to Iniss podLice thiCIi A etiloplhid wasp Icmihlaol sp effctiicly regushy

lates the leatf lcloalapolm2a spioanoiiitr heat andc rice antid should be collsctvcd

leaf beetles whCllose e am] eggs ate laid in the grutindhirO ould be motC difficutlt to control by niatulral enelV Mnll ipuhltiOn But species sUici IS c_gsthispid beetles ith

and larVle oilleascs ate vtilcratble to iparaites and pledators

Natural enemisc of tlirips otl rice are little known

28 IRPS No 123 January 1987

Table S Parasites recorded from upland rice insect pests in Tanauan Batangas Philippines 1977-84

Order Family Species Host stage

I lymttenoptera Braconidae Leaffolders Cnaphalocrocis Marasinia spy

Cardiochilesphilipplinensis Ashmead a Larva

Cotesia angustillasis(Gahan)Cotesia or cypris (Nixon)

Larva Larva

Cotesia or taeniaticornis(Nixon) Larva Cilonusinotakatae Munakata Chelonits SppAlacrocetrts philippinensis (Ashtmead)

Egg to larva Egg to larva Larva

Iclineumonidac Orgilts spp Isclutojppa hireator(Fabricius)

Larva Larva

Itoplectis naraigav(Ashmcad) Larva Tenelhtcha philippiensis (Ashmead) Larva Tenelchha stangli (Ashmead) a

irichomma cnaphalocrosis Uchida Larva Larva

Bethylidac Elasmidac

Goniozms or trianguli r Kieffer lasmtusalhopictus Crawford

Larva Larva

Diptera

Encyrtidac Chalcididac

Tachinidac

Elasmtus spp Copidosomopsisnacohiae (Fidy) Brach nteria evcarinata Gahan Biachyieria lasus (Walker) Ztgohothria ciliata (Wulp)

Larva Egg to larva Pupa Pupa Larva

Argyrophylx nigrotihialis(Blaranov) Larva

l-ymenoptera Braconidac Yellow stern borer Scirpopiaga incertulas

Braco chinensisSzeplegeti Larva CIthelotis spp Chelonus niunakatae Munakata Cotesi flaripes Cameron Stenoracon nicerilli (Bingham) a

Larva Egg to larva Larva Larva

Ichneumonidac Tropobracon schoenobii (Viereck) Amatroinorpiaaccepta metathoracica(Ashmiead)Frihorus sinicus (I lolmgren)

Larva Larva Larva

Ischnojoppaluteator (Fabricius) Larva Isotima nr dammeratiti Rohwer Larva Tenelucha philippinensis (Ashmead) a

Larva Tenehicha stangli (Ashmcad)Trichomma enaphalocrosis Uchida

Larva Lara

Chalcididac Eulophidac Trichogranmmatidae

Xanthopimnpla stemmator (Thunberg) Charops hrachYpterumn Gupta and Malteswary Brachtm eria sppTetrasticttus schoenohii Ferriere Trichorigranm chihunis Ishi

Larva to pupa Larva Pupa Egg Egg

Diptera

Scelionidac Pteromalidae Tachinidae

Tt ichograintnta aponiicuitt (Ashniead) Telenomits rowani (Gahan) a

Trichoinalopsis apattelctena (Crawford) Jeirhaea spp

Egg Fgg Egg pupa Larva

Z4gobothria ciliata (Wulp) Larva

Ilymenoptera lecneumonirlac Gol fringed borer (hilo auricilius

Trichomm enoptalocrosisUchida Larva

Ieioenoptera Braconidae

Aantholimlh steiniator (Thtinberg) a

Pink stet Ioret Sesamia inferens StoInacoil nicrilhi (Bingh am)

Loarva to pupa

larva

Diptera Ilyinenoptera

Armvtnyorms Mythimna separata Spodoptera mauritia acronyetoidesTachinidae Zvgohothriaatropiora( Robinea i -Desvoid y)Eulophidae It-phctrtiscitapadoe (Ashimead)

Larva Larva

(halcididae Braconidac

Brachpin ria spip (oesia spp

Larva larva

IHyntenoptera I)iptera

Rice skipper Ielop ida (halcididae Tachinidae

math ias aidgreen iortedci ciliarNIelan itis eda Brat lhincria sp iii matuqitatt Cameron Irgyrolhirlaxi trotlhiais(marumtv)a1

ismtne Larva Larva

Diptera Ilymenoptera

Tachnidae lelineutionidae

fuown se tilo per Mois frugalisIrgtrophrlax 1igrothiahiv 3 a rat ov) anthopimpa pmnctata (Iabricius)

Larva Larva to pupa

aMost dominant

IRPS No 123 January 1987 29

Normally predatory thrips prey on herbivorous thrips strepsiptera on both Spiders dominate tilenymphaladultNatural enemies of stem bugs --Scolinoplara and Tibraa predators Spittle bugs move to rice from pasture grasses - are in order of importance egg par~isites and predators and biocontrol would be successful only ifcarried out before fungal pathogens and nymphal adult parasites Egg tiley immigrate into ricefields parasites could either be introduced fron other areas or Biological control tactics for hemipterous seed bugs mass produced and released ould follow those outlined for stein bugs

Armyworms are normally lwld in hcck by the activities Coml)arison of environnewtns Natural enemies are of egg and larval parasites When these natural enemies fail perhaps faced with even greaterconstraints than insect pestsusually because of drought armyworns become epidemic to survive ind reproduce in upland environments TheyIt may be worthwhile therefore to release parasites during have adapted ani survived by having wide host ranges the the rainy months following a drought Virus diseases of ability to aestivate or hibernate over unfavorable seasonslarvae would beanotheravnue to exploreforarnyworms and the ability to disperse or by other mechanisms

Parasites have ben traditionallyconsidered for biolegical Comparing the most dominant and therefore the most control attempts against stein borers usually by rekasing adapted species of natural enemies collected in various exotic species( 166) This approach has not met with success environments in the Philippines we can understand how and new avenues should be explored The role of predators different rice environments are arid pathogens is little inderstood and deserves greater Anagqrms olptahiisa mymarid planthopper egg parasiteattention Egg parasitism rates are normally reasonably is cquallyadapted to all Philippine rice environments (Tablehigh but should be supplemented with effective egg 6) It is selective for planthoppers and even parasitizes eggspredators Orthopterans are potential egg predators of the n-iize planthopper Peregrinitsmaidis itt upland areas Metioche vittaticollis (Sta ) Anavipha nifiwnnis (Saus- Alavohis also parasitizes maize planthopper eggs but is sure) and Conocephahs hongipnni (de i-laan) feed on most prevalent on BPH in the irrigated wetlands (onashystem borer eggs etioche and Anaxil)ha specialize in eggs tocerus spp like A optalhilis on BPH and WBPI Iattacks with no hair such as Chilo spp The omnivorous (ono- the eggs of GLH in ell rie environments It is also found cephahsspecializes in eggs cov red with hairs it is a proven parasitizing the eggs ef the white rice licahopper (oimna egg predator of Scirlouphaga spp but has the discouraging spetra Oirosita naias a trichogrammatid egg parasitehabit of eating rice guin (65) attacks BIPH and WBPH in the wetlands and has no other

Grasshoppers have egg (scelionid) and nymphalad ult host A related species 0 aesopi specializes in G1-H in (nemestrinid tachinid and sarcophagid) v irasites Among wetland environments Dryinid strepsipt-an and pipunshypathogens protozoans have been recorded most frequently culid nymphal and adult parasites of planthoppers and Control of locust species by natural enemies could con- leafhoppers also specialize either in hopper species or centrate on their habitual breeding areas environments Among the dryinids Pseuhdgonatoipsflavi-

Leaffolders have rich complexes of natural enemies some feimur prefers the lowlands and BPH while P nuch has no adapted to upland rice (22 27) ranging from egg predators environmental prefirences but is adapted only to WBPH (gryllids coccinellids) to larval parasites (braconids ichncu- The main d ryinid species on GLH aptogonatop)usspp is monids) to larval prcdators (ants and carabids) to larval most dominant in rainfed lowlands and also attacks other pathogeiis (vira and fungal diseases) Efforts to augment leafhoppers within Amnrasca Cicadulinaand Bachaltuathese natural enemies should focus on the egg predators Strepsiptera are more dominant on plarithoppers than (rearing releasing)and viral or fungal pathogens (culturing lealhoppers The Elenchus species in the Philippines attack spraying) all species of planthoppers within the genera Sogawilla

Parasitization of the large conspicuous larvae of Sogatodels NilaparsataHarualia and Opiconsiva Elenshyskippers green horned caterpillars and other polyphagous cusrvasuntaisuiis adapted to the major rice environments Lepidoptera by tachinids and chalcids is normally low ELg but prefers BPH in rainfed arid WBPIH in wetland environshyparasites and predators are perhaps the key to their control ments Ilalictoph(utsmtnroeiand HVpectrus attack G LH

Gall midges are parasitized by pterormalids and platy- and COfimravcctra respectivelygasterids that aestivate within their larval host between Pipunculids are mainly parasitic oii lealhoppers Each seasons Shoot aphids are preyed upon by coccinellids species has a unique environmental preference Tomosshysyrphids and chrysopids varireIa oraetoraprefers Ne7 otetltix nigropictrs and

Planthoppers arid leafhoppers have egg parasites and Pilunwulhtsjavanensisattacks Dehoceplhahls spp which predators as well as nymphialad nlt parasitesand predators are more abundant in upland than irrigated lowland their rich complex of natural enemies includes fungal environments T suhvirescensattacks Nephotettix virescens pathogens The principal egg parasites are myrnarids and and prefers the lowlands Pilpuiulus nttillattls is an trichogramniatids Egg predators are norinally mirid plant upland rice parasite arid attacks only Nephotetix speciesbugs that suck out the yolk Nymphal ad nIt parasites are More than 17 recorded species of leaffolder parasites in dryinids on planthoppers pipunculids on lealhoppers and the Philippines attack species of C(naplwocrocis and

30 IRPS No 123 January 1987

Table 6 Dominant natural enemies of rice pests in upland rainfed lowland and irrigated lowland rice environments of the Philippines

Natural enemy

Host stage Species

Brown planthop-Parasite Egg

MymaridaeAnagrus optabilis(Perkins) Anagrusflaveolus Waterhouse

TrichogrammatidacOligosita naiasGirault

NymphAdult Dryinidac

Pseudogotatopuslaiifemttr Esaki et Hashimoto Elenchidae

Elenchus yasurnatsui Kifune and Hirashima

Environment

Upand Rainfed lowland Irrigated lowland

Ailaparata lugens

x x X x

x

x x

x X X Whitebacked planthoppers Sogatella furcifera and Sogatodes pusanusEgg

MymaridaeAnagrus optabilis(Perkins)

TrichogrammatidacOl~gositanaias Girault

NymphAdult Dryinidae

Pseudogonatopusnudus Perkins Elenchidae

Elenchus yasutatsui Kifune et Hirashima

Parasite Egg

MymaridaeGonatocerus spp

TrichogrammatidaeOligositaaesopi Girault

NymphAdult Pipunculidac

Tomosvaryella subviresccns Loew Tomosvaryella oryzaetora (Koizumi) Pipunclusmutillatus (de Meijere)

DryinidaeIlaplogonatopusspp

Egg Copidosomopsisnacoleiae (Eady)

Larva BraLonidae

Cardiochiles philipphnensisAshnead lchneumonidac

Trichiomnia cnaphalocrosisUchida Temelucha stangli (Ashmead)

Pupa Chalcididae

BrachymeriaexcarinataGahan Brachymerialasus (Walker) Brachymeria nr tachardiaegroup

Parasite Egg

Scelionidae Telenomus rowani (Gahan)

EulophidacTettasticusschoenobhi Ferriere

Larva Braconidae

Stenobraconnicevillei (Bingham) Iclhncurnonidac

Temelucha phiiippinensis(Ashinead)Temelucha stangli (Ashmead)

x x x

x x

X x X

x x Green leafhopper Nephotettix virescens

X x x

x x

X X X X

X

X Rice leaffolders CnaphalocrocisMarasmia

x X x

X

X X

x x x x x X x x x

Yellow stem borer Scirpophaga incertulas

X X x

x

x

x X XAinauromorphaaccepta metathoracica(Ashmead) x

continued on next page

IRPS No 123 January 1987 31

Table 6 cntinued

Natural enemy Environment

Host stage Species Upland Rainfed lowland Irrigated lowland

Larva-pupa Gold fringed borer Chilo attriciiuslchneumonidae

Xanthopimpla stemmator (Thunberg) x aPupa x

Pteromalidae Trichoinalopsis apanteloctena (Crawford) ax

Pink stem borerb Sesamnia inferensParasite

Larva Braconidae

Stenobracon nicevillei (Bingham) x b b Armyworms Mvthinna SpodopteraLarva

Tachinidac Zygobothriaarroptvora (Robineau-Desvoidy) x x

Braconidac Cotesia spp

X Skipper Pelopidas mathias and green horned caterpillar Melanitis leda ismeneF-gg

TrichogramrnatidacTrichogrannaspp x

Scelionidae Undetermined Scelionid

x Larva

Tachinidae Argyroph lax nitrotibialis (Baranov) X x

Pteromalidae Trichomnalopsis apanteloctena (Crawford)

X Predator

CoenagrionidaeAgriocnenis femina femina Brauer x xIschnura senegalensis tambur

Miridae CrtorhinsliidipennisReuter x x

Nabidac Stcnonabis tagalicus (Stal) x

VeliidaeAlicrovelia douglasi atrolineata (Bergroth) x

Coccinellidae Micraspis crocea (Mulsant) X x x

Vespidae Eumenes campaniformis (Fabricius) x

Formicidac Diaecamna spp xOdontoponera transversa (F Smith) x

Carabidae Chaenius spp OphioneaW- aJasciataSchmidt-Goebel

x

Ophionea ishii ishii Habu x x X

Gryllidav Anaxipta longipennis (Saussur-) x xMetioche ittaticollis (Sial) x x x

Tettigoniidae Conocephalus longipennis (de Haan) x x x

aNo rearing made on rainfed lowland bNo rearing made on rainfed lowland or Irrigated lowland

MrasmiaAmong the Laval parasites the braconid Cardio- lowlands and parasitizes oriental maize borer OsIriniachilesphilijpphensis is most adapted to an upland environ- furnacalis (Guenee) larvae The leaffolder pupal parasitesment It also parasitizes Hydelepta indicata a common not only occur equally in all environments but also haveleaffolder of legumes The ichneumonids Trichomma cna- wvide host ranges- parasitizing species of OsIrinia Marucaphalocrosis and Temehcha stangli occur more in the Hydelepta and Honiona - in maize and legumeslowlands T stangli is most adapted to irrigated lowlands The principal egg parasite of yeltow stem borer (YSB)and also parasitizes Chilo and Scirpophaga stem borer Telenomus rowani is widely adapted to all environmentslarvae T cnaphalocrosis is most prevalent in rainfed and has an unusual alternate host -Tabanus eggs (25)

32 IRPS No 123 January 1987

Tetrastictu schoenobii specializes only in YSB and occurs mainly in irrigated lowlands -- further proof that YSB is most adapted to the wetter environments

Larval parasites of YSB are highly environment-specific The braconid Stenohracon nicevillei occurs only in upland rice but also attacks the pink stem borer (PSB) Tieinchwa phiiiplPwnsis an ichneumonid occurs in upland rice and also parasitizes the larvae of (7hiu stqpresaisinatro-morpha accepta melaihoracica like 7 stangi is mnost adapted to the irrigated lowlands

The dark-headed stem borer(l)HSB) (ih pochit3sus was not collected in all rice environments and the gold fringed borer Chilo auricilius nd PSB were collected only in upland rice Xaantiholiplastenrnaloran ichneumonid larval parasite appears to be widely adapted whereas TrihomaslopisapanitI)cteiiaa pteromalid pupal parasite occurs mostly in wetlands and also parasitizes larvae of Chilo suppressais and Pelhuidas mathias

Armyworms Myit hinin and Spodujipera have two main larval parasites - a tachinid Zrgohothriaatropivora in rainfed environments and a braconid Cotesia spp in irrigated lowlands

The butterfly pests Pelopidas and 4elanitishave tricho-grammatid egg parasites found only in irrigated lowland environments

Two larval parasites show environmental preferences --Argrroph av nigrotihialis a tachinid in rainfed habitats and Trichonialopsisapantelkcnela in irrigated ricefields A limgrotibialisalsoattacks the sweet potato hornwormAgrius convolvtdi and T apanteloctena parasitizes DH S1B

Predators as a group show more distinct environmental preferences The aquatic damselflies Coenagrionidae are most abundant in the lowlands Cvriorhinus is also most adapted to the lowlands The nabid Stenonabis tagalicus occurs only in the uplands and isalso prevalent on legumes The coccinellid lady beetles wasps and ants are more prevalent in upland areas AlicrasIpis cit-ea larvae prey ona variety of aphids attacking legumes and maize Etunienes campanonia vespid mud wasp makes nests in trees and is therefore most adapted to the more botanically diverse uplands Soil-dwelling ants cannot tolerate flooding

Upland environments are habitats to arboreal carabid beetles Three species of Chlaeniusbeetles prey on leaffolder larvae and are perhaps more important than parasites (27)

The gryllids and tettigoniids are widely adapted egg predators not only on rice but also on maize and legumes They feed on eggs of most insects that are laid on leaves

The spidercommunity of upland rice environments isrich in species At one site (Batangas Philippines) 31 species have been recorded (23) Of the 176 spider species recorded in Philippine ricefields about one-half (82) occur in upland rice The spiderspecies of upland rice environments overlap more with those of rainfed lowland than with those of irrigated lowland rice (24) The same study showed that of the three environments irrigated rice has (ie greatest spider species diversity foiiowed by rainfed lowland and upland

However of the 10 most prevalent ricefield spiders 9 were abundant in upland rice showing wide adaptation All three spider guilds -- orb-web space-web and huting spiders shywere prevalent in upland ricefields There were however differences in environmental prelerences for some spider species

Of the orb-wcavers Tetragniathamanlihulatawas partishycila rlv abundant 7 japonica was low and Ltucatge dehcorata was absent in upland rice

As in other environments Aiypeiwnaifirtosana was the most dominant space-web spider however its relative numbers were lowerthan for lowland sites Among hunting spiders Ox|opes javanus and Lcosa leatcost fgla were more abundant and Lvcosa lpseudoanmnlata was less abundant in upland compared with lowland rice envishyroniments

Extensive rearing of rice insect pests in upland rainfed lowland and irrigated sites over an 8-yr period in the Philippines provided insight into the effectiveness of parasites as natural enemies (Table 7 8) Most of toe data are from large sample sizes taken over at least one crop which overcomes the pitfalls expressed by Van Driesche (238)

VSB egg parasitization was surprisingly similar across all environments Larval parasitization was highly dynamic among sites and even years within the same site but showed somewhat lowerlevels in uplandsthan in irrigated wetlands This would have been predicted for egg parasites as well since YSB is adapted to the lowlands and so must be its parasites This isevidence that Telenomus rowani is as good at dispersing as its main hosts Trowani clings onto the anal tufts of female YSB and parasitizes eggs as they are laid This phoresy may explain its wide adaptation The more specialized lowland YSB larval parasites therefore appear more effective than their upland counterparts with more alternate hosts

Low collection levels of PSB larvae especially in the wetlands make comparisons between environments difficult Low levels of larval parasifization were recorded in both rainfed environments Except in the Cagayan rainfed wetland site in 1980 low levels ef larval parasitization were evident at irrigated and rainfed sitesalike No differences by environment were evident Higher levels of parasitization occurred in Sarawak during a 1967 outbreak also from tachinids Rothschild (197) indicated that higher parasishytization occurred in upland rice areas

Skipper and green horned caterpillar larval parasitization rates were generally low and showed no environmental effect htighest activity occurred in Cagayan in 1980 (48 parasitization) Although larval parasitization levels of RLF were variable year to year activity appeared greater in the irrigated lowlands than in the uplands Again the highest levels were recorded in Cagayan in 1980 Cagayan has extensive grasslands and fallow around the rice areas and a relatively long rainfall period Rainfall in 1930 was good for crop growth and apparently for increase of larval parasites

IRPS No 123 January 1987 33

Table 7 Parasitization rates of rice planthoppers and green leafhopper in upland rainfed lowland and irrigated lowland environments Philipshypines 1976-840

Environment and site Year Egg parasitization () Nymphaladult parasitization ()

BPH W13P GLH BPII WBPH GLIH

UplandBatangas

Rainfed lowland

1976 1979 1980 1982

15 17 12

2(47)8(39)

4(51)

9(1800)14 (212)

12(304)

8 (101)6 (127)

9(116)

Iloilo

Cagayan

1976 1977 1978 1981 1980 1981 1982

0 (69)

11

44 48 40

-

-

46 55 44

0 (471)

11

52 56 49

0 (200)9 (214)

10 (1259)8(235)6 (900) 3(800)

-7 (462)

11 (270)3 (313)5 (900) 1 (800)

4 (600)10 (725)10 (362)12(870)5 (900) 3 (800)

Pangasinan

Irrigatedlowland

1984 1976 1978 1979 1982

13 (436) 13(379) 14 (457)

6(14)7 (197)

17(198) 11 (486) 18 (275)

3 (81)10 (274)3 (116)

16(871) 19(88)

28(211)9 (764)5 (647)7 (1204) 8(305)

Laguna 1977 11 7 21 1978 31 1979 14 4 4 28 1980 7 8 20 1981 1983

56 7(278)

26 16 (1650)

16 27(1921)

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R I Apostol unpublished 1lo1lo and Pangasinan 136 Cagayan BCanapi unpublished Liuna 175 I C Pantua unpublished 118 234 43 Carino and Shepard unpublished BPH = brown planthopper Nila parvara htgens W131Pi =whitebacked planthopper Sogatelia furcifera GLH = green leafhopper Nephotettrix virescens Figiires in parentheses are sample sizes

Tauher et al (230) reported that internal external and genetic factors influence the seasonal activity of parasitoids

Hopper egg parasites but not nymphaladult parasites were also parlicularly abundant at the Cagayan site possibly the egg parasites have more alternative hopper prey in the weedy areas than nymphaladult paisites

The seasonal dynamics of hopper egg parasites can be seen for upland rice in Batangas(Fig I 2) and for rainfed lowland lice in Pangasinan( 135)and Cagayan( 105) Levels of BPHWBPI-L and GLH egg parasitization were corn-parable in upland and irrigated lowland environments Except for some high rates of paraMsitization early in the season the parasitization rate tended to be steady in all envilonments However hopper nymphaladult parasi-tizution levels were more similar in the rainfed environments and lower than in irrigated rice

Overall parasites do not appear highly effective by themselves against upland or even lowland pests Among natural enemies parasites have attracted the most attention because they can be more readily assesscd

Predators and pathogens lend themselves better to management practices Predators the most important group of natural enemies in riccfields are difficult to quantify On the other hand pathogens appear less im-portant but can be readily cultured and disseminated

The natural enemy commulnity of upland rice is rich in species and differs significantly from that of lowland rice

These beneficial organisms must be conserved by applying insecticides judiciously particularly sprays Seed or soil placement of chemicals minimizes exposure to natural enemies The strategy to derive the greatest benefit from natural enemies isto allow the greatest number of beneficial species to thrive Then perhaps at least one species will be effective against each pest at any tine overcoming the variable abundance of each species within and between years

Programs to introduce exotic species or mass produce indigenous ones are ambitious and expensive because they require trained people on a sustained not ad hoc basis

Chemical control Insecticides offer rapid and efficient control of upland insect pests (190) but-should be used only after other control measures have been considered Insecticides are rarely used on upland rice because of cost (32) Upland rice yields are normally too low to justify the expenditure Also most upland rice is grown as a cash crop Spraying upland rice is more difficult than lowland rice because water is less accessible Government subsidy programs to provide pestishycides to tribal peoples for upland rice production have created problems of toxicity to humans because isolated people have had no experience to be able to handle pesticides safely

However ins-eticide use has beenjustified when a crop is

0Z

Table 8 Parasitization rates of lepidopterous pests in upland rainfed lowland and irrigated lowland environments Philippines 1976-84a

Parasitization (5)

Environment and site Year YSB GFB PSB Earcutting Swarming Skipper Greenborned RLF Brown larva larva caterpillar caterpillar caterpillar caterpillar larva semilooperEgg Larva larva larva larva larva larva

UplandBatangas 1977 - 0 (57) - 0 (63) - 0 (71) - 0 (106) 11(142) 2 (101)1978 43 (4697) 8(451) 5 (38) 3 (40) 6 (61) 2(58) 15 (57) 13 (129) 13 (1286) 4 (30)1979 59(4135) 12(320) 13 (11) 5(28) 9(45) 6(46) 7(23) 8(160) 9(1152) 8(26)

Rainfed lowland Iloilo 1976 - 0 (10) 0(10) 0 (7) 0 (19) 2 13) 15 (180) 5 (76)1978 0 (54) 0 (3) 0 (17) 9 (64) 4 (78) 3 (73) 0 (121) 0 (191)1979 64 (2350) 16 (84) 6 (17) 0(6) 18 (68) 25 (104)

1981 56 (1322) 18(304) 0(5) 12(51) 18(138)Pangasinan 1976 60 (7 7 )b 62(82) 0(111) 5 (39)

1978 58 (53)b 5(51) Cagayan 1979 59(713) 8 (101) 6 (34) 0(7)

1980 0(11) 4 (28) 15 (286)

62 ( 48(177) 61(140)1981 0(5) 12 (2) 13(45) 14 (92)

Irrigatedlowland Laguna 1974 62 8

1974-75 50 41 1975 43 72 1978 96 (623) 21(24) 33 (60)1979 83(2017) 18(110) 8(62) 11 (89) 22(400)1980 78 (1625) 20(75) 0 (16) 4 (72)1983 60

9 (80) 7 (270) 30

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R F Apostol unpublished Iloilo and Pangasnan 136 Cagayan B Canapi unpublished Laguna 175 P CPantua unpublished 118 234 143 Carino and Shepard unpublished YSB = yellow stem borer ScirpophagaincertulasGFB = gold fringed borer Chilo auricilius PSB = pink stem borerSesamia inferens earcutting caterpillar Alythimna separata swarming caterpillar Spcdoptera mauritia acronyctoides skipper Pelopidas mathias reen horned caterpillar Melanitls ledaismene RLF = rice leaffoldersCnaphalocrocismedinalis and Marasmiaspp brown semilooper Mocisfrugalis Figures in parentheses are sample sizes Egg masses

IRPS No 123 January 1987 35

Table 9 Effect of seeding rate and insecticide on plant density and Yield losses from insect pests in upland rice often areyield of Dagge upland rice Batangas Philippines 1976a comparable to those of lowland rainfed or irrigated rice

Seeding rate Insecticide Dosage Plant density Yield (0-Y30o) but usually from different pests The principal (kg aiha) seed treatment (kg aiha) (noIra-row) (tha) groups are soil pests feeding on sown seed and rootsat 14 DE followed by seedling pests These groups have largely gone

50 None - 35 a 20 a unnoticed leading researchers to conclude that stem borers50 Carbofuran F 10 38 ab 23 a and seed bugs are the most important pests (17) Yield loss 10 beCabofuranF100 None - 4551 c 23 a trials that pinpoint major pests need to includc seed

100 Dieldrin WP 10 52 cd 22 a treatments Granules applied in the furrow may not give150 None - 59 d 22 a adequate protection from many seed pests

aAv of 8 fields DE = days after crop emergence In a column means The floristic and pedological diversity of upland rice followed by a common letter are not significantly different at the environments is matched by the diversity of arthropods5level by D)MRT F = flowable Wi = wettable powder both pests and their natural enemies Irrigation has homoshy

genized lowland rice environments Flooded soils tend tothreatened by epidemics of locusts or armyworms New have the same properties and to eliminate soil pests The technology stressing efficiency has minimized insecticide spatial and temporal dominance of cultivated rice has cost benefited specialist pests

Seed treatment is normally inexpensive and can be Upland rice has no specialist arthropod species The economicallyjustified in -nanycases(66) Treating seed with monophagous pests on upland rice prefer the lowlands systemic insecticide (216) but not chlorinated hydrocarbons where rice evolved Rice in the uplands exists only because (247) is most effective in controlling seedling maggots of man and even though lowland rice predates upland riceFoliar sprays have to be repeated to obtain control and specialist species evolved Wheat maize and sorghum do often are washed off by frequent early season rains not have monophagous pests leading us to conclude that Granular insecticides are effective but required dosages are there is no evolutionary advantage for inssects to have too expensive to justify except under heavy infestation narrow host ranges in upland graminaceae(247) When directed against first-instar larvae banding Pests have adapted to the uplands not only by having granules in seed furrows (177) can be effective against white wider plant host ranges but also by having long life cyclesgrubs at low dosages (025 kg aii ha) (134) Seed treatments undergoing dormancy or being dispersive Each pest has a are not promising for white grub control (248) unique life history strategy suited to highly unpredictable

Seed treatment is also effective ainst ants and may be environments In agivenyear someinsect specics an better cheaper than increasing the seeding rate In Batangas exploit upland rice as a food source but no pest is a senious treating 50 kg seed ha was as effective as increasing the one every year on upland rice seeding rate to 100 kgha and using untreated seeds (Table The low yielding potential of upland rice means less 9) In Brazil seed treatments control termites and other soil quantity and lower quality of food for pests to exploit but insects including the lessercornstalk borer(66) Domiciano also less incentive for farmers to undertake control (58) however found seed treatments inconsistent in their The most effective control would be early and synchroshyeffect against high populations of lesser corn stalk borer nous planting of early-maturing upland rice varieties to

Baiting can be an inexpensive way of controlling ants prevent insect pest buildup Breeding for insect resistance seedling maggot flies mole crickets and field crickets Baits should initially focus on stem borers because they are verymade of locally available material and impregnated with difficult to control by other means Biocontrol efforts insecticide can be sparingly distributed taking adantage of should first stress conservation of natural enemies preshythe pests ability to disperse and encounter bait sites dators and pathogens rather than parasites Efficient

The new low-volume hand-held controlled-droplet chemical control includes seed treatment or baiting for seed sprayers may offer an advantage over high-volume knap- seedling and some root pests and using controlled dropletsack sprayes for controlling foliar pests Upland rice sprayers for spot treatments against foliar pests and seed canopies are more open than those in lowland rice and bugs when economic thresholds are reached better droplet penetration should result Controlled droplet applicators would be ideal for seedbugs and defoliators

CONCLUSION

Upland rice is attacked by a wider array of insect pests than is lowland rice mainly beca use of the addition of soil pests Generally population levels of lowland rice pests are lower in upland than lowland rice leading many people to conclude that insect pests are not important on upland rice

36 IRPS No 123 January 1987

REFERENCES CITED

1Abraham C C and K S R Mony 1978 Occurrence ofLeptocorisa acuta Fabr (Coreidae Hemiptera) as a pest of nutmeg trees J Bombay Nat ttistSoc 74553

2 Agyen-Sampong M 1975 HlierogrlphusdiaganensisKrauss (Orthop-tera Acrididae) a new pest of rice in Northern Ghana Ghana J Agric Sci 8249-253

3 Agyen-Samnpong M 1976 Varietal resistance and other inle-ratcd pest control components as factors of rice varietal improvement in West Africa Prcsented at the Second varietal improvement serinar 14-18 Sep 1976 Bouak Ivory Coast West Africa Rice Development Association Monrovia Liberia 18 p

4 Agyen-Sampong M 1977 Outbreak of Ihccrorrdnisorae in rice seedlings near Rokupr Sierra Leone lit Rice Res Newsl 2(6)16-17

5 Agyen-Sampong M 1980 Dirty panicles and rice yield reduction caused by hugs IntRice Res Newsl 5(l)11-12

6 Agyen-Sampong M 1982 The major pest problems of irrigated upland and mangrove swanip rice ecosystems ir the humid tropical Guinea savanna and Salel climate zones Pages 318-328 in Pro-ceedings training course cn concepts techniques and application of irtegrated pest management in rice inWest Africa 10-20 Jan 1982 Fendall Liberia West Africa Rice l)evelopioent Association Monrovia Liberia 505 p

7 Akibo-Betts V I and S A Raymundo 1978 Aphids as rice pests in Sierra Leone lit Rice Res Newsl 3(6)15-16

8 Akinsola F A 1975 Present status of different rice stemborers in parts of Nigeria iicL Entomo Newsl 328

9 Akinsola FA 1979 Ihe biology and ecology of rice stemborers in Nigeria Ph 1) thesis University ol lbadan Nigeria 217 p

10 Akinsola E A 1979 Guidelines of integrated management of insect pests of upland rice in West Africa Presented at the Seminar on integrated management of rice diseases and insect pests at Bobo-Dioulasso Upper Volta 17-22 Sep 1979 West Africa Rice Develop-ment Association Monrovia Liberia

II Akinsola IA 1984 Insect pests of upland rice in Africa Pages 301-305 in An overview of upland rice research Proceedings of the 1982 lIouak Iwsrv Coast upland rice workshop International Rice Research Institute 13O Box 933 Manila Philippines

12 Alam M S V TJohn and Kaung Zan 1985 Insect pests and diseases of rice in Africa Pages 67-82 in Rice improvement in Eastern Central and Southern Africa proceedings of the international rice workshop at Iusaka Zambia 9-19 April 1984

13 Alticri M A A van Schoonhovcn and J D Doll 1977 The ecological role of weeds in insect pest management systems a review illustiated with bean (Phaseohsvulgaris L)cropping systems PANS (Pest Artie News Summ) 23195-205

14 Anmante F 1967 The leaf-cutting ant Atia capiguaraa pest of pastures [in Portuguese] Biol6gico (Sao Paulo) 33113-120 (Rev Appl Entomol A56207)

15Andrade A C and D Puzzi 1951 Preliminary results of experi-merits on the control of Sculptcoris iastantus on sugarcane [in Portuguese] Bioulgico (Sao Paulo) 1744-49 (Rev Appl Entomol A41421422)

16 Appert J 1970 Maliarplia separaelha (rice stem borer) new observations of entomological problems of rice in Madagascar [in French] Agron Trop (Paris) 25329-334

17 Ariadeau Mand Z Harahap 1986 Relevant upland rice breeding objectives Pages 189-197 in Progress in upland rice research Proceedings of the 1985 Jakarta conference International Rice Research Institute 1O Box 933 Manila Philippines

18Asar S and Kisimoto 1975 Seasonal occurrence of the smaller brown planthopper in upland rice [in Japanese] Proc Kanto-Tosan Plant Prot Soc 2228

19 Bakker W 1974 Characterization and ecological aspects of rice yellow mottle virus in Kenya Meded Landbouwhogesh Wage-ningen 152 p

20 Balut F F 1970 White grubs attacking rice Oryza ativa L [in Portuguese] Biol6gico (So Paulo) 36321-322

21 Banerjee S N and L M Pramanik 1967 The lepidopterous stalk borers of rice and their life cycles in the tropics pages 103-124 in The major insect pests ofthe rice plant Proceedings ofa symposium at the International Rice Research Institute September 1964 The Johns Hopkins Press Baltimore Maryland USA

22 Barrion A T and J A Litsinger 1980 Ants a natural enemy of cliaiarac in dryland rice IntRice Res Newsl

5(4)22-23 23 Barrion A T and J A litsinger 1981 The spider fauna of

Philippine rice agroecosystem 1Dryland Philipp Entomnol 5139-166

24 Barrion A T and 1 A Iitsinger 1984 The spider fauna of Philippine rice agroccosystcms II Welland Philipp Eiitomol 611-37

25 Barrion A T1 and J A Litsinger 1984 lihanus (Diptera Tabanidae) eggs an alternative host of rice stei borer (SI3) egg parasite Telenoonis digns (Iliymenoptcra Scelionidae) lit Rice Res Newvsl 9(6)19

26 Barrion A T and J A Litsinger 1985 Identilication of rice leaffolders by wing markings Int Rice Res Newsl 10( 1)24

27 Barrion A T and J A Litsinger 1985 Chctiosspp (Coleoptera Carabidae) a leaffolder ILF)predator lit Rice Res Newsl 10(1)21

28 Bertels A 1970 Insect pests of rice and theircontrol [in Portuguese] Pelotas Brazil Instituto de Expcrimentacao Agropccuararias do Sul Circular 43 24 p

29 Bonzi S M 1982 Ctilo (flisilineus J de Joannis (Lepidoptera Pyralidae) stem borer of irrigated cereals in Upper Volta LioFrench] Agron Trop Riu Rizic Cult Vivrieres Trop 37207-209

30 Bourke T V T L Fenner J N LStihick G Baker EHassan D F OSullivan and C S li 1977 Insect pest survey for the year ending 30th June 1969 Papta New Guinea )epartment of Agrishyculture Stock and Fisheries Port Moresby 57 p

31 Bowden 1 1973 Migration of pests in the tropics Meded Rijksfac Landbouwwet Gent 38785-796

32 Breniere J 1973 The main rice parasites in West Africa and their control Pages 27-42 in Proceedings seminar on plant protection for the rice crop West Africa Rice Development Association Monrovia Liberia

33 Breniere J 1976 The principal insect pests of rice in West Africa and their control West Africa Rice Development Association Monrovia Liberia 52 p

34 Brown ES 1962 The African arinyworn Spoiolptera eXI)Ina (Walker) (Lepidoptera Noctuidae) a review of the literature Cornshymonwealth Institute of Entomology London UK57 p

35 Brown F B 1968 Upland rice in Latin America FAO-International Rice Commission 12th Session of the Working Party on Rice Production and Protection 2-14 Sept Peradeniya Sri lanka Food and Agriculture Organization Rome Italy 6 p

36 Buyckx F J 1 1962 Disease and insect pests of crops inCongo Rwanda and Burundi [iil French] Publications dc LInstilut National pour lestude Agrononiquedu Congo M Weissenbruck S A Brussels Belgium 708 p

37 Cariiio F 0 1981 Role of natural enentics inpopulation suppression and pest management of green rice lealhoppers Ph D thesis University ofthe Philippines at Los Bafitos PhilippinesLaguna 136 -

38 Carpente A J 1983 The development of rice cultivation and extension of vegetable productin in Zanzibar Final Report Vol II1 October 1983 Food and Agricultme Organization Ronte Italy

39 Carroll C P and S1 Risch 1983 Tropical annual cropping systems ant ecology Environ Manage 751-57

4(0 Cates R G 198 1 Host plant predictability and the feeding patterns of monophagous oligophagous and polyphagous insect herbivores Oecologia (Berl) 48319-326

IRPS No 123 January 1987 37

41 Cendcafia S M and F B Caora 1967 Insect pests of rice in the 62 Ene 1 C 1981 I)iiorldhol)ierusihilis Slater (licteroplera Philippines Pages 591-616 in The najor insect pests ofthe rice plant lygacidue Illissinac) a new rice pest front Nigeria lit Rice Res Proceedings of a symposium at the International Rice Research Newsl N3)15 Institute September 1964 The Johns Hopkins Press Baltimore 63 Fugenio T S 1965 rhe effect oi e-vCgetation on rice production Maryland USA 729 p and the possibility of overcoming it by leaf pruning MS thesis

42 Centro Ihternacional de Agricklittira Tropical 1984 Upland rice in University ol the Philippines at Los Baijos ILaguna Ihilippins 69 p Iatin Anerica Pages 93-119 hi An os ervies of uplaind rice research 04 Farrow R A 1975 The Alrican nmigratory locust ii is main Proceedings of the 1982 Boinakamp Ivory Coast upland rice workshop outbreak area of the middle Niger qlinlitatise studies of solitary International Rice Research Institute P0 Blox 933 Manila populations in relation to environmental factors Locusla No II Philippines 198 p

43 Chandra G 1979 Taxonomy ad hn riononlics ofthe imect parasites of 65 Fennal R G 1943 Food crop investigation Windward and rice Icalloppers and planthoppers in the Pihilippines and their Ieeward Islands First report Report onl work dunring the periodiiiportairce iii natural biol)gical cointrol Philipp Fntoltol 4119-139 April 1942-April 1943 Imperial College of Tropical Agriculture

44 Chang T T 1976 The originii colut ion enlIivation dissemination Trinidad 16 p (Rev of Appl Fintornol A3222-25) and diersificition of Asian and African rices -uphytica 25425-44 1 66 Ferreira F 1983 Integrated pest ritaageiret of upland rice [ill

45 Chatierjee P B 1979 Rice leaffolder attacks in India lit Rice Res Portuguese English summary] Pages 323-341 in Upland riceculttre Newsl 4(3)21 factors affecting production E Ferreira T Yamada and F-

46 Cheng Cbien-pan 19(4 Upland rice in Taiwan International Rice Malmolta eds Institutoda Potassae Foslato Institnto Internacional Cnmmission Wiorking Party oit Rice Production ard Protection I()th da Potassa Piracicaha Sao Paulo lraid Meeting 3-10 March 1964 Manila Philippines Food and Agricul- 67 Ferreira E arid F dl Silva Martins 1979 Occurrence of ture Organiaion Rome Italy 5 p Sogatohs orvziola (Muir) inupland rice in Goias Ilrail lit Rice

47 Chcrrett 1 M and 1) 1Peregrine 1976 A review of the status of Rcs Ncwssl 4(4)18 leaf-cutting ants and their control Ann Appl Biol 84124-128 68 Ferreira E and 1F da Silva Martiris 1984 Insect pestof rice ill

48 Chipeta F M 1974 Pests of rice in Malawi Pages 90-95 in Brazil and their control Jill Portugucsej FMIlRAtPA-CNPAF Proceedings Sith eastern Africa cereals research conference 10-15 Goiania Brmiil 67 p March 1974 DRB Mialda ed Ministry of Agriculture lilongwe 69 Ferreira FI F ca Silha Miartins S S Nero and 1 It P de Malawi Carvallio 1982 M tltiplc regression analysis todeterimine the effect of

49 Chu Yau-i 1979 The d atiaging period of the rice seedling fly insect populatiois ott upland rice yield [inIPortuguese Pesqui (Atlterigotiavxigu Stcin) (NItiscide Diptera) oil cori) at Fast Java Agropecu Bras 17671-675 Indonesia Plant Prot Bill 21403413 70 Ferreira F 1F da Silva Martins and F 1PZimmermann 1979

50 Conklin H C 1957 IHaninoo agriculture a report on aln integral Rice varietal resistance to tle lesser corn stalk borer [in Port uguese] systent of shi fti rig cult ivationi in the Pltilippincs Food and Agriculture Pesqui Agropccu Biras 14317-321 Organization Rome Italy 209 p 71 Fleming W F 1968 Biological control of lie hapanese beetle U S

51 Cramer H II 1967 Plant proection and world crop production Dep Agric RLs Sers [ech Bull 1383 78 p Pilaniensclhutz-Nachr 201-524 72 Food and Agriculture Organiation of the United Nations 1982

52 l)ani R C and Majunidar 1978 Prelininary observations on tie Country report surmimary Pages 27-37 in Report on the first incidence of root aphids on differeit rice cultnrcs Sci Cult 4488-89 FAOIUN)IPhail and regional training coiuirse in iniproved ctltural

53 Davidson R I 1979 lffects of root feeding on foliage yield Pages practices for upland rice 5-23 Oct 1981 Northern Region Agrculturd117-120 in T K Ci osb and R B Pottinger eds Proceedings of the Developririt Center Chiang Mai Food and Agricuilture Organiiashy2nd Australian conlerence on grassland invertebrate ecology Govt tion Rome Italy 37 n Printer Wellington New Zealand 73 Franco do Ariaral S 195 1 Residual action of insecticides in turfy

54 l)ean G 1W 1976 Rice insect pests inlaos Int Rice Res Newsl soil--the results of a field experiment [in Portuguese] Biol6gico (Sao 1(2)15 Paulo) 17209-214 (Rev Appl Entomol A41423)

55 I)ean G J W 1978 Insect pests of rice in Laos PANS 24(3)280-289 74 Freeman ) 1970 Report onithe Iban london School of -coroiics 56 )ingle H 1984 Behavior genes and lifc histories conplex adapta- Monographs on Social Anthropology No 41 Athlane Press

tions to uncertain environments Pages 169-194 in A new ecology London UK 317 p novel approaches to interactive systems P W Price C N Slobod- 75 Fuchs T W1 A Harding and 1W Smithlr 1979 Induction and chikoff and W S Gaud cdsl ohn Wiley New York USA termination oldiapause in the sugarcane borer Ann Flitonlol Soc

57 Dobelmann-JP and M Falais 1968 Upland rice cultivation ill tie Am 72271-274 North-western part of Madagascar in 1966-1967 [in French English 76 (hbriel B1) 1975 A review tf the major insect pests of some tpland summary] Trop Agron 2313-21 crops in tlc Philippines Pages 86-111 in Ad Htoc panel of experts on

58 Doniiciano N L 1979 Chemical control of tie lesser corn stalk pest disease arnd weed problems in some rainfed crops Bangkok (lolsonno~paloIgnowslhts) (Zeller 1848) (I-epid(ptera Ihycitid ae) Thailand Food aid Agriculture Organization Roire Italy

on rice in Ponta Gressa Parara 1977-1978 [in Portuguese] Pages 77 Garg D K and N K Shal 1983 White grub outbreak on rainfed 291-303 in Anais 1 Reuniao de tecnicos cmiirizicultura do estado de dryland rice in Uttar Pradeslh Int Rice Res Ncwsl 81(4)16-17 Sao Paulo-Campinas Cati Brazil 78 Garg I) K and IP Tandon 1982 Major insect pests of rice on hilly

59 Dunsmore 1 R 1970 Investigations on tlie varieties pests and tracts of Uttar Pradesh India lilt Rice Res Newsl 7(11)11-12diseases of upland rice (hill padi) in Sarawak Malaysia Int Rice 79 Garrity 1) P 1983 Upland rice research an international bibliograpy Comm Newsl 1929-35 1975-1982 Iternational Rice Research hnstitnte PO Box 933

60 Elias 1R1965 The new decline of rice seedlings is not caused by Manila Philippines 121 p parasites [in Portuguese] Biol6gico (Sao Paulo) 313-6 80 Ghauri M S K 1982 A new subspecies of Iinorh)rone bull

61 Empresa Brasiliera de Pesquisa Agropecuaria 1984 Upland rice itt cornlus Slater (Hemiptera Iygaeidae) front Papua New Guinea on Brazil Pages 121-134 it Av overview of upland rice research rice and carpet grass Ihll lEntoniol Res 72133-137 Proceedings ofthe 1982 Blouak Ivory Coast Upland rice workshop 81 Gonzales F 0 PArregoces R I ihernandef and OT Parada International Rice Research Institute PO Box 933 Manila 1983 Insect and imite pests of rice arid their control in Latin America Philippines [in Spanish] la Federacion Nacional de Arrocerts (Fedearroz)

Bogota Colombia 60 n

38 IRPS No 123 January 1987

82 Grist D H and R J A W Lever 1969 Pests of rice Longmans London UK520 p

83 Hale P R 1979 Two subterranean pests of upland rice in Papua New Guirnca IntRice Res Newsl 4(2)16-17

84 Hale P R and J T Hale 1975 Insect pests of rice in the New Guinea Islands Rice Entomol Newsl 34

85 Harris W V 1969 Termites as pests of crops and trees Common-wealth Institute of Entomology London UK 41 p

86 Hattori 1 1968 A method to distinguish between grass leaf roller (Ctaplha rocis medinalisGuenee) and the rice leaf roller (Susurtia exigua Butler) [in Japanese] Plant Prot (Japan) 23167-170

87 Henry I E and 1A Onsager 1984 Experimental control of the mormon cricket Iahrosshnph by NJsena locustae(Aficrospora Microsporidae) a protozoan parasite of grasshoppers (Ort Acri-didae) Entomophaga 27197-201

88 Hidaka 1 P Vungsilabulr and SKadkao 1974 Studies on ecology and control of the rice gall midge in Thailand Tech Bull 6 TropicalAgricultural Research Center Tokyo Japan 113 p

89 Hincklcy A 1) 1963 Ecology and control of rice planthoppers in Fiji Bull Entomrol Res 53467 481

90 Hirao J 1982 Ecology and chtInical control of the rice leaffolder Jpn Pestic Inf 4114-17

91 Ho DT 1984 Sten borer incifience in rice ecosystems in KenyaEast Africa Pages 307-320 in A overview of upland rice research Proceedings of the 1982 Bouac Ivory Coast upland rice workshop International Rice Research Institute PO Box 933 Manila Philippines

92 Ho )T J 1Njokah and J GKibuka 1983 Studies on rice sten borers in Kenya with emphasis on ifaliarpiaipiraellRag Insect Sci Appl 465-73

93 Ichihara I T Ito A NIatsuo and M Sawada 1971 Chemical control of the rice striped stem borer in upland rice grown in plastic mulch [iii Japanese] Proc Kanto-Tosan Plant Prot Soc 1882

94 IclhiharaIand A Matsuo 1969 Damage caused by rice striped stett borer to upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1686-87

95 Ichhara I atid A MIatsun11970 Conside ration oft lie extraordinary prevalence of rice striped stem borer in upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1778-79

96 Institut de Recherches Agrolnliqes Iropicale et des Cultures Vivrieres 1978 Insect resistance Pages 12-13 in Upland rice 1978 atnual report Bouak Ivory Coast 27 p

97 International Institute for Tropical Agriculture 1983 Aphids itt rice roots I ITA Res Briefs 4(1-3)4-5

98 International Institite fi Tropical Agriculture 1983 Hopperburn in rice caused by brown plathopper IITA Res Briefs 4(l-3)6-7

99 International Rice Research Institute 1975 Annual report for 1974 PO Box 933 Manila Philippines p 216

100 International Rice Restarch Institute 1976 Annual repiort for 1975 PO Box 933 Manila Philippines p 228

101 International Rice Research Institute 1979 Cropping systems program Component technology development and evaluation Page 404 i Annual report for 1978 11O Box 933 Manila Philippines

102 International Rice Research Institute Department of Entomology 1981) Insecticid evaluation 1979 Tables 73 74 PO Box 933Manila Philipes

103 International Rice Research Institute Department of Entonmology 1981 Insecticide evalomtion 1980 Table 81 POBox 933 Manila Philippines

104 International Rice Research Institute 1981 Report of a meeting of Asian rice-based cropping systems entomologists Eleventh cropping systems working group meeting 18-22 May 1981 PO Box 933 Manila Philippines p 132 133 144

105 International Rice Research Institute 1984 Cropping systems program Pages 368-373 in Annual report for 1983 PO Box 933 Manila Philippines

106 International Rice Research Institutc 1984 Upland rice in Asia Pages 45-68 in An overview of upland rice research Proceedings of the 1982 Bouakamp Ivory Coast upland rice workiiop loternational Rice Research Institute 11O Box 933 Manila Philippines

107 Irshad M 1977 Grasshoppers in rice in Pakistan lit Rice Res Newsl 2(3)8-9

108 Ito R and TOno 1964 Upland lice its culture and varieties in Japan lIternational Rice Cormmission loth Meeting of the Working Party on Rice Production and Protection 3-10 March 1964 Manila Philippines Food and Agriculture Organization Rome Italy 12 p

109 Jeffrey M I and V A Dyck 1983 Effect of moonlight on lightshytrap catches of brown planthopper lit Rice Res Newsl 8(6) 13

110 Jennings P R and T Alicia Pineda 1983 Management ofthe rice plnthoppers Sogatodsorivi(coIl in Latin America Presented at the meeting on stable plant resistance FAO and lIndocsian I)inectorate of Food Crops Protection Bali lndonesia 29 Novcniber-2 l)ccemshyber 9 p

IllJimenez Gomez J A 1979 Rice insect pests i lla ios ()riCntales lil Spanish] Pages 99-103 in Rice ciltivation Instituto (olotibiatto Agropecuario Compndio No 29

112 Jongeleen F JJ Soesanto and A Mockidjo 1979 Studies oti pathogens of two species of Scarabacidac froinIndonesia Z Angew Enlomnol 87412-417

113 Jouvenaz 1) PC S lofgren and W A Baiks 1981 Biological control of imported fire antsa review of cirrent knowledge Bull Entomol Soc Am 27203-208

114 Kainoh Y K Shiniiu S MarUani Y Taiaki 1980 lost finding behavior of tie rice bu lplotori+JchinvittjI)allas (liciniptera Corcidac) with special reference to diet pattern ol aggregation and feeding on the rice plant Appl Entomol Zool 15225-233

115 Kakkar K 0 P Sharma and P I Sharma 1977 Note on Ihteronyrchus liodresRedt (Coleopicra I)ynastidae) as asporadic pest of paddy and maie inHimachal Pradesh Corr Agric 156-57

116 Kalode M131 79 Major insect pest problems and their control in upland rice (Unpublishcd mineogriph CWI oJNaaiat lni Chandaratat (198 1)original not seen)

117 Kalshoven L i 98 I The pests of crops in I ldoniesia (translated by P A vatn der Laan) 1 T Ichtiar B3aru-vati Iloeve lakarta Indonesia 701 p

118 Kamal N Q 1981 Suppression of whitebacked planto ner Sogatella fircaifra (Horvath) aLId rice lealfolder (nhal -cis licdb lis (Guenee) populations by natural cnclmie hlI) tiesis Grcgorio Araneta University Foundation Victoncta Park Metto Manila Philippines 179 Ip

119 Karim A N M Reaul 1975 A species ofliealy bug Trioni i70s ceres new oii rice in the Philippines Rice Entoniol Newsl 247

120 Katanyukul Wand C Chandaratat 1981 Insect pestsof uplaid rice and their control Report firstFAOUNlFi Tliailand training course on improved cultural practices for tplad rice Chiang Mai Thailand 5-23 Oct 1981 Food and Agriculture Organiation Rome Italy 22 p

121 Kern F 1955 lhletwioa(I)uninemts) bhhit Biturm(Coleoptera) a new pest of sugarcane inVenezuela [inSpanish with Etnglish sunmary] Agron Trop (Maracay) 527-29 (Rev Appl Eitoniol A4572)

122 Khan K M and S Ghai 1974 White grubs ald their control in India Pesticides (1Bombay) 819-25

123 Kiritani K 1975 Integrated pest management of rice pests Pages 66-70 itt Proceedings platning workshop ott cooperative hIeld research in pest managementP SMotooka ed East-West Center Honololu Hawaii USA

124 Kisimoto R 1967 Ecology of insect vectors forecasting and chemical control Pages 243-255 in The virus diseases of tle rice plant Proceedings of a symposium atthe International Rice Research Institute April 1967 The Johns Hopkins Press altimore Maryshyland USA

125 Kisimoto R and V A Dyck 1976 Climate and rice insects Pages 367-391 in Climate and rice International Rice Research Institute PO Box 933 Manila Philippines

126 Kureha S S Kobayashi SAsari and T Ishihara 1977 Life history and control of the mole cricket Grrllotalpa africana Palisot de Beauvois [in Japanese English translation 7520 1RRI library] Proc Kanto-Tosan Plant Prot Soc 21192-197

127 Kushwaha K S 1981 The black beetle as a pest of rice in Haryana India Int Rice Res Newsl 6(4)14

128 Lawani S M 1982 A review of tile effects of various agronomic practices on cereal stem borers Trop Pest Manage 28266-276

129 Leuck D B 1966 Biology of the lesser cornstalk borer in South Georgia J Econ Entomol 59 797-801

130 lever R J A W 1955 Insect pests of the rice plant in Malaya Int Rice Comm Newsl 1612-21

131 Li Ching-Sing and Shin-Foon Chiu 1951 The study ofthe rice gall midge Iachydiplosis orvza Wood Mason [in Chinese with English summary] 1Taiwan Agric Res 21-13

132 Iitsinger J A 1984 Assesament of the need-based application for rice Proceedings TechnologyTransfer Workshop 15-16 March 1984 International Rice Research Institute 1PO Box 933 Manila Philippines 6 p

133 Litsinger J A 1984 Environmental diversity and insect pest abundance with reference to the Pacific Pages 285-307 in Proceedings sub-regional training course on methods of controlling diseases insects and other pests of plants in the South Pacific 4-20 Oct 1982 Vaini Kingdom of Tonga Nukualofa

134 Litsinger J A RF Apostol and M BObusan 1983 White grub Leucopholis irrorata (Coleoptera Scarabaeidae) pest status popula-tion dynamics and chemical conllol in arice-maize cropping pattern in the Philippines J Econ Entoinol 761133-1138

135 Litsinger J A C de la Cruz F Raymundo A T Barrion M D Lumaban J P Bandong M S Venugopal F Paragna and E Balete 1982 Insect pests and insecticide reponse of dry-seeded bunded rice Pages 179-193 in Report of workshop on cropping systems research in Asia International Rice Research Institute Los Bafos Philippine

136 Litsinger J A M 1) Luinaban J P Bandong A T Barrion W Katanyukul and N Panda 1978 Insect pest management in rainfcd lowland rice agroecosystms of tropical Asia International Rice Research Conference 17-21 April 1978 International Rice Research Institute Los Bafios Philippines

137 Litsinger J A and K Moody 1976 Integrated pest management in multiple cropping systems Pages 293-317 in R I Papendick P A Sanchez and G B Triplett eds Multiple cropping American Society of Agronomy Madison Wisconsin USA

138 Loevinsohn M E 1984 The ecology and control of rice pests in relation to the intensity and synchrony of cultivation Ph D thesis University of London 36) p

139 Loevinsohn M E J A Litsinger N Panda M A Austria R F Apostol C Andrion and J Bandong 1982 A basis for predicting rice insect pest abundance in cropping systems Paper presented at the 1982 International rice research conference on rice and cropping systems 19-23 April 1982 International Rice Research Institute Los Bafios Iaguna Philippines 25 p

140 Lopez A W 1930 Repot of the Entomology Department Pages 145-172 in Annual report of the Philippine Sugar Association 1929-30 Manila

141 Lopez AW 1931 Entomology Department Pages 227-266 in Annual report of the director or research 1930-31 Philippine Sugar Association Manila

142 Lopez A W 1932 Entomology Deparment Pages 252-279 in Annual report of the Philippine Sugar Association 1931-32 Manila

143 Lungibill P and G G Ainslee 1917 The lessercornstalk borer US Dep Agric Agric Res Serv Tech Bull 539 27 p

IRPS No 123 January 1987 39

144 McGuire J U Jr and BS Crandall 1967 Survey of insect pests and plant diseases ofselected food crops of Mexico Central America and Panama U SDepartment of Agriculture International Agriculshytural Development Service Agricultuial Researrh Service October 1967 11 p

145 McNeill Sand T R ESouthwood 1978 The role of nitrogen in the development of insectplant relations Pages 77-9 in lliochemical aspects of plant and animal coevolution J B Ilarbornu ed Academic Press London UK

146 Maebara H 1955 Spatial distribution of lastinstiarlanaeof purplish stem borer (Sesamia iqferens Walker) hibernating in an upland rice field Bull Kagoshima Univ Fac Agric 438-41

147 Mallick E H and N G Hajra 1977 Response of rice plant to defoliation treatments at different developmental stages Indian Agric 2151-56

148 Mansingh A 1971 Physiological classification of dormancies in insects Canl Entomol 103983-10)9

149 Martell C G 1974 Catalogue of plant feeding insects of Mexico [in Spanish] Fitofilo 27(69)1-176

150 Martinez 0 J 1982 Principal rice pests in Ilanos orientalis [in Spanish] P~ages 80-94 in Curso deact nalization de conocimientos en arroz Villaniencis Instituto (olonmbiano Agropecuario

151 Martins J F da Silva F Ferrcira A S Prabhu F 1 P Zimmermann 1980 Use of pesticides for the control of the main subterranean upland rice pests esqui Agropecu Bras 1553-u

152 Martins I F da Silva F Fe eira rod BdamSilveira Pinheiro I Q2 Damage simulation caused by defoliators of upland rice [in Portuguese with English summary] Pesqui Agropecu Bras 171113-1119

153 Martins 1F da Silva N(iyuen van Tan and 1Ida Silveira Pinheiro 1981 Stem borer resistance in upland rice associated with plant morphological characters [in Portuuce Iesqui Agropecu Bras 16187-192

154 Masaki S 1980 Sumnmer diapause Ann Rev Entomiol 251-25 155 Masuzawa T 11 Suwa and F Nakasoji 1983 l)ifkleences of

oviposition preferencc and survival rate of two skipper butterflies Parnara guttata and PIchlpidai iathiais (Iepidoptera HIesperiidac) on rice plant and cogon grass New Entomol 3247-56

156 Maurya D M and C P Vaish 1984 Upland rice in India Pages 69-92 in An overview oftpland rice research Proceedings ofthe 1982 Bouakamp Ivory Coast upland rice workshop International Rice Research Institute IO Box 933 Mnila Philippines

157 Menon M G R R N Katiyai S Kuar and S S Misra 1970 Termites attacking the paddy crop Sci Cult 36294-295

158 Miyahara Y and 1 Nishikubo 1967 Behavior of young nymphs of the red rice root aphid Iholahsipiun ri liabhdontinalisSasaki in relation to movement from the leaf to the roots of upland rice [in Japanese] Pioc Assoc Plant Prot Kyushu 1363-66

159 Mo ljoa Tjien 1952 White grub of upland rice (Holotrichiahelleri Brsk) [in Indonesian with English summary] Contrib Agric Res Stn Bogor No 13134 p

160 Mochida 0 and T Suryana 1979 The occurrence of the brown planthopper and ragged stunt on lowland and upland rice ill Central Java and logyakarta in tile late of January 1979 Entomnolgy Survey Report No 15 Central Research Institute of Agriculture (CRIA) Sukamandi Indonesia I February 1979 9 p

161Monet C 1979 Varietal resistance to upland rice pests [inI French] Pages 35-39 in Annual report 1977 Institut des Savanes Department des Cultures Vivrieres Bouak6 Ivory Coast

162 Monte 0 1942 A caterpillar ofrice-fields[in Portuguese] Biol6gico (Sao Paulo) 8161-163 (Rev Appl Entomol A31414)

163 Munroe I) D 1979 Pest management strategy for hill rice in four resettlement areas in Sarawak Malaysia Pages 89-102 i1 Guidelines for integrated control of rice insect pests Food and Agriculture Organization Plant Production and Protection Paper 14 Rome Italy 115 p

40 IRPS No 123 January 1987

164 Nakasuji F M Ishii and Jen-Zon Ito 1984 Rice skippers in Taiwan and their life histories ZZrC (Taipei) Tech Bull 82 10 p

165 Navas I) 1976 Fall armyworum in rice Pages 99-106 il Ecological animal control by habitat management Proc Tall Timbers Con-ference 28 Feb-I Mar 1974 Fall Timbers Res Stn Misc Publ 6 Tallahassee Horida USA

166 Nickel 1 1 1961 Biological control of rice stem borer A feasibility study Tcch Bull 2 International Rice Rcarch Institute PO Box 933 Manila Philippines IIl p

167 Nigan P M and R A Vernia 1985 Insect pests of uplaild rice in Uttar Pradesh Int Rice Res Newsl 10(6)22

168 Nilakhe S S 1985 -cological observations on spittle bugs with emphasis on their occirence in rice Pcsqtui Agropecu Bras 20407-414

169 Nixon G EJ 195 1 lie association of ats with aphids and coccids Commonwealth Iusti lcte of Entonology Iondon UK 36 p

170 Nurdin F an Ishak Manti 1985 Study oit seedling fly of upland rice Interlational Upland Rice Moitoring Tour Report 26 lebruary-2 MarLI Sukaranti West Sumatra

171 Oka 1 N 1979 Ciltural conttol of the brown planthopper Pages 357-369 in Brown planthopper threat to rice production in Asia International Rice Research Institutc 1p0 Box 933 Manila Philippines

172 Ohni M and A Vilnaii 1975 Rice insect pests of Italy biology of Sipha grcria (Kaltenbach) and its control [in Italian] Riso (Milan) 2459-71

173 Otanes F Q atnd 1 I Sison 194 1 Pests of rice Philipp J Agric 12211-259

174 Painter R I 1951 Insect icsistance in crop plants University Press of Kansits Lawrence Kansas USA 520 p

175 Pantua P C and 1A litsinger 1981 Arthropod abundance in continuous vs biannual rice cropping systeis Int Rice Res Ncwsl 6(2)20-21

176 Pathak M ) and V A l)yck 1975 Sttudics ott insect pests of upland rice Pages 186-191 in Major research in upland rice International Rice Research Institute PO Box 933 Ilanila Philippines

177 Patil B R B M IHasabe and 1) S Ajri 198 I Chemical control of white gritb lIditriihinhorraia Fabr intfesting groundntut paddy atnd sorghum Indian 1Entomol 43232-236

178 Pawar A 1) 1974 (ollection and identification of insect pests of rice rlerminal repirt lInternational Rice Research Institute PO Box 933 Manila Philippines 72 p (tnimco)

179 IerrajU A aid A S Reedy 1965 Observations ott diapausc itt tie steti borer ohrice Rice News Teller (lai-Mar)9-13

180 Pont A C 1972 A review of the Oriental species of theri ona Rondani (I)iptera Muscidac) of econtomic importance Pages 27-102 in Control of sorglium shoot fly M G lomtani and W R Young eds Oxford atd IH1 Ptthl Co New I)clhi India 324 p

181 Prasad S C 1PChand aid M 1PSingh 1983 Incidence of rice gall midge in the upland plateau region of Hihar Oryza 20235-236

182 Rai P S 198 0 Nisaga simpex damage to rice in the hill tracts of South India Int Rice kles News[ 5(I)12

183 Rathorc Y S 1969 Gonocplhahli dcprssunt Fabr as itpredator of (tilo zonwIlhis (Swinhoc) and other moths in field cage at Patnagar Uttar Pradesh Indian 1Entotnol 3127

184 Reed W 1983 Crop losses causcd by insect pists in the developing world Proceedings 10th Internatial Congress of Plant Protection lrighton UK 174-80

185 Reedy M V 1981 Ateactogasterinhens(Walker)a new root-grub pest of tipland paddy and inaize Curr Sci 501076

186 Reveno M A 1968 Catalogue of the insect pests of economic crops in Colombia [in Spaiish] Asociacion Latino Americana de Entomo-logia (AIAE) 129-33

187 Reyes G M 1957 Rice dwarf disease in the Philippines Food and Agriculture Organiiation Plant Prot Bull 617-19

188 Rice S E A A Grigarick and M 0 Way 1982 Effect of leaf and panicle feeding byarmyworm (Lepidoptera Noctuidae) larvae on rice grain yield Environ Entomol 7593-595

189 Risch S J 1) Andow and M A Altieri 1983 Agioccosystem diversity arnd pest control data tentative conclusions and nizw research dirctions Environ Entomol 12625-629

190 Rivera C T 1956 Three organic insecticides in thecontrol of insects affecting upland rice Philipp Agric 39465-472

191 Roffrey 1 1972 Locusta outbreaks in the Philippines Acrida 1177-188

192 Rojas R 1) 1941 The effect of pruning at varying ages upon the yield of Ramairio Phipp Agric 29851-860

193 Roonwal M i 1979 Ternite life and termite control in tropical South Asia Scientilic Pnbl odhpr India 139 p

194 Rosetto C1 F da Silva Martins N C Schmidt and L EAzzini 1978 Damage caused by pasturespittle bugs 4Dvoi)lavopictaandDJ schach) on rice lBragantia Nota 6 (Matcli)25-27

195 Rosztto C1- S S Neto I- Amantc I) Iink 1Garcia-Vieira D M de SoLza N V Bansa t and A M de Olivcira 1971 Rice pests of Brazil [in Portuguese] Pages 149-238 inrFA()-International Rice Commission 2nd meeting of the Rice Committee for the Americas 6-11 l)ecember 1971 Pelotas Rio Grande do Sul Brazil

196 Rothschild G H L 1969 Obseratiois on the armyworm Spodoptera matritia acronrctoiiles Gn (Lep Noctuidae) Bull Entomol Res 59143-160

197 Rothschild G 1- L 1974 Parasites of the armworm Spolopiera mauritiaacroncitoihes in Malaysia Fntornopliaga 19293-299

198 Sachan i N S K Gangwar and 1N Katujar 1980 Field crickets damaging cages in North Eastern IHill Region Indian J Entomol 42526-527

199 Sands 1) P A 1977 The biology and ecology of Lepocorisa (Hemiptera Alydidac) in Papua New Guinea Res Bull 18 Departshyment of Primary Industry Port Moresby Papua New Guinea 104 p

200 Sarawak 1968 titomology division Pages 97-106 in Report of the research branch of the Department of Altriculture 1966 Kuching (Rev AIppl Eintomol A56603)

201 Satpathy JM M S Das and K Naik 1977 Effect of multiple and mixed cropping on the incidence of scine important pestsI Entomol Res 178-85

202 Sauer IHF G 1939 Notes Ott Elasmpnlpualtsligniosellths Zeller (Lep Pyr) a serious pest ofcereals in Sao Paolo [in Portuguese] Arq Inst Biol 10199-206 (Rev Appl Entotnol A28378)

203 Satinders 1L A 1 SKing and C L Vargas S 1983 Crop pests of Central Anerica with t bibliography [in Spanish] Centro Agroshynotnico Tropical de Investigacion y Ensenanza Turrialba Costa Rica (CATIE) Tec Bol 9 90 p

204 Sauphanor B 1985 Some factors of upland rice tolerttce to stem borers in West Africa Inseu Sci Appl 6429434

205 Sen P atnd S Chakravorty 1976 Stentflics in Haritighata Nadia West Bengal India [it Rice Res Newsl 1(12) 15

206 Senapai B and 1M Satpathy 1983 The seedling fly Atherigona orrzae Malloch is a pest of rice in Orissa (India) J Entomol Res 777-79

207 Shhjahan M S Alatn M Ah nad and BN Islam 1983 A new scarabaeid beetle pest of dryland rice in Bangladesh Int Rice Res Ncwsl 8(I)16-17

208 Sharma G L S C Modgol and R C Gautam 1979 Effect of depth of submergence ott incidence of bacterial blight and white grub infestation in transplanted rice Int Rice Res Newsl 4(6)12-13

209 Sharma S 1985 Scarabaeid beetle outbreak itt upland rice Int Rice Res Newsl 1(1(l)24

210 Shiraki T 1917 The paddy borer Schoenobhits incertulas Walker [achaku Agric Exp Stn Formosa 256 p

211 Silva A de Brito and 11P Magalhaes 1981 Insect pests of rice [in Portuguese] Circular Tecnica No 22 EMBRAPA Centro de Pesquisa t ecuaria de Tropico Unido (CPAJU) Belem Brazil 14 p

212 Singh S A 1979 Emergence of rice leaf butterfly as a rice pest in Manipur India int Rice Res Newsl 4(4)16-17

213 SinghS A 1983 Changing trends of insect pestsof rice in Manipur Int Rice Res Newsl 8(f)12

214 Siwi S S and P H van Doesburg 1984 Lenovorisa Latreille in Indonesia (Heteroptera Coreidae Alydinae) Zool Meded (Leiden) 58117-129

215 Soejitno J 1977 Reaction of some upland rice varieties toseedling fly eltherikonaexigua Stein [in Indonesian] Siri Hana I Penyakit 630-35 Lembga I osat Fenclilian Pertanian Bogor Indonesia

216 Socnardi 1967 Insect pests of rice in Indonesia Pages675-683 in The major inse pests of thu rice plant Proceedings of a symposium at the Internatital Rice Rec Wi itt S pternber 1964 The Johns Hopkins Press Baltimore Maryland USA

217 Soto P E A T Petcz and I W Buddenhagen 1976 Survey of insect pests and diseases of rice in different ecological zones in Nigeria Rice Entomol Ncwsl 435-46

218 Soto 11E and Z Ziddiqi 1976 Screening for resistance to African rice insect pests Varietal improsetnent sCmi n r Bouak Ivory Coast 13-17 Sept 1976 West Africa Rice l)evelopment Association Monrovia Iiberia

219 Squire F A 1972 Entomological problems in Bolivia PANS (PestArtic News Sumr) 18249-268

220 Stapely 1 II 1978 The rice leaf roller in the Solomon Islands Int Rice Res Newsl 3(l)13

221 Suenaga H 1937 On aphids attacking the roots of rice Oyo-I)oubts Zasshi 9150-152 [in Japanese] (Rev Appl Entomol A25745)

222 Suenaga H 1952 On the seasonal abundance of three subterranean aphids on upland rice Bull Kyushu Agric Exp Sti 1249-262

223 Sutynso Yas and Dandi Soekaria 1964 Beberapa hana yang ttenperang tana man padi tanah kering [in Indonesian] Rapat kerja Padi Tanah Kering 6 sd 8 Juily 1964 Kaliurang 9 p

224 Tallamy I) W and R F Denno 1983 Alternative life history patterns in risky environments in example from lace bugs Pages 129-147 in Insect life history patterns R F l)cnio and If Dingle eds Spring-Verlag 225 p

225 Tamura 1 1939 Observations on some insect pests of field crops [in Japanese] 1 Plant Prot 26414-420 (Rev Appl EntomolA27-556)

226 Tamura and Suzuki 1964 Influences of the early-transplanting practice in rice culture upon development of the rice stem borer Chilo siql)ressalisWalker Bull Ilokuriku Natil Agric Exp Stn 761-94

227 Tan J P 1924 The rice root aphid Drowhi ihirsoia Philipp Agric 13277-288

228 Tanaka 1 1957 Taxonomy and distribution of some subterranean aphids injurious to upland rice in Japan with description of a new species [in Japanese English summary] Botyu-Kagaku 22168-176

229 Tanaka T 1961 The rice root aphids their ecology and control Spec [full College of Agriculture Utsunomiya University Japan F3 p

230 Tauber M IC A Tauber R Nechols and 1Obrycki 1983 Seasonal activity of parasitoids control by external internal and genetic factors Pages 87-108 itt I)iapause and life cycle strategies in insects V K Brown and I Ilodek eds )r W Junk The Hague Netherlands

231 Taylor W F 1972 Effects of artificial defoliation (simulating pest damage) on varieties of upland rice Exp Agric 879-83

232 feotia T P S and S Natud 1966 Biotoomics of the rice skipper Parnaraniathiis Fabricius (Lepidoptera Hesperiidac) Indian J Entomol 28181-186

233 Teran F 0 1971 Rice pests of Santa Cruz [in Spanish Ministerio de Agricultura Sita Cruz Bolivia Bull 45 8 p

234 Than Itun 1976 Population dynamics of the yellow rice borer Tryporza itcertulas (Walker) and its damage to the rice plant MS thesis University of the Philippines at Ios Bafios Laguna Philippines 92 p

235 Uichanco L B 1911 Pests Pages 55-74 it Rice Philipp Agric 11-200

IRPS No 123 January 1987 41

236 van der Laan P A 195 1 Life history of the rice seedling fly Contrib Gen Agric Res Stn Boor Indonesia 118 15 p

237 van der Laan 11A 1959 Correlation between rainfall in the dry season and the occurrence of the white rice borer (Scirpolhaga innotala WIk) in lava Entomol Fxp Appl 212-20

238 Van Driesche R G 1983 Meaning of percent parasitismin studies of insect parasitism Fnviron Entomol 121 I1- 122

239 Venugopal M S and A litsingcr 1984 Fffect ofcarbofuran on rice growth Prot Ecol 7313-317

240 Vietnam Directorate of Rural Affairs 1962 Annual work progress report on crop improvement program of rice sugarcane vegetable and field crops (lor the period from July 1961 to June 1962) (11) 299 p (Rev Appl Entomol A51526-527)

241 Wan M T K 1972 Observations on rice leaf- and planthoppers in Sarawak (Malaysian Borneo) Malay Agric 148308-335

242 West Africa Rice I)esclopnient Association 1978 Annual research report Vol 2 Statistics Monrovia Liberia

243 West Africa Rice Development Association 1979 Annual research report p47 Monrovia Liberia

244 Williams ) F and CS lofgren 1981 Fli I illy l-468 anew bait toxicant for control of the red imported lire at Fla Entomol 64472477

245 Williams I) 1 1970 The mealybugs OFlomoptera Coccoidea Iscudococcidac) of sugar-cane rice and sorghum Bull Entomol Res 0109-188

246 Williams D) 1 1982 Pulinaria icervi (Signoret) and its allies ott sugar cane and other grasses Btll Entoinol Res 72111-117

247 Wirjosuiardjo S 1976 Study ofchenmical cont rot of insect pests ott upland rice Imiu Pertanian (Jogjakarta) 1323-329

248 Wirjosuhardjo S and N Wirarta 1976 Chemical control of the white grub tIolorithiatlltriin upland rice in M uneng Irobolinggo Indonesia) [in Indonesian] Imut Pertanian (Jogjakarta) 1330-332

249 Yano K T Miyake and V F Eastop 1983 The biology and economic importance of rice aphids (Henuiptera Aphid idae) a review Bull Entomol Res 73539-566

250 Yelp D and P Rhodes 19(4 lhe success of drilled dryland rice iii the Fiji Islands International Rice Commission Working Party on Rice Production and Protection Manila Philippines 3-10 March1964 Food and Agriculture Organization Rome Italy 13 p

251 Young W R 1981 Fifty-five years of research on the sorghum shootfly Insect Sci Appi 23-9

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No 104 Faridpur acomputer-assisted instruction nmodel forrainfed lowland random coefficient model No 116 Consumer demand for rice grain quality in Thailand Indonesia andrice

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No 106 Rice grassy stunt virus 2 a new strain of rice grassy stunt in tire literature

Philippines No 118 IRRI-Korca collaborative project for the development of cold-No 107 Physical losses and quality deterioration in rice plstprodnCtion tolerant lines through anthei culture

systemis No 119 Problen soils as potenial areas for adverse soils-tolerant rice No 108 (olptblication ofIRRI materials a survey of translators and varieties in South and Ssuthcasi Asia

publishers No 120 Changes isinall-farm iLtireliing techii)hgy in Thailand and [he No 109 Classification of Philippine rainfall patterns Philippines No 110 Contributions of modern rice varieties to rutrition in Asia No 121 Landforms and modern rice varieties No Ill Changes in rice breeding in 10 Asian countries 1965-84 No 122 Yield stability and modern rice technology No 112 Design parameters affecting the performance of the IRRI-designed

axial-flow pump

Page 23: UPLANDRICE. INSECTl--1i M T -E!R.EOLOU t RTANCE ,AND …delphacid.s3.amazonaws.com › 6958.pdf · The significance of abiotic factors in upland rice insect ecology will be apparent

Table 4 Yield losses to insect pests and yield responses to insecticide in upland rice in the Philippines Brazil and Upper Volta

Yield (tha) Yield loss (5) Location Year Cultivar Protected

with Protected

without Check Total Vegetative Reproductive Ripening

Basal N (kgha) Reference

carbofuran carbofuran Protected No with carbofuran

carbofuran

Philippines Pili Camarines Sur Tanauan Batangas

Pangantucan Bukidnon

Dumarao t-apiz

BrazilGoias Minas Ge~ais

Upper Volta6

1975 1974 1974 1975 1975 1976 1977 1978

1979

1980

1979 1980 1979

1977 1978 1979

Bursiging puti Kinanda Dagge Kinanda Dagge Dagge Dagge Dagge UPL Ri5 Dagge UPL Ri5 Dagge UPL RiS UPL Ri5 UPL Ri5 UPL RiS

IAC47 IAC47 IAC47

14 33 11 36 35 31 30 27 37 26 40 30 40 42 46 35

19 21 09

--

-34 24 24 27 --

29 43 31 42 -

47 25

---

10 28 05 27 27 30 26 29 33 28 33 29 43 23 36 11

15 15 06

29 15 55 25 23

3 13 -7 11

-7 18

3 -7 45 22 69

24 28 35

--

21 -11 -20

4 -

-3

23 7

-2 -23 56

-

-

-__0

6 --

-

-23 14

5 0 8 2

22

-

--

0

-

--18 11

3 -10

4 10

7

--7 -

14 -

-1 6

10 -5

8 9 7

-

--

0 0 0 0 0 0 0 0 0 0 0

30 30 30

12 12 12

99 99

100 100 101 101 102 102 102 102 103 103 104 104 104

66 66 66

Farako-Ba

Mean

1977 1978

IRAT10 IRATIO

38 66

--

40 43

-5 35

21

-

10

--

--

--

- 242 243

Z

tJ

-w

24 IRPS No 123 January 1987

Philippines which had two treatments offering protection In the 9 Philippine trials where yield losses were measuredwith and without carbofuran granules relied on further by growth stage 10 of the yield loss (21) occurred at thecontrol with foliar spray and not on seed dressings The vegetative stage and 517 occurred at each of the reproshycarbofuran-protected plots had an 18yield los compared ductive and ripening stages Ibis result is similar to thewith only 10 with foliar sprayS This yield difference is pattern of yield loss in lowland rice measured by successiveprobably due to differences in protection against sown-seed treatments where the crop is unprotected at each growthpests (foliar spray do not control sown-seed pests) rather stage but proticted during the other growth stagesthan phytotonic effects became basal fertilizer wa used in However the insect complexes r sponsible for vegetativeonly 2 of the 10 comparisons stage yield losses in upland rice are entirely diftfrent than in

The recorded yield losses were not related to yield lowland rice The key vegetative stage pests in lowland ricepotential (Fig 3) One might expect higheryielding crops to arc aquatic whereas those in upland rice are sown--seedhave higher percentage of yield losses but this ws not tile (sown in soil) root and seedling pests This dift rence maycase One reason is that insect damage exacerbates plant explain why agricul torists have overlooked tile ilmportanccinjury from drought stress low yielding fields from d ought of upland rice insect pests Many reports of upland rice pestsstress would show disproportionate differences between in Asia ha Ve focused on major lowland species (46 108)insect-protected and -unprotected treatments Shoot and It isalso surprising that tei vegctative stage of upland rice root pests would be more responsible for losses from would record the highest yield losses becatuseseeral studiesdrought stress than sown-scd pests A second reason have shown that upland rice can readily recover from foliageconcerns seed pests which would have a greater impact on reinoval during that stage (117 152 165 188) Upland ricelow tillering low yielding varieties such as )aggc Low poscsses a hitgh obilitV to compensate for ca rly loss oftillering varieties nnot fill in the space created by removal foliage and will even be stinulated by foliage removal toof seeds oweer 1) R i 5 is high yielding because it tillersP1 produce a higher viel than plants ithout any foliageactively which can close the canopy to provide greater weed removed (63 192 231) [ie reason for this apparentcontrol discrepancy hovce is that yield loss at the vegetative

stage does not octUr frol leat arca loss but fron pestsYield loss () removing sown seed and feeding on devcloping tillers and roots -- damage that cannot be readily conpLtnsatCd for

70 - Measuring yield losses in upland rice cnvironrnents is further complicated by a report from Brazil that greater yield losses occurred in response to better insect control60 - =-09 as Insecticide seed treatments resuled in denser plant stands

(n=21) which in turn created a selectively more favorable environshyment for blast (69) This resuIt was parlticurla rlxexacerbated50 when cirbofuran granules were used in seed furrows Aside

ffrom a greater plant stand ca rbofuran iay have auigmented the N lecl and stimulated blast If so ca rbofuran should be40 - replaced with another insecticide or the seeding or fertilizerI rates decreased

Other methods to determine the impact of upland rice30 shy pests wvcrc to interview farners or to develop single speciesbull bull correlations of population levels and yield loss Feneira (66)2 NIM)st (75(1) Golas farmers reporteddid both in Brazil

20- insect losses in ufpland rice and 251 said those lossesS avcragcd 37(1 [his result is consistent with tie yield losses 0 0 measured by field trials in the same statc(Table4) For every

I0 4 of seedlings removC d by leaf-cutting ants a I 7yield loss is predicted Similarly Ferreira (66) found that ai

0 - average of 53 thrips per panicle before panicle initiation meant 3 times more unfillh d grains

-IO0 I 1 1 I I-101 0 2 3 4 5 6 7

Yield (tho) C)N [ROt IM FI IODS 3 Correlation between upland rice yield and yield lols caused by Upland rice insect pests may be controlled by culturalinsects measured in 21 trials in the Philippines Ivory Coast andBrazil genetic biological and chemical methods

IRPS No 123 January 1987 25

Cultural control dry season plowing will have little effect on them Plowing Crop husbandry to reduce upland insect pest populations when the soil is wet from the first monsoon rains will expose can be classified into practices effective in a single field and soil insects that tunnel close to the soil surface to predatory those effective only at the community level Cultural birds chickens dogs and even man (field crickets nole controls should come first in a pest management program crickets and white grubs a re delicacies in man tra ditional They have broad stable effects as pests have little pos- diets) sibility of overcoming them through selection of biotyps The greatest degree i1 inIscct control fron tillage comes (138) On the other hand some cultu ral controls work only from plowing soon after rice ha rvest when most soil insects with high labor or power inputs and connunity-wilt lie close to the soil surface anrd stern borers are in the rice methods require coordination of iany farniters stuble Whitegrubsare mature and tillage exposesthem to

Pantin time The luxury of being able to shift the time of predators Tirnelv control will prtec the crop following pla ntring upland rice is open onl hr la rniers in areas with a rice and lowr the populationIfor succeedinig years Plowingprolonged rainifall pa ttern lI copping intensity or underthe ice stulble helps it decompose killing sten borer rnechanied land preparatiot Gcnerally planlting as soon larvae inside as the rFiinv season begins will lower populatiois of 1110st [requent tillage destroys ait nests bit not termites insect pests Insect populationsarelo aftcra dlvseoll or Tillage and clean culture howes er rcliove food for winter fallow delayed plaiting lets them build tip on termites which then mtay attack a young rice crop Scarab alIternate hosts that gi vigorotsly with the first seasonial beetles also prefer to miposit in eceitly tilled fields Noshyratiris This first flush iof weds cros lItxuttriartlv with iceCult is dibbling faors the survival oftillace Clre Such rineralied N released during the dr season aid made stils i aels tnd insects thtt pttpate or lay eggs- in the soil

available to the plants with the rains It iliso igrtmitcally Zero tilla hio eerl conserves predators ofthe GIH (37) advanttgeous for the rice ciop to tap this trlLItlaI fertility nt it mcttoi lo lessen the impact o0 soil insects and grow vigorotsly to compete with wrus and to tolerate brotdcasting seed or planting in ftltrrows is prefc ired over pest infestation dihlirig in hills Crickets are large enotlgh to eat seedlings in

Larly plantngs terd to escape seedling maggolts Upland a hill atrd tillering normnally ca iriot fill in tlhe missing space rice in Bitangas is planted in Ma and ine itli the Iirst Oftcn dibbll rd is not well covered and iseasily found by rains Fairners are highly mtotiated to plant early tocscape foraging insets ioIdents and birds drlitght and typlotns ricar harct Arn Atrgtst planting Plan uit sii Incrcatsini sced density protects tile cropwould be evcely attacked by It riim orivzo At tti againrst seed and seelling pests especially ants particularly other Philippiie tipland sites Rea l Qteot antd (laelit if the crop will germinate and ermerge quickly M isainis Oriental seedlig mnaggots arnrige ric boe t se hr trftinll itI India rice intcrclopped witi cotton or farnes have a longer growing season and plnit 2 rin or pigeon pci had lower 6IIt and WBIlII poptrlations than mote after the ortset of the rainy season liaaiangas ftrmners rice alone (211) NfailC arid Urplald ir_ 2re coilltori in the plant ca rly becia use ofa shorter raiinfall period arnd the desi re Philippines he chatngC in pest status folii iniercropping is to lliarvet before the typhoor sea son peaks ltighly locatioll-specific (133) cmianarid the net elfc be either

Early plantings cornbind with early-iatiuring varictics nil bcnelficial ortdetriential (189) For example BIatangias provide higher crp tolerartce for white grubs Aestiatior farmers intercrip Itro in their ipland ricefields A liornshyterminates with the first rains but 5-6 moare ineeded for the nrii iotd-Ahis i (I iiiacius) which norlially feeds resting larvae to develop into beetles emerge and lay eggs on taro ill feed or miarbv rice plants Intercropping and for larvae to reach the danriigii g ird-instar stage probably would not afft0 seed pests A crluanion crop (134 In lataigas this normually iseiough tiite for harvest planted ith rice and haing gleater tolerarnce for root

It Brazil howecver delayed planting is a suggested dariage or being toxic nayact isi trapcrop to root-feeditig cultural control for the lesser cornstalk borer and insect pests Acial-fei i ig pests voultId be most affected by pests favored l drought (66 195) Cropping internsity in intrer ropping Shoot aphids leaf beetles sten borers and Brail is low and delayed plraing places the rice crop in the hopplers would be likely ca ntidates to note effcts On the niost stable rainfall period reducing likelihood old rought othlt halnd the corlipalior crop stch~ sitniaie Iiiv be more

In rilniy arca s rinmirsare roi atl to plant other crops benehited than rice from initecropping ( 137) before rice and althNgh it woultl be desirabh to avoid 11echi Weeuling luring the first month after clop infestations by planting rice cuatl lariners lack the labor cstablishent will force pcts such is anyvworris which and time to do so ol a low priority crp prefer grassy cetds ontor tle vorling rice Soil iampsecs will

77age Uplild rice Stils ae light textured and Call be mvnye to the rice crop Cleani citrltirc also lay lore territes plowed duiring the d rv seasor notl only to grin time for to attack I u cetincti inrice ceds i ay he le optinal planting with lie fitst Inilsoon riiains but also to desiccate solition 13) soil insects arid weeds such is ntsedge that foim rhizomes Irtlizr Itneatsing N Iciiliier oin hlwland rice has Htowever becta use most soil insects lie below die plow layer fAtwvored Irigher populains of pliithoppeiv Imlhoppers

26 IRPS No 123 January 1987

and leaffolde rs However because of the need to temper fertilizer use on upland rice o prevent blast fIrtilizer rates will be lower than is comonia r for lowlad rice causing relatixely less hopper and IaIfTfoide r population buildup However inupland rice areas without a history of blast such Is Batangas farmers apply high rates of N and conscqtrently the crop has high infcstation levels of WBPI]and RIF The fhrmrcrs leec insect buildup by splitting the fertili11r in0to two to three applicatiois bit do not applV fertilizer at the vegetati estage because this prodrces a tall crop which illeasily lodge during typhoons Greater crop fertility leads to greater insect survival (better nutrition) larger insects (to cat no re foliage) arid greater fec titd t (to lav nmre eggs) (128 145 171) Not all insect species respondequally hovr Increasing rates (if N l anrid K increased Dialraa s-acchralis and (iaeiurema flea beetle but reduced thrips (66) Zn rcdLuced deadheart density from the lesser cornIstalk borer

Fnldingi Although most Upland rice areas do nIot hawxcaccess to wa ter floodiig the fields is rRrrnriierdcd in areas of Brazil or root bug and white gib control ( 195)

Crop rotatioti Tillage before planting a crop after rc harvest will control stern boreis and other pests renain n the stubble arid will unearth soil pests Planting a non-gratriiaceous crop after rice is recommncrded for termite control (66) Crop rotation is less nttdcd in upland rice culture than in lowlands because atcrop-free period is assured even after harvest of crops following rice

t1ulchilt iryl plastic irli irig designed for weedcoitrol ii upland rice inJapan resulted in ntr highanri liy infestation of (ihilo sulytresvais (93 9-1 95) NIulched Upland rice grew more luxuriantlv and attracted more (Avipositing mothis than u lched rice Also the rice plant and stein borers imatoured 10rC quickly and a partialgeneratiori af vouing ista iNoverwintcred in the stubbleLotili--malitrt varieti Quickly maturing crops reduce the tmbCr of pCst generations that Calt build up Rapid crop establishment olan ca rl v-natu ring vaitvty if)pa 11icLa r will lessen white gib and stem borer damiage White grub larvae intle last iistar are highly destIruCtive so aii early-iiatri ng variety will escape serious root loss Stem borers build 1ip slowlyx in rice to ecormic abundant in a late-Maturing crop anrd caiuse whi tlieads Numibers of somespecies increase exponeitially with each succeeding genera-ti so a xarietyexhat atures I trowh earier thian another will hac less insect damage Early-naturing vaiicties however will not lessen seed pest daiage either ii the soil or oina standing crop

Strwhoi hAIJttam Inrscts tcd irig on thlie aerial portions of rice plats disperse from carly to late plantedfields AS iiiibCirs increase exponentially with eachinsect

gtscnrattm e eiglbrig f rmc~s1(uil upp wh rc generation pests build tIP where neighboring fihrTC e9r1sstagger their plantings II Batangas farmers plant uplandrice within I iro use 120-d s-arieties and plant at the start of the rainy season Ini Misarnis Oriental laricrs stagger their

plantings up to 3 mo apart and use a set of atieties that mature in 4-7 mo Insect pest danage particularly from stem borers is greater in Misarmis Oriental Also seedling maggots damage rice there because rainfall favors frequent planting of maic In Batangas rice escapes seedling maggots bCca usC of distinct drs season and synchron0us early planting with the onset of ains

Therefore combinations of cultural practices ea rlyplanting synchronous planing crop rotation and earlyshymaturing vaneties protect the rice crop against most insect pests

Ilmant resistance Beca tse upland rice has few specialized insect pests and many insect types attack the crop worldwide regional breeding and strong internatiornal cooperation are required

Sown-sced pests arc economically controlled by insectishycide seed treatment and should the relore receive lowxpriori ty inbreeding objectives

Root pests are normally cost ly to control with insecticides but finding resista t sonres mayvbe di fficult Tanaka (229) for exa -ple fi led to find varieties resistant to root aphids Selecting for large root bioniass perhaps should be the strategy for pests sLuch as Whlite grubs that renliove roots Fainter(174) howcvcr cites varietiesof crops tolerant ofor resistant t(i root pests Some sorghun varieties are high in cyanide so rice sorghun might incorporate broadshyspcctruri chemical resistance

Seedling vigor and drought tolerance are high prioritybreeding objectives so tupland rices should have large r root systernsand be able to tolerate ligherlevels of root loss Sonie root loss inolder plants is beneficial if it stimulates new root developnrut and therefore crihances intake of soil nutrients (53) The rice seedling inaggot rtiLotia spp may be

controlled thirough resistant aittics A hi rgc breeding effort on sorghniu has had fair success inideveloping arreties esrstait to Asoccata Rodani (251) A field trial comparing 10 upland ricc varieties showed differences aniong varieties va ring from II to 4(- damaged tilleis (215) Hovver resistarce to 1Iill-0r gtia species may rot cross ox-rti otier

Shiraki (210) reported that upland i-ice stens wcrc harder than lows-land rice stemsperhaps becaIusc (f a higher silicoricontent and thus were more resistant to yellow stern borer hc tortia Iow tillering (f uplaid iice xarneties prodtres a

higher percentage of infested tillers than would occor in higher tillering lowland iice under a siiilar egg density The corollary therefore is to select for high tilleriig upland rices

Stemii bor amcstheh main targes dobrecders inAfrica (571 204 )but nticl oifth e sc reen irig is dotie unde r lo wland co d t osrm s wih A h r~ o l pe i s n acorditions (11 96 218) As with lthrinrnwspecies mayhave to be dealt with independently however initial results show cross resistance between African and Asian species(92)

In Brazil screening Ior insect resistance focttses oil lou I pest problems Varieties tolerant of lasmolwalpus ligi-sellus and resistant to )iatraea saccharalis have been identified (70 153) Current work also emphasies 7iraca limativetntris and Thmialspis (=Iosis) flavouicta (66) These insect pests Ilt onlV citise IIigh yield losses tihey also ire difficult to control chemically ofr by other ilealls

ManV upland rice vaieties aic early maturing and are resistant to pests such as gall midge (I Xl ) Early mIturity is often a highly desirable trait to esuilpe pest buildup but it is not true resista nce bca usc ilthose va reties are plinted late they will be itsdamaged as ain early planted susc ible variety

P~erhaps tile cosiiiopolitall of all upland rice pestsimost are the sced pests (ripenling stagc) which pose a particular problem beta use scuds re ised IsIO for hitnit4ans Ayltoxic substatice introd tcld geticticllyilmay also be toxic to nal iniless it caln he placed old in thfe IdhuffSelectmi fo hairiness or lotng wns kill mIlct sthilrohleiiis dutilig threshing pa rtictfarl maitial threshinfl [hireshers ill complain of itcfimin atid itteascd (fust levels

Attempts itIRRI to fitid tcistatncc to the rice bug plorisa have ilotbeCnl prmitisilg lIIetIattonlids iorc

through the lenilna or palca hillhairs sedstVIor seeds wilh awns that deter bugs promote complaints from threshers Seed bugs liptocorisa and Stcnocoris enter the seed through the opening between tile lemna and palC but varieties with narrow openings are diifficult to mill

Breeding priorities should concentrate ott inotCI gruLtLIps that are most difficult to c nttrol by other meains Steil) horers appear to be the most widespread group that sigiiificantly dalag s rice aind is hard to manage by other Ilnca us

Biological control Very fe stutdic of mitatt~l eneties of rice pests in upland envirollnletits wee in tile wefouiid literature Therefore have not compiled a list of species Ve present a list of natural enemies iOtid in upland ricefields tuic latialln Batangas for some conitno iOldiar pest species ([itble 5)

Muich ofl thc lollowinle itfor ittiol cotlliCs rol Stutdies in the lowlatnds or roi crops otithcr thlin tice

Qu(ailtr o 11ll t11t( Nat trrtf lii hcoeits ight Ile used agaitist ile social insects tilt and termites Sollie predator specice live ill theirf nests n ilnic theirnllt ip-pearanIlCe aMRidbehavior IJtrouMdCing a mote aggressise ant species to displace pest ant codonies is a possibility (169) Patihogens as potential natural encnies have reccived little atteition (I13) Natural ncnuics asa group however hac not been toIl iporta it in Iegulating aintshown be and ternite numbers

The nocturnal behavior aid subteitmien habitat of lole crickets and field crickets protect them from many ntllral enemies There are only limited records of natural cnelics for these groups of pests some pathogens (87) but mainly

IRPS No 123 JattuIny I 271987

parasites - scelionids on eggsand sphecids on ilymphs and adults Sphecid wasps Larra carutmaria (SImitI hI L huzwuensis Rohwer and lsaltiuhta Willi ams speciali in mole crickets pa ralyzing them nd dragging the ninto thcir nests as food lot their voting ()tier sphCcid species Lins aurulenoa (Iabricius) lot maiac (Ashmead ) 1 lhshylssellaus (Smith) and Ml lal)oriosu (Snithf specialie in field clickets

White grUbsO the other hand hae Icti scriuns enuigh to have been extensively studied for cotttroi 1W nati al enemies( 112 140 141) Scoliid parasites C(ajsoMcispp have beetn intrtduced itto tilePhilippites for control of Ielxl ]udisiro la The rcsearchcts w Intttrdced themle during ill era claimed that the white gttionthreak as controlled (142) howevcr it isnormal for pest epidemics to subside and there was no diect evidence to indictte that tfe parasite was responsible A tmore promisitg titctllod might he to identify iso ltcttass prt~f tcartd apply hiactelaln fingal pathogens to tile soil Ihe lalattese beetle in North America has hcci coitrolled hv tite conmnleciall aailable hactcrial prcpa ration itacihlpu pav ofr milksdiscas (71) Spote prtyefott the soil itloted ilrtier reitiain actic fr Veas

Root aphid sardtnctly bugs readilslic tilib to predatots and parlsites once their tetndine ants are controlled The larvac and adtilts of a coccinellid beetle 1Scynms sp prey on llriadlouais in slash and burn upland rice Baits treated with insecticide may be used to control tending ants by killing their ourig (244)

Root-tkcditig blugs false wireoris wireorms and root wccils aic r ofsuch little ifilpoltatice that thenomttll role of liatural Cticniics hias tiot beei asscssed A possibility hoevrl tiglt he tit ItaIIitic niaCIMtodcs lintt1g11en Itese soil-itlhlfbitine mCtitodcs W5uhtfhave to be imiass Produiced

large miltibcr of niatltrll cnemics ircknown ft CltIoriS IC IpblCeii is thtt clttvortnlls ofonlie the crop soott alter land ptel)tritiOti a1tithe bcgit iniig of the raittv

Citsoln hfien tiatllral leniiiy populations aric lo Colltloil WOtuld iii ole mass prodtticing andirclcaslng key spccies

le egg stage is the fpirt (olthe Cedlitgt Ialot life cycle tI hich to f)ctis ftor tittlril elctit cottntrol hplraites o1 pleditors ttsit elleggs highly expoIsYCl I Sgg pailsites or predators stolld probahly e ittass andtbe produced released Littlh i toruialioll is a ailable ott what species to trV o ho to Iniss podLice thiCIi A etiloplhid wasp Icmihlaol sp effctiicly regushy

lates the leatf lcloalapolm2a spioanoiiitr heat andc rice antid should be collsctvcd

leaf beetles whCllose e am] eggs ate laid in the grutindhirO ould be motC difficutlt to control by niatulral enelV Mnll ipuhltiOn But species sUici IS c_gsthispid beetles ith

and larVle oilleascs ate vtilcratble to iparaites and pledators

Natural enemisc of tlirips otl rice are little known

28 IRPS No 123 January 1987

Table S Parasites recorded from upland rice insect pests in Tanauan Batangas Philippines 1977-84

Order Family Species Host stage

I lymttenoptera Braconidae Leaffolders Cnaphalocrocis Marasinia spy

Cardiochilesphilipplinensis Ashmead a Larva

Cotesia angustillasis(Gahan)Cotesia or cypris (Nixon)

Larva Larva

Cotesia or taeniaticornis(Nixon) Larva Cilonusinotakatae Munakata Chelonits SppAlacrocetrts philippinensis (Ashtmead)

Egg to larva Egg to larva Larva

Iclineumonidac Orgilts spp Isclutojppa hireator(Fabricius)

Larva Larva

Itoplectis naraigav(Ashmcad) Larva Tenelhtcha philippiensis (Ashmead) Larva Tenelchha stangli (Ashmead) a

irichomma cnaphalocrosis Uchida Larva Larva

Bethylidac Elasmidac

Goniozms or trianguli r Kieffer lasmtusalhopictus Crawford

Larva Larva

Diptera

Encyrtidac Chalcididac

Tachinidac

Elasmtus spp Copidosomopsisnacohiae (Fidy) Brach nteria evcarinata Gahan Biachyieria lasus (Walker) Ztgohothria ciliata (Wulp)

Larva Egg to larva Pupa Pupa Larva

Argyrophylx nigrotihialis(Blaranov) Larva

l-ymenoptera Braconidac Yellow stern borer Scirpopiaga incertulas

Braco chinensisSzeplegeti Larva CIthelotis spp Chelonus niunakatae Munakata Cotesi flaripes Cameron Stenoracon nicerilli (Bingham) a

Larva Egg to larva Larva Larva

Ichneumonidac Tropobracon schoenobii (Viereck) Amatroinorpiaaccepta metathoracica(Ashmiead)Frihorus sinicus (I lolmgren)

Larva Larva Larva

Ischnojoppaluteator (Fabricius) Larva Isotima nr dammeratiti Rohwer Larva Tenelucha philippinensis (Ashmead) a

Larva Tenehicha stangli (Ashmcad)Trichomma enaphalocrosis Uchida

Larva Lara

Chalcididac Eulophidac Trichogranmmatidae

Xanthopimnpla stemmator (Thunberg) Charops hrachYpterumn Gupta and Malteswary Brachtm eria sppTetrasticttus schoenohii Ferriere Trichorigranm chihunis Ishi

Larva to pupa Larva Pupa Egg Egg

Diptera

Scelionidac Pteromalidae Tachinidae

Tt ichograintnta aponiicuitt (Ashniead) Telenomits rowani (Gahan) a

Trichoinalopsis apattelctena (Crawford) Jeirhaea spp

Egg Fgg Egg pupa Larva

Z4gobothria ciliata (Wulp) Larva

Ilymenoptera lecneumonirlac Gol fringed borer (hilo auricilius

Trichomm enoptalocrosisUchida Larva

Ieioenoptera Braconidae

Aantholimlh steiniator (Thtinberg) a

Pink stet Ioret Sesamia inferens StoInacoil nicrilhi (Bingh am)

Loarva to pupa

larva

Diptera Ilyinenoptera

Armvtnyorms Mythimna separata Spodoptera mauritia acronyetoidesTachinidae Zvgohothriaatropiora( Robinea i -Desvoid y)Eulophidae It-phctrtiscitapadoe (Ashimead)

Larva Larva

(halcididae Braconidac

Brachpin ria spip (oesia spp

Larva larva

IHyntenoptera I)iptera

Rice skipper Ielop ida (halcididae Tachinidae

math ias aidgreen iortedci ciliarNIelan itis eda Brat lhincria sp iii matuqitatt Cameron Irgyrolhirlaxi trotlhiais(marumtv)a1

ismtne Larva Larva

Diptera Ilymenoptera

Tachnidae lelineutionidae

fuown se tilo per Mois frugalisIrgtrophrlax 1igrothiahiv 3 a rat ov) anthopimpa pmnctata (Iabricius)

Larva Larva to pupa

aMost dominant

IRPS No 123 January 1987 29

Normally predatory thrips prey on herbivorous thrips strepsiptera on both Spiders dominate tilenymphaladultNatural enemies of stem bugs --Scolinoplara and Tibraa predators Spittle bugs move to rice from pasture grasses - are in order of importance egg par~isites and predators and biocontrol would be successful only ifcarried out before fungal pathogens and nymphal adult parasites Egg tiley immigrate into ricefields parasites could either be introduced fron other areas or Biological control tactics for hemipterous seed bugs mass produced and released ould follow those outlined for stein bugs

Armyworms are normally lwld in hcck by the activities Coml)arison of environnewtns Natural enemies are of egg and larval parasites When these natural enemies fail perhaps faced with even greaterconstraints than insect pestsusually because of drought armyworns become epidemic to survive ind reproduce in upland environments TheyIt may be worthwhile therefore to release parasites during have adapted ani survived by having wide host ranges the the rainy months following a drought Virus diseases of ability to aestivate or hibernate over unfavorable seasonslarvae would beanotheravnue to exploreforarnyworms and the ability to disperse or by other mechanisms

Parasites have ben traditionallyconsidered for biolegical Comparing the most dominant and therefore the most control attempts against stein borers usually by rekasing adapted species of natural enemies collected in various exotic species( 166) This approach has not met with success environments in the Philippines we can understand how and new avenues should be explored The role of predators different rice environments are arid pathogens is little inderstood and deserves greater Anagqrms olptahiisa mymarid planthopper egg parasiteattention Egg parasitism rates are normally reasonably is cquallyadapted to all Philippine rice environments (Tablehigh but should be supplemented with effective egg 6) It is selective for planthoppers and even parasitizes eggspredators Orthopterans are potential egg predators of the n-iize planthopper Peregrinitsmaidis itt upland areas Metioche vittaticollis (Sta ) Anavipha nifiwnnis (Saus- Alavohis also parasitizes maize planthopper eggs but is sure) and Conocephahs hongipnni (de i-laan) feed on most prevalent on BPH in the irrigated wetlands (onashystem borer eggs etioche and Anaxil)ha specialize in eggs tocerus spp like A optalhilis on BPH and WBPI Iattacks with no hair such as Chilo spp The omnivorous (ono- the eggs of GLH in ell rie environments It is also found cephahsspecializes in eggs cov red with hairs it is a proven parasitizing the eggs ef the white rice licahopper (oimna egg predator of Scirlouphaga spp but has the discouraging spetra Oirosita naias a trichogrammatid egg parasitehabit of eating rice guin (65) attacks BIPH and WBPH in the wetlands and has no other

Grasshoppers have egg (scelionid) and nymphalad ult host A related species 0 aesopi specializes in G1-H in (nemestrinid tachinid and sarcophagid) v irasites Among wetland environments Dryinid strepsipt-an and pipunshypathogens protozoans have been recorded most frequently culid nymphal and adult parasites of planthoppers and Control of locust species by natural enemies could con- leafhoppers also specialize either in hopper species or centrate on their habitual breeding areas environments Among the dryinids Pseuhdgonatoipsflavi-

Leaffolders have rich complexes of natural enemies some feimur prefers the lowlands and BPH while P nuch has no adapted to upland rice (22 27) ranging from egg predators environmental prefirences but is adapted only to WBPH (gryllids coccinellids) to larval parasites (braconids ichncu- The main d ryinid species on GLH aptogonatop)usspp is monids) to larval prcdators (ants and carabids) to larval most dominant in rainfed lowlands and also attacks other pathogeiis (vira and fungal diseases) Efforts to augment leafhoppers within Amnrasca Cicadulinaand Bachaltuathese natural enemies should focus on the egg predators Strepsiptera are more dominant on plarithoppers than (rearing releasing)and viral or fungal pathogens (culturing lealhoppers The Elenchus species in the Philippines attack spraying) all species of planthoppers within the genera Sogawilla

Parasitization of the large conspicuous larvae of Sogatodels NilaparsataHarualia and Opiconsiva Elenshyskippers green horned caterpillars and other polyphagous cusrvasuntaisuiis adapted to the major rice environments Lepidoptera by tachinids and chalcids is normally low ELg but prefers BPH in rainfed arid WBPIH in wetland environshyparasites and predators are perhaps the key to their control ments Ilalictoph(utsmtnroeiand HVpectrus attack G LH

Gall midges are parasitized by pterormalids and platy- and COfimravcctra respectivelygasterids that aestivate within their larval host between Pipunculids are mainly parasitic oii lealhoppers Each seasons Shoot aphids are preyed upon by coccinellids species has a unique environmental preference Tomosshysyrphids and chrysopids varireIa oraetoraprefers Ne7 otetltix nigropictrs and

Planthoppers arid leafhoppers have egg parasites and Pilunwulhtsjavanensisattacks Dehoceplhahls spp which predators as well as nymphialad nlt parasitesand predators are more abundant in upland than irrigated lowland their rich complex of natural enemies includes fungal environments T suhvirescensattacks Nephotettix virescens pathogens The principal egg parasites are myrnarids and and prefers the lowlands Pilpuiulus nttillattls is an trichogramniatids Egg predators are norinally mirid plant upland rice parasite arid attacks only Nephotetix speciesbugs that suck out the yolk Nymphal ad nIt parasites are More than 17 recorded species of leaffolder parasites in dryinids on planthoppers pipunculids on lealhoppers and the Philippines attack species of C(naplwocrocis and

30 IRPS No 123 January 1987

Table 6 Dominant natural enemies of rice pests in upland rainfed lowland and irrigated lowland rice environments of the Philippines

Natural enemy

Host stage Species

Brown planthop-Parasite Egg

MymaridaeAnagrus optabilis(Perkins) Anagrusflaveolus Waterhouse

TrichogrammatidacOligosita naiasGirault

NymphAdult Dryinidac

Pseudogotatopuslaiifemttr Esaki et Hashimoto Elenchidae

Elenchus yasurnatsui Kifune and Hirashima

Environment

Upand Rainfed lowland Irrigated lowland

Ailaparata lugens

x x X x

x

x x

x X X Whitebacked planthoppers Sogatella furcifera and Sogatodes pusanusEgg

MymaridaeAnagrus optabilis(Perkins)

TrichogrammatidacOl~gositanaias Girault

NymphAdult Dryinidae

Pseudogonatopusnudus Perkins Elenchidae

Elenchus yasutatsui Kifune et Hirashima

Parasite Egg

MymaridaeGonatocerus spp

TrichogrammatidaeOligositaaesopi Girault

NymphAdult Pipunculidac

Tomosvaryella subviresccns Loew Tomosvaryella oryzaetora (Koizumi) Pipunclusmutillatus (de Meijere)

DryinidaeIlaplogonatopusspp

Egg Copidosomopsisnacoleiae (Eady)

Larva BraLonidae

Cardiochiles philipphnensisAshnead lchneumonidac

Trichiomnia cnaphalocrosisUchida Temelucha stangli (Ashmead)

Pupa Chalcididae

BrachymeriaexcarinataGahan Brachymerialasus (Walker) Brachymeria nr tachardiaegroup

Parasite Egg

Scelionidae Telenomus rowani (Gahan)

EulophidacTettasticusschoenobhi Ferriere

Larva Braconidae

Stenobraconnicevillei (Bingham) Iclhncurnonidac

Temelucha phiiippinensis(Ashinead)Temelucha stangli (Ashmead)

x x x

x x

X x X

x x Green leafhopper Nephotettix virescens

X x x

x x

X X X X

X

X Rice leaffolders CnaphalocrocisMarasmia

x X x

X

X X

x x x x x X x x x

Yellow stem borer Scirpophaga incertulas

X X x

x

x

x X XAinauromorphaaccepta metathoracica(Ashmead) x

continued on next page

IRPS No 123 January 1987 31

Table 6 cntinued

Natural enemy Environment

Host stage Species Upland Rainfed lowland Irrigated lowland

Larva-pupa Gold fringed borer Chilo attriciiuslchneumonidae

Xanthopimpla stemmator (Thunberg) x aPupa x

Pteromalidae Trichoinalopsis apanteloctena (Crawford) ax

Pink stem borerb Sesamnia inferensParasite

Larva Braconidae

Stenobracon nicevillei (Bingham) x b b Armyworms Mvthinna SpodopteraLarva

Tachinidac Zygobothriaarroptvora (Robineau-Desvoidy) x x

Braconidac Cotesia spp

X Skipper Pelopidas mathias and green horned caterpillar Melanitis leda ismeneF-gg

TrichogramrnatidacTrichogrannaspp x

Scelionidae Undetermined Scelionid

x Larva

Tachinidae Argyroph lax nitrotibialis (Baranov) X x

Pteromalidae Trichomnalopsis apanteloctena (Crawford)

X Predator

CoenagrionidaeAgriocnenis femina femina Brauer x xIschnura senegalensis tambur

Miridae CrtorhinsliidipennisReuter x x

Nabidac Stcnonabis tagalicus (Stal) x

VeliidaeAlicrovelia douglasi atrolineata (Bergroth) x

Coccinellidae Micraspis crocea (Mulsant) X x x

Vespidae Eumenes campaniformis (Fabricius) x

Formicidac Diaecamna spp xOdontoponera transversa (F Smith) x

Carabidae Chaenius spp OphioneaW- aJasciataSchmidt-Goebel

x

Ophionea ishii ishii Habu x x X

Gryllidav Anaxipta longipennis (Saussur-) x xMetioche ittaticollis (Sial) x x x

Tettigoniidae Conocephalus longipennis (de Haan) x x x

aNo rearing made on rainfed lowland bNo rearing made on rainfed lowland or Irrigated lowland

MrasmiaAmong the Laval parasites the braconid Cardio- lowlands and parasitizes oriental maize borer OsIriniachilesphilijpphensis is most adapted to an upland environ- furnacalis (Guenee) larvae The leaffolder pupal parasitesment It also parasitizes Hydelepta indicata a common not only occur equally in all environments but also haveleaffolder of legumes The ichneumonids Trichomma cna- wvide host ranges- parasitizing species of OsIrinia Marucaphalocrosis and Temehcha stangli occur more in the Hydelepta and Honiona - in maize and legumeslowlands T stangli is most adapted to irrigated lowlands The principal egg parasite of yeltow stem borer (YSB)and also parasitizes Chilo and Scirpophaga stem borer Telenomus rowani is widely adapted to all environmentslarvae T cnaphalocrosis is most prevalent in rainfed and has an unusual alternate host -Tabanus eggs (25)

32 IRPS No 123 January 1987

Tetrastictu schoenobii specializes only in YSB and occurs mainly in irrigated lowlands -- further proof that YSB is most adapted to the wetter environments

Larval parasites of YSB are highly environment-specific The braconid Stenohracon nicevillei occurs only in upland rice but also attacks the pink stem borer (PSB) Tieinchwa phiiiplPwnsis an ichneumonid occurs in upland rice and also parasitizes the larvae of (7hiu stqpresaisinatro-morpha accepta melaihoracica like 7 stangi is mnost adapted to the irrigated lowlands

The dark-headed stem borer(l)HSB) (ih pochit3sus was not collected in all rice environments and the gold fringed borer Chilo auricilius nd PSB were collected only in upland rice Xaantiholiplastenrnaloran ichneumonid larval parasite appears to be widely adapted whereas TrihomaslopisapanitI)cteiiaa pteromalid pupal parasite occurs mostly in wetlands and also parasitizes larvae of Chilo suppressais and Pelhuidas mathias

Armyworms Myit hinin and Spodujipera have two main larval parasites - a tachinid Zrgohothriaatropivora in rainfed environments and a braconid Cotesia spp in irrigated lowlands

The butterfly pests Pelopidas and 4elanitishave tricho-grammatid egg parasites found only in irrigated lowland environments

Two larval parasites show environmental preferences --Argrroph av nigrotihialis a tachinid in rainfed habitats and Trichonialopsisapantelkcnela in irrigated ricefields A limgrotibialisalsoattacks the sweet potato hornwormAgrius convolvtdi and T apanteloctena parasitizes DH S1B

Predators as a group show more distinct environmental preferences The aquatic damselflies Coenagrionidae are most abundant in the lowlands Cvriorhinus is also most adapted to the lowlands The nabid Stenonabis tagalicus occurs only in the uplands and isalso prevalent on legumes The coccinellid lady beetles wasps and ants are more prevalent in upland areas AlicrasIpis cit-ea larvae prey ona variety of aphids attacking legumes and maize Etunienes campanonia vespid mud wasp makes nests in trees and is therefore most adapted to the more botanically diverse uplands Soil-dwelling ants cannot tolerate flooding

Upland environments are habitats to arboreal carabid beetles Three species of Chlaeniusbeetles prey on leaffolder larvae and are perhaps more important than parasites (27)

The gryllids and tettigoniids are widely adapted egg predators not only on rice but also on maize and legumes They feed on eggs of most insects that are laid on leaves

The spidercommunity of upland rice environments isrich in species At one site (Batangas Philippines) 31 species have been recorded (23) Of the 176 spider species recorded in Philippine ricefields about one-half (82) occur in upland rice The spiderspecies of upland rice environments overlap more with those of rainfed lowland than with those of irrigated lowland rice (24) The same study showed that of the three environments irrigated rice has (ie greatest spider species diversity foiiowed by rainfed lowland and upland

However of the 10 most prevalent ricefield spiders 9 were abundant in upland rice showing wide adaptation All three spider guilds -- orb-web space-web and huting spiders shywere prevalent in upland ricefields There were however differences in environmental prelerences for some spider species

Of the orb-wcavers Tetragniathamanlihulatawas partishycila rlv abundant 7 japonica was low and Ltucatge dehcorata was absent in upland rice

As in other environments Aiypeiwnaifirtosana was the most dominant space-web spider however its relative numbers were lowerthan for lowland sites Among hunting spiders Ox|opes javanus and Lcosa leatcost fgla were more abundant and Lvcosa lpseudoanmnlata was less abundant in upland compared with lowland rice envishyroniments

Extensive rearing of rice insect pests in upland rainfed lowland and irrigated sites over an 8-yr period in the Philippines provided insight into the effectiveness of parasites as natural enemies (Table 7 8) Most of toe data are from large sample sizes taken over at least one crop which overcomes the pitfalls expressed by Van Driesche (238)

VSB egg parasitization was surprisingly similar across all environments Larval parasitization was highly dynamic among sites and even years within the same site but showed somewhat lowerlevels in uplandsthan in irrigated wetlands This would have been predicted for egg parasites as well since YSB is adapted to the lowlands and so must be its parasites This isevidence that Telenomus rowani is as good at dispersing as its main hosts Trowani clings onto the anal tufts of female YSB and parasitizes eggs as they are laid This phoresy may explain its wide adaptation The more specialized lowland YSB larval parasites therefore appear more effective than their upland counterparts with more alternate hosts

Low collection levels of PSB larvae especially in the wetlands make comparisons between environments difficult Low levels of larval parasifization were recorded in both rainfed environments Except in the Cagayan rainfed wetland site in 1980 low levels ef larval parasitization were evident at irrigated and rainfed sitesalike No differences by environment were evident Higher levels of parasitization occurred in Sarawak during a 1967 outbreak also from tachinids Rothschild (197) indicated that higher parasishytization occurred in upland rice areas

Skipper and green horned caterpillar larval parasitization rates were generally low and showed no environmental effect htighest activity occurred in Cagayan in 1980 (48 parasitization) Although larval parasitization levels of RLF were variable year to year activity appeared greater in the irrigated lowlands than in the uplands Again the highest levels were recorded in Cagayan in 1980 Cagayan has extensive grasslands and fallow around the rice areas and a relatively long rainfall period Rainfall in 1930 was good for crop growth and apparently for increase of larval parasites

IRPS No 123 January 1987 33

Table 7 Parasitization rates of rice planthoppers and green leafhopper in upland rainfed lowland and irrigated lowland environments Philipshypines 1976-840

Environment and site Year Egg parasitization () Nymphaladult parasitization ()

BPH W13P GLH BPII WBPH GLIH

UplandBatangas

Rainfed lowland

1976 1979 1980 1982

15 17 12

2(47)8(39)

4(51)

9(1800)14 (212)

12(304)

8 (101)6 (127)

9(116)

Iloilo

Cagayan

1976 1977 1978 1981 1980 1981 1982

0 (69)

11

44 48 40

-

-

46 55 44

0 (471)

11

52 56 49

0 (200)9 (214)

10 (1259)8(235)6 (900) 3(800)

-7 (462)

11 (270)3 (313)5 (900) 1 (800)

4 (600)10 (725)10 (362)12(870)5 (900) 3 (800)

Pangasinan

Irrigatedlowland

1984 1976 1978 1979 1982

13 (436) 13(379) 14 (457)

6(14)7 (197)

17(198) 11 (486) 18 (275)

3 (81)10 (274)3 (116)

16(871) 19(88)

28(211)9 (764)5 (647)7 (1204) 8(305)

Laguna 1977 11 7 21 1978 31 1979 14 4 4 28 1980 7 8 20 1981 1983

56 7(278)

26 16 (1650)

16 27(1921)

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R I Apostol unpublished 1lo1lo and Pangasinan 136 Cagayan BCanapi unpublished Liuna 175 I C Pantua unpublished 118 234 43 Carino and Shepard unpublished BPH = brown planthopper Nila parvara htgens W131Pi =whitebacked planthopper Sogatelia furcifera GLH = green leafhopper Nephotettrix virescens Figiires in parentheses are sample sizes

Tauher et al (230) reported that internal external and genetic factors influence the seasonal activity of parasitoids

Hopper egg parasites but not nymphaladult parasites were also parlicularly abundant at the Cagayan site possibly the egg parasites have more alternative hopper prey in the weedy areas than nymphaladult paisites

The seasonal dynamics of hopper egg parasites can be seen for upland rice in Batangas(Fig I 2) and for rainfed lowland lice in Pangasinan( 135)and Cagayan( 105) Levels of BPHWBPI-L and GLH egg parasitization were corn-parable in upland and irrigated lowland environments Except for some high rates of paraMsitization early in the season the parasitization rate tended to be steady in all envilonments However hopper nymphaladult parasi-tizution levels were more similar in the rainfed environments and lower than in irrigated rice

Overall parasites do not appear highly effective by themselves against upland or even lowland pests Among natural enemies parasites have attracted the most attention because they can be more readily assesscd

Predators and pathogens lend themselves better to management practices Predators the most important group of natural enemies in riccfields are difficult to quantify On the other hand pathogens appear less im-portant but can be readily cultured and disseminated

The natural enemy commulnity of upland rice is rich in species and differs significantly from that of lowland rice

These beneficial organisms must be conserved by applying insecticides judiciously particularly sprays Seed or soil placement of chemicals minimizes exposure to natural enemies The strategy to derive the greatest benefit from natural enemies isto allow the greatest number of beneficial species to thrive Then perhaps at least one species will be effective against each pest at any tine overcoming the variable abundance of each species within and between years

Programs to introduce exotic species or mass produce indigenous ones are ambitious and expensive because they require trained people on a sustained not ad hoc basis

Chemical control Insecticides offer rapid and efficient control of upland insect pests (190) but-should be used only after other control measures have been considered Insecticides are rarely used on upland rice because of cost (32) Upland rice yields are normally too low to justify the expenditure Also most upland rice is grown as a cash crop Spraying upland rice is more difficult than lowland rice because water is less accessible Government subsidy programs to provide pestishycides to tribal peoples for upland rice production have created problems of toxicity to humans because isolated people have had no experience to be able to handle pesticides safely

However ins-eticide use has beenjustified when a crop is

0Z

Table 8 Parasitization rates of lepidopterous pests in upland rainfed lowland and irrigated lowland environments Philippines 1976-84a

Parasitization (5)

Environment and site Year YSB GFB PSB Earcutting Swarming Skipper Greenborned RLF Brown larva larva caterpillar caterpillar caterpillar caterpillar larva semilooperEgg Larva larva larva larva larva larva

UplandBatangas 1977 - 0 (57) - 0 (63) - 0 (71) - 0 (106) 11(142) 2 (101)1978 43 (4697) 8(451) 5 (38) 3 (40) 6 (61) 2(58) 15 (57) 13 (129) 13 (1286) 4 (30)1979 59(4135) 12(320) 13 (11) 5(28) 9(45) 6(46) 7(23) 8(160) 9(1152) 8(26)

Rainfed lowland Iloilo 1976 - 0 (10) 0(10) 0 (7) 0 (19) 2 13) 15 (180) 5 (76)1978 0 (54) 0 (3) 0 (17) 9 (64) 4 (78) 3 (73) 0 (121) 0 (191)1979 64 (2350) 16 (84) 6 (17) 0(6) 18 (68) 25 (104)

1981 56 (1322) 18(304) 0(5) 12(51) 18(138)Pangasinan 1976 60 (7 7 )b 62(82) 0(111) 5 (39)

1978 58 (53)b 5(51) Cagayan 1979 59(713) 8 (101) 6 (34) 0(7)

1980 0(11) 4 (28) 15 (286)

62 ( 48(177) 61(140)1981 0(5) 12 (2) 13(45) 14 (92)

Irrigatedlowland Laguna 1974 62 8

1974-75 50 41 1975 43 72 1978 96 (623) 21(24) 33 (60)1979 83(2017) 18(110) 8(62) 11 (89) 22(400)1980 78 (1625) 20(75) 0 (16) 4 (72)1983 60

9 (80) 7 (270) 30

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R F Apostol unpublished Iloilo and Pangasnan 136 Cagayan B Canapi unpublished Laguna 175 P CPantua unpublished 118 234 143 Carino and Shepard unpublished YSB = yellow stem borer ScirpophagaincertulasGFB = gold fringed borer Chilo auricilius PSB = pink stem borerSesamia inferens earcutting caterpillar Alythimna separata swarming caterpillar Spcdoptera mauritia acronyctoides skipper Pelopidas mathias reen horned caterpillar Melanitls ledaismene RLF = rice leaffoldersCnaphalocrocismedinalis and Marasmiaspp brown semilooper Mocisfrugalis Figures in parentheses are sample sizes Egg masses

IRPS No 123 January 1987 35

Table 9 Effect of seeding rate and insecticide on plant density and Yield losses from insect pests in upland rice often areyield of Dagge upland rice Batangas Philippines 1976a comparable to those of lowland rainfed or irrigated rice

Seeding rate Insecticide Dosage Plant density Yield (0-Y30o) but usually from different pests The principal (kg aiha) seed treatment (kg aiha) (noIra-row) (tha) groups are soil pests feeding on sown seed and rootsat 14 DE followed by seedling pests These groups have largely gone

50 None - 35 a 20 a unnoticed leading researchers to conclude that stem borers50 Carbofuran F 10 38 ab 23 a and seed bugs are the most important pests (17) Yield loss 10 beCabofuranF100 None - 4551 c 23 a trials that pinpoint major pests need to includc seed

100 Dieldrin WP 10 52 cd 22 a treatments Granules applied in the furrow may not give150 None - 59 d 22 a adequate protection from many seed pests

aAv of 8 fields DE = days after crop emergence In a column means The floristic and pedological diversity of upland rice followed by a common letter are not significantly different at the environments is matched by the diversity of arthropods5level by D)MRT F = flowable Wi = wettable powder both pests and their natural enemies Irrigation has homoshy

genized lowland rice environments Flooded soils tend tothreatened by epidemics of locusts or armyworms New have the same properties and to eliminate soil pests The technology stressing efficiency has minimized insecticide spatial and temporal dominance of cultivated rice has cost benefited specialist pests

Seed treatment is normally inexpensive and can be Upland rice has no specialist arthropod species The economicallyjustified in -nanycases(66) Treating seed with monophagous pests on upland rice prefer the lowlands systemic insecticide (216) but not chlorinated hydrocarbons where rice evolved Rice in the uplands exists only because (247) is most effective in controlling seedling maggots of man and even though lowland rice predates upland riceFoliar sprays have to be repeated to obtain control and specialist species evolved Wheat maize and sorghum do often are washed off by frequent early season rains not have monophagous pests leading us to conclude that Granular insecticides are effective but required dosages are there is no evolutionary advantage for inssects to have too expensive to justify except under heavy infestation narrow host ranges in upland graminaceae(247) When directed against first-instar larvae banding Pests have adapted to the uplands not only by having granules in seed furrows (177) can be effective against white wider plant host ranges but also by having long life cyclesgrubs at low dosages (025 kg aii ha) (134) Seed treatments undergoing dormancy or being dispersive Each pest has a are not promising for white grub control (248) unique life history strategy suited to highly unpredictable

Seed treatment is also effective ainst ants and may be environments In agivenyear someinsect specics an better cheaper than increasing the seeding rate In Batangas exploit upland rice as a food source but no pest is a senious treating 50 kg seed ha was as effective as increasing the one every year on upland rice seeding rate to 100 kgha and using untreated seeds (Table The low yielding potential of upland rice means less 9) In Brazil seed treatments control termites and other soil quantity and lower quality of food for pests to exploit but insects including the lessercornstalk borer(66) Domiciano also less incentive for farmers to undertake control (58) however found seed treatments inconsistent in their The most effective control would be early and synchroshyeffect against high populations of lesser corn stalk borer nous planting of early-maturing upland rice varieties to

Baiting can be an inexpensive way of controlling ants prevent insect pest buildup Breeding for insect resistance seedling maggot flies mole crickets and field crickets Baits should initially focus on stem borers because they are verymade of locally available material and impregnated with difficult to control by other means Biocontrol efforts insecticide can be sparingly distributed taking adantage of should first stress conservation of natural enemies preshythe pests ability to disperse and encounter bait sites dators and pathogens rather than parasites Efficient

The new low-volume hand-held controlled-droplet chemical control includes seed treatment or baiting for seed sprayers may offer an advantage over high-volume knap- seedling and some root pests and using controlled dropletsack sprayes for controlling foliar pests Upland rice sprayers for spot treatments against foliar pests and seed canopies are more open than those in lowland rice and bugs when economic thresholds are reached better droplet penetration should result Controlled droplet applicators would be ideal for seedbugs and defoliators

CONCLUSION

Upland rice is attacked by a wider array of insect pests than is lowland rice mainly beca use of the addition of soil pests Generally population levels of lowland rice pests are lower in upland than lowland rice leading many people to conclude that insect pests are not important on upland rice

36 IRPS No 123 January 1987

REFERENCES CITED

1Abraham C C and K S R Mony 1978 Occurrence ofLeptocorisa acuta Fabr (Coreidae Hemiptera) as a pest of nutmeg trees J Bombay Nat ttistSoc 74553

2 Agyen-Sampong M 1975 HlierogrlphusdiaganensisKrauss (Orthop-tera Acrididae) a new pest of rice in Northern Ghana Ghana J Agric Sci 8249-253

3 Agyen-Samnpong M 1976 Varietal resistance and other inle-ratcd pest control components as factors of rice varietal improvement in West Africa Prcsented at the Second varietal improvement serinar 14-18 Sep 1976 Bouak Ivory Coast West Africa Rice Development Association Monrovia Liberia 18 p

4 Agyen-Sampong M 1977 Outbreak of Ihccrorrdnisorae in rice seedlings near Rokupr Sierra Leone lit Rice Res Newsl 2(6)16-17

5 Agyen-Sampong M 1980 Dirty panicles and rice yield reduction caused by hugs IntRice Res Newsl 5(l)11-12

6 Agyen-Sampong M 1982 The major pest problems of irrigated upland and mangrove swanip rice ecosystems ir the humid tropical Guinea savanna and Salel climate zones Pages 318-328 in Pro-ceedings training course cn concepts techniques and application of irtegrated pest management in rice inWest Africa 10-20 Jan 1982 Fendall Liberia West Africa Rice l)evelopioent Association Monrovia Liberia 505 p

7 Akibo-Betts V I and S A Raymundo 1978 Aphids as rice pests in Sierra Leone lit Rice Res Newsl 3(6)15-16

8 Akinsola F A 1975 Present status of different rice stemborers in parts of Nigeria iicL Entomo Newsl 328

9 Akinsola FA 1979 Ihe biology and ecology of rice stemborers in Nigeria Ph 1) thesis University ol lbadan Nigeria 217 p

10 Akinsola E A 1979 Guidelines of integrated management of insect pests of upland rice in West Africa Presented at the Seminar on integrated management of rice diseases and insect pests at Bobo-Dioulasso Upper Volta 17-22 Sep 1979 West Africa Rice Develop-ment Association Monrovia Liberia

II Akinsola IA 1984 Insect pests of upland rice in Africa Pages 301-305 in An overview of upland rice research Proceedings of the 1982 lIouak Iwsrv Coast upland rice workshop International Rice Research Institute 13O Box 933 Manila Philippines

12 Alam M S V TJohn and Kaung Zan 1985 Insect pests and diseases of rice in Africa Pages 67-82 in Rice improvement in Eastern Central and Southern Africa proceedings of the international rice workshop at Iusaka Zambia 9-19 April 1984

13 Alticri M A A van Schoonhovcn and J D Doll 1977 The ecological role of weeds in insect pest management systems a review illustiated with bean (Phaseohsvulgaris L)cropping systems PANS (Pest Artie News Summ) 23195-205

14 Anmante F 1967 The leaf-cutting ant Atia capiguaraa pest of pastures [in Portuguese] Biol6gico (Sao Paulo) 33113-120 (Rev Appl Entomol A56207)

15Andrade A C and D Puzzi 1951 Preliminary results of experi-merits on the control of Sculptcoris iastantus on sugarcane [in Portuguese] Bioulgico (Sao Paulo) 1744-49 (Rev Appl Entomol A41421422)

16 Appert J 1970 Maliarplia separaelha (rice stem borer) new observations of entomological problems of rice in Madagascar [in French] Agron Trop (Paris) 25329-334

17 Ariadeau Mand Z Harahap 1986 Relevant upland rice breeding objectives Pages 189-197 in Progress in upland rice research Proceedings of the 1985 Jakarta conference International Rice Research Institute 1O Box 933 Manila Philippines

18Asar S and Kisimoto 1975 Seasonal occurrence of the smaller brown planthopper in upland rice [in Japanese] Proc Kanto-Tosan Plant Prot Soc 2228

19 Bakker W 1974 Characterization and ecological aspects of rice yellow mottle virus in Kenya Meded Landbouwhogesh Wage-ningen 152 p

20 Balut F F 1970 White grubs attacking rice Oryza ativa L [in Portuguese] Biol6gico (So Paulo) 36321-322

21 Banerjee S N and L M Pramanik 1967 The lepidopterous stalk borers of rice and their life cycles in the tropics pages 103-124 in The major insect pests ofthe rice plant Proceedings ofa symposium at the International Rice Research Institute September 1964 The Johns Hopkins Press Baltimore Maryland USA

22 Barrion A T and J A Litsinger 1980 Ants a natural enemy of cliaiarac in dryland rice IntRice Res Newsl

5(4)22-23 23 Barrion A T and J A litsinger 1981 The spider fauna of

Philippine rice agroecosystem 1Dryland Philipp Entomnol 5139-166

24 Barrion A T and 1 A Iitsinger 1984 The spider fauna of Philippine rice agroccosystcms II Welland Philipp Eiitomol 611-37

25 Barrion A T1 and J A Litsinger 1984 lihanus (Diptera Tabanidae) eggs an alternative host of rice stei borer (SI3) egg parasite Telenoonis digns (Iliymenoptcra Scelionidae) lit Rice Res Newvsl 9(6)19

26 Barrion A T and J A Litsinger 1985 Identilication of rice leaffolders by wing markings Int Rice Res Newsl 10( 1)24

27 Barrion A T and J A Litsinger 1985 Chctiosspp (Coleoptera Carabidae) a leaffolder ILF)predator lit Rice Res Newsl 10(1)21

28 Bertels A 1970 Insect pests of rice and theircontrol [in Portuguese] Pelotas Brazil Instituto de Expcrimentacao Agropccuararias do Sul Circular 43 24 p

29 Bonzi S M 1982 Ctilo (flisilineus J de Joannis (Lepidoptera Pyralidae) stem borer of irrigated cereals in Upper Volta LioFrench] Agron Trop Riu Rizic Cult Vivrieres Trop 37207-209

30 Bourke T V T L Fenner J N LStihick G Baker EHassan D F OSullivan and C S li 1977 Insect pest survey for the year ending 30th June 1969 Papta New Guinea )epartment of Agrishyculture Stock and Fisheries Port Moresby 57 p

31 Bowden 1 1973 Migration of pests in the tropics Meded Rijksfac Landbouwwet Gent 38785-796

32 Breniere J 1973 The main rice parasites in West Africa and their control Pages 27-42 in Proceedings seminar on plant protection for the rice crop West Africa Rice Development Association Monrovia Liberia

33 Breniere J 1976 The principal insect pests of rice in West Africa and their control West Africa Rice Development Association Monrovia Liberia 52 p

34 Brown ES 1962 The African arinyworn Spoiolptera eXI)Ina (Walker) (Lepidoptera Noctuidae) a review of the literature Cornshymonwealth Institute of Entomology London UK57 p

35 Brown F B 1968 Upland rice in Latin America FAO-International Rice Commission 12th Session of the Working Party on Rice Production and Protection 2-14 Sept Peradeniya Sri lanka Food and Agriculture Organization Rome Italy 6 p

36 Buyckx F J 1 1962 Disease and insect pests of crops inCongo Rwanda and Burundi [iil French] Publications dc LInstilut National pour lestude Agrononiquedu Congo M Weissenbruck S A Brussels Belgium 708 p

37 Cariiio F 0 1981 Role of natural enentics inpopulation suppression and pest management of green rice lealhoppers Ph D thesis University ofthe Philippines at Los Bafitos PhilippinesLaguna 136 -

38 Carpente A J 1983 The development of rice cultivation and extension of vegetable productin in Zanzibar Final Report Vol II1 October 1983 Food and Agricultme Organization Ronte Italy

39 Carroll C P and S1 Risch 1983 Tropical annual cropping systems ant ecology Environ Manage 751-57

4(0 Cates R G 198 1 Host plant predictability and the feeding patterns of monophagous oligophagous and polyphagous insect herbivores Oecologia (Berl) 48319-326

IRPS No 123 January 1987 37

41 Cendcafia S M and F B Caora 1967 Insect pests of rice in the 62 Ene 1 C 1981 I)iiorldhol)ierusihilis Slater (licteroplera Philippines Pages 591-616 in The najor insect pests ofthe rice plant lygacidue Illissinac) a new rice pest front Nigeria lit Rice Res Proceedings of a symposium at the International Rice Research Newsl N3)15 Institute September 1964 The Johns Hopkins Press Baltimore 63 Fugenio T S 1965 rhe effect oi e-vCgetation on rice production Maryland USA 729 p and the possibility of overcoming it by leaf pruning MS thesis

42 Centro Ihternacional de Agricklittira Tropical 1984 Upland rice in University ol the Philippines at Los Baijos ILaguna Ihilippins 69 p Iatin Anerica Pages 93-119 hi An os ervies of uplaind rice research 04 Farrow R A 1975 The Alrican nmigratory locust ii is main Proceedings of the 1982 Boinakamp Ivory Coast upland rice workshop outbreak area of the middle Niger qlinlitatise studies of solitary International Rice Research Institute P0 Blox 933 Manila populations in relation to environmental factors Locusla No II Philippines 198 p

43 Chandra G 1979 Taxonomy ad hn riononlics ofthe imect parasites of 65 Fennal R G 1943 Food crop investigation Windward and rice Icalloppers and planthoppers in the Pihilippines and their Ieeward Islands First report Report onl work dunring the periodiiiportairce iii natural biol)gical cointrol Philipp Fntoltol 4119-139 April 1942-April 1943 Imperial College of Tropical Agriculture

44 Chang T T 1976 The originii colut ion enlIivation dissemination Trinidad 16 p (Rev of Appl Fintornol A3222-25) and diersificition of Asian and African rices -uphytica 25425-44 1 66 Ferreira F 1983 Integrated pest ritaageiret of upland rice [ill

45 Chatierjee P B 1979 Rice leaffolder attacks in India lit Rice Res Portuguese English summary] Pages 323-341 in Upland riceculttre Newsl 4(3)21 factors affecting production E Ferreira T Yamada and F-

46 Cheng Cbien-pan 19(4 Upland rice in Taiwan International Rice Malmolta eds Institutoda Potassae Foslato Institnto Internacional Cnmmission Wiorking Party oit Rice Production ard Protection I()th da Potassa Piracicaha Sao Paulo lraid Meeting 3-10 March 1964 Manila Philippines Food and Agricul- 67 Ferreira E arid F dl Silva Martins 1979 Occurrence of ture Organiaion Rome Italy 5 p Sogatohs orvziola (Muir) inupland rice in Goias Ilrail lit Rice

47 Chcrrett 1 M and 1) 1Peregrine 1976 A review of the status of Rcs Ncwssl 4(4)18 leaf-cutting ants and their control Ann Appl Biol 84124-128 68 Ferreira E and 1F da Silva Martiris 1984 Insect pestof rice ill

48 Chipeta F M 1974 Pests of rice in Malawi Pages 90-95 in Brazil and their control Jill Portugucsej FMIlRAtPA-CNPAF Proceedings Sith eastern Africa cereals research conference 10-15 Goiania Brmiil 67 p March 1974 DRB Mialda ed Ministry of Agriculture lilongwe 69 Ferreira FI F ca Silha Miartins S S Nero and 1 It P de Malawi Carvallio 1982 M tltiplc regression analysis todeterimine the effect of

49 Chu Yau-i 1979 The d atiaging period of the rice seedling fly insect populatiois ott upland rice yield [inIPortuguese Pesqui (Atlterigotiavxigu Stcin) (NItiscide Diptera) oil cori) at Fast Java Agropecu Bras 17671-675 Indonesia Plant Prot Bill 21403413 70 Ferreira F 1F da Silva Martins and F 1PZimmermann 1979

50 Conklin H C 1957 IHaninoo agriculture a report on aln integral Rice varietal resistance to tle lesser corn stalk borer [in Port uguese] systent of shi fti rig cult ivationi in the Pltilippincs Food and Agriculture Pesqui Agropccu Biras 14317-321 Organization Rome Italy 209 p 71 Fleming W F 1968 Biological control of lie hapanese beetle U S

51 Cramer H II 1967 Plant proection and world crop production Dep Agric RLs Sers [ech Bull 1383 78 p Pilaniensclhutz-Nachr 201-524 72 Food and Agriculture Organiation of the United Nations 1982

52 l)ani R C and Majunidar 1978 Prelininary observations on tie Country report surmimary Pages 27-37 in Report on the first incidence of root aphids on differeit rice cultnrcs Sci Cult 4488-89 FAOIUN)IPhail and regional training coiuirse in iniproved ctltural

53 Davidson R I 1979 lffects of root feeding on foliage yield Pages practices for upland rice 5-23 Oct 1981 Northern Region Agrculturd117-120 in T K Ci osb and R B Pottinger eds Proceedings of the Developririt Center Chiang Mai Food and Agricuilture Organiiashy2nd Australian conlerence on grassland invertebrate ecology Govt tion Rome Italy 37 n Printer Wellington New Zealand 73 Franco do Ariaral S 195 1 Residual action of insecticides in turfy

54 l)ean G 1W 1976 Rice insect pests inlaos Int Rice Res Newsl soil--the results of a field experiment [in Portuguese] Biol6gico (Sao 1(2)15 Paulo) 17209-214 (Rev Appl Entomol A41423)

55 I)ean G J W 1978 Insect pests of rice in Laos PANS 24(3)280-289 74 Freeman ) 1970 Report onithe Iban london School of -coroiics 56 )ingle H 1984 Behavior genes and lifc histories conplex adapta- Monographs on Social Anthropology No 41 Athlane Press

tions to uncertain environments Pages 169-194 in A new ecology London UK 317 p novel approaches to interactive systems P W Price C N Slobod- 75 Fuchs T W1 A Harding and 1W Smithlr 1979 Induction and chikoff and W S Gaud cdsl ohn Wiley New York USA termination oldiapause in the sugarcane borer Ann Flitonlol Soc

57 Dobelmann-JP and M Falais 1968 Upland rice cultivation ill tie Am 72271-274 North-western part of Madagascar in 1966-1967 [in French English 76 (hbriel B1) 1975 A review tf the major insect pests of some tpland summary] Trop Agron 2313-21 crops in tlc Philippines Pages 86-111 in Ad Htoc panel of experts on

58 Doniiciano N L 1979 Chemical control of tie lesser corn stalk pest disease arnd weed problems in some rainfed crops Bangkok (lolsonno~paloIgnowslhts) (Zeller 1848) (I-epid(ptera Ihycitid ae) Thailand Food aid Agriculture Organization Roire Italy

on rice in Ponta Gressa Parara 1977-1978 [in Portuguese] Pages 77 Garg D K and N K Shal 1983 White grub outbreak on rainfed 291-303 in Anais 1 Reuniao de tecnicos cmiirizicultura do estado de dryland rice in Uttar Pradeslh Int Rice Res Ncwsl 81(4)16-17 Sao Paulo-Campinas Cati Brazil 78 Garg I) K and IP Tandon 1982 Major insect pests of rice on hilly

59 Dunsmore 1 R 1970 Investigations on tlie varieties pests and tracts of Uttar Pradesh India lilt Rice Res Newsl 7(11)11-12diseases of upland rice (hill padi) in Sarawak Malaysia Int Rice 79 Garrity 1) P 1983 Upland rice research an international bibliograpy Comm Newsl 1929-35 1975-1982 Iternational Rice Research hnstitnte PO Box 933

60 Elias 1R1965 The new decline of rice seedlings is not caused by Manila Philippines 121 p parasites [in Portuguese] Biol6gico (Sao Paulo) 313-6 80 Ghauri M S K 1982 A new subspecies of Iinorh)rone bull

61 Empresa Brasiliera de Pesquisa Agropecuaria 1984 Upland rice itt cornlus Slater (Hemiptera Iygaeidae) front Papua New Guinea on Brazil Pages 121-134 it Av overview of upland rice research rice and carpet grass Ihll lEntoniol Res 72133-137 Proceedings ofthe 1982 Blouak Ivory Coast Upland rice workshop 81 Gonzales F 0 PArregoces R I ihernandef and OT Parada International Rice Research Institute PO Box 933 Manila 1983 Insect and imite pests of rice arid their control in Latin America Philippines [in Spanish] la Federacion Nacional de Arrocerts (Fedearroz)

Bogota Colombia 60 n

38 IRPS No 123 January 1987

82 Grist D H and R J A W Lever 1969 Pests of rice Longmans London UK520 p

83 Hale P R 1979 Two subterranean pests of upland rice in Papua New Guirnca IntRice Res Newsl 4(2)16-17

84 Hale P R and J T Hale 1975 Insect pests of rice in the New Guinea Islands Rice Entomol Newsl 34

85 Harris W V 1969 Termites as pests of crops and trees Common-wealth Institute of Entomology London UK 41 p

86 Hattori 1 1968 A method to distinguish between grass leaf roller (Ctaplha rocis medinalisGuenee) and the rice leaf roller (Susurtia exigua Butler) [in Japanese] Plant Prot (Japan) 23167-170

87 Henry I E and 1A Onsager 1984 Experimental control of the mormon cricket Iahrosshnph by NJsena locustae(Aficrospora Microsporidae) a protozoan parasite of grasshoppers (Ort Acri-didae) Entomophaga 27197-201

88 Hidaka 1 P Vungsilabulr and SKadkao 1974 Studies on ecology and control of the rice gall midge in Thailand Tech Bull 6 TropicalAgricultural Research Center Tokyo Japan 113 p

89 Hincklcy A 1) 1963 Ecology and control of rice planthoppers in Fiji Bull Entomrol Res 53467 481

90 Hirao J 1982 Ecology and chtInical control of the rice leaffolder Jpn Pestic Inf 4114-17

91 Ho DT 1984 Sten borer incifience in rice ecosystems in KenyaEast Africa Pages 307-320 in A overview of upland rice research Proceedings of the 1982 Bouac Ivory Coast upland rice workshop International Rice Research Institute PO Box 933 Manila Philippines

92 Ho )T J 1Njokah and J GKibuka 1983 Studies on rice sten borers in Kenya with emphasis on ifaliarpiaipiraellRag Insect Sci Appl 465-73

93 Ichihara I T Ito A NIatsuo and M Sawada 1971 Chemical control of the rice striped stem borer in upland rice grown in plastic mulch [iii Japanese] Proc Kanto-Tosan Plant Prot Soc 1882

94 IclhiharaIand A Matsuo 1969 Damage caused by rice striped stett borer to upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1686-87

95 Ichhara I atid A MIatsun11970 Conside ration oft lie extraordinary prevalence of rice striped stem borer in upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1778-79

96 Institut de Recherches Agrolnliqes Iropicale et des Cultures Vivrieres 1978 Insect resistance Pages 12-13 in Upland rice 1978 atnual report Bouak Ivory Coast 27 p

97 International Institute for Tropical Agriculture 1983 Aphids itt rice roots I ITA Res Briefs 4(1-3)4-5

98 International Institite fi Tropical Agriculture 1983 Hopperburn in rice caused by brown plathopper IITA Res Briefs 4(l-3)6-7

99 International Rice Research Institute 1975 Annual report for 1974 PO Box 933 Manila Philippines p 216

100 International Rice Restarch Institute 1976 Annual repiort for 1975 PO Box 933 Manila Philippines p 228

101 International Rice Research Institute 1979 Cropping systems program Component technology development and evaluation Page 404 i Annual report for 1978 11O Box 933 Manila Philippines

102 International Rice Research Institute Department of Entomology 1981) Insecticid evaluation 1979 Tables 73 74 PO Box 933Manila Philipes

103 International Rice Research Institute Department of Entonmology 1981 Insecticide evalomtion 1980 Table 81 POBox 933 Manila Philippines

104 International Rice Research Institute 1981 Report of a meeting of Asian rice-based cropping systems entomologists Eleventh cropping systems working group meeting 18-22 May 1981 PO Box 933 Manila Philippines p 132 133 144

105 International Rice Research Institute 1984 Cropping systems program Pages 368-373 in Annual report for 1983 PO Box 933 Manila Philippines

106 International Rice Research Institutc 1984 Upland rice in Asia Pages 45-68 in An overview of upland rice research Proceedings of the 1982 Bouakamp Ivory Coast upland rice workiiop loternational Rice Research Institute 11O Box 933 Manila Philippines

107 Irshad M 1977 Grasshoppers in rice in Pakistan lit Rice Res Newsl 2(3)8-9

108 Ito R and TOno 1964 Upland lice its culture and varieties in Japan lIternational Rice Cormmission loth Meeting of the Working Party on Rice Production and Protection 3-10 March 1964 Manila Philippines Food and Agriculture Organization Rome Italy 12 p

109 Jeffrey M I and V A Dyck 1983 Effect of moonlight on lightshytrap catches of brown planthopper lit Rice Res Newsl 8(6) 13

110 Jennings P R and T Alicia Pineda 1983 Management ofthe rice plnthoppers Sogatodsorivi(coIl in Latin America Presented at the meeting on stable plant resistance FAO and lIndocsian I)inectorate of Food Crops Protection Bali lndonesia 29 Novcniber-2 l)ccemshyber 9 p

IllJimenez Gomez J A 1979 Rice insect pests i lla ios ()riCntales lil Spanish] Pages 99-103 in Rice ciltivation Instituto (olotibiatto Agropecuario Compndio No 29

112 Jongeleen F JJ Soesanto and A Mockidjo 1979 Studies oti pathogens of two species of Scarabacidac froinIndonesia Z Angew Enlomnol 87412-417

113 Jouvenaz 1) PC S lofgren and W A Baiks 1981 Biological control of imported fire antsa review of cirrent knowledge Bull Entomol Soc Am 27203-208

114 Kainoh Y K Shiniiu S MarUani Y Taiaki 1980 lost finding behavior of tie rice bu lplotori+JchinvittjI)allas (liciniptera Corcidac) with special reference to diet pattern ol aggregation and feeding on the rice plant Appl Entomol Zool 15225-233

115 Kakkar K 0 P Sharma and P I Sharma 1977 Note on Ihteronyrchus liodresRedt (Coleopicra I)ynastidae) as asporadic pest of paddy and maie inHimachal Pradesh Corr Agric 156-57

116 Kalode M131 79 Major insect pest problems and their control in upland rice (Unpublishcd mineogriph CWI oJNaaiat lni Chandaratat (198 1)original not seen)

117 Kalshoven L i 98 I The pests of crops in I ldoniesia (translated by P A vatn der Laan) 1 T Ichtiar B3aru-vati Iloeve lakarta Indonesia 701 p

118 Kamal N Q 1981 Suppression of whitebacked planto ner Sogatella fircaifra (Horvath) aLId rice lealfolder (nhal -cis licdb lis (Guenee) populations by natural cnclmie hlI) tiesis Grcgorio Araneta University Foundation Victoncta Park Metto Manila Philippines 179 Ip

119 Karim A N M Reaul 1975 A species ofliealy bug Trioni i70s ceres new oii rice in the Philippines Rice Entoniol Newsl 247

120 Katanyukul Wand C Chandaratat 1981 Insect pestsof uplaid rice and their control Report firstFAOUNlFi Tliailand training course on improved cultural practices for tplad rice Chiang Mai Thailand 5-23 Oct 1981 Food and Agriculture Organiation Rome Italy 22 p

121 Kern F 1955 lhletwioa(I)uninemts) bhhit Biturm(Coleoptera) a new pest of sugarcane inVenezuela [inSpanish with Etnglish sunmary] Agron Trop (Maracay) 527-29 (Rev Appl Eitoniol A4572)

122 Khan K M and S Ghai 1974 White grubs ald their control in India Pesticides (1Bombay) 819-25

123 Kiritani K 1975 Integrated pest management of rice pests Pages 66-70 itt Proceedings platning workshop ott cooperative hIeld research in pest managementP SMotooka ed East-West Center Honololu Hawaii USA

124 Kisimoto R 1967 Ecology of insect vectors forecasting and chemical control Pages 243-255 in The virus diseases of tle rice plant Proceedings of a symposium atthe International Rice Research Institute April 1967 The Johns Hopkins Press altimore Maryshyland USA

125 Kisimoto R and V A Dyck 1976 Climate and rice insects Pages 367-391 in Climate and rice International Rice Research Institute PO Box 933 Manila Philippines

126 Kureha S S Kobayashi SAsari and T Ishihara 1977 Life history and control of the mole cricket Grrllotalpa africana Palisot de Beauvois [in Japanese English translation 7520 1RRI library] Proc Kanto-Tosan Plant Prot Soc 21192-197

127 Kushwaha K S 1981 The black beetle as a pest of rice in Haryana India Int Rice Res Newsl 6(4)14

128 Lawani S M 1982 A review of tile effects of various agronomic practices on cereal stem borers Trop Pest Manage 28266-276

129 Leuck D B 1966 Biology of the lesser cornstalk borer in South Georgia J Econ Entomol 59 797-801

130 lever R J A W 1955 Insect pests of the rice plant in Malaya Int Rice Comm Newsl 1612-21

131 Li Ching-Sing and Shin-Foon Chiu 1951 The study ofthe rice gall midge Iachydiplosis orvza Wood Mason [in Chinese with English summary] 1Taiwan Agric Res 21-13

132 Iitsinger J A 1984 Assesament of the need-based application for rice Proceedings TechnologyTransfer Workshop 15-16 March 1984 International Rice Research Institute 1PO Box 933 Manila Philippines 6 p

133 Litsinger J A 1984 Environmental diversity and insect pest abundance with reference to the Pacific Pages 285-307 in Proceedings sub-regional training course on methods of controlling diseases insects and other pests of plants in the South Pacific 4-20 Oct 1982 Vaini Kingdom of Tonga Nukualofa

134 Litsinger J A RF Apostol and M BObusan 1983 White grub Leucopholis irrorata (Coleoptera Scarabaeidae) pest status popula-tion dynamics and chemical conllol in arice-maize cropping pattern in the Philippines J Econ Entoinol 761133-1138

135 Litsinger J A C de la Cruz F Raymundo A T Barrion M D Lumaban J P Bandong M S Venugopal F Paragna and E Balete 1982 Insect pests and insecticide reponse of dry-seeded bunded rice Pages 179-193 in Report of workshop on cropping systems research in Asia International Rice Research Institute Los Bafos Philippine

136 Litsinger J A M 1) Luinaban J P Bandong A T Barrion W Katanyukul and N Panda 1978 Insect pest management in rainfcd lowland rice agroecosystms of tropical Asia International Rice Research Conference 17-21 April 1978 International Rice Research Institute Los Bafios Philippines

137 Litsinger J A and K Moody 1976 Integrated pest management in multiple cropping systems Pages 293-317 in R I Papendick P A Sanchez and G B Triplett eds Multiple cropping American Society of Agronomy Madison Wisconsin USA

138 Loevinsohn M E 1984 The ecology and control of rice pests in relation to the intensity and synchrony of cultivation Ph D thesis University of London 36) p

139 Loevinsohn M E J A Litsinger N Panda M A Austria R F Apostol C Andrion and J Bandong 1982 A basis for predicting rice insect pest abundance in cropping systems Paper presented at the 1982 International rice research conference on rice and cropping systems 19-23 April 1982 International Rice Research Institute Los Bafios Iaguna Philippines 25 p

140 Lopez A W 1930 Repot of the Entomology Department Pages 145-172 in Annual report of the Philippine Sugar Association 1929-30 Manila

141 Lopez AW 1931 Entomology Department Pages 227-266 in Annual report of the director or research 1930-31 Philippine Sugar Association Manila

142 Lopez A W 1932 Entomology Deparment Pages 252-279 in Annual report of the Philippine Sugar Association 1931-32 Manila

143 Lungibill P and G G Ainslee 1917 The lessercornstalk borer US Dep Agric Agric Res Serv Tech Bull 539 27 p

IRPS No 123 January 1987 39

144 McGuire J U Jr and BS Crandall 1967 Survey of insect pests and plant diseases ofselected food crops of Mexico Central America and Panama U SDepartment of Agriculture International Agriculshytural Development Service Agricultuial Researrh Service October 1967 11 p

145 McNeill Sand T R ESouthwood 1978 The role of nitrogen in the development of insectplant relations Pages 77-9 in lliochemical aspects of plant and animal coevolution J B Ilarbornu ed Academic Press London UK

146 Maebara H 1955 Spatial distribution of lastinstiarlanaeof purplish stem borer (Sesamia iqferens Walker) hibernating in an upland rice field Bull Kagoshima Univ Fac Agric 438-41

147 Mallick E H and N G Hajra 1977 Response of rice plant to defoliation treatments at different developmental stages Indian Agric 2151-56

148 Mansingh A 1971 Physiological classification of dormancies in insects Canl Entomol 103983-10)9

149 Martell C G 1974 Catalogue of plant feeding insects of Mexico [in Spanish] Fitofilo 27(69)1-176

150 Martinez 0 J 1982 Principal rice pests in Ilanos orientalis [in Spanish] P~ages 80-94 in Curso deact nalization de conocimientos en arroz Villaniencis Instituto (olonmbiano Agropecuario

151 Martins J F da Silva F Ferrcira A S Prabhu F 1 P Zimmermann 1980 Use of pesticides for the control of the main subterranean upland rice pests esqui Agropecu Bras 1553-u

152 Martins I F da Silva F Fe eira rod BdamSilveira Pinheiro I Q2 Damage simulation caused by defoliators of upland rice [in Portuguese with English summary] Pesqui Agropecu Bras 171113-1119

153 Martins 1F da Silva N(iyuen van Tan and 1Ida Silveira Pinheiro 1981 Stem borer resistance in upland rice associated with plant morphological characters [in Portuuce Iesqui Agropecu Bras 16187-192

154 Masaki S 1980 Sumnmer diapause Ann Rev Entomiol 251-25 155 Masuzawa T 11 Suwa and F Nakasoji 1983 l)ifkleences of

oviposition preferencc and survival rate of two skipper butterflies Parnara guttata and PIchlpidai iathiais (Iepidoptera HIesperiidac) on rice plant and cogon grass New Entomol 3247-56

156 Maurya D M and C P Vaish 1984 Upland rice in India Pages 69-92 in An overview oftpland rice research Proceedings ofthe 1982 Bouakamp Ivory Coast upland rice workshop International Rice Research Institute IO Box 933 Mnila Philippines

157 Menon M G R R N Katiyai S Kuar and S S Misra 1970 Termites attacking the paddy crop Sci Cult 36294-295

158 Miyahara Y and 1 Nishikubo 1967 Behavior of young nymphs of the red rice root aphid Iholahsipiun ri liabhdontinalisSasaki in relation to movement from the leaf to the roots of upland rice [in Japanese] Pioc Assoc Plant Prot Kyushu 1363-66

159 Mo ljoa Tjien 1952 White grub of upland rice (Holotrichiahelleri Brsk) [in Indonesian with English summary] Contrib Agric Res Stn Bogor No 13134 p

160 Mochida 0 and T Suryana 1979 The occurrence of the brown planthopper and ragged stunt on lowland and upland rice ill Central Java and logyakarta in tile late of January 1979 Entomnolgy Survey Report No 15 Central Research Institute of Agriculture (CRIA) Sukamandi Indonesia I February 1979 9 p

161Monet C 1979 Varietal resistance to upland rice pests [inI French] Pages 35-39 in Annual report 1977 Institut des Savanes Department des Cultures Vivrieres Bouak6 Ivory Coast

162 Monte 0 1942 A caterpillar ofrice-fields[in Portuguese] Biol6gico (Sao Paulo) 8161-163 (Rev Appl Entomol A31414)

163 Munroe I) D 1979 Pest management strategy for hill rice in four resettlement areas in Sarawak Malaysia Pages 89-102 i1 Guidelines for integrated control of rice insect pests Food and Agriculture Organization Plant Production and Protection Paper 14 Rome Italy 115 p

40 IRPS No 123 January 1987

164 Nakasuji F M Ishii and Jen-Zon Ito 1984 Rice skippers in Taiwan and their life histories ZZrC (Taipei) Tech Bull 82 10 p

165 Navas I) 1976 Fall armyworum in rice Pages 99-106 il Ecological animal control by habitat management Proc Tall Timbers Con-ference 28 Feb-I Mar 1974 Fall Timbers Res Stn Misc Publ 6 Tallahassee Horida USA

166 Nickel 1 1 1961 Biological control of rice stem borer A feasibility study Tcch Bull 2 International Rice Rcarch Institute PO Box 933 Manila Philippines IIl p

167 Nigan P M and R A Vernia 1985 Insect pests of uplaild rice in Uttar Pradesh Int Rice Res Newsl 10(6)22

168 Nilakhe S S 1985 -cological observations on spittle bugs with emphasis on their occirence in rice Pcsqtui Agropecu Bras 20407-414

169 Nixon G EJ 195 1 lie association of ats with aphids and coccids Commonwealth Iusti lcte of Entonology Iondon UK 36 p

170 Nurdin F an Ishak Manti 1985 Study oit seedling fly of upland rice Interlational Upland Rice Moitoring Tour Report 26 lebruary-2 MarLI Sukaranti West Sumatra

171 Oka 1 N 1979 Ciltural conttol of the brown planthopper Pages 357-369 in Brown planthopper threat to rice production in Asia International Rice Research Institutc 1p0 Box 933 Manila Philippines

172 Ohni M and A Vilnaii 1975 Rice insect pests of Italy biology of Sipha grcria (Kaltenbach) and its control [in Italian] Riso (Milan) 2459-71

173 Otanes F Q atnd 1 I Sison 194 1 Pests of rice Philipp J Agric 12211-259

174 Painter R I 1951 Insect icsistance in crop plants University Press of Kansits Lawrence Kansas USA 520 p

175 Pantua P C and 1A litsinger 1981 Arthropod abundance in continuous vs biannual rice cropping systeis Int Rice Res Ncwsl 6(2)20-21

176 Pathak M ) and V A l)yck 1975 Sttudics ott insect pests of upland rice Pages 186-191 in Major research in upland rice International Rice Research Institute PO Box 933 Ilanila Philippines

177 Patil B R B M IHasabe and 1) S Ajri 198 I Chemical control of white gritb lIditriihinhorraia Fabr intfesting groundntut paddy atnd sorghum Indian 1Entomol 43232-236

178 Pawar A 1) 1974 (ollection and identification of insect pests of rice rlerminal repirt lInternational Rice Research Institute PO Box 933 Manila Philippines 72 p (tnimco)

179 IerrajU A aid A S Reedy 1965 Observations ott diapausc itt tie steti borer ohrice Rice News Teller (lai-Mar)9-13

180 Pont A C 1972 A review of the Oriental species of theri ona Rondani (I)iptera Muscidac) of econtomic importance Pages 27-102 in Control of sorglium shoot fly M G lomtani and W R Young eds Oxford atd IH1 Ptthl Co New I)clhi India 324 p

181 Prasad S C 1PChand aid M 1PSingh 1983 Incidence of rice gall midge in the upland plateau region of Hihar Oryza 20235-236

182 Rai P S 198 0 Nisaga simpex damage to rice in the hill tracts of South India Int Rice kles News[ 5(I)12

183 Rathorc Y S 1969 Gonocplhahli dcprssunt Fabr as itpredator of (tilo zonwIlhis (Swinhoc) and other moths in field cage at Patnagar Uttar Pradesh Indian 1Entotnol 3127

184 Reed W 1983 Crop losses causcd by insect pists in the developing world Proceedings 10th Internatial Congress of Plant Protection lrighton UK 174-80

185 Reedy M V 1981 Ateactogasterinhens(Walker)a new root-grub pest of tipland paddy and inaize Curr Sci 501076

186 Reveno M A 1968 Catalogue of the insect pests of economic crops in Colombia [in Spaiish] Asociacion Latino Americana de Entomo-logia (AIAE) 129-33

187 Reyes G M 1957 Rice dwarf disease in the Philippines Food and Agriculture Organiiation Plant Prot Bull 617-19

188 Rice S E A A Grigarick and M 0 Way 1982 Effect of leaf and panicle feeding byarmyworm (Lepidoptera Noctuidae) larvae on rice grain yield Environ Entomol 7593-595

189 Risch S J 1) Andow and M A Altieri 1983 Agioccosystem diversity arnd pest control data tentative conclusions and nizw research dirctions Environ Entomol 12625-629

190 Rivera C T 1956 Three organic insecticides in thecontrol of insects affecting upland rice Philipp Agric 39465-472

191 Roffrey 1 1972 Locusta outbreaks in the Philippines Acrida 1177-188

192 Rojas R 1) 1941 The effect of pruning at varying ages upon the yield of Ramairio Phipp Agric 29851-860

193 Roonwal M i 1979 Ternite life and termite control in tropical South Asia Scientilic Pnbl odhpr India 139 p

194 Rosetto C1 F da Silva Martins N C Schmidt and L EAzzini 1978 Damage caused by pasturespittle bugs 4Dvoi)lavopictaandDJ schach) on rice lBragantia Nota 6 (Matcli)25-27

195 Rosztto C1- S S Neto I- Amantc I) Iink 1Garcia-Vieira D M de SoLza N V Bansa t and A M de Olivcira 1971 Rice pests of Brazil [in Portuguese] Pages 149-238 inrFA()-International Rice Commission 2nd meeting of the Rice Committee for the Americas 6-11 l)ecember 1971 Pelotas Rio Grande do Sul Brazil

196 Rothschild G H L 1969 Obseratiois on the armyworm Spodoptera matritia acronrctoiiles Gn (Lep Noctuidae) Bull Entomol Res 59143-160

197 Rothschild G 1- L 1974 Parasites of the armworm Spolopiera mauritiaacroncitoihes in Malaysia Fntornopliaga 19293-299

198 Sachan i N S K Gangwar and 1N Katujar 1980 Field crickets damaging cages in North Eastern IHill Region Indian J Entomol 42526-527

199 Sands 1) P A 1977 The biology and ecology of Lepocorisa (Hemiptera Alydidac) in Papua New Guinea Res Bull 18 Departshyment of Primary Industry Port Moresby Papua New Guinea 104 p

200 Sarawak 1968 titomology division Pages 97-106 in Report of the research branch of the Department of Altriculture 1966 Kuching (Rev AIppl Eintomol A56603)

201 Satpathy JM M S Das and K Naik 1977 Effect of multiple and mixed cropping on the incidence of scine important pestsI Entomol Res 178-85

202 Sauer IHF G 1939 Notes Ott Elasmpnlpualtsligniosellths Zeller (Lep Pyr) a serious pest ofcereals in Sao Paolo [in Portuguese] Arq Inst Biol 10199-206 (Rev Appl Entotnol A28378)

203 Satinders 1L A 1 SKing and C L Vargas S 1983 Crop pests of Central Anerica with t bibliography [in Spanish] Centro Agroshynotnico Tropical de Investigacion y Ensenanza Turrialba Costa Rica (CATIE) Tec Bol 9 90 p

204 Sauphanor B 1985 Some factors of upland rice tolerttce to stem borers in West Africa Inseu Sci Appl 6429434

205 Sen P atnd S Chakravorty 1976 Stentflics in Haritighata Nadia West Bengal India [it Rice Res Newsl 1(12) 15

206 Senapai B and 1M Satpathy 1983 The seedling fly Atherigona orrzae Malloch is a pest of rice in Orissa (India) J Entomol Res 777-79

207 Shhjahan M S Alatn M Ah nad and BN Islam 1983 A new scarabaeid beetle pest of dryland rice in Bangladesh Int Rice Res Ncwsl 8(I)16-17

208 Sharma G L S C Modgol and R C Gautam 1979 Effect of depth of submergence ott incidence of bacterial blight and white grub infestation in transplanted rice Int Rice Res Newsl 4(6)12-13

209 Sharma S 1985 Scarabaeid beetle outbreak itt upland rice Int Rice Res Newsl 1(1(l)24

210 Shiraki T 1917 The paddy borer Schoenobhits incertulas Walker [achaku Agric Exp Stn Formosa 256 p

211 Silva A de Brito and 11P Magalhaes 1981 Insect pests of rice [in Portuguese] Circular Tecnica No 22 EMBRAPA Centro de Pesquisa t ecuaria de Tropico Unido (CPAJU) Belem Brazil 14 p

212 Singh S A 1979 Emergence of rice leaf butterfly as a rice pest in Manipur India int Rice Res Newsl 4(4)16-17

213 SinghS A 1983 Changing trends of insect pestsof rice in Manipur Int Rice Res Newsl 8(f)12

214 Siwi S S and P H van Doesburg 1984 Lenovorisa Latreille in Indonesia (Heteroptera Coreidae Alydinae) Zool Meded (Leiden) 58117-129

215 Soejitno J 1977 Reaction of some upland rice varieties toseedling fly eltherikonaexigua Stein [in Indonesian] Siri Hana I Penyakit 630-35 Lembga I osat Fenclilian Pertanian Bogor Indonesia

216 Socnardi 1967 Insect pests of rice in Indonesia Pages675-683 in The major inse pests of thu rice plant Proceedings of a symposium at the Internatital Rice Rec Wi itt S pternber 1964 The Johns Hopkins Press Baltimore Maryland USA

217 Soto P E A T Petcz and I W Buddenhagen 1976 Survey of insect pests and diseases of rice in different ecological zones in Nigeria Rice Entomol Ncwsl 435-46

218 Soto 11E and Z Ziddiqi 1976 Screening for resistance to African rice insect pests Varietal improsetnent sCmi n r Bouak Ivory Coast 13-17 Sept 1976 West Africa Rice l)evelopment Association Monrovia Iiberia

219 Squire F A 1972 Entomological problems in Bolivia PANS (PestArtic News Sumr) 18249-268

220 Stapely 1 II 1978 The rice leaf roller in the Solomon Islands Int Rice Res Newsl 3(l)13

221 Suenaga H 1937 On aphids attacking the roots of rice Oyo-I)oubts Zasshi 9150-152 [in Japanese] (Rev Appl Entomol A25745)

222 Suenaga H 1952 On the seasonal abundance of three subterranean aphids on upland rice Bull Kyushu Agric Exp Sti 1249-262

223 Sutynso Yas and Dandi Soekaria 1964 Beberapa hana yang ttenperang tana man padi tanah kering [in Indonesian] Rapat kerja Padi Tanah Kering 6 sd 8 Juily 1964 Kaliurang 9 p

224 Tallamy I) W and R F Denno 1983 Alternative life history patterns in risky environments in example from lace bugs Pages 129-147 in Insect life history patterns R F l)cnio and If Dingle eds Spring-Verlag 225 p

225 Tamura 1 1939 Observations on some insect pests of field crops [in Japanese] 1 Plant Prot 26414-420 (Rev Appl EntomolA27-556)

226 Tamura and Suzuki 1964 Influences of the early-transplanting practice in rice culture upon development of the rice stem borer Chilo siql)ressalisWalker Bull Ilokuriku Natil Agric Exp Stn 761-94

227 Tan J P 1924 The rice root aphid Drowhi ihirsoia Philipp Agric 13277-288

228 Tanaka 1 1957 Taxonomy and distribution of some subterranean aphids injurious to upland rice in Japan with description of a new species [in Japanese English summary] Botyu-Kagaku 22168-176

229 Tanaka T 1961 The rice root aphids their ecology and control Spec [full College of Agriculture Utsunomiya University Japan F3 p

230 Tauber M IC A Tauber R Nechols and 1Obrycki 1983 Seasonal activity of parasitoids control by external internal and genetic factors Pages 87-108 itt I)iapause and life cycle strategies in insects V K Brown and I Ilodek eds )r W Junk The Hague Netherlands

231 Taylor W F 1972 Effects of artificial defoliation (simulating pest damage) on varieties of upland rice Exp Agric 879-83

232 feotia T P S and S Natud 1966 Biotoomics of the rice skipper Parnaraniathiis Fabricius (Lepidoptera Hesperiidac) Indian J Entomol 28181-186

233 Teran F 0 1971 Rice pests of Santa Cruz [in Spanish Ministerio de Agricultura Sita Cruz Bolivia Bull 45 8 p

234 Than Itun 1976 Population dynamics of the yellow rice borer Tryporza itcertulas (Walker) and its damage to the rice plant MS thesis University of the Philippines at Ios Bafios Laguna Philippines 92 p

235 Uichanco L B 1911 Pests Pages 55-74 it Rice Philipp Agric 11-200

IRPS No 123 January 1987 41

236 van der Laan P A 195 1 Life history of the rice seedling fly Contrib Gen Agric Res Stn Boor Indonesia 118 15 p

237 van der Laan 11A 1959 Correlation between rainfall in the dry season and the occurrence of the white rice borer (Scirpolhaga innotala WIk) in lava Entomol Fxp Appl 212-20

238 Van Driesche R G 1983 Meaning of percent parasitismin studies of insect parasitism Fnviron Entomol 121 I1- 122

239 Venugopal M S and A litsingcr 1984 Fffect ofcarbofuran on rice growth Prot Ecol 7313-317

240 Vietnam Directorate of Rural Affairs 1962 Annual work progress report on crop improvement program of rice sugarcane vegetable and field crops (lor the period from July 1961 to June 1962) (11) 299 p (Rev Appl Entomol A51526-527)

241 Wan M T K 1972 Observations on rice leaf- and planthoppers in Sarawak (Malaysian Borneo) Malay Agric 148308-335

242 West Africa Rice I)esclopnient Association 1978 Annual research report Vol 2 Statistics Monrovia Liberia

243 West Africa Rice Development Association 1979 Annual research report p47 Monrovia Liberia

244 Williams ) F and CS lofgren 1981 Fli I illy l-468 anew bait toxicant for control of the red imported lire at Fla Entomol 64472477

245 Williams I) 1 1970 The mealybugs OFlomoptera Coccoidea Iscudococcidac) of sugar-cane rice and sorghum Bull Entomol Res 0109-188

246 Williams D) 1 1982 Pulinaria icervi (Signoret) and its allies ott sugar cane and other grasses Btll Entoinol Res 72111-117

247 Wirjosuiardjo S 1976 Study ofchenmical cont rot of insect pests ott upland rice Imiu Pertanian (Jogjakarta) 1323-329

248 Wirjosuhardjo S and N Wirarta 1976 Chemical control of the white grub tIolorithiatlltriin upland rice in M uneng Irobolinggo Indonesia) [in Indonesian] Imut Pertanian (Jogjakarta) 1330-332

249 Yano K T Miyake and V F Eastop 1983 The biology and economic importance of rice aphids (Henuiptera Aphid idae) a review Bull Entomol Res 73539-566

250 Yelp D and P Rhodes 19(4 lhe success of drilled dryland rice iii the Fiji Islands International Rice Commission Working Party on Rice Production and Protection Manila Philippines 3-10 March1964 Food and Agriculture Organization Rome Italy 13 p

251 Young W R 1981 Fifty-five years of research on the sorghum shootfly Insect Sci Appi 23-9

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No 101The economics of hybrid rice production in (hina No 113 Boron toxicity in rice No 102 Rice ratooning No 114 Energy analysis rice production systems and rice research No 103 Growth and development of the (Jeep water rice plant No 115 Production risk and optimal fertilizer rates an application of the

No 104 Faridpur acomputer-assisted instruction nmodel forrainfed lowland random coefficient model No 116 Consumer demand for rice grain quality in Thailand Indonesia andrice

No 105 A readingand listening comprehcnsion test inEnglish for non native the Philippines speakers applying for tiraining at IRIRI No 117 Morphological changes in rice panicle development a review of

No 106 Rice grassy stunt virus 2 a new strain of rice grassy stunt in tire literature

Philippines No 118 IRRI-Korca collaborative project for the development of cold-No 107 Physical losses and quality deterioration in rice plstprodnCtion tolerant lines through anthei culture

systemis No 119 Problen soils as potenial areas for adverse soils-tolerant rice No 108 (olptblication ofIRRI materials a survey of translators and varieties in South and Ssuthcasi Asia

publishers No 120 Changes isinall-farm iLtireliing techii)hgy in Thailand and [he No 109 Classification of Philippine rainfall patterns Philippines No 110 Contributions of modern rice varieties to rutrition in Asia No 121 Landforms and modern rice varieties No Ill Changes in rice breeding in 10 Asian countries 1965-84 No 122 Yield stability and modern rice technology No 112 Design parameters affecting the performance of the IRRI-designed

axial-flow pump

Page 24: UPLANDRICE. INSECTl--1i M T -E!R.EOLOU t RTANCE ,AND …delphacid.s3.amazonaws.com › 6958.pdf · The significance of abiotic factors in upland rice insect ecology will be apparent

24 IRPS No 123 January 1987

Philippines which had two treatments offering protection In the 9 Philippine trials where yield losses were measuredwith and without carbofuran granules relied on further by growth stage 10 of the yield loss (21) occurred at thecontrol with foliar spray and not on seed dressings The vegetative stage and 517 occurred at each of the reproshycarbofuran-protected plots had an 18yield los compared ductive and ripening stages Ibis result is similar to thewith only 10 with foliar sprayS This yield difference is pattern of yield loss in lowland rice measured by successiveprobably due to differences in protection against sown-seed treatments where the crop is unprotected at each growthpests (foliar spray do not control sown-seed pests) rather stage but proticted during the other growth stagesthan phytotonic effects became basal fertilizer wa used in However the insect complexes r sponsible for vegetativeonly 2 of the 10 comparisons stage yield losses in upland rice are entirely diftfrent than in

The recorded yield losses were not related to yield lowland rice The key vegetative stage pests in lowland ricepotential (Fig 3) One might expect higheryielding crops to arc aquatic whereas those in upland rice are sown--seedhave higher percentage of yield losses but this ws not tile (sown in soil) root and seedling pests This dift rence maycase One reason is that insect damage exacerbates plant explain why agricul torists have overlooked tile ilmportanccinjury from drought stress low yielding fields from d ought of upland rice insect pests Many reports of upland rice pestsstress would show disproportionate differences between in Asia ha Ve focused on major lowland species (46 108)insect-protected and -unprotected treatments Shoot and It isalso surprising that tei vegctative stage of upland rice root pests would be more responsible for losses from would record the highest yield losses becatuseseeral studiesdrought stress than sown-scd pests A second reason have shown that upland rice can readily recover from foliageconcerns seed pests which would have a greater impact on reinoval during that stage (117 152 165 188) Upland ricelow tillering low yielding varieties such as )aggc Low poscsses a hitgh obilitV to compensate for ca rly loss oftillering varieties nnot fill in the space created by removal foliage and will even be stinulated by foliage removal toof seeds oweer 1) R i 5 is high yielding because it tillersP1 produce a higher viel than plants ithout any foliageactively which can close the canopy to provide greater weed removed (63 192 231) [ie reason for this apparentcontrol discrepancy hovce is that yield loss at the vegetative

stage does not octUr frol leat arca loss but fron pestsYield loss () removing sown seed and feeding on devcloping tillers and roots -- damage that cannot be readily conpLtnsatCd for

70 - Measuring yield losses in upland rice cnvironrnents is further complicated by a report from Brazil that greater yield losses occurred in response to better insect control60 - =-09 as Insecticide seed treatments resuled in denser plant stands

(n=21) which in turn created a selectively more favorable environshyment for blast (69) This resuIt was parlticurla rlxexacerbated50 when cirbofuran granules were used in seed furrows Aside

ffrom a greater plant stand ca rbofuran iay have auigmented the N lecl and stimulated blast If so ca rbofuran should be40 - replaced with another insecticide or the seeding or fertilizerI rates decreased

Other methods to determine the impact of upland rice30 shy pests wvcrc to interview farners or to develop single speciesbull bull correlations of population levels and yield loss Feneira (66)2 NIM)st (75(1) Golas farmers reporteddid both in Brazil

20- insect losses in ufpland rice and 251 said those lossesS avcragcd 37(1 [his result is consistent with tie yield losses 0 0 measured by field trials in the same statc(Table4) For every

I0 4 of seedlings removC d by leaf-cutting ants a I 7yield loss is predicted Similarly Ferreira (66) found that ai

0 - average of 53 thrips per panicle before panicle initiation meant 3 times more unfillh d grains

-IO0 I 1 1 I I-101 0 2 3 4 5 6 7

Yield (tho) C)N [ROt IM FI IODS 3 Correlation between upland rice yield and yield lols caused by Upland rice insect pests may be controlled by culturalinsects measured in 21 trials in the Philippines Ivory Coast andBrazil genetic biological and chemical methods

IRPS No 123 January 1987 25

Cultural control dry season plowing will have little effect on them Plowing Crop husbandry to reduce upland insect pest populations when the soil is wet from the first monsoon rains will expose can be classified into practices effective in a single field and soil insects that tunnel close to the soil surface to predatory those effective only at the community level Cultural birds chickens dogs and even man (field crickets nole controls should come first in a pest management program crickets and white grubs a re delicacies in man tra ditional They have broad stable effects as pests have little pos- diets) sibility of overcoming them through selection of biotyps The greatest degree i1 inIscct control fron tillage comes (138) On the other hand some cultu ral controls work only from plowing soon after rice ha rvest when most soil insects with high labor or power inputs and connunity-wilt lie close to the soil surface anrd stern borers are in the rice methods require coordination of iany farniters stuble Whitegrubsare mature and tillage exposesthem to

Pantin time The luxury of being able to shift the time of predators Tirnelv control will prtec the crop following pla ntring upland rice is open onl hr la rniers in areas with a rice and lowr the populationIfor succeedinig years Plowingprolonged rainifall pa ttern lI copping intensity or underthe ice stulble helps it decompose killing sten borer rnechanied land preparatiot Gcnerally planlting as soon larvae inside as the rFiinv season begins will lower populatiois of 1110st [requent tillage destroys ait nests bit not termites insect pests Insect populationsarelo aftcra dlvseoll or Tillage and clean culture howes er rcliove food for winter fallow delayed plaiting lets them build tip on termites which then mtay attack a young rice crop Scarab alIternate hosts that gi vigorotsly with the first seasonial beetles also prefer to miposit in eceitly tilled fields Noshyratiris This first flush iof weds cros lItxuttriartlv with iceCult is dibbling faors the survival oftillace Clre Such rineralied N released during the dr season aid made stils i aels tnd insects thtt pttpate or lay eggs- in the soil

available to the plants with the rains It iliso igrtmitcally Zero tilla hio eerl conserves predators ofthe GIH (37) advanttgeous for the rice ciop to tap this trlLItlaI fertility nt it mcttoi lo lessen the impact o0 soil insects and grow vigorotsly to compete with wrus and to tolerate brotdcasting seed or planting in ftltrrows is prefc ired over pest infestation dihlirig in hills Crickets are large enotlgh to eat seedlings in

Larly plantngs terd to escape seedling maggolts Upland a hill atrd tillering normnally ca iriot fill in tlhe missing space rice in Bitangas is planted in Ma and ine itli the Iirst Oftcn dibbll rd is not well covered and iseasily found by rains Fairners are highly mtotiated to plant early tocscape foraging insets ioIdents and birds drlitght and typlotns ricar harct Arn Atrgtst planting Plan uit sii Incrcatsini sced density protects tile cropwould be evcely attacked by It riim orivzo At tti againrst seed and seelling pests especially ants particularly other Philippiie tipland sites Rea l Qteot antd (laelit if the crop will germinate and ermerge quickly M isainis Oriental seedlig mnaggots arnrige ric boe t se hr trftinll itI India rice intcrclopped witi cotton or farnes have a longer growing season and plnit 2 rin or pigeon pci had lower 6IIt and WBIlII poptrlations than mote after the ortset of the rainy season liaaiangas ftrmners rice alone (211) NfailC arid Urplald ir_ 2re coilltori in the plant ca rly becia use ofa shorter raiinfall period arnd the desi re Philippines he chatngC in pest status folii iniercropping is to lliarvet before the typhoor sea son peaks ltighly locatioll-specific (133) cmianarid the net elfc be either

Early plantings cornbind with early-iatiuring varictics nil bcnelficial ortdetriential (189) For example BIatangias provide higher crp tolerartce for white grubs Aestiatior farmers intercrip Itro in their ipland ricefields A liornshyterminates with the first rains but 5-6 moare ineeded for the nrii iotd-Ahis i (I iiiacius) which norlially feeds resting larvae to develop into beetles emerge and lay eggs on taro ill feed or miarbv rice plants Intercropping and for larvae to reach the danriigii g ird-instar stage probably would not afft0 seed pests A crluanion crop (134 In lataigas this normually iseiough tiite for harvest planted ith rice and haing gleater tolerarnce for root

It Brazil howecver delayed planting is a suggested dariage or being toxic nayact isi trapcrop to root-feeditig cultural control for the lesser cornstalk borer and insect pests Acial-fei i ig pests voultId be most affected by pests favored l drought (66 195) Cropping internsity in intrer ropping Shoot aphids leaf beetles sten borers and Brail is low and delayed plraing places the rice crop in the hopplers would be likely ca ntidates to note effcts On the niost stable rainfall period reducing likelihood old rought othlt halnd the corlipalior crop stch~ sitniaie Iiiv be more

In rilniy arca s rinmirsare roi atl to plant other crops benehited than rice from initecropping ( 137) before rice and althNgh it woultl be desirabh to avoid 11echi Weeuling luring the first month after clop infestations by planting rice cuatl lariners lack the labor cstablishent will force pcts such is anyvworris which and time to do so ol a low priority crp prefer grassy cetds ontor tle vorling rice Soil iampsecs will

77age Uplild rice Stils ae light textured and Call be mvnye to the rice crop Cleani citrltirc also lay lore territes plowed duiring the d rv seasor notl only to grin time for to attack I u cetincti inrice ceds i ay he le optinal planting with lie fitst Inilsoon riiains but also to desiccate solition 13) soil insects arid weeds such is ntsedge that foim rhizomes Irtlizr Itneatsing N Iciiliier oin hlwland rice has Htowever becta use most soil insects lie below die plow layer fAtwvored Irigher populains of pliithoppeiv Imlhoppers

26 IRPS No 123 January 1987

and leaffolde rs However because of the need to temper fertilizer use on upland rice o prevent blast fIrtilizer rates will be lower than is comonia r for lowlad rice causing relatixely less hopper and IaIfTfoide r population buildup However inupland rice areas without a history of blast such Is Batangas farmers apply high rates of N and conscqtrently the crop has high infcstation levels of WBPI]and RIF The fhrmrcrs leec insect buildup by splitting the fertili11r in0to two to three applicatiois bit do not applV fertilizer at the vegetati estage because this prodrces a tall crop which illeasily lodge during typhoons Greater crop fertility leads to greater insect survival (better nutrition) larger insects (to cat no re foliage) arid greater fec titd t (to lav nmre eggs) (128 145 171) Not all insect species respondequally hovr Increasing rates (if N l anrid K increased Dialraa s-acchralis and (iaeiurema flea beetle but reduced thrips (66) Zn rcdLuced deadheart density from the lesser cornIstalk borer

Fnldingi Although most Upland rice areas do nIot hawxcaccess to wa ter floodiig the fields is rRrrnriierdcd in areas of Brazil or root bug and white gib control ( 195)

Crop rotatioti Tillage before planting a crop after rc harvest will control stern boreis and other pests renain n the stubble arid will unearth soil pests Planting a non-gratriiaceous crop after rice is recommncrded for termite control (66) Crop rotation is less nttdcd in upland rice culture than in lowlands because atcrop-free period is assured even after harvest of crops following rice

t1ulchilt iryl plastic irli irig designed for weedcoitrol ii upland rice inJapan resulted in ntr highanri liy infestation of (ihilo sulytresvais (93 9-1 95) NIulched Upland rice grew more luxuriantlv and attracted more (Avipositing mothis than u lched rice Also the rice plant and stein borers imatoured 10rC quickly and a partialgeneratiori af vouing ista iNoverwintcred in the stubbleLotili--malitrt varieti Quickly maturing crops reduce the tmbCr of pCst generations that Calt build up Rapid crop establishment olan ca rl v-natu ring vaitvty if)pa 11icLa r will lessen white gib and stem borer damiage White grub larvae intle last iistar are highly destIruCtive so aii early-iiatri ng variety will escape serious root loss Stem borers build 1ip slowlyx in rice to ecormic abundant in a late-Maturing crop anrd caiuse whi tlieads Numibers of somespecies increase exponeitially with each succeeding genera-ti so a xarietyexhat atures I trowh earier thian another will hac less insect damage Early-naturing vaiicties however will not lessen seed pest daiage either ii the soil or oina standing crop

Strwhoi hAIJttam Inrscts tcd irig on thlie aerial portions of rice plats disperse from carly to late plantedfields AS iiiibCirs increase exponentially with eachinsect

gtscnrattm e eiglbrig f rmc~s1(uil upp wh rc generation pests build tIP where neighboring fihrTC e9r1sstagger their plantings II Batangas farmers plant uplandrice within I iro use 120-d s-arieties and plant at the start of the rainy season Ini Misarnis Oriental laricrs stagger their

plantings up to 3 mo apart and use a set of atieties that mature in 4-7 mo Insect pest danage particularly from stem borers is greater in Misarmis Oriental Also seedling maggots damage rice there because rainfall favors frequent planting of maic In Batangas rice escapes seedling maggots bCca usC of distinct drs season and synchron0us early planting with the onset of ains

Therefore combinations of cultural practices ea rlyplanting synchronous planing crop rotation and earlyshymaturing vaneties protect the rice crop against most insect pests

Ilmant resistance Beca tse upland rice has few specialized insect pests and many insect types attack the crop worldwide regional breeding and strong internatiornal cooperation are required

Sown-sced pests arc economically controlled by insectishycide seed treatment and should the relore receive lowxpriori ty inbreeding objectives

Root pests are normally cost ly to control with insecticides but finding resista t sonres mayvbe di fficult Tanaka (229) for exa -ple fi led to find varieties resistant to root aphids Selecting for large root bioniass perhaps should be the strategy for pests sLuch as Whlite grubs that renliove roots Fainter(174) howcvcr cites varietiesof crops tolerant ofor resistant t(i root pests Some sorghun varieties are high in cyanide so rice sorghun might incorporate broadshyspcctruri chemical resistance

Seedling vigor and drought tolerance are high prioritybreeding objectives so tupland rices should have large r root systernsand be able to tolerate ligherlevels of root loss Sonie root loss inolder plants is beneficial if it stimulates new root developnrut and therefore crihances intake of soil nutrients (53) The rice seedling inaggot rtiLotia spp may be

controlled thirough resistant aittics A hi rgc breeding effort on sorghniu has had fair success inideveloping arreties esrstait to Asoccata Rodani (251) A field trial comparing 10 upland ricc varieties showed differences aniong varieties va ring from II to 4(- damaged tilleis (215) Hovver resistarce to 1Iill-0r gtia species may rot cross ox-rti otier

Shiraki (210) reported that upland i-ice stens wcrc harder than lows-land rice stemsperhaps becaIusc (f a higher silicoricontent and thus were more resistant to yellow stern borer hc tortia Iow tillering (f uplaid iice xarneties prodtres a

higher percentage of infested tillers than would occor in higher tillering lowland iice under a siiilar egg density The corollary therefore is to select for high tilleriig upland rices

Stemii bor amcstheh main targes dobrecders inAfrica (571 204 )but nticl oifth e sc reen irig is dotie unde r lo wland co d t osrm s wih A h r~ o l pe i s n acorditions (11 96 218) As with lthrinrnwspecies mayhave to be dealt with independently however initial results show cross resistance between African and Asian species(92)

In Brazil screening Ior insect resistance focttses oil lou I pest problems Varieties tolerant of lasmolwalpus ligi-sellus and resistant to )iatraea saccharalis have been identified (70 153) Current work also emphasies 7iraca limativetntris and Thmialspis (=Iosis) flavouicta (66) These insect pests Ilt onlV citise IIigh yield losses tihey also ire difficult to control chemically ofr by other ilealls

ManV upland rice vaieties aic early maturing and are resistant to pests such as gall midge (I Xl ) Early mIturity is often a highly desirable trait to esuilpe pest buildup but it is not true resista nce bca usc ilthose va reties are plinted late they will be itsdamaged as ain early planted susc ible variety

P~erhaps tile cosiiiopolitall of all upland rice pestsimost are the sced pests (ripenling stagc) which pose a particular problem beta use scuds re ised IsIO for hitnit4ans Ayltoxic substatice introd tcld geticticllyilmay also be toxic to nal iniless it caln he placed old in thfe IdhuffSelectmi fo hairiness or lotng wns kill mIlct sthilrohleiiis dutilig threshing pa rtictfarl maitial threshinfl [hireshers ill complain of itcfimin atid itteascd (fust levels

Attempts itIRRI to fitid tcistatncc to the rice bug plorisa have ilotbeCnl prmitisilg lIIetIattonlids iorc

through the lenilna or palca hillhairs sedstVIor seeds wilh awns that deter bugs promote complaints from threshers Seed bugs liptocorisa and Stcnocoris enter the seed through the opening between tile lemna and palC but varieties with narrow openings are diifficult to mill

Breeding priorities should concentrate ott inotCI gruLtLIps that are most difficult to c nttrol by other meains Steil) horers appear to be the most widespread group that sigiiificantly dalag s rice aind is hard to manage by other Ilnca us

Biological control Very fe stutdic of mitatt~l eneties of rice pests in upland envirollnletits wee in tile wefouiid literature Therefore have not compiled a list of species Ve present a list of natural enemies iOtid in upland ricefields tuic latialln Batangas for some conitno iOldiar pest species ([itble 5)

Muich ofl thc lollowinle itfor ittiol cotlliCs rol Stutdies in the lowlatnds or roi crops otithcr thlin tice

Qu(ailtr o 11ll t11t( Nat trrtf lii hcoeits ight Ile used agaitist ile social insects tilt and termites Sollie predator specice live ill theirf nests n ilnic theirnllt ip-pearanIlCe aMRidbehavior IJtrouMdCing a mote aggressise ant species to displace pest ant codonies is a possibility (169) Patihogens as potential natural encnies have reccived little atteition (I13) Natural ncnuics asa group however hac not been toIl iporta it in Iegulating aintshown be and ternite numbers

The nocturnal behavior aid subteitmien habitat of lole crickets and field crickets protect them from many ntllral enemies There are only limited records of natural cnelics for these groups of pests some pathogens (87) but mainly

IRPS No 123 JattuIny I 271987

parasites - scelionids on eggsand sphecids on ilymphs and adults Sphecid wasps Larra carutmaria (SImitI hI L huzwuensis Rohwer and lsaltiuhta Willi ams speciali in mole crickets pa ralyzing them nd dragging the ninto thcir nests as food lot their voting ()tier sphCcid species Lins aurulenoa (Iabricius) lot maiac (Ashmead ) 1 lhshylssellaus (Smith) and Ml lal)oriosu (Snithf specialie in field clickets

White grUbsO the other hand hae Icti scriuns enuigh to have been extensively studied for cotttroi 1W nati al enemies( 112 140 141) Scoliid parasites C(ajsoMcispp have beetn intrtduced itto tilePhilippites for control of Ielxl ]udisiro la The rcsearchcts w Intttrdced themle during ill era claimed that the white gttionthreak as controlled (142) howevcr it isnormal for pest epidemics to subside and there was no diect evidence to indictte that tfe parasite was responsible A tmore promisitg titctllod might he to identify iso ltcttass prt~f tcartd apply hiactelaln fingal pathogens to tile soil Ihe lalattese beetle in North America has hcci coitrolled hv tite conmnleciall aailable hactcrial prcpa ration itacihlpu pav ofr milksdiscas (71) Spote prtyefott the soil itloted ilrtier reitiain actic fr Veas

Root aphid sardtnctly bugs readilslic tilib to predatots and parlsites once their tetndine ants are controlled The larvac and adtilts of a coccinellid beetle 1Scynms sp prey on llriadlouais in slash and burn upland rice Baits treated with insecticide may be used to control tending ants by killing their ourig (244)

Root-tkcditig blugs false wireoris wireorms and root wccils aic r ofsuch little ifilpoltatice that thenomttll role of liatural Cticniics hias tiot beei asscssed A possibility hoevrl tiglt he tit ItaIIitic niaCIMtodcs lintt1g11en Itese soil-itlhlfbitine mCtitodcs W5uhtfhave to be imiass Produiced

large miltibcr of niatltrll cnemics ircknown ft CltIoriS IC IpblCeii is thtt clttvortnlls ofonlie the crop soott alter land ptel)tritiOti a1tithe bcgit iniig of the raittv

Citsoln hfien tiatllral leniiiy populations aric lo Colltloil WOtuld iii ole mass prodtticing andirclcaslng key spccies

le egg stage is the fpirt (olthe Cedlitgt Ialot life cycle tI hich to f)ctis ftor tittlril elctit cottntrol hplraites o1 pleditors ttsit elleggs highly expoIsYCl I Sgg pailsites or predators stolld probahly e ittass andtbe produced released Littlh i toruialioll is a ailable ott what species to trV o ho to Iniss podLice thiCIi A etiloplhid wasp Icmihlaol sp effctiicly regushy

lates the leatf lcloalapolm2a spioanoiiitr heat andc rice antid should be collsctvcd

leaf beetles whCllose e am] eggs ate laid in the grutindhirO ould be motC difficutlt to control by niatulral enelV Mnll ipuhltiOn But species sUici IS c_gsthispid beetles ith

and larVle oilleascs ate vtilcratble to iparaites and pledators

Natural enemisc of tlirips otl rice are little known

28 IRPS No 123 January 1987

Table S Parasites recorded from upland rice insect pests in Tanauan Batangas Philippines 1977-84

Order Family Species Host stage

I lymttenoptera Braconidae Leaffolders Cnaphalocrocis Marasinia spy

Cardiochilesphilipplinensis Ashmead a Larva

Cotesia angustillasis(Gahan)Cotesia or cypris (Nixon)

Larva Larva

Cotesia or taeniaticornis(Nixon) Larva Cilonusinotakatae Munakata Chelonits SppAlacrocetrts philippinensis (Ashtmead)

Egg to larva Egg to larva Larva

Iclineumonidac Orgilts spp Isclutojppa hireator(Fabricius)

Larva Larva

Itoplectis naraigav(Ashmcad) Larva Tenelhtcha philippiensis (Ashmead) Larva Tenelchha stangli (Ashmead) a

irichomma cnaphalocrosis Uchida Larva Larva

Bethylidac Elasmidac

Goniozms or trianguli r Kieffer lasmtusalhopictus Crawford

Larva Larva

Diptera

Encyrtidac Chalcididac

Tachinidac

Elasmtus spp Copidosomopsisnacohiae (Fidy) Brach nteria evcarinata Gahan Biachyieria lasus (Walker) Ztgohothria ciliata (Wulp)

Larva Egg to larva Pupa Pupa Larva

Argyrophylx nigrotihialis(Blaranov) Larva

l-ymenoptera Braconidac Yellow stern borer Scirpopiaga incertulas

Braco chinensisSzeplegeti Larva CIthelotis spp Chelonus niunakatae Munakata Cotesi flaripes Cameron Stenoracon nicerilli (Bingham) a

Larva Egg to larva Larva Larva

Ichneumonidac Tropobracon schoenobii (Viereck) Amatroinorpiaaccepta metathoracica(Ashmiead)Frihorus sinicus (I lolmgren)

Larva Larva Larva

Ischnojoppaluteator (Fabricius) Larva Isotima nr dammeratiti Rohwer Larva Tenelucha philippinensis (Ashmead) a

Larva Tenehicha stangli (Ashmcad)Trichomma enaphalocrosis Uchida

Larva Lara

Chalcididac Eulophidac Trichogranmmatidae

Xanthopimnpla stemmator (Thunberg) Charops hrachYpterumn Gupta and Malteswary Brachtm eria sppTetrasticttus schoenohii Ferriere Trichorigranm chihunis Ishi

Larva to pupa Larva Pupa Egg Egg

Diptera

Scelionidac Pteromalidae Tachinidae

Tt ichograintnta aponiicuitt (Ashniead) Telenomits rowani (Gahan) a

Trichoinalopsis apattelctena (Crawford) Jeirhaea spp

Egg Fgg Egg pupa Larva

Z4gobothria ciliata (Wulp) Larva

Ilymenoptera lecneumonirlac Gol fringed borer (hilo auricilius

Trichomm enoptalocrosisUchida Larva

Ieioenoptera Braconidae

Aantholimlh steiniator (Thtinberg) a

Pink stet Ioret Sesamia inferens StoInacoil nicrilhi (Bingh am)

Loarva to pupa

larva

Diptera Ilyinenoptera

Armvtnyorms Mythimna separata Spodoptera mauritia acronyetoidesTachinidae Zvgohothriaatropiora( Robinea i -Desvoid y)Eulophidae It-phctrtiscitapadoe (Ashimead)

Larva Larva

(halcididae Braconidac

Brachpin ria spip (oesia spp

Larva larva

IHyntenoptera I)iptera

Rice skipper Ielop ida (halcididae Tachinidae

math ias aidgreen iortedci ciliarNIelan itis eda Brat lhincria sp iii matuqitatt Cameron Irgyrolhirlaxi trotlhiais(marumtv)a1

ismtne Larva Larva

Diptera Ilymenoptera

Tachnidae lelineutionidae

fuown se tilo per Mois frugalisIrgtrophrlax 1igrothiahiv 3 a rat ov) anthopimpa pmnctata (Iabricius)

Larva Larva to pupa

aMost dominant

IRPS No 123 January 1987 29

Normally predatory thrips prey on herbivorous thrips strepsiptera on both Spiders dominate tilenymphaladultNatural enemies of stem bugs --Scolinoplara and Tibraa predators Spittle bugs move to rice from pasture grasses - are in order of importance egg par~isites and predators and biocontrol would be successful only ifcarried out before fungal pathogens and nymphal adult parasites Egg tiley immigrate into ricefields parasites could either be introduced fron other areas or Biological control tactics for hemipterous seed bugs mass produced and released ould follow those outlined for stein bugs

Armyworms are normally lwld in hcck by the activities Coml)arison of environnewtns Natural enemies are of egg and larval parasites When these natural enemies fail perhaps faced with even greaterconstraints than insect pestsusually because of drought armyworns become epidemic to survive ind reproduce in upland environments TheyIt may be worthwhile therefore to release parasites during have adapted ani survived by having wide host ranges the the rainy months following a drought Virus diseases of ability to aestivate or hibernate over unfavorable seasonslarvae would beanotheravnue to exploreforarnyworms and the ability to disperse or by other mechanisms

Parasites have ben traditionallyconsidered for biolegical Comparing the most dominant and therefore the most control attempts against stein borers usually by rekasing adapted species of natural enemies collected in various exotic species( 166) This approach has not met with success environments in the Philippines we can understand how and new avenues should be explored The role of predators different rice environments are arid pathogens is little inderstood and deserves greater Anagqrms olptahiisa mymarid planthopper egg parasiteattention Egg parasitism rates are normally reasonably is cquallyadapted to all Philippine rice environments (Tablehigh but should be supplemented with effective egg 6) It is selective for planthoppers and even parasitizes eggspredators Orthopterans are potential egg predators of the n-iize planthopper Peregrinitsmaidis itt upland areas Metioche vittaticollis (Sta ) Anavipha nifiwnnis (Saus- Alavohis also parasitizes maize planthopper eggs but is sure) and Conocephahs hongipnni (de i-laan) feed on most prevalent on BPH in the irrigated wetlands (onashystem borer eggs etioche and Anaxil)ha specialize in eggs tocerus spp like A optalhilis on BPH and WBPI Iattacks with no hair such as Chilo spp The omnivorous (ono- the eggs of GLH in ell rie environments It is also found cephahsspecializes in eggs cov red with hairs it is a proven parasitizing the eggs ef the white rice licahopper (oimna egg predator of Scirlouphaga spp but has the discouraging spetra Oirosita naias a trichogrammatid egg parasitehabit of eating rice guin (65) attacks BIPH and WBPH in the wetlands and has no other

Grasshoppers have egg (scelionid) and nymphalad ult host A related species 0 aesopi specializes in G1-H in (nemestrinid tachinid and sarcophagid) v irasites Among wetland environments Dryinid strepsipt-an and pipunshypathogens protozoans have been recorded most frequently culid nymphal and adult parasites of planthoppers and Control of locust species by natural enemies could con- leafhoppers also specialize either in hopper species or centrate on their habitual breeding areas environments Among the dryinids Pseuhdgonatoipsflavi-

Leaffolders have rich complexes of natural enemies some feimur prefers the lowlands and BPH while P nuch has no adapted to upland rice (22 27) ranging from egg predators environmental prefirences but is adapted only to WBPH (gryllids coccinellids) to larval parasites (braconids ichncu- The main d ryinid species on GLH aptogonatop)usspp is monids) to larval prcdators (ants and carabids) to larval most dominant in rainfed lowlands and also attacks other pathogeiis (vira and fungal diseases) Efforts to augment leafhoppers within Amnrasca Cicadulinaand Bachaltuathese natural enemies should focus on the egg predators Strepsiptera are more dominant on plarithoppers than (rearing releasing)and viral or fungal pathogens (culturing lealhoppers The Elenchus species in the Philippines attack spraying) all species of planthoppers within the genera Sogawilla

Parasitization of the large conspicuous larvae of Sogatodels NilaparsataHarualia and Opiconsiva Elenshyskippers green horned caterpillars and other polyphagous cusrvasuntaisuiis adapted to the major rice environments Lepidoptera by tachinids and chalcids is normally low ELg but prefers BPH in rainfed arid WBPIH in wetland environshyparasites and predators are perhaps the key to their control ments Ilalictoph(utsmtnroeiand HVpectrus attack G LH

Gall midges are parasitized by pterormalids and platy- and COfimravcctra respectivelygasterids that aestivate within their larval host between Pipunculids are mainly parasitic oii lealhoppers Each seasons Shoot aphids are preyed upon by coccinellids species has a unique environmental preference Tomosshysyrphids and chrysopids varireIa oraetoraprefers Ne7 otetltix nigropictrs and

Planthoppers arid leafhoppers have egg parasites and Pilunwulhtsjavanensisattacks Dehoceplhahls spp which predators as well as nymphialad nlt parasitesand predators are more abundant in upland than irrigated lowland their rich complex of natural enemies includes fungal environments T suhvirescensattacks Nephotettix virescens pathogens The principal egg parasites are myrnarids and and prefers the lowlands Pilpuiulus nttillattls is an trichogramniatids Egg predators are norinally mirid plant upland rice parasite arid attacks only Nephotetix speciesbugs that suck out the yolk Nymphal ad nIt parasites are More than 17 recorded species of leaffolder parasites in dryinids on planthoppers pipunculids on lealhoppers and the Philippines attack species of C(naplwocrocis and

30 IRPS No 123 January 1987

Table 6 Dominant natural enemies of rice pests in upland rainfed lowland and irrigated lowland rice environments of the Philippines

Natural enemy

Host stage Species

Brown planthop-Parasite Egg

MymaridaeAnagrus optabilis(Perkins) Anagrusflaveolus Waterhouse

TrichogrammatidacOligosita naiasGirault

NymphAdult Dryinidac

Pseudogotatopuslaiifemttr Esaki et Hashimoto Elenchidae

Elenchus yasurnatsui Kifune and Hirashima

Environment

Upand Rainfed lowland Irrigated lowland

Ailaparata lugens

x x X x

x

x x

x X X Whitebacked planthoppers Sogatella furcifera and Sogatodes pusanusEgg

MymaridaeAnagrus optabilis(Perkins)

TrichogrammatidacOl~gositanaias Girault

NymphAdult Dryinidae

Pseudogonatopusnudus Perkins Elenchidae

Elenchus yasutatsui Kifune et Hirashima

Parasite Egg

MymaridaeGonatocerus spp

TrichogrammatidaeOligositaaesopi Girault

NymphAdult Pipunculidac

Tomosvaryella subviresccns Loew Tomosvaryella oryzaetora (Koizumi) Pipunclusmutillatus (de Meijere)

DryinidaeIlaplogonatopusspp

Egg Copidosomopsisnacoleiae (Eady)

Larva BraLonidae

Cardiochiles philipphnensisAshnead lchneumonidac

Trichiomnia cnaphalocrosisUchida Temelucha stangli (Ashmead)

Pupa Chalcididae

BrachymeriaexcarinataGahan Brachymerialasus (Walker) Brachymeria nr tachardiaegroup

Parasite Egg

Scelionidae Telenomus rowani (Gahan)

EulophidacTettasticusschoenobhi Ferriere

Larva Braconidae

Stenobraconnicevillei (Bingham) Iclhncurnonidac

Temelucha phiiippinensis(Ashinead)Temelucha stangli (Ashmead)

x x x

x x

X x X

x x Green leafhopper Nephotettix virescens

X x x

x x

X X X X

X

X Rice leaffolders CnaphalocrocisMarasmia

x X x

X

X X

x x x x x X x x x

Yellow stem borer Scirpophaga incertulas

X X x

x

x

x X XAinauromorphaaccepta metathoracica(Ashmead) x

continued on next page

IRPS No 123 January 1987 31

Table 6 cntinued

Natural enemy Environment

Host stage Species Upland Rainfed lowland Irrigated lowland

Larva-pupa Gold fringed borer Chilo attriciiuslchneumonidae

Xanthopimpla stemmator (Thunberg) x aPupa x

Pteromalidae Trichoinalopsis apanteloctena (Crawford) ax

Pink stem borerb Sesamnia inferensParasite

Larva Braconidae

Stenobracon nicevillei (Bingham) x b b Armyworms Mvthinna SpodopteraLarva

Tachinidac Zygobothriaarroptvora (Robineau-Desvoidy) x x

Braconidac Cotesia spp

X Skipper Pelopidas mathias and green horned caterpillar Melanitis leda ismeneF-gg

TrichogramrnatidacTrichogrannaspp x

Scelionidae Undetermined Scelionid

x Larva

Tachinidae Argyroph lax nitrotibialis (Baranov) X x

Pteromalidae Trichomnalopsis apanteloctena (Crawford)

X Predator

CoenagrionidaeAgriocnenis femina femina Brauer x xIschnura senegalensis tambur

Miridae CrtorhinsliidipennisReuter x x

Nabidac Stcnonabis tagalicus (Stal) x

VeliidaeAlicrovelia douglasi atrolineata (Bergroth) x

Coccinellidae Micraspis crocea (Mulsant) X x x

Vespidae Eumenes campaniformis (Fabricius) x

Formicidac Diaecamna spp xOdontoponera transversa (F Smith) x

Carabidae Chaenius spp OphioneaW- aJasciataSchmidt-Goebel

x

Ophionea ishii ishii Habu x x X

Gryllidav Anaxipta longipennis (Saussur-) x xMetioche ittaticollis (Sial) x x x

Tettigoniidae Conocephalus longipennis (de Haan) x x x

aNo rearing made on rainfed lowland bNo rearing made on rainfed lowland or Irrigated lowland

MrasmiaAmong the Laval parasites the braconid Cardio- lowlands and parasitizes oriental maize borer OsIriniachilesphilijpphensis is most adapted to an upland environ- furnacalis (Guenee) larvae The leaffolder pupal parasitesment It also parasitizes Hydelepta indicata a common not only occur equally in all environments but also haveleaffolder of legumes The ichneumonids Trichomma cna- wvide host ranges- parasitizing species of OsIrinia Marucaphalocrosis and Temehcha stangli occur more in the Hydelepta and Honiona - in maize and legumeslowlands T stangli is most adapted to irrigated lowlands The principal egg parasite of yeltow stem borer (YSB)and also parasitizes Chilo and Scirpophaga stem borer Telenomus rowani is widely adapted to all environmentslarvae T cnaphalocrosis is most prevalent in rainfed and has an unusual alternate host -Tabanus eggs (25)

32 IRPS No 123 January 1987

Tetrastictu schoenobii specializes only in YSB and occurs mainly in irrigated lowlands -- further proof that YSB is most adapted to the wetter environments

Larval parasites of YSB are highly environment-specific The braconid Stenohracon nicevillei occurs only in upland rice but also attacks the pink stem borer (PSB) Tieinchwa phiiiplPwnsis an ichneumonid occurs in upland rice and also parasitizes the larvae of (7hiu stqpresaisinatro-morpha accepta melaihoracica like 7 stangi is mnost adapted to the irrigated lowlands

The dark-headed stem borer(l)HSB) (ih pochit3sus was not collected in all rice environments and the gold fringed borer Chilo auricilius nd PSB were collected only in upland rice Xaantiholiplastenrnaloran ichneumonid larval parasite appears to be widely adapted whereas TrihomaslopisapanitI)cteiiaa pteromalid pupal parasite occurs mostly in wetlands and also parasitizes larvae of Chilo suppressais and Pelhuidas mathias

Armyworms Myit hinin and Spodujipera have two main larval parasites - a tachinid Zrgohothriaatropivora in rainfed environments and a braconid Cotesia spp in irrigated lowlands

The butterfly pests Pelopidas and 4elanitishave tricho-grammatid egg parasites found only in irrigated lowland environments

Two larval parasites show environmental preferences --Argrroph av nigrotihialis a tachinid in rainfed habitats and Trichonialopsisapantelkcnela in irrigated ricefields A limgrotibialisalsoattacks the sweet potato hornwormAgrius convolvtdi and T apanteloctena parasitizes DH S1B

Predators as a group show more distinct environmental preferences The aquatic damselflies Coenagrionidae are most abundant in the lowlands Cvriorhinus is also most adapted to the lowlands The nabid Stenonabis tagalicus occurs only in the uplands and isalso prevalent on legumes The coccinellid lady beetles wasps and ants are more prevalent in upland areas AlicrasIpis cit-ea larvae prey ona variety of aphids attacking legumes and maize Etunienes campanonia vespid mud wasp makes nests in trees and is therefore most adapted to the more botanically diverse uplands Soil-dwelling ants cannot tolerate flooding

Upland environments are habitats to arboreal carabid beetles Three species of Chlaeniusbeetles prey on leaffolder larvae and are perhaps more important than parasites (27)

The gryllids and tettigoniids are widely adapted egg predators not only on rice but also on maize and legumes They feed on eggs of most insects that are laid on leaves

The spidercommunity of upland rice environments isrich in species At one site (Batangas Philippines) 31 species have been recorded (23) Of the 176 spider species recorded in Philippine ricefields about one-half (82) occur in upland rice The spiderspecies of upland rice environments overlap more with those of rainfed lowland than with those of irrigated lowland rice (24) The same study showed that of the three environments irrigated rice has (ie greatest spider species diversity foiiowed by rainfed lowland and upland

However of the 10 most prevalent ricefield spiders 9 were abundant in upland rice showing wide adaptation All three spider guilds -- orb-web space-web and huting spiders shywere prevalent in upland ricefields There were however differences in environmental prelerences for some spider species

Of the orb-wcavers Tetragniathamanlihulatawas partishycila rlv abundant 7 japonica was low and Ltucatge dehcorata was absent in upland rice

As in other environments Aiypeiwnaifirtosana was the most dominant space-web spider however its relative numbers were lowerthan for lowland sites Among hunting spiders Ox|opes javanus and Lcosa leatcost fgla were more abundant and Lvcosa lpseudoanmnlata was less abundant in upland compared with lowland rice envishyroniments

Extensive rearing of rice insect pests in upland rainfed lowland and irrigated sites over an 8-yr period in the Philippines provided insight into the effectiveness of parasites as natural enemies (Table 7 8) Most of toe data are from large sample sizes taken over at least one crop which overcomes the pitfalls expressed by Van Driesche (238)

VSB egg parasitization was surprisingly similar across all environments Larval parasitization was highly dynamic among sites and even years within the same site but showed somewhat lowerlevels in uplandsthan in irrigated wetlands This would have been predicted for egg parasites as well since YSB is adapted to the lowlands and so must be its parasites This isevidence that Telenomus rowani is as good at dispersing as its main hosts Trowani clings onto the anal tufts of female YSB and parasitizes eggs as they are laid This phoresy may explain its wide adaptation The more specialized lowland YSB larval parasites therefore appear more effective than their upland counterparts with more alternate hosts

Low collection levels of PSB larvae especially in the wetlands make comparisons between environments difficult Low levels of larval parasifization were recorded in both rainfed environments Except in the Cagayan rainfed wetland site in 1980 low levels ef larval parasitization were evident at irrigated and rainfed sitesalike No differences by environment were evident Higher levels of parasitization occurred in Sarawak during a 1967 outbreak also from tachinids Rothschild (197) indicated that higher parasishytization occurred in upland rice areas

Skipper and green horned caterpillar larval parasitization rates were generally low and showed no environmental effect htighest activity occurred in Cagayan in 1980 (48 parasitization) Although larval parasitization levels of RLF were variable year to year activity appeared greater in the irrigated lowlands than in the uplands Again the highest levels were recorded in Cagayan in 1980 Cagayan has extensive grasslands and fallow around the rice areas and a relatively long rainfall period Rainfall in 1930 was good for crop growth and apparently for increase of larval parasites

IRPS No 123 January 1987 33

Table 7 Parasitization rates of rice planthoppers and green leafhopper in upland rainfed lowland and irrigated lowland environments Philipshypines 1976-840

Environment and site Year Egg parasitization () Nymphaladult parasitization ()

BPH W13P GLH BPII WBPH GLIH

UplandBatangas

Rainfed lowland

1976 1979 1980 1982

15 17 12

2(47)8(39)

4(51)

9(1800)14 (212)

12(304)

8 (101)6 (127)

9(116)

Iloilo

Cagayan

1976 1977 1978 1981 1980 1981 1982

0 (69)

11

44 48 40

-

-

46 55 44

0 (471)

11

52 56 49

0 (200)9 (214)

10 (1259)8(235)6 (900) 3(800)

-7 (462)

11 (270)3 (313)5 (900) 1 (800)

4 (600)10 (725)10 (362)12(870)5 (900) 3 (800)

Pangasinan

Irrigatedlowland

1984 1976 1978 1979 1982

13 (436) 13(379) 14 (457)

6(14)7 (197)

17(198) 11 (486) 18 (275)

3 (81)10 (274)3 (116)

16(871) 19(88)

28(211)9 (764)5 (647)7 (1204) 8(305)

Laguna 1977 11 7 21 1978 31 1979 14 4 4 28 1980 7 8 20 1981 1983

56 7(278)

26 16 (1650)

16 27(1921)

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R I Apostol unpublished 1lo1lo and Pangasinan 136 Cagayan BCanapi unpublished Liuna 175 I C Pantua unpublished 118 234 43 Carino and Shepard unpublished BPH = brown planthopper Nila parvara htgens W131Pi =whitebacked planthopper Sogatelia furcifera GLH = green leafhopper Nephotettrix virescens Figiires in parentheses are sample sizes

Tauher et al (230) reported that internal external and genetic factors influence the seasonal activity of parasitoids

Hopper egg parasites but not nymphaladult parasites were also parlicularly abundant at the Cagayan site possibly the egg parasites have more alternative hopper prey in the weedy areas than nymphaladult paisites

The seasonal dynamics of hopper egg parasites can be seen for upland rice in Batangas(Fig I 2) and for rainfed lowland lice in Pangasinan( 135)and Cagayan( 105) Levels of BPHWBPI-L and GLH egg parasitization were corn-parable in upland and irrigated lowland environments Except for some high rates of paraMsitization early in the season the parasitization rate tended to be steady in all envilonments However hopper nymphaladult parasi-tizution levels were more similar in the rainfed environments and lower than in irrigated rice

Overall parasites do not appear highly effective by themselves against upland or even lowland pests Among natural enemies parasites have attracted the most attention because they can be more readily assesscd

Predators and pathogens lend themselves better to management practices Predators the most important group of natural enemies in riccfields are difficult to quantify On the other hand pathogens appear less im-portant but can be readily cultured and disseminated

The natural enemy commulnity of upland rice is rich in species and differs significantly from that of lowland rice

These beneficial organisms must be conserved by applying insecticides judiciously particularly sprays Seed or soil placement of chemicals minimizes exposure to natural enemies The strategy to derive the greatest benefit from natural enemies isto allow the greatest number of beneficial species to thrive Then perhaps at least one species will be effective against each pest at any tine overcoming the variable abundance of each species within and between years

Programs to introduce exotic species or mass produce indigenous ones are ambitious and expensive because they require trained people on a sustained not ad hoc basis

Chemical control Insecticides offer rapid and efficient control of upland insect pests (190) but-should be used only after other control measures have been considered Insecticides are rarely used on upland rice because of cost (32) Upland rice yields are normally too low to justify the expenditure Also most upland rice is grown as a cash crop Spraying upland rice is more difficult than lowland rice because water is less accessible Government subsidy programs to provide pestishycides to tribal peoples for upland rice production have created problems of toxicity to humans because isolated people have had no experience to be able to handle pesticides safely

However ins-eticide use has beenjustified when a crop is

0Z

Table 8 Parasitization rates of lepidopterous pests in upland rainfed lowland and irrigated lowland environments Philippines 1976-84a

Parasitization (5)

Environment and site Year YSB GFB PSB Earcutting Swarming Skipper Greenborned RLF Brown larva larva caterpillar caterpillar caterpillar caterpillar larva semilooperEgg Larva larva larva larva larva larva

UplandBatangas 1977 - 0 (57) - 0 (63) - 0 (71) - 0 (106) 11(142) 2 (101)1978 43 (4697) 8(451) 5 (38) 3 (40) 6 (61) 2(58) 15 (57) 13 (129) 13 (1286) 4 (30)1979 59(4135) 12(320) 13 (11) 5(28) 9(45) 6(46) 7(23) 8(160) 9(1152) 8(26)

Rainfed lowland Iloilo 1976 - 0 (10) 0(10) 0 (7) 0 (19) 2 13) 15 (180) 5 (76)1978 0 (54) 0 (3) 0 (17) 9 (64) 4 (78) 3 (73) 0 (121) 0 (191)1979 64 (2350) 16 (84) 6 (17) 0(6) 18 (68) 25 (104)

1981 56 (1322) 18(304) 0(5) 12(51) 18(138)Pangasinan 1976 60 (7 7 )b 62(82) 0(111) 5 (39)

1978 58 (53)b 5(51) Cagayan 1979 59(713) 8 (101) 6 (34) 0(7)

1980 0(11) 4 (28) 15 (286)

62 ( 48(177) 61(140)1981 0(5) 12 (2) 13(45) 14 (92)

Irrigatedlowland Laguna 1974 62 8

1974-75 50 41 1975 43 72 1978 96 (623) 21(24) 33 (60)1979 83(2017) 18(110) 8(62) 11 (89) 22(400)1980 78 (1625) 20(75) 0 (16) 4 (72)1983 60

9 (80) 7 (270) 30

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R F Apostol unpublished Iloilo and Pangasnan 136 Cagayan B Canapi unpublished Laguna 175 P CPantua unpublished 118 234 143 Carino and Shepard unpublished YSB = yellow stem borer ScirpophagaincertulasGFB = gold fringed borer Chilo auricilius PSB = pink stem borerSesamia inferens earcutting caterpillar Alythimna separata swarming caterpillar Spcdoptera mauritia acronyctoides skipper Pelopidas mathias reen horned caterpillar Melanitls ledaismene RLF = rice leaffoldersCnaphalocrocismedinalis and Marasmiaspp brown semilooper Mocisfrugalis Figures in parentheses are sample sizes Egg masses

IRPS No 123 January 1987 35

Table 9 Effect of seeding rate and insecticide on plant density and Yield losses from insect pests in upland rice often areyield of Dagge upland rice Batangas Philippines 1976a comparable to those of lowland rainfed or irrigated rice

Seeding rate Insecticide Dosage Plant density Yield (0-Y30o) but usually from different pests The principal (kg aiha) seed treatment (kg aiha) (noIra-row) (tha) groups are soil pests feeding on sown seed and rootsat 14 DE followed by seedling pests These groups have largely gone

50 None - 35 a 20 a unnoticed leading researchers to conclude that stem borers50 Carbofuran F 10 38 ab 23 a and seed bugs are the most important pests (17) Yield loss 10 beCabofuranF100 None - 4551 c 23 a trials that pinpoint major pests need to includc seed

100 Dieldrin WP 10 52 cd 22 a treatments Granules applied in the furrow may not give150 None - 59 d 22 a adequate protection from many seed pests

aAv of 8 fields DE = days after crop emergence In a column means The floristic and pedological diversity of upland rice followed by a common letter are not significantly different at the environments is matched by the diversity of arthropods5level by D)MRT F = flowable Wi = wettable powder both pests and their natural enemies Irrigation has homoshy

genized lowland rice environments Flooded soils tend tothreatened by epidemics of locusts or armyworms New have the same properties and to eliminate soil pests The technology stressing efficiency has minimized insecticide spatial and temporal dominance of cultivated rice has cost benefited specialist pests

Seed treatment is normally inexpensive and can be Upland rice has no specialist arthropod species The economicallyjustified in -nanycases(66) Treating seed with monophagous pests on upland rice prefer the lowlands systemic insecticide (216) but not chlorinated hydrocarbons where rice evolved Rice in the uplands exists only because (247) is most effective in controlling seedling maggots of man and even though lowland rice predates upland riceFoliar sprays have to be repeated to obtain control and specialist species evolved Wheat maize and sorghum do often are washed off by frequent early season rains not have monophagous pests leading us to conclude that Granular insecticides are effective but required dosages are there is no evolutionary advantage for inssects to have too expensive to justify except under heavy infestation narrow host ranges in upland graminaceae(247) When directed against first-instar larvae banding Pests have adapted to the uplands not only by having granules in seed furrows (177) can be effective against white wider plant host ranges but also by having long life cyclesgrubs at low dosages (025 kg aii ha) (134) Seed treatments undergoing dormancy or being dispersive Each pest has a are not promising for white grub control (248) unique life history strategy suited to highly unpredictable

Seed treatment is also effective ainst ants and may be environments In agivenyear someinsect specics an better cheaper than increasing the seeding rate In Batangas exploit upland rice as a food source but no pest is a senious treating 50 kg seed ha was as effective as increasing the one every year on upland rice seeding rate to 100 kgha and using untreated seeds (Table The low yielding potential of upland rice means less 9) In Brazil seed treatments control termites and other soil quantity and lower quality of food for pests to exploit but insects including the lessercornstalk borer(66) Domiciano also less incentive for farmers to undertake control (58) however found seed treatments inconsistent in their The most effective control would be early and synchroshyeffect against high populations of lesser corn stalk borer nous planting of early-maturing upland rice varieties to

Baiting can be an inexpensive way of controlling ants prevent insect pest buildup Breeding for insect resistance seedling maggot flies mole crickets and field crickets Baits should initially focus on stem borers because they are verymade of locally available material and impregnated with difficult to control by other means Biocontrol efforts insecticide can be sparingly distributed taking adantage of should first stress conservation of natural enemies preshythe pests ability to disperse and encounter bait sites dators and pathogens rather than parasites Efficient

The new low-volume hand-held controlled-droplet chemical control includes seed treatment or baiting for seed sprayers may offer an advantage over high-volume knap- seedling and some root pests and using controlled dropletsack sprayes for controlling foliar pests Upland rice sprayers for spot treatments against foliar pests and seed canopies are more open than those in lowland rice and bugs when economic thresholds are reached better droplet penetration should result Controlled droplet applicators would be ideal for seedbugs and defoliators

CONCLUSION

Upland rice is attacked by a wider array of insect pests than is lowland rice mainly beca use of the addition of soil pests Generally population levels of lowland rice pests are lower in upland than lowland rice leading many people to conclude that insect pests are not important on upland rice

36 IRPS No 123 January 1987

REFERENCES CITED

1Abraham C C and K S R Mony 1978 Occurrence ofLeptocorisa acuta Fabr (Coreidae Hemiptera) as a pest of nutmeg trees J Bombay Nat ttistSoc 74553

2 Agyen-Sampong M 1975 HlierogrlphusdiaganensisKrauss (Orthop-tera Acrididae) a new pest of rice in Northern Ghana Ghana J Agric Sci 8249-253

3 Agyen-Samnpong M 1976 Varietal resistance and other inle-ratcd pest control components as factors of rice varietal improvement in West Africa Prcsented at the Second varietal improvement serinar 14-18 Sep 1976 Bouak Ivory Coast West Africa Rice Development Association Monrovia Liberia 18 p

4 Agyen-Sampong M 1977 Outbreak of Ihccrorrdnisorae in rice seedlings near Rokupr Sierra Leone lit Rice Res Newsl 2(6)16-17

5 Agyen-Sampong M 1980 Dirty panicles and rice yield reduction caused by hugs IntRice Res Newsl 5(l)11-12

6 Agyen-Sampong M 1982 The major pest problems of irrigated upland and mangrove swanip rice ecosystems ir the humid tropical Guinea savanna and Salel climate zones Pages 318-328 in Pro-ceedings training course cn concepts techniques and application of irtegrated pest management in rice inWest Africa 10-20 Jan 1982 Fendall Liberia West Africa Rice l)evelopioent Association Monrovia Liberia 505 p

7 Akibo-Betts V I and S A Raymundo 1978 Aphids as rice pests in Sierra Leone lit Rice Res Newsl 3(6)15-16

8 Akinsola F A 1975 Present status of different rice stemborers in parts of Nigeria iicL Entomo Newsl 328

9 Akinsola FA 1979 Ihe biology and ecology of rice stemborers in Nigeria Ph 1) thesis University ol lbadan Nigeria 217 p

10 Akinsola E A 1979 Guidelines of integrated management of insect pests of upland rice in West Africa Presented at the Seminar on integrated management of rice diseases and insect pests at Bobo-Dioulasso Upper Volta 17-22 Sep 1979 West Africa Rice Develop-ment Association Monrovia Liberia

II Akinsola IA 1984 Insect pests of upland rice in Africa Pages 301-305 in An overview of upland rice research Proceedings of the 1982 lIouak Iwsrv Coast upland rice workshop International Rice Research Institute 13O Box 933 Manila Philippines

12 Alam M S V TJohn and Kaung Zan 1985 Insect pests and diseases of rice in Africa Pages 67-82 in Rice improvement in Eastern Central and Southern Africa proceedings of the international rice workshop at Iusaka Zambia 9-19 April 1984

13 Alticri M A A van Schoonhovcn and J D Doll 1977 The ecological role of weeds in insect pest management systems a review illustiated with bean (Phaseohsvulgaris L)cropping systems PANS (Pest Artie News Summ) 23195-205

14 Anmante F 1967 The leaf-cutting ant Atia capiguaraa pest of pastures [in Portuguese] Biol6gico (Sao Paulo) 33113-120 (Rev Appl Entomol A56207)

15Andrade A C and D Puzzi 1951 Preliminary results of experi-merits on the control of Sculptcoris iastantus on sugarcane [in Portuguese] Bioulgico (Sao Paulo) 1744-49 (Rev Appl Entomol A41421422)

16 Appert J 1970 Maliarplia separaelha (rice stem borer) new observations of entomological problems of rice in Madagascar [in French] Agron Trop (Paris) 25329-334

17 Ariadeau Mand Z Harahap 1986 Relevant upland rice breeding objectives Pages 189-197 in Progress in upland rice research Proceedings of the 1985 Jakarta conference International Rice Research Institute 1O Box 933 Manila Philippines

18Asar S and Kisimoto 1975 Seasonal occurrence of the smaller brown planthopper in upland rice [in Japanese] Proc Kanto-Tosan Plant Prot Soc 2228

19 Bakker W 1974 Characterization and ecological aspects of rice yellow mottle virus in Kenya Meded Landbouwhogesh Wage-ningen 152 p

20 Balut F F 1970 White grubs attacking rice Oryza ativa L [in Portuguese] Biol6gico (So Paulo) 36321-322

21 Banerjee S N and L M Pramanik 1967 The lepidopterous stalk borers of rice and their life cycles in the tropics pages 103-124 in The major insect pests ofthe rice plant Proceedings ofa symposium at the International Rice Research Institute September 1964 The Johns Hopkins Press Baltimore Maryland USA

22 Barrion A T and J A Litsinger 1980 Ants a natural enemy of cliaiarac in dryland rice IntRice Res Newsl

5(4)22-23 23 Barrion A T and J A litsinger 1981 The spider fauna of

Philippine rice agroecosystem 1Dryland Philipp Entomnol 5139-166

24 Barrion A T and 1 A Iitsinger 1984 The spider fauna of Philippine rice agroccosystcms II Welland Philipp Eiitomol 611-37

25 Barrion A T1 and J A Litsinger 1984 lihanus (Diptera Tabanidae) eggs an alternative host of rice stei borer (SI3) egg parasite Telenoonis digns (Iliymenoptcra Scelionidae) lit Rice Res Newvsl 9(6)19

26 Barrion A T and J A Litsinger 1985 Identilication of rice leaffolders by wing markings Int Rice Res Newsl 10( 1)24

27 Barrion A T and J A Litsinger 1985 Chctiosspp (Coleoptera Carabidae) a leaffolder ILF)predator lit Rice Res Newsl 10(1)21

28 Bertels A 1970 Insect pests of rice and theircontrol [in Portuguese] Pelotas Brazil Instituto de Expcrimentacao Agropccuararias do Sul Circular 43 24 p

29 Bonzi S M 1982 Ctilo (flisilineus J de Joannis (Lepidoptera Pyralidae) stem borer of irrigated cereals in Upper Volta LioFrench] Agron Trop Riu Rizic Cult Vivrieres Trop 37207-209

30 Bourke T V T L Fenner J N LStihick G Baker EHassan D F OSullivan and C S li 1977 Insect pest survey for the year ending 30th June 1969 Papta New Guinea )epartment of Agrishyculture Stock and Fisheries Port Moresby 57 p

31 Bowden 1 1973 Migration of pests in the tropics Meded Rijksfac Landbouwwet Gent 38785-796

32 Breniere J 1973 The main rice parasites in West Africa and their control Pages 27-42 in Proceedings seminar on plant protection for the rice crop West Africa Rice Development Association Monrovia Liberia

33 Breniere J 1976 The principal insect pests of rice in West Africa and their control West Africa Rice Development Association Monrovia Liberia 52 p

34 Brown ES 1962 The African arinyworn Spoiolptera eXI)Ina (Walker) (Lepidoptera Noctuidae) a review of the literature Cornshymonwealth Institute of Entomology London UK57 p

35 Brown F B 1968 Upland rice in Latin America FAO-International Rice Commission 12th Session of the Working Party on Rice Production and Protection 2-14 Sept Peradeniya Sri lanka Food and Agriculture Organization Rome Italy 6 p

36 Buyckx F J 1 1962 Disease and insect pests of crops inCongo Rwanda and Burundi [iil French] Publications dc LInstilut National pour lestude Agrononiquedu Congo M Weissenbruck S A Brussels Belgium 708 p

37 Cariiio F 0 1981 Role of natural enentics inpopulation suppression and pest management of green rice lealhoppers Ph D thesis University ofthe Philippines at Los Bafitos PhilippinesLaguna 136 -

38 Carpente A J 1983 The development of rice cultivation and extension of vegetable productin in Zanzibar Final Report Vol II1 October 1983 Food and Agricultme Organization Ronte Italy

39 Carroll C P and S1 Risch 1983 Tropical annual cropping systems ant ecology Environ Manage 751-57

4(0 Cates R G 198 1 Host plant predictability and the feeding patterns of monophagous oligophagous and polyphagous insect herbivores Oecologia (Berl) 48319-326

IRPS No 123 January 1987 37

41 Cendcafia S M and F B Caora 1967 Insect pests of rice in the 62 Ene 1 C 1981 I)iiorldhol)ierusihilis Slater (licteroplera Philippines Pages 591-616 in The najor insect pests ofthe rice plant lygacidue Illissinac) a new rice pest front Nigeria lit Rice Res Proceedings of a symposium at the International Rice Research Newsl N3)15 Institute September 1964 The Johns Hopkins Press Baltimore 63 Fugenio T S 1965 rhe effect oi e-vCgetation on rice production Maryland USA 729 p and the possibility of overcoming it by leaf pruning MS thesis

42 Centro Ihternacional de Agricklittira Tropical 1984 Upland rice in University ol the Philippines at Los Baijos ILaguna Ihilippins 69 p Iatin Anerica Pages 93-119 hi An os ervies of uplaind rice research 04 Farrow R A 1975 The Alrican nmigratory locust ii is main Proceedings of the 1982 Boinakamp Ivory Coast upland rice workshop outbreak area of the middle Niger qlinlitatise studies of solitary International Rice Research Institute P0 Blox 933 Manila populations in relation to environmental factors Locusla No II Philippines 198 p

43 Chandra G 1979 Taxonomy ad hn riononlics ofthe imect parasites of 65 Fennal R G 1943 Food crop investigation Windward and rice Icalloppers and planthoppers in the Pihilippines and their Ieeward Islands First report Report onl work dunring the periodiiiportairce iii natural biol)gical cointrol Philipp Fntoltol 4119-139 April 1942-April 1943 Imperial College of Tropical Agriculture

44 Chang T T 1976 The originii colut ion enlIivation dissemination Trinidad 16 p (Rev of Appl Fintornol A3222-25) and diersificition of Asian and African rices -uphytica 25425-44 1 66 Ferreira F 1983 Integrated pest ritaageiret of upland rice [ill

45 Chatierjee P B 1979 Rice leaffolder attacks in India lit Rice Res Portuguese English summary] Pages 323-341 in Upland riceculttre Newsl 4(3)21 factors affecting production E Ferreira T Yamada and F-

46 Cheng Cbien-pan 19(4 Upland rice in Taiwan International Rice Malmolta eds Institutoda Potassae Foslato Institnto Internacional Cnmmission Wiorking Party oit Rice Production ard Protection I()th da Potassa Piracicaha Sao Paulo lraid Meeting 3-10 March 1964 Manila Philippines Food and Agricul- 67 Ferreira E arid F dl Silva Martins 1979 Occurrence of ture Organiaion Rome Italy 5 p Sogatohs orvziola (Muir) inupland rice in Goias Ilrail lit Rice

47 Chcrrett 1 M and 1) 1Peregrine 1976 A review of the status of Rcs Ncwssl 4(4)18 leaf-cutting ants and their control Ann Appl Biol 84124-128 68 Ferreira E and 1F da Silva Martiris 1984 Insect pestof rice ill

48 Chipeta F M 1974 Pests of rice in Malawi Pages 90-95 in Brazil and their control Jill Portugucsej FMIlRAtPA-CNPAF Proceedings Sith eastern Africa cereals research conference 10-15 Goiania Brmiil 67 p March 1974 DRB Mialda ed Ministry of Agriculture lilongwe 69 Ferreira FI F ca Silha Miartins S S Nero and 1 It P de Malawi Carvallio 1982 M tltiplc regression analysis todeterimine the effect of

49 Chu Yau-i 1979 The d atiaging period of the rice seedling fly insect populatiois ott upland rice yield [inIPortuguese Pesqui (Atlterigotiavxigu Stcin) (NItiscide Diptera) oil cori) at Fast Java Agropecu Bras 17671-675 Indonesia Plant Prot Bill 21403413 70 Ferreira F 1F da Silva Martins and F 1PZimmermann 1979

50 Conklin H C 1957 IHaninoo agriculture a report on aln integral Rice varietal resistance to tle lesser corn stalk borer [in Port uguese] systent of shi fti rig cult ivationi in the Pltilippincs Food and Agriculture Pesqui Agropccu Biras 14317-321 Organization Rome Italy 209 p 71 Fleming W F 1968 Biological control of lie hapanese beetle U S

51 Cramer H II 1967 Plant proection and world crop production Dep Agric RLs Sers [ech Bull 1383 78 p Pilaniensclhutz-Nachr 201-524 72 Food and Agriculture Organiation of the United Nations 1982

52 l)ani R C and Majunidar 1978 Prelininary observations on tie Country report surmimary Pages 27-37 in Report on the first incidence of root aphids on differeit rice cultnrcs Sci Cult 4488-89 FAOIUN)IPhail and regional training coiuirse in iniproved ctltural

53 Davidson R I 1979 lffects of root feeding on foliage yield Pages practices for upland rice 5-23 Oct 1981 Northern Region Agrculturd117-120 in T K Ci osb and R B Pottinger eds Proceedings of the Developririt Center Chiang Mai Food and Agricuilture Organiiashy2nd Australian conlerence on grassland invertebrate ecology Govt tion Rome Italy 37 n Printer Wellington New Zealand 73 Franco do Ariaral S 195 1 Residual action of insecticides in turfy

54 l)ean G 1W 1976 Rice insect pests inlaos Int Rice Res Newsl soil--the results of a field experiment [in Portuguese] Biol6gico (Sao 1(2)15 Paulo) 17209-214 (Rev Appl Entomol A41423)

55 I)ean G J W 1978 Insect pests of rice in Laos PANS 24(3)280-289 74 Freeman ) 1970 Report onithe Iban london School of -coroiics 56 )ingle H 1984 Behavior genes and lifc histories conplex adapta- Monographs on Social Anthropology No 41 Athlane Press

tions to uncertain environments Pages 169-194 in A new ecology London UK 317 p novel approaches to interactive systems P W Price C N Slobod- 75 Fuchs T W1 A Harding and 1W Smithlr 1979 Induction and chikoff and W S Gaud cdsl ohn Wiley New York USA termination oldiapause in the sugarcane borer Ann Flitonlol Soc

57 Dobelmann-JP and M Falais 1968 Upland rice cultivation ill tie Am 72271-274 North-western part of Madagascar in 1966-1967 [in French English 76 (hbriel B1) 1975 A review tf the major insect pests of some tpland summary] Trop Agron 2313-21 crops in tlc Philippines Pages 86-111 in Ad Htoc panel of experts on

58 Doniiciano N L 1979 Chemical control of tie lesser corn stalk pest disease arnd weed problems in some rainfed crops Bangkok (lolsonno~paloIgnowslhts) (Zeller 1848) (I-epid(ptera Ihycitid ae) Thailand Food aid Agriculture Organization Roire Italy

on rice in Ponta Gressa Parara 1977-1978 [in Portuguese] Pages 77 Garg D K and N K Shal 1983 White grub outbreak on rainfed 291-303 in Anais 1 Reuniao de tecnicos cmiirizicultura do estado de dryland rice in Uttar Pradeslh Int Rice Res Ncwsl 81(4)16-17 Sao Paulo-Campinas Cati Brazil 78 Garg I) K and IP Tandon 1982 Major insect pests of rice on hilly

59 Dunsmore 1 R 1970 Investigations on tlie varieties pests and tracts of Uttar Pradesh India lilt Rice Res Newsl 7(11)11-12diseases of upland rice (hill padi) in Sarawak Malaysia Int Rice 79 Garrity 1) P 1983 Upland rice research an international bibliograpy Comm Newsl 1929-35 1975-1982 Iternational Rice Research hnstitnte PO Box 933

60 Elias 1R1965 The new decline of rice seedlings is not caused by Manila Philippines 121 p parasites [in Portuguese] Biol6gico (Sao Paulo) 313-6 80 Ghauri M S K 1982 A new subspecies of Iinorh)rone bull

61 Empresa Brasiliera de Pesquisa Agropecuaria 1984 Upland rice itt cornlus Slater (Hemiptera Iygaeidae) front Papua New Guinea on Brazil Pages 121-134 it Av overview of upland rice research rice and carpet grass Ihll lEntoniol Res 72133-137 Proceedings ofthe 1982 Blouak Ivory Coast Upland rice workshop 81 Gonzales F 0 PArregoces R I ihernandef and OT Parada International Rice Research Institute PO Box 933 Manila 1983 Insect and imite pests of rice arid their control in Latin America Philippines [in Spanish] la Federacion Nacional de Arrocerts (Fedearroz)

Bogota Colombia 60 n

38 IRPS No 123 January 1987

82 Grist D H and R J A W Lever 1969 Pests of rice Longmans London UK520 p

83 Hale P R 1979 Two subterranean pests of upland rice in Papua New Guirnca IntRice Res Newsl 4(2)16-17

84 Hale P R and J T Hale 1975 Insect pests of rice in the New Guinea Islands Rice Entomol Newsl 34

85 Harris W V 1969 Termites as pests of crops and trees Common-wealth Institute of Entomology London UK 41 p

86 Hattori 1 1968 A method to distinguish between grass leaf roller (Ctaplha rocis medinalisGuenee) and the rice leaf roller (Susurtia exigua Butler) [in Japanese] Plant Prot (Japan) 23167-170

87 Henry I E and 1A Onsager 1984 Experimental control of the mormon cricket Iahrosshnph by NJsena locustae(Aficrospora Microsporidae) a protozoan parasite of grasshoppers (Ort Acri-didae) Entomophaga 27197-201

88 Hidaka 1 P Vungsilabulr and SKadkao 1974 Studies on ecology and control of the rice gall midge in Thailand Tech Bull 6 TropicalAgricultural Research Center Tokyo Japan 113 p

89 Hincklcy A 1) 1963 Ecology and control of rice planthoppers in Fiji Bull Entomrol Res 53467 481

90 Hirao J 1982 Ecology and chtInical control of the rice leaffolder Jpn Pestic Inf 4114-17

91 Ho DT 1984 Sten borer incifience in rice ecosystems in KenyaEast Africa Pages 307-320 in A overview of upland rice research Proceedings of the 1982 Bouac Ivory Coast upland rice workshop International Rice Research Institute PO Box 933 Manila Philippines

92 Ho )T J 1Njokah and J GKibuka 1983 Studies on rice sten borers in Kenya with emphasis on ifaliarpiaipiraellRag Insect Sci Appl 465-73

93 Ichihara I T Ito A NIatsuo and M Sawada 1971 Chemical control of the rice striped stem borer in upland rice grown in plastic mulch [iii Japanese] Proc Kanto-Tosan Plant Prot Soc 1882

94 IclhiharaIand A Matsuo 1969 Damage caused by rice striped stett borer to upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1686-87

95 Ichhara I atid A MIatsun11970 Conside ration oft lie extraordinary prevalence of rice striped stem borer in upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1778-79

96 Institut de Recherches Agrolnliqes Iropicale et des Cultures Vivrieres 1978 Insect resistance Pages 12-13 in Upland rice 1978 atnual report Bouak Ivory Coast 27 p

97 International Institute for Tropical Agriculture 1983 Aphids itt rice roots I ITA Res Briefs 4(1-3)4-5

98 International Institite fi Tropical Agriculture 1983 Hopperburn in rice caused by brown plathopper IITA Res Briefs 4(l-3)6-7

99 International Rice Research Institute 1975 Annual report for 1974 PO Box 933 Manila Philippines p 216

100 International Rice Restarch Institute 1976 Annual repiort for 1975 PO Box 933 Manila Philippines p 228

101 International Rice Research Institute 1979 Cropping systems program Component technology development and evaluation Page 404 i Annual report for 1978 11O Box 933 Manila Philippines

102 International Rice Research Institute Department of Entomology 1981) Insecticid evaluation 1979 Tables 73 74 PO Box 933Manila Philipes

103 International Rice Research Institute Department of Entonmology 1981 Insecticide evalomtion 1980 Table 81 POBox 933 Manila Philippines

104 International Rice Research Institute 1981 Report of a meeting of Asian rice-based cropping systems entomologists Eleventh cropping systems working group meeting 18-22 May 1981 PO Box 933 Manila Philippines p 132 133 144

105 International Rice Research Institute 1984 Cropping systems program Pages 368-373 in Annual report for 1983 PO Box 933 Manila Philippines

106 International Rice Research Institutc 1984 Upland rice in Asia Pages 45-68 in An overview of upland rice research Proceedings of the 1982 Bouakamp Ivory Coast upland rice workiiop loternational Rice Research Institute 11O Box 933 Manila Philippines

107 Irshad M 1977 Grasshoppers in rice in Pakistan lit Rice Res Newsl 2(3)8-9

108 Ito R and TOno 1964 Upland lice its culture and varieties in Japan lIternational Rice Cormmission loth Meeting of the Working Party on Rice Production and Protection 3-10 March 1964 Manila Philippines Food and Agriculture Organization Rome Italy 12 p

109 Jeffrey M I and V A Dyck 1983 Effect of moonlight on lightshytrap catches of brown planthopper lit Rice Res Newsl 8(6) 13

110 Jennings P R and T Alicia Pineda 1983 Management ofthe rice plnthoppers Sogatodsorivi(coIl in Latin America Presented at the meeting on stable plant resistance FAO and lIndocsian I)inectorate of Food Crops Protection Bali lndonesia 29 Novcniber-2 l)ccemshyber 9 p

IllJimenez Gomez J A 1979 Rice insect pests i lla ios ()riCntales lil Spanish] Pages 99-103 in Rice ciltivation Instituto (olotibiatto Agropecuario Compndio No 29

112 Jongeleen F JJ Soesanto and A Mockidjo 1979 Studies oti pathogens of two species of Scarabacidac froinIndonesia Z Angew Enlomnol 87412-417

113 Jouvenaz 1) PC S lofgren and W A Baiks 1981 Biological control of imported fire antsa review of cirrent knowledge Bull Entomol Soc Am 27203-208

114 Kainoh Y K Shiniiu S MarUani Y Taiaki 1980 lost finding behavior of tie rice bu lplotori+JchinvittjI)allas (liciniptera Corcidac) with special reference to diet pattern ol aggregation and feeding on the rice plant Appl Entomol Zool 15225-233

115 Kakkar K 0 P Sharma and P I Sharma 1977 Note on Ihteronyrchus liodresRedt (Coleopicra I)ynastidae) as asporadic pest of paddy and maie inHimachal Pradesh Corr Agric 156-57

116 Kalode M131 79 Major insect pest problems and their control in upland rice (Unpublishcd mineogriph CWI oJNaaiat lni Chandaratat (198 1)original not seen)

117 Kalshoven L i 98 I The pests of crops in I ldoniesia (translated by P A vatn der Laan) 1 T Ichtiar B3aru-vati Iloeve lakarta Indonesia 701 p

118 Kamal N Q 1981 Suppression of whitebacked planto ner Sogatella fircaifra (Horvath) aLId rice lealfolder (nhal -cis licdb lis (Guenee) populations by natural cnclmie hlI) tiesis Grcgorio Araneta University Foundation Victoncta Park Metto Manila Philippines 179 Ip

119 Karim A N M Reaul 1975 A species ofliealy bug Trioni i70s ceres new oii rice in the Philippines Rice Entoniol Newsl 247

120 Katanyukul Wand C Chandaratat 1981 Insect pestsof uplaid rice and their control Report firstFAOUNlFi Tliailand training course on improved cultural practices for tplad rice Chiang Mai Thailand 5-23 Oct 1981 Food and Agriculture Organiation Rome Italy 22 p

121 Kern F 1955 lhletwioa(I)uninemts) bhhit Biturm(Coleoptera) a new pest of sugarcane inVenezuela [inSpanish with Etnglish sunmary] Agron Trop (Maracay) 527-29 (Rev Appl Eitoniol A4572)

122 Khan K M and S Ghai 1974 White grubs ald their control in India Pesticides (1Bombay) 819-25

123 Kiritani K 1975 Integrated pest management of rice pests Pages 66-70 itt Proceedings platning workshop ott cooperative hIeld research in pest managementP SMotooka ed East-West Center Honololu Hawaii USA

124 Kisimoto R 1967 Ecology of insect vectors forecasting and chemical control Pages 243-255 in The virus diseases of tle rice plant Proceedings of a symposium atthe International Rice Research Institute April 1967 The Johns Hopkins Press altimore Maryshyland USA

125 Kisimoto R and V A Dyck 1976 Climate and rice insects Pages 367-391 in Climate and rice International Rice Research Institute PO Box 933 Manila Philippines

126 Kureha S S Kobayashi SAsari and T Ishihara 1977 Life history and control of the mole cricket Grrllotalpa africana Palisot de Beauvois [in Japanese English translation 7520 1RRI library] Proc Kanto-Tosan Plant Prot Soc 21192-197

127 Kushwaha K S 1981 The black beetle as a pest of rice in Haryana India Int Rice Res Newsl 6(4)14

128 Lawani S M 1982 A review of tile effects of various agronomic practices on cereal stem borers Trop Pest Manage 28266-276

129 Leuck D B 1966 Biology of the lesser cornstalk borer in South Georgia J Econ Entomol 59 797-801

130 lever R J A W 1955 Insect pests of the rice plant in Malaya Int Rice Comm Newsl 1612-21

131 Li Ching-Sing and Shin-Foon Chiu 1951 The study ofthe rice gall midge Iachydiplosis orvza Wood Mason [in Chinese with English summary] 1Taiwan Agric Res 21-13

132 Iitsinger J A 1984 Assesament of the need-based application for rice Proceedings TechnologyTransfer Workshop 15-16 March 1984 International Rice Research Institute 1PO Box 933 Manila Philippines 6 p

133 Litsinger J A 1984 Environmental diversity and insect pest abundance with reference to the Pacific Pages 285-307 in Proceedings sub-regional training course on methods of controlling diseases insects and other pests of plants in the South Pacific 4-20 Oct 1982 Vaini Kingdom of Tonga Nukualofa

134 Litsinger J A RF Apostol and M BObusan 1983 White grub Leucopholis irrorata (Coleoptera Scarabaeidae) pest status popula-tion dynamics and chemical conllol in arice-maize cropping pattern in the Philippines J Econ Entoinol 761133-1138

135 Litsinger J A C de la Cruz F Raymundo A T Barrion M D Lumaban J P Bandong M S Venugopal F Paragna and E Balete 1982 Insect pests and insecticide reponse of dry-seeded bunded rice Pages 179-193 in Report of workshop on cropping systems research in Asia International Rice Research Institute Los Bafos Philippine

136 Litsinger J A M 1) Luinaban J P Bandong A T Barrion W Katanyukul and N Panda 1978 Insect pest management in rainfcd lowland rice agroecosystms of tropical Asia International Rice Research Conference 17-21 April 1978 International Rice Research Institute Los Bafios Philippines

137 Litsinger J A and K Moody 1976 Integrated pest management in multiple cropping systems Pages 293-317 in R I Papendick P A Sanchez and G B Triplett eds Multiple cropping American Society of Agronomy Madison Wisconsin USA

138 Loevinsohn M E 1984 The ecology and control of rice pests in relation to the intensity and synchrony of cultivation Ph D thesis University of London 36) p

139 Loevinsohn M E J A Litsinger N Panda M A Austria R F Apostol C Andrion and J Bandong 1982 A basis for predicting rice insect pest abundance in cropping systems Paper presented at the 1982 International rice research conference on rice and cropping systems 19-23 April 1982 International Rice Research Institute Los Bafios Iaguna Philippines 25 p

140 Lopez A W 1930 Repot of the Entomology Department Pages 145-172 in Annual report of the Philippine Sugar Association 1929-30 Manila

141 Lopez AW 1931 Entomology Department Pages 227-266 in Annual report of the director or research 1930-31 Philippine Sugar Association Manila

142 Lopez A W 1932 Entomology Deparment Pages 252-279 in Annual report of the Philippine Sugar Association 1931-32 Manila

143 Lungibill P and G G Ainslee 1917 The lessercornstalk borer US Dep Agric Agric Res Serv Tech Bull 539 27 p

IRPS No 123 January 1987 39

144 McGuire J U Jr and BS Crandall 1967 Survey of insect pests and plant diseases ofselected food crops of Mexico Central America and Panama U SDepartment of Agriculture International Agriculshytural Development Service Agricultuial Researrh Service October 1967 11 p

145 McNeill Sand T R ESouthwood 1978 The role of nitrogen in the development of insectplant relations Pages 77-9 in lliochemical aspects of plant and animal coevolution J B Ilarbornu ed Academic Press London UK

146 Maebara H 1955 Spatial distribution of lastinstiarlanaeof purplish stem borer (Sesamia iqferens Walker) hibernating in an upland rice field Bull Kagoshima Univ Fac Agric 438-41

147 Mallick E H and N G Hajra 1977 Response of rice plant to defoliation treatments at different developmental stages Indian Agric 2151-56

148 Mansingh A 1971 Physiological classification of dormancies in insects Canl Entomol 103983-10)9

149 Martell C G 1974 Catalogue of plant feeding insects of Mexico [in Spanish] Fitofilo 27(69)1-176

150 Martinez 0 J 1982 Principal rice pests in Ilanos orientalis [in Spanish] P~ages 80-94 in Curso deact nalization de conocimientos en arroz Villaniencis Instituto (olonmbiano Agropecuario

151 Martins J F da Silva F Ferrcira A S Prabhu F 1 P Zimmermann 1980 Use of pesticides for the control of the main subterranean upland rice pests esqui Agropecu Bras 1553-u

152 Martins I F da Silva F Fe eira rod BdamSilveira Pinheiro I Q2 Damage simulation caused by defoliators of upland rice [in Portuguese with English summary] Pesqui Agropecu Bras 171113-1119

153 Martins 1F da Silva N(iyuen van Tan and 1Ida Silveira Pinheiro 1981 Stem borer resistance in upland rice associated with plant morphological characters [in Portuuce Iesqui Agropecu Bras 16187-192

154 Masaki S 1980 Sumnmer diapause Ann Rev Entomiol 251-25 155 Masuzawa T 11 Suwa and F Nakasoji 1983 l)ifkleences of

oviposition preferencc and survival rate of two skipper butterflies Parnara guttata and PIchlpidai iathiais (Iepidoptera HIesperiidac) on rice plant and cogon grass New Entomol 3247-56

156 Maurya D M and C P Vaish 1984 Upland rice in India Pages 69-92 in An overview oftpland rice research Proceedings ofthe 1982 Bouakamp Ivory Coast upland rice workshop International Rice Research Institute IO Box 933 Mnila Philippines

157 Menon M G R R N Katiyai S Kuar and S S Misra 1970 Termites attacking the paddy crop Sci Cult 36294-295

158 Miyahara Y and 1 Nishikubo 1967 Behavior of young nymphs of the red rice root aphid Iholahsipiun ri liabhdontinalisSasaki in relation to movement from the leaf to the roots of upland rice [in Japanese] Pioc Assoc Plant Prot Kyushu 1363-66

159 Mo ljoa Tjien 1952 White grub of upland rice (Holotrichiahelleri Brsk) [in Indonesian with English summary] Contrib Agric Res Stn Bogor No 13134 p

160 Mochida 0 and T Suryana 1979 The occurrence of the brown planthopper and ragged stunt on lowland and upland rice ill Central Java and logyakarta in tile late of January 1979 Entomnolgy Survey Report No 15 Central Research Institute of Agriculture (CRIA) Sukamandi Indonesia I February 1979 9 p

161Monet C 1979 Varietal resistance to upland rice pests [inI French] Pages 35-39 in Annual report 1977 Institut des Savanes Department des Cultures Vivrieres Bouak6 Ivory Coast

162 Monte 0 1942 A caterpillar ofrice-fields[in Portuguese] Biol6gico (Sao Paulo) 8161-163 (Rev Appl Entomol A31414)

163 Munroe I) D 1979 Pest management strategy for hill rice in four resettlement areas in Sarawak Malaysia Pages 89-102 i1 Guidelines for integrated control of rice insect pests Food and Agriculture Organization Plant Production and Protection Paper 14 Rome Italy 115 p

40 IRPS No 123 January 1987

164 Nakasuji F M Ishii and Jen-Zon Ito 1984 Rice skippers in Taiwan and their life histories ZZrC (Taipei) Tech Bull 82 10 p

165 Navas I) 1976 Fall armyworum in rice Pages 99-106 il Ecological animal control by habitat management Proc Tall Timbers Con-ference 28 Feb-I Mar 1974 Fall Timbers Res Stn Misc Publ 6 Tallahassee Horida USA

166 Nickel 1 1 1961 Biological control of rice stem borer A feasibility study Tcch Bull 2 International Rice Rcarch Institute PO Box 933 Manila Philippines IIl p

167 Nigan P M and R A Vernia 1985 Insect pests of uplaild rice in Uttar Pradesh Int Rice Res Newsl 10(6)22

168 Nilakhe S S 1985 -cological observations on spittle bugs with emphasis on their occirence in rice Pcsqtui Agropecu Bras 20407-414

169 Nixon G EJ 195 1 lie association of ats with aphids and coccids Commonwealth Iusti lcte of Entonology Iondon UK 36 p

170 Nurdin F an Ishak Manti 1985 Study oit seedling fly of upland rice Interlational Upland Rice Moitoring Tour Report 26 lebruary-2 MarLI Sukaranti West Sumatra

171 Oka 1 N 1979 Ciltural conttol of the brown planthopper Pages 357-369 in Brown planthopper threat to rice production in Asia International Rice Research Institutc 1p0 Box 933 Manila Philippines

172 Ohni M and A Vilnaii 1975 Rice insect pests of Italy biology of Sipha grcria (Kaltenbach) and its control [in Italian] Riso (Milan) 2459-71

173 Otanes F Q atnd 1 I Sison 194 1 Pests of rice Philipp J Agric 12211-259

174 Painter R I 1951 Insect icsistance in crop plants University Press of Kansits Lawrence Kansas USA 520 p

175 Pantua P C and 1A litsinger 1981 Arthropod abundance in continuous vs biannual rice cropping systeis Int Rice Res Ncwsl 6(2)20-21

176 Pathak M ) and V A l)yck 1975 Sttudics ott insect pests of upland rice Pages 186-191 in Major research in upland rice International Rice Research Institute PO Box 933 Ilanila Philippines

177 Patil B R B M IHasabe and 1) S Ajri 198 I Chemical control of white gritb lIditriihinhorraia Fabr intfesting groundntut paddy atnd sorghum Indian 1Entomol 43232-236

178 Pawar A 1) 1974 (ollection and identification of insect pests of rice rlerminal repirt lInternational Rice Research Institute PO Box 933 Manila Philippines 72 p (tnimco)

179 IerrajU A aid A S Reedy 1965 Observations ott diapausc itt tie steti borer ohrice Rice News Teller (lai-Mar)9-13

180 Pont A C 1972 A review of the Oriental species of theri ona Rondani (I)iptera Muscidac) of econtomic importance Pages 27-102 in Control of sorglium shoot fly M G lomtani and W R Young eds Oxford atd IH1 Ptthl Co New I)clhi India 324 p

181 Prasad S C 1PChand aid M 1PSingh 1983 Incidence of rice gall midge in the upland plateau region of Hihar Oryza 20235-236

182 Rai P S 198 0 Nisaga simpex damage to rice in the hill tracts of South India Int Rice kles News[ 5(I)12

183 Rathorc Y S 1969 Gonocplhahli dcprssunt Fabr as itpredator of (tilo zonwIlhis (Swinhoc) and other moths in field cage at Patnagar Uttar Pradesh Indian 1Entotnol 3127

184 Reed W 1983 Crop losses causcd by insect pists in the developing world Proceedings 10th Internatial Congress of Plant Protection lrighton UK 174-80

185 Reedy M V 1981 Ateactogasterinhens(Walker)a new root-grub pest of tipland paddy and inaize Curr Sci 501076

186 Reveno M A 1968 Catalogue of the insect pests of economic crops in Colombia [in Spaiish] Asociacion Latino Americana de Entomo-logia (AIAE) 129-33

187 Reyes G M 1957 Rice dwarf disease in the Philippines Food and Agriculture Organiiation Plant Prot Bull 617-19

188 Rice S E A A Grigarick and M 0 Way 1982 Effect of leaf and panicle feeding byarmyworm (Lepidoptera Noctuidae) larvae on rice grain yield Environ Entomol 7593-595

189 Risch S J 1) Andow and M A Altieri 1983 Agioccosystem diversity arnd pest control data tentative conclusions and nizw research dirctions Environ Entomol 12625-629

190 Rivera C T 1956 Three organic insecticides in thecontrol of insects affecting upland rice Philipp Agric 39465-472

191 Roffrey 1 1972 Locusta outbreaks in the Philippines Acrida 1177-188

192 Rojas R 1) 1941 The effect of pruning at varying ages upon the yield of Ramairio Phipp Agric 29851-860

193 Roonwal M i 1979 Ternite life and termite control in tropical South Asia Scientilic Pnbl odhpr India 139 p

194 Rosetto C1 F da Silva Martins N C Schmidt and L EAzzini 1978 Damage caused by pasturespittle bugs 4Dvoi)lavopictaandDJ schach) on rice lBragantia Nota 6 (Matcli)25-27

195 Rosztto C1- S S Neto I- Amantc I) Iink 1Garcia-Vieira D M de SoLza N V Bansa t and A M de Olivcira 1971 Rice pests of Brazil [in Portuguese] Pages 149-238 inrFA()-International Rice Commission 2nd meeting of the Rice Committee for the Americas 6-11 l)ecember 1971 Pelotas Rio Grande do Sul Brazil

196 Rothschild G H L 1969 Obseratiois on the armyworm Spodoptera matritia acronrctoiiles Gn (Lep Noctuidae) Bull Entomol Res 59143-160

197 Rothschild G 1- L 1974 Parasites of the armworm Spolopiera mauritiaacroncitoihes in Malaysia Fntornopliaga 19293-299

198 Sachan i N S K Gangwar and 1N Katujar 1980 Field crickets damaging cages in North Eastern IHill Region Indian J Entomol 42526-527

199 Sands 1) P A 1977 The biology and ecology of Lepocorisa (Hemiptera Alydidac) in Papua New Guinea Res Bull 18 Departshyment of Primary Industry Port Moresby Papua New Guinea 104 p

200 Sarawak 1968 titomology division Pages 97-106 in Report of the research branch of the Department of Altriculture 1966 Kuching (Rev AIppl Eintomol A56603)

201 Satpathy JM M S Das and K Naik 1977 Effect of multiple and mixed cropping on the incidence of scine important pestsI Entomol Res 178-85

202 Sauer IHF G 1939 Notes Ott Elasmpnlpualtsligniosellths Zeller (Lep Pyr) a serious pest ofcereals in Sao Paolo [in Portuguese] Arq Inst Biol 10199-206 (Rev Appl Entotnol A28378)

203 Satinders 1L A 1 SKing and C L Vargas S 1983 Crop pests of Central Anerica with t bibliography [in Spanish] Centro Agroshynotnico Tropical de Investigacion y Ensenanza Turrialba Costa Rica (CATIE) Tec Bol 9 90 p

204 Sauphanor B 1985 Some factors of upland rice tolerttce to stem borers in West Africa Inseu Sci Appl 6429434

205 Sen P atnd S Chakravorty 1976 Stentflics in Haritighata Nadia West Bengal India [it Rice Res Newsl 1(12) 15

206 Senapai B and 1M Satpathy 1983 The seedling fly Atherigona orrzae Malloch is a pest of rice in Orissa (India) J Entomol Res 777-79

207 Shhjahan M S Alatn M Ah nad and BN Islam 1983 A new scarabaeid beetle pest of dryland rice in Bangladesh Int Rice Res Ncwsl 8(I)16-17

208 Sharma G L S C Modgol and R C Gautam 1979 Effect of depth of submergence ott incidence of bacterial blight and white grub infestation in transplanted rice Int Rice Res Newsl 4(6)12-13

209 Sharma S 1985 Scarabaeid beetle outbreak itt upland rice Int Rice Res Newsl 1(1(l)24

210 Shiraki T 1917 The paddy borer Schoenobhits incertulas Walker [achaku Agric Exp Stn Formosa 256 p

211 Silva A de Brito and 11P Magalhaes 1981 Insect pests of rice [in Portuguese] Circular Tecnica No 22 EMBRAPA Centro de Pesquisa t ecuaria de Tropico Unido (CPAJU) Belem Brazil 14 p

212 Singh S A 1979 Emergence of rice leaf butterfly as a rice pest in Manipur India int Rice Res Newsl 4(4)16-17

213 SinghS A 1983 Changing trends of insect pestsof rice in Manipur Int Rice Res Newsl 8(f)12

214 Siwi S S and P H van Doesburg 1984 Lenovorisa Latreille in Indonesia (Heteroptera Coreidae Alydinae) Zool Meded (Leiden) 58117-129

215 Soejitno J 1977 Reaction of some upland rice varieties toseedling fly eltherikonaexigua Stein [in Indonesian] Siri Hana I Penyakit 630-35 Lembga I osat Fenclilian Pertanian Bogor Indonesia

216 Socnardi 1967 Insect pests of rice in Indonesia Pages675-683 in The major inse pests of thu rice plant Proceedings of a symposium at the Internatital Rice Rec Wi itt S pternber 1964 The Johns Hopkins Press Baltimore Maryland USA

217 Soto P E A T Petcz and I W Buddenhagen 1976 Survey of insect pests and diseases of rice in different ecological zones in Nigeria Rice Entomol Ncwsl 435-46

218 Soto 11E and Z Ziddiqi 1976 Screening for resistance to African rice insect pests Varietal improsetnent sCmi n r Bouak Ivory Coast 13-17 Sept 1976 West Africa Rice l)evelopment Association Monrovia Iiberia

219 Squire F A 1972 Entomological problems in Bolivia PANS (PestArtic News Sumr) 18249-268

220 Stapely 1 II 1978 The rice leaf roller in the Solomon Islands Int Rice Res Newsl 3(l)13

221 Suenaga H 1937 On aphids attacking the roots of rice Oyo-I)oubts Zasshi 9150-152 [in Japanese] (Rev Appl Entomol A25745)

222 Suenaga H 1952 On the seasonal abundance of three subterranean aphids on upland rice Bull Kyushu Agric Exp Sti 1249-262

223 Sutynso Yas and Dandi Soekaria 1964 Beberapa hana yang ttenperang tana man padi tanah kering [in Indonesian] Rapat kerja Padi Tanah Kering 6 sd 8 Juily 1964 Kaliurang 9 p

224 Tallamy I) W and R F Denno 1983 Alternative life history patterns in risky environments in example from lace bugs Pages 129-147 in Insect life history patterns R F l)cnio and If Dingle eds Spring-Verlag 225 p

225 Tamura 1 1939 Observations on some insect pests of field crops [in Japanese] 1 Plant Prot 26414-420 (Rev Appl EntomolA27-556)

226 Tamura and Suzuki 1964 Influences of the early-transplanting practice in rice culture upon development of the rice stem borer Chilo siql)ressalisWalker Bull Ilokuriku Natil Agric Exp Stn 761-94

227 Tan J P 1924 The rice root aphid Drowhi ihirsoia Philipp Agric 13277-288

228 Tanaka 1 1957 Taxonomy and distribution of some subterranean aphids injurious to upland rice in Japan with description of a new species [in Japanese English summary] Botyu-Kagaku 22168-176

229 Tanaka T 1961 The rice root aphids their ecology and control Spec [full College of Agriculture Utsunomiya University Japan F3 p

230 Tauber M IC A Tauber R Nechols and 1Obrycki 1983 Seasonal activity of parasitoids control by external internal and genetic factors Pages 87-108 itt I)iapause and life cycle strategies in insects V K Brown and I Ilodek eds )r W Junk The Hague Netherlands

231 Taylor W F 1972 Effects of artificial defoliation (simulating pest damage) on varieties of upland rice Exp Agric 879-83

232 feotia T P S and S Natud 1966 Biotoomics of the rice skipper Parnaraniathiis Fabricius (Lepidoptera Hesperiidac) Indian J Entomol 28181-186

233 Teran F 0 1971 Rice pests of Santa Cruz [in Spanish Ministerio de Agricultura Sita Cruz Bolivia Bull 45 8 p

234 Than Itun 1976 Population dynamics of the yellow rice borer Tryporza itcertulas (Walker) and its damage to the rice plant MS thesis University of the Philippines at Ios Bafios Laguna Philippines 92 p

235 Uichanco L B 1911 Pests Pages 55-74 it Rice Philipp Agric 11-200

IRPS No 123 January 1987 41

236 van der Laan P A 195 1 Life history of the rice seedling fly Contrib Gen Agric Res Stn Boor Indonesia 118 15 p

237 van der Laan 11A 1959 Correlation between rainfall in the dry season and the occurrence of the white rice borer (Scirpolhaga innotala WIk) in lava Entomol Fxp Appl 212-20

238 Van Driesche R G 1983 Meaning of percent parasitismin studies of insect parasitism Fnviron Entomol 121 I1- 122

239 Venugopal M S and A litsingcr 1984 Fffect ofcarbofuran on rice growth Prot Ecol 7313-317

240 Vietnam Directorate of Rural Affairs 1962 Annual work progress report on crop improvement program of rice sugarcane vegetable and field crops (lor the period from July 1961 to June 1962) (11) 299 p (Rev Appl Entomol A51526-527)

241 Wan M T K 1972 Observations on rice leaf- and planthoppers in Sarawak (Malaysian Borneo) Malay Agric 148308-335

242 West Africa Rice I)esclopnient Association 1978 Annual research report Vol 2 Statistics Monrovia Liberia

243 West Africa Rice Development Association 1979 Annual research report p47 Monrovia Liberia

244 Williams ) F and CS lofgren 1981 Fli I illy l-468 anew bait toxicant for control of the red imported lire at Fla Entomol 64472477

245 Williams I) 1 1970 The mealybugs OFlomoptera Coccoidea Iscudococcidac) of sugar-cane rice and sorghum Bull Entomol Res 0109-188

246 Williams D) 1 1982 Pulinaria icervi (Signoret) and its allies ott sugar cane and other grasses Btll Entoinol Res 72111-117

247 Wirjosuiardjo S 1976 Study ofchenmical cont rot of insect pests ott upland rice Imiu Pertanian (Jogjakarta) 1323-329

248 Wirjosuhardjo S and N Wirarta 1976 Chemical control of the white grub tIolorithiatlltriin upland rice in M uneng Irobolinggo Indonesia) [in Indonesian] Imut Pertanian (Jogjakarta) 1330-332

249 Yano K T Miyake and V F Eastop 1983 The biology and economic importance of rice aphids (Henuiptera Aphid idae) a review Bull Entomol Res 73539-566

250 Yelp D and P Rhodes 19(4 lhe success of drilled dryland rice iii the Fiji Islands International Rice Commission Working Party on Rice Production and Protection Manila Philippines 3-10 March1964 Food and Agriculture Organization Rome Italy 13 p

251 Young W R 1981 Fifty-five years of research on the sorghum shootfly Insect Sci Appi 23-9

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No 104 Faridpur acomputer-assisted instruction nmodel forrainfed lowland random coefficient model No 116 Consumer demand for rice grain quality in Thailand Indonesia andrice

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No 106 Rice grassy stunt virus 2 a new strain of rice grassy stunt in tire literature

Philippines No 118 IRRI-Korca collaborative project for the development of cold-No 107 Physical losses and quality deterioration in rice plstprodnCtion tolerant lines through anthei culture

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publishers No 120 Changes isinall-farm iLtireliing techii)hgy in Thailand and [he No 109 Classification of Philippine rainfall patterns Philippines No 110 Contributions of modern rice varieties to rutrition in Asia No 121 Landforms and modern rice varieties No Ill Changes in rice breeding in 10 Asian countries 1965-84 No 122 Yield stability and modern rice technology No 112 Design parameters affecting the performance of the IRRI-designed

axial-flow pump

Page 25: UPLANDRICE. INSECTl--1i M T -E!R.EOLOU t RTANCE ,AND …delphacid.s3.amazonaws.com › 6958.pdf · The significance of abiotic factors in upland rice insect ecology will be apparent

IRPS No 123 January 1987 25

Cultural control dry season plowing will have little effect on them Plowing Crop husbandry to reduce upland insect pest populations when the soil is wet from the first monsoon rains will expose can be classified into practices effective in a single field and soil insects that tunnel close to the soil surface to predatory those effective only at the community level Cultural birds chickens dogs and even man (field crickets nole controls should come first in a pest management program crickets and white grubs a re delicacies in man tra ditional They have broad stable effects as pests have little pos- diets) sibility of overcoming them through selection of biotyps The greatest degree i1 inIscct control fron tillage comes (138) On the other hand some cultu ral controls work only from plowing soon after rice ha rvest when most soil insects with high labor or power inputs and connunity-wilt lie close to the soil surface anrd stern borers are in the rice methods require coordination of iany farniters stuble Whitegrubsare mature and tillage exposesthem to

Pantin time The luxury of being able to shift the time of predators Tirnelv control will prtec the crop following pla ntring upland rice is open onl hr la rniers in areas with a rice and lowr the populationIfor succeedinig years Plowingprolonged rainifall pa ttern lI copping intensity or underthe ice stulble helps it decompose killing sten borer rnechanied land preparatiot Gcnerally planlting as soon larvae inside as the rFiinv season begins will lower populatiois of 1110st [requent tillage destroys ait nests bit not termites insect pests Insect populationsarelo aftcra dlvseoll or Tillage and clean culture howes er rcliove food for winter fallow delayed plaiting lets them build tip on termites which then mtay attack a young rice crop Scarab alIternate hosts that gi vigorotsly with the first seasonial beetles also prefer to miposit in eceitly tilled fields Noshyratiris This first flush iof weds cros lItxuttriartlv with iceCult is dibbling faors the survival oftillace Clre Such rineralied N released during the dr season aid made stils i aels tnd insects thtt pttpate or lay eggs- in the soil

available to the plants with the rains It iliso igrtmitcally Zero tilla hio eerl conserves predators ofthe GIH (37) advanttgeous for the rice ciop to tap this trlLItlaI fertility nt it mcttoi lo lessen the impact o0 soil insects and grow vigorotsly to compete with wrus and to tolerate brotdcasting seed or planting in ftltrrows is prefc ired over pest infestation dihlirig in hills Crickets are large enotlgh to eat seedlings in

Larly plantngs terd to escape seedling maggolts Upland a hill atrd tillering normnally ca iriot fill in tlhe missing space rice in Bitangas is planted in Ma and ine itli the Iirst Oftcn dibbll rd is not well covered and iseasily found by rains Fairners are highly mtotiated to plant early tocscape foraging insets ioIdents and birds drlitght and typlotns ricar harct Arn Atrgtst planting Plan uit sii Incrcatsini sced density protects tile cropwould be evcely attacked by It riim orivzo At tti againrst seed and seelling pests especially ants particularly other Philippiie tipland sites Rea l Qteot antd (laelit if the crop will germinate and ermerge quickly M isainis Oriental seedlig mnaggots arnrige ric boe t se hr trftinll itI India rice intcrclopped witi cotton or farnes have a longer growing season and plnit 2 rin or pigeon pci had lower 6IIt and WBIlII poptrlations than mote after the ortset of the rainy season liaaiangas ftrmners rice alone (211) NfailC arid Urplald ir_ 2re coilltori in the plant ca rly becia use ofa shorter raiinfall period arnd the desi re Philippines he chatngC in pest status folii iniercropping is to lliarvet before the typhoor sea son peaks ltighly locatioll-specific (133) cmianarid the net elfc be either

Early plantings cornbind with early-iatiuring varictics nil bcnelficial ortdetriential (189) For example BIatangias provide higher crp tolerartce for white grubs Aestiatior farmers intercrip Itro in their ipland ricefields A liornshyterminates with the first rains but 5-6 moare ineeded for the nrii iotd-Ahis i (I iiiacius) which norlially feeds resting larvae to develop into beetles emerge and lay eggs on taro ill feed or miarbv rice plants Intercropping and for larvae to reach the danriigii g ird-instar stage probably would not afft0 seed pests A crluanion crop (134 In lataigas this normually iseiough tiite for harvest planted ith rice and haing gleater tolerarnce for root

It Brazil howecver delayed planting is a suggested dariage or being toxic nayact isi trapcrop to root-feeditig cultural control for the lesser cornstalk borer and insect pests Acial-fei i ig pests voultId be most affected by pests favored l drought (66 195) Cropping internsity in intrer ropping Shoot aphids leaf beetles sten borers and Brail is low and delayed plraing places the rice crop in the hopplers would be likely ca ntidates to note effcts On the niost stable rainfall period reducing likelihood old rought othlt halnd the corlipalior crop stch~ sitniaie Iiiv be more

In rilniy arca s rinmirsare roi atl to plant other crops benehited than rice from initecropping ( 137) before rice and althNgh it woultl be desirabh to avoid 11echi Weeuling luring the first month after clop infestations by planting rice cuatl lariners lack the labor cstablishent will force pcts such is anyvworris which and time to do so ol a low priority crp prefer grassy cetds ontor tle vorling rice Soil iampsecs will

77age Uplild rice Stils ae light textured and Call be mvnye to the rice crop Cleani citrltirc also lay lore territes plowed duiring the d rv seasor notl only to grin time for to attack I u cetincti inrice ceds i ay he le optinal planting with lie fitst Inilsoon riiains but also to desiccate solition 13) soil insects arid weeds such is ntsedge that foim rhizomes Irtlizr Itneatsing N Iciiliier oin hlwland rice has Htowever becta use most soil insects lie below die plow layer fAtwvored Irigher populains of pliithoppeiv Imlhoppers

26 IRPS No 123 January 1987

and leaffolde rs However because of the need to temper fertilizer use on upland rice o prevent blast fIrtilizer rates will be lower than is comonia r for lowlad rice causing relatixely less hopper and IaIfTfoide r population buildup However inupland rice areas without a history of blast such Is Batangas farmers apply high rates of N and conscqtrently the crop has high infcstation levels of WBPI]and RIF The fhrmrcrs leec insect buildup by splitting the fertili11r in0to two to three applicatiois bit do not applV fertilizer at the vegetati estage because this prodrces a tall crop which illeasily lodge during typhoons Greater crop fertility leads to greater insect survival (better nutrition) larger insects (to cat no re foliage) arid greater fec titd t (to lav nmre eggs) (128 145 171) Not all insect species respondequally hovr Increasing rates (if N l anrid K increased Dialraa s-acchralis and (iaeiurema flea beetle but reduced thrips (66) Zn rcdLuced deadheart density from the lesser cornIstalk borer

Fnldingi Although most Upland rice areas do nIot hawxcaccess to wa ter floodiig the fields is rRrrnriierdcd in areas of Brazil or root bug and white gib control ( 195)

Crop rotatioti Tillage before planting a crop after rc harvest will control stern boreis and other pests renain n the stubble arid will unearth soil pests Planting a non-gratriiaceous crop after rice is recommncrded for termite control (66) Crop rotation is less nttdcd in upland rice culture than in lowlands because atcrop-free period is assured even after harvest of crops following rice

t1ulchilt iryl plastic irli irig designed for weedcoitrol ii upland rice inJapan resulted in ntr highanri liy infestation of (ihilo sulytresvais (93 9-1 95) NIulched Upland rice grew more luxuriantlv and attracted more (Avipositing mothis than u lched rice Also the rice plant and stein borers imatoured 10rC quickly and a partialgeneratiori af vouing ista iNoverwintcred in the stubbleLotili--malitrt varieti Quickly maturing crops reduce the tmbCr of pCst generations that Calt build up Rapid crop establishment olan ca rl v-natu ring vaitvty if)pa 11icLa r will lessen white gib and stem borer damiage White grub larvae intle last iistar are highly destIruCtive so aii early-iiatri ng variety will escape serious root loss Stem borers build 1ip slowlyx in rice to ecormic abundant in a late-Maturing crop anrd caiuse whi tlieads Numibers of somespecies increase exponeitially with each succeeding genera-ti so a xarietyexhat atures I trowh earier thian another will hac less insect damage Early-naturing vaiicties however will not lessen seed pest daiage either ii the soil or oina standing crop

Strwhoi hAIJttam Inrscts tcd irig on thlie aerial portions of rice plats disperse from carly to late plantedfields AS iiiibCirs increase exponentially with eachinsect

gtscnrattm e eiglbrig f rmc~s1(uil upp wh rc generation pests build tIP where neighboring fihrTC e9r1sstagger their plantings II Batangas farmers plant uplandrice within I iro use 120-d s-arieties and plant at the start of the rainy season Ini Misarnis Oriental laricrs stagger their

plantings up to 3 mo apart and use a set of atieties that mature in 4-7 mo Insect pest danage particularly from stem borers is greater in Misarmis Oriental Also seedling maggots damage rice there because rainfall favors frequent planting of maic In Batangas rice escapes seedling maggots bCca usC of distinct drs season and synchron0us early planting with the onset of ains

Therefore combinations of cultural practices ea rlyplanting synchronous planing crop rotation and earlyshymaturing vaneties protect the rice crop against most insect pests

Ilmant resistance Beca tse upland rice has few specialized insect pests and many insect types attack the crop worldwide regional breeding and strong internatiornal cooperation are required

Sown-sced pests arc economically controlled by insectishycide seed treatment and should the relore receive lowxpriori ty inbreeding objectives

Root pests are normally cost ly to control with insecticides but finding resista t sonres mayvbe di fficult Tanaka (229) for exa -ple fi led to find varieties resistant to root aphids Selecting for large root bioniass perhaps should be the strategy for pests sLuch as Whlite grubs that renliove roots Fainter(174) howcvcr cites varietiesof crops tolerant ofor resistant t(i root pests Some sorghun varieties are high in cyanide so rice sorghun might incorporate broadshyspcctruri chemical resistance

Seedling vigor and drought tolerance are high prioritybreeding objectives so tupland rices should have large r root systernsand be able to tolerate ligherlevels of root loss Sonie root loss inolder plants is beneficial if it stimulates new root developnrut and therefore crihances intake of soil nutrients (53) The rice seedling inaggot rtiLotia spp may be

controlled thirough resistant aittics A hi rgc breeding effort on sorghniu has had fair success inideveloping arreties esrstait to Asoccata Rodani (251) A field trial comparing 10 upland ricc varieties showed differences aniong varieties va ring from II to 4(- damaged tilleis (215) Hovver resistarce to 1Iill-0r gtia species may rot cross ox-rti otier

Shiraki (210) reported that upland i-ice stens wcrc harder than lows-land rice stemsperhaps becaIusc (f a higher silicoricontent and thus were more resistant to yellow stern borer hc tortia Iow tillering (f uplaid iice xarneties prodtres a

higher percentage of infested tillers than would occor in higher tillering lowland iice under a siiilar egg density The corollary therefore is to select for high tilleriig upland rices

Stemii bor amcstheh main targes dobrecders inAfrica (571 204 )but nticl oifth e sc reen irig is dotie unde r lo wland co d t osrm s wih A h r~ o l pe i s n acorditions (11 96 218) As with lthrinrnwspecies mayhave to be dealt with independently however initial results show cross resistance between African and Asian species(92)

In Brazil screening Ior insect resistance focttses oil lou I pest problems Varieties tolerant of lasmolwalpus ligi-sellus and resistant to )iatraea saccharalis have been identified (70 153) Current work also emphasies 7iraca limativetntris and Thmialspis (=Iosis) flavouicta (66) These insect pests Ilt onlV citise IIigh yield losses tihey also ire difficult to control chemically ofr by other ilealls

ManV upland rice vaieties aic early maturing and are resistant to pests such as gall midge (I Xl ) Early mIturity is often a highly desirable trait to esuilpe pest buildup but it is not true resista nce bca usc ilthose va reties are plinted late they will be itsdamaged as ain early planted susc ible variety

P~erhaps tile cosiiiopolitall of all upland rice pestsimost are the sced pests (ripenling stagc) which pose a particular problem beta use scuds re ised IsIO for hitnit4ans Ayltoxic substatice introd tcld geticticllyilmay also be toxic to nal iniless it caln he placed old in thfe IdhuffSelectmi fo hairiness or lotng wns kill mIlct sthilrohleiiis dutilig threshing pa rtictfarl maitial threshinfl [hireshers ill complain of itcfimin atid itteascd (fust levels

Attempts itIRRI to fitid tcistatncc to the rice bug plorisa have ilotbeCnl prmitisilg lIIetIattonlids iorc

through the lenilna or palca hillhairs sedstVIor seeds wilh awns that deter bugs promote complaints from threshers Seed bugs liptocorisa and Stcnocoris enter the seed through the opening between tile lemna and palC but varieties with narrow openings are diifficult to mill

Breeding priorities should concentrate ott inotCI gruLtLIps that are most difficult to c nttrol by other meains Steil) horers appear to be the most widespread group that sigiiificantly dalag s rice aind is hard to manage by other Ilnca us

Biological control Very fe stutdic of mitatt~l eneties of rice pests in upland envirollnletits wee in tile wefouiid literature Therefore have not compiled a list of species Ve present a list of natural enemies iOtid in upland ricefields tuic latialln Batangas for some conitno iOldiar pest species ([itble 5)

Muich ofl thc lollowinle itfor ittiol cotlliCs rol Stutdies in the lowlatnds or roi crops otithcr thlin tice

Qu(ailtr o 11ll t11t( Nat trrtf lii hcoeits ight Ile used agaitist ile social insects tilt and termites Sollie predator specice live ill theirf nests n ilnic theirnllt ip-pearanIlCe aMRidbehavior IJtrouMdCing a mote aggressise ant species to displace pest ant codonies is a possibility (169) Patihogens as potential natural encnies have reccived little atteition (I13) Natural ncnuics asa group however hac not been toIl iporta it in Iegulating aintshown be and ternite numbers

The nocturnal behavior aid subteitmien habitat of lole crickets and field crickets protect them from many ntllral enemies There are only limited records of natural cnelics for these groups of pests some pathogens (87) but mainly

IRPS No 123 JattuIny I 271987

parasites - scelionids on eggsand sphecids on ilymphs and adults Sphecid wasps Larra carutmaria (SImitI hI L huzwuensis Rohwer and lsaltiuhta Willi ams speciali in mole crickets pa ralyzing them nd dragging the ninto thcir nests as food lot their voting ()tier sphCcid species Lins aurulenoa (Iabricius) lot maiac (Ashmead ) 1 lhshylssellaus (Smith) and Ml lal)oriosu (Snithf specialie in field clickets

White grUbsO the other hand hae Icti scriuns enuigh to have been extensively studied for cotttroi 1W nati al enemies( 112 140 141) Scoliid parasites C(ajsoMcispp have beetn intrtduced itto tilePhilippites for control of Ielxl ]udisiro la The rcsearchcts w Intttrdced themle during ill era claimed that the white gttionthreak as controlled (142) howevcr it isnormal for pest epidemics to subside and there was no diect evidence to indictte that tfe parasite was responsible A tmore promisitg titctllod might he to identify iso ltcttass prt~f tcartd apply hiactelaln fingal pathogens to tile soil Ihe lalattese beetle in North America has hcci coitrolled hv tite conmnleciall aailable hactcrial prcpa ration itacihlpu pav ofr milksdiscas (71) Spote prtyefott the soil itloted ilrtier reitiain actic fr Veas

Root aphid sardtnctly bugs readilslic tilib to predatots and parlsites once their tetndine ants are controlled The larvac and adtilts of a coccinellid beetle 1Scynms sp prey on llriadlouais in slash and burn upland rice Baits treated with insecticide may be used to control tending ants by killing their ourig (244)

Root-tkcditig blugs false wireoris wireorms and root wccils aic r ofsuch little ifilpoltatice that thenomttll role of liatural Cticniics hias tiot beei asscssed A possibility hoevrl tiglt he tit ItaIIitic niaCIMtodcs lintt1g11en Itese soil-itlhlfbitine mCtitodcs W5uhtfhave to be imiass Produiced

large miltibcr of niatltrll cnemics ircknown ft CltIoriS IC IpblCeii is thtt clttvortnlls ofonlie the crop soott alter land ptel)tritiOti a1tithe bcgit iniig of the raittv

Citsoln hfien tiatllral leniiiy populations aric lo Colltloil WOtuld iii ole mass prodtticing andirclcaslng key spccies

le egg stage is the fpirt (olthe Cedlitgt Ialot life cycle tI hich to f)ctis ftor tittlril elctit cottntrol hplraites o1 pleditors ttsit elleggs highly expoIsYCl I Sgg pailsites or predators stolld probahly e ittass andtbe produced released Littlh i toruialioll is a ailable ott what species to trV o ho to Iniss podLice thiCIi A etiloplhid wasp Icmihlaol sp effctiicly regushy

lates the leatf lcloalapolm2a spioanoiiitr heat andc rice antid should be collsctvcd

leaf beetles whCllose e am] eggs ate laid in the grutindhirO ould be motC difficutlt to control by niatulral enelV Mnll ipuhltiOn But species sUici IS c_gsthispid beetles ith

and larVle oilleascs ate vtilcratble to iparaites and pledators

Natural enemisc of tlirips otl rice are little known

28 IRPS No 123 January 1987

Table S Parasites recorded from upland rice insect pests in Tanauan Batangas Philippines 1977-84

Order Family Species Host stage

I lymttenoptera Braconidae Leaffolders Cnaphalocrocis Marasinia spy

Cardiochilesphilipplinensis Ashmead a Larva

Cotesia angustillasis(Gahan)Cotesia or cypris (Nixon)

Larva Larva

Cotesia or taeniaticornis(Nixon) Larva Cilonusinotakatae Munakata Chelonits SppAlacrocetrts philippinensis (Ashtmead)

Egg to larva Egg to larva Larva

Iclineumonidac Orgilts spp Isclutojppa hireator(Fabricius)

Larva Larva

Itoplectis naraigav(Ashmcad) Larva Tenelhtcha philippiensis (Ashmead) Larva Tenelchha stangli (Ashmead) a

irichomma cnaphalocrosis Uchida Larva Larva

Bethylidac Elasmidac

Goniozms or trianguli r Kieffer lasmtusalhopictus Crawford

Larva Larva

Diptera

Encyrtidac Chalcididac

Tachinidac

Elasmtus spp Copidosomopsisnacohiae (Fidy) Brach nteria evcarinata Gahan Biachyieria lasus (Walker) Ztgohothria ciliata (Wulp)

Larva Egg to larva Pupa Pupa Larva

Argyrophylx nigrotihialis(Blaranov) Larva

l-ymenoptera Braconidac Yellow stern borer Scirpopiaga incertulas

Braco chinensisSzeplegeti Larva CIthelotis spp Chelonus niunakatae Munakata Cotesi flaripes Cameron Stenoracon nicerilli (Bingham) a

Larva Egg to larva Larva Larva

Ichneumonidac Tropobracon schoenobii (Viereck) Amatroinorpiaaccepta metathoracica(Ashmiead)Frihorus sinicus (I lolmgren)

Larva Larva Larva

Ischnojoppaluteator (Fabricius) Larva Isotima nr dammeratiti Rohwer Larva Tenelucha philippinensis (Ashmead) a

Larva Tenehicha stangli (Ashmcad)Trichomma enaphalocrosis Uchida

Larva Lara

Chalcididac Eulophidac Trichogranmmatidae

Xanthopimnpla stemmator (Thunberg) Charops hrachYpterumn Gupta and Malteswary Brachtm eria sppTetrasticttus schoenohii Ferriere Trichorigranm chihunis Ishi

Larva to pupa Larva Pupa Egg Egg

Diptera

Scelionidac Pteromalidae Tachinidae

Tt ichograintnta aponiicuitt (Ashniead) Telenomits rowani (Gahan) a

Trichoinalopsis apattelctena (Crawford) Jeirhaea spp

Egg Fgg Egg pupa Larva

Z4gobothria ciliata (Wulp) Larva

Ilymenoptera lecneumonirlac Gol fringed borer (hilo auricilius

Trichomm enoptalocrosisUchida Larva

Ieioenoptera Braconidae

Aantholimlh steiniator (Thtinberg) a

Pink stet Ioret Sesamia inferens StoInacoil nicrilhi (Bingh am)

Loarva to pupa

larva

Diptera Ilyinenoptera

Armvtnyorms Mythimna separata Spodoptera mauritia acronyetoidesTachinidae Zvgohothriaatropiora( Robinea i -Desvoid y)Eulophidae It-phctrtiscitapadoe (Ashimead)

Larva Larva

(halcididae Braconidac

Brachpin ria spip (oesia spp

Larva larva

IHyntenoptera I)iptera

Rice skipper Ielop ida (halcididae Tachinidae

math ias aidgreen iortedci ciliarNIelan itis eda Brat lhincria sp iii matuqitatt Cameron Irgyrolhirlaxi trotlhiais(marumtv)a1

ismtne Larva Larva

Diptera Ilymenoptera

Tachnidae lelineutionidae

fuown se tilo per Mois frugalisIrgtrophrlax 1igrothiahiv 3 a rat ov) anthopimpa pmnctata (Iabricius)

Larva Larva to pupa

aMost dominant

IRPS No 123 January 1987 29

Normally predatory thrips prey on herbivorous thrips strepsiptera on both Spiders dominate tilenymphaladultNatural enemies of stem bugs --Scolinoplara and Tibraa predators Spittle bugs move to rice from pasture grasses - are in order of importance egg par~isites and predators and biocontrol would be successful only ifcarried out before fungal pathogens and nymphal adult parasites Egg tiley immigrate into ricefields parasites could either be introduced fron other areas or Biological control tactics for hemipterous seed bugs mass produced and released ould follow those outlined for stein bugs

Armyworms are normally lwld in hcck by the activities Coml)arison of environnewtns Natural enemies are of egg and larval parasites When these natural enemies fail perhaps faced with even greaterconstraints than insect pestsusually because of drought armyworns become epidemic to survive ind reproduce in upland environments TheyIt may be worthwhile therefore to release parasites during have adapted ani survived by having wide host ranges the the rainy months following a drought Virus diseases of ability to aestivate or hibernate over unfavorable seasonslarvae would beanotheravnue to exploreforarnyworms and the ability to disperse or by other mechanisms

Parasites have ben traditionallyconsidered for biolegical Comparing the most dominant and therefore the most control attempts against stein borers usually by rekasing adapted species of natural enemies collected in various exotic species( 166) This approach has not met with success environments in the Philippines we can understand how and new avenues should be explored The role of predators different rice environments are arid pathogens is little inderstood and deserves greater Anagqrms olptahiisa mymarid planthopper egg parasiteattention Egg parasitism rates are normally reasonably is cquallyadapted to all Philippine rice environments (Tablehigh but should be supplemented with effective egg 6) It is selective for planthoppers and even parasitizes eggspredators Orthopterans are potential egg predators of the n-iize planthopper Peregrinitsmaidis itt upland areas Metioche vittaticollis (Sta ) Anavipha nifiwnnis (Saus- Alavohis also parasitizes maize planthopper eggs but is sure) and Conocephahs hongipnni (de i-laan) feed on most prevalent on BPH in the irrigated wetlands (onashystem borer eggs etioche and Anaxil)ha specialize in eggs tocerus spp like A optalhilis on BPH and WBPI Iattacks with no hair such as Chilo spp The omnivorous (ono- the eggs of GLH in ell rie environments It is also found cephahsspecializes in eggs cov red with hairs it is a proven parasitizing the eggs ef the white rice licahopper (oimna egg predator of Scirlouphaga spp but has the discouraging spetra Oirosita naias a trichogrammatid egg parasitehabit of eating rice guin (65) attacks BIPH and WBPH in the wetlands and has no other

Grasshoppers have egg (scelionid) and nymphalad ult host A related species 0 aesopi specializes in G1-H in (nemestrinid tachinid and sarcophagid) v irasites Among wetland environments Dryinid strepsipt-an and pipunshypathogens protozoans have been recorded most frequently culid nymphal and adult parasites of planthoppers and Control of locust species by natural enemies could con- leafhoppers also specialize either in hopper species or centrate on their habitual breeding areas environments Among the dryinids Pseuhdgonatoipsflavi-

Leaffolders have rich complexes of natural enemies some feimur prefers the lowlands and BPH while P nuch has no adapted to upland rice (22 27) ranging from egg predators environmental prefirences but is adapted only to WBPH (gryllids coccinellids) to larval parasites (braconids ichncu- The main d ryinid species on GLH aptogonatop)usspp is monids) to larval prcdators (ants and carabids) to larval most dominant in rainfed lowlands and also attacks other pathogeiis (vira and fungal diseases) Efforts to augment leafhoppers within Amnrasca Cicadulinaand Bachaltuathese natural enemies should focus on the egg predators Strepsiptera are more dominant on plarithoppers than (rearing releasing)and viral or fungal pathogens (culturing lealhoppers The Elenchus species in the Philippines attack spraying) all species of planthoppers within the genera Sogawilla

Parasitization of the large conspicuous larvae of Sogatodels NilaparsataHarualia and Opiconsiva Elenshyskippers green horned caterpillars and other polyphagous cusrvasuntaisuiis adapted to the major rice environments Lepidoptera by tachinids and chalcids is normally low ELg but prefers BPH in rainfed arid WBPIH in wetland environshyparasites and predators are perhaps the key to their control ments Ilalictoph(utsmtnroeiand HVpectrus attack G LH

Gall midges are parasitized by pterormalids and platy- and COfimravcctra respectivelygasterids that aestivate within their larval host between Pipunculids are mainly parasitic oii lealhoppers Each seasons Shoot aphids are preyed upon by coccinellids species has a unique environmental preference Tomosshysyrphids and chrysopids varireIa oraetoraprefers Ne7 otetltix nigropictrs and

Planthoppers arid leafhoppers have egg parasites and Pilunwulhtsjavanensisattacks Dehoceplhahls spp which predators as well as nymphialad nlt parasitesand predators are more abundant in upland than irrigated lowland their rich complex of natural enemies includes fungal environments T suhvirescensattacks Nephotettix virescens pathogens The principal egg parasites are myrnarids and and prefers the lowlands Pilpuiulus nttillattls is an trichogramniatids Egg predators are norinally mirid plant upland rice parasite arid attacks only Nephotetix speciesbugs that suck out the yolk Nymphal ad nIt parasites are More than 17 recorded species of leaffolder parasites in dryinids on planthoppers pipunculids on lealhoppers and the Philippines attack species of C(naplwocrocis and

30 IRPS No 123 January 1987

Table 6 Dominant natural enemies of rice pests in upland rainfed lowland and irrigated lowland rice environments of the Philippines

Natural enemy

Host stage Species

Brown planthop-Parasite Egg

MymaridaeAnagrus optabilis(Perkins) Anagrusflaveolus Waterhouse

TrichogrammatidacOligosita naiasGirault

NymphAdult Dryinidac

Pseudogotatopuslaiifemttr Esaki et Hashimoto Elenchidae

Elenchus yasurnatsui Kifune and Hirashima

Environment

Upand Rainfed lowland Irrigated lowland

Ailaparata lugens

x x X x

x

x x

x X X Whitebacked planthoppers Sogatella furcifera and Sogatodes pusanusEgg

MymaridaeAnagrus optabilis(Perkins)

TrichogrammatidacOl~gositanaias Girault

NymphAdult Dryinidae

Pseudogonatopusnudus Perkins Elenchidae

Elenchus yasutatsui Kifune et Hirashima

Parasite Egg

MymaridaeGonatocerus spp

TrichogrammatidaeOligositaaesopi Girault

NymphAdult Pipunculidac

Tomosvaryella subviresccns Loew Tomosvaryella oryzaetora (Koizumi) Pipunclusmutillatus (de Meijere)

DryinidaeIlaplogonatopusspp

Egg Copidosomopsisnacoleiae (Eady)

Larva BraLonidae

Cardiochiles philipphnensisAshnead lchneumonidac

Trichiomnia cnaphalocrosisUchida Temelucha stangli (Ashmead)

Pupa Chalcididae

BrachymeriaexcarinataGahan Brachymerialasus (Walker) Brachymeria nr tachardiaegroup

Parasite Egg

Scelionidae Telenomus rowani (Gahan)

EulophidacTettasticusschoenobhi Ferriere

Larva Braconidae

Stenobraconnicevillei (Bingham) Iclhncurnonidac

Temelucha phiiippinensis(Ashinead)Temelucha stangli (Ashmead)

x x x

x x

X x X

x x Green leafhopper Nephotettix virescens

X x x

x x

X X X X

X

X Rice leaffolders CnaphalocrocisMarasmia

x X x

X

X X

x x x x x X x x x

Yellow stem borer Scirpophaga incertulas

X X x

x

x

x X XAinauromorphaaccepta metathoracica(Ashmead) x

continued on next page

IRPS No 123 January 1987 31

Table 6 cntinued

Natural enemy Environment

Host stage Species Upland Rainfed lowland Irrigated lowland

Larva-pupa Gold fringed borer Chilo attriciiuslchneumonidae

Xanthopimpla stemmator (Thunberg) x aPupa x

Pteromalidae Trichoinalopsis apanteloctena (Crawford) ax

Pink stem borerb Sesamnia inferensParasite

Larva Braconidae

Stenobracon nicevillei (Bingham) x b b Armyworms Mvthinna SpodopteraLarva

Tachinidac Zygobothriaarroptvora (Robineau-Desvoidy) x x

Braconidac Cotesia spp

X Skipper Pelopidas mathias and green horned caterpillar Melanitis leda ismeneF-gg

TrichogramrnatidacTrichogrannaspp x

Scelionidae Undetermined Scelionid

x Larva

Tachinidae Argyroph lax nitrotibialis (Baranov) X x

Pteromalidae Trichomnalopsis apanteloctena (Crawford)

X Predator

CoenagrionidaeAgriocnenis femina femina Brauer x xIschnura senegalensis tambur

Miridae CrtorhinsliidipennisReuter x x

Nabidac Stcnonabis tagalicus (Stal) x

VeliidaeAlicrovelia douglasi atrolineata (Bergroth) x

Coccinellidae Micraspis crocea (Mulsant) X x x

Vespidae Eumenes campaniformis (Fabricius) x

Formicidac Diaecamna spp xOdontoponera transversa (F Smith) x

Carabidae Chaenius spp OphioneaW- aJasciataSchmidt-Goebel

x

Ophionea ishii ishii Habu x x X

Gryllidav Anaxipta longipennis (Saussur-) x xMetioche ittaticollis (Sial) x x x

Tettigoniidae Conocephalus longipennis (de Haan) x x x

aNo rearing made on rainfed lowland bNo rearing made on rainfed lowland or Irrigated lowland

MrasmiaAmong the Laval parasites the braconid Cardio- lowlands and parasitizes oriental maize borer OsIriniachilesphilijpphensis is most adapted to an upland environ- furnacalis (Guenee) larvae The leaffolder pupal parasitesment It also parasitizes Hydelepta indicata a common not only occur equally in all environments but also haveleaffolder of legumes The ichneumonids Trichomma cna- wvide host ranges- parasitizing species of OsIrinia Marucaphalocrosis and Temehcha stangli occur more in the Hydelepta and Honiona - in maize and legumeslowlands T stangli is most adapted to irrigated lowlands The principal egg parasite of yeltow stem borer (YSB)and also parasitizes Chilo and Scirpophaga stem borer Telenomus rowani is widely adapted to all environmentslarvae T cnaphalocrosis is most prevalent in rainfed and has an unusual alternate host -Tabanus eggs (25)

32 IRPS No 123 January 1987

Tetrastictu schoenobii specializes only in YSB and occurs mainly in irrigated lowlands -- further proof that YSB is most adapted to the wetter environments

Larval parasites of YSB are highly environment-specific The braconid Stenohracon nicevillei occurs only in upland rice but also attacks the pink stem borer (PSB) Tieinchwa phiiiplPwnsis an ichneumonid occurs in upland rice and also parasitizes the larvae of (7hiu stqpresaisinatro-morpha accepta melaihoracica like 7 stangi is mnost adapted to the irrigated lowlands

The dark-headed stem borer(l)HSB) (ih pochit3sus was not collected in all rice environments and the gold fringed borer Chilo auricilius nd PSB were collected only in upland rice Xaantiholiplastenrnaloran ichneumonid larval parasite appears to be widely adapted whereas TrihomaslopisapanitI)cteiiaa pteromalid pupal parasite occurs mostly in wetlands and also parasitizes larvae of Chilo suppressais and Pelhuidas mathias

Armyworms Myit hinin and Spodujipera have two main larval parasites - a tachinid Zrgohothriaatropivora in rainfed environments and a braconid Cotesia spp in irrigated lowlands

The butterfly pests Pelopidas and 4elanitishave tricho-grammatid egg parasites found only in irrigated lowland environments

Two larval parasites show environmental preferences --Argrroph av nigrotihialis a tachinid in rainfed habitats and Trichonialopsisapantelkcnela in irrigated ricefields A limgrotibialisalsoattacks the sweet potato hornwormAgrius convolvtdi and T apanteloctena parasitizes DH S1B

Predators as a group show more distinct environmental preferences The aquatic damselflies Coenagrionidae are most abundant in the lowlands Cvriorhinus is also most adapted to the lowlands The nabid Stenonabis tagalicus occurs only in the uplands and isalso prevalent on legumes The coccinellid lady beetles wasps and ants are more prevalent in upland areas AlicrasIpis cit-ea larvae prey ona variety of aphids attacking legumes and maize Etunienes campanonia vespid mud wasp makes nests in trees and is therefore most adapted to the more botanically diverse uplands Soil-dwelling ants cannot tolerate flooding

Upland environments are habitats to arboreal carabid beetles Three species of Chlaeniusbeetles prey on leaffolder larvae and are perhaps more important than parasites (27)

The gryllids and tettigoniids are widely adapted egg predators not only on rice but also on maize and legumes They feed on eggs of most insects that are laid on leaves

The spidercommunity of upland rice environments isrich in species At one site (Batangas Philippines) 31 species have been recorded (23) Of the 176 spider species recorded in Philippine ricefields about one-half (82) occur in upland rice The spiderspecies of upland rice environments overlap more with those of rainfed lowland than with those of irrigated lowland rice (24) The same study showed that of the three environments irrigated rice has (ie greatest spider species diversity foiiowed by rainfed lowland and upland

However of the 10 most prevalent ricefield spiders 9 were abundant in upland rice showing wide adaptation All three spider guilds -- orb-web space-web and huting spiders shywere prevalent in upland ricefields There were however differences in environmental prelerences for some spider species

Of the orb-wcavers Tetragniathamanlihulatawas partishycila rlv abundant 7 japonica was low and Ltucatge dehcorata was absent in upland rice

As in other environments Aiypeiwnaifirtosana was the most dominant space-web spider however its relative numbers were lowerthan for lowland sites Among hunting spiders Ox|opes javanus and Lcosa leatcost fgla were more abundant and Lvcosa lpseudoanmnlata was less abundant in upland compared with lowland rice envishyroniments

Extensive rearing of rice insect pests in upland rainfed lowland and irrigated sites over an 8-yr period in the Philippines provided insight into the effectiveness of parasites as natural enemies (Table 7 8) Most of toe data are from large sample sizes taken over at least one crop which overcomes the pitfalls expressed by Van Driesche (238)

VSB egg parasitization was surprisingly similar across all environments Larval parasitization was highly dynamic among sites and even years within the same site but showed somewhat lowerlevels in uplandsthan in irrigated wetlands This would have been predicted for egg parasites as well since YSB is adapted to the lowlands and so must be its parasites This isevidence that Telenomus rowani is as good at dispersing as its main hosts Trowani clings onto the anal tufts of female YSB and parasitizes eggs as they are laid This phoresy may explain its wide adaptation The more specialized lowland YSB larval parasites therefore appear more effective than their upland counterparts with more alternate hosts

Low collection levels of PSB larvae especially in the wetlands make comparisons between environments difficult Low levels of larval parasifization were recorded in both rainfed environments Except in the Cagayan rainfed wetland site in 1980 low levels ef larval parasitization were evident at irrigated and rainfed sitesalike No differences by environment were evident Higher levels of parasitization occurred in Sarawak during a 1967 outbreak also from tachinids Rothschild (197) indicated that higher parasishytization occurred in upland rice areas

Skipper and green horned caterpillar larval parasitization rates were generally low and showed no environmental effect htighest activity occurred in Cagayan in 1980 (48 parasitization) Although larval parasitization levels of RLF were variable year to year activity appeared greater in the irrigated lowlands than in the uplands Again the highest levels were recorded in Cagayan in 1980 Cagayan has extensive grasslands and fallow around the rice areas and a relatively long rainfall period Rainfall in 1930 was good for crop growth and apparently for increase of larval parasites

IRPS No 123 January 1987 33

Table 7 Parasitization rates of rice planthoppers and green leafhopper in upland rainfed lowland and irrigated lowland environments Philipshypines 1976-840

Environment and site Year Egg parasitization () Nymphaladult parasitization ()

BPH W13P GLH BPII WBPH GLIH

UplandBatangas

Rainfed lowland

1976 1979 1980 1982

15 17 12

2(47)8(39)

4(51)

9(1800)14 (212)

12(304)

8 (101)6 (127)

9(116)

Iloilo

Cagayan

1976 1977 1978 1981 1980 1981 1982

0 (69)

11

44 48 40

-

-

46 55 44

0 (471)

11

52 56 49

0 (200)9 (214)

10 (1259)8(235)6 (900) 3(800)

-7 (462)

11 (270)3 (313)5 (900) 1 (800)

4 (600)10 (725)10 (362)12(870)5 (900) 3 (800)

Pangasinan

Irrigatedlowland

1984 1976 1978 1979 1982

13 (436) 13(379) 14 (457)

6(14)7 (197)

17(198) 11 (486) 18 (275)

3 (81)10 (274)3 (116)

16(871) 19(88)

28(211)9 (764)5 (647)7 (1204) 8(305)

Laguna 1977 11 7 21 1978 31 1979 14 4 4 28 1980 7 8 20 1981 1983

56 7(278)

26 16 (1650)

16 27(1921)

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R I Apostol unpublished 1lo1lo and Pangasinan 136 Cagayan BCanapi unpublished Liuna 175 I C Pantua unpublished 118 234 43 Carino and Shepard unpublished BPH = brown planthopper Nila parvara htgens W131Pi =whitebacked planthopper Sogatelia furcifera GLH = green leafhopper Nephotettrix virescens Figiires in parentheses are sample sizes

Tauher et al (230) reported that internal external and genetic factors influence the seasonal activity of parasitoids

Hopper egg parasites but not nymphaladult parasites were also parlicularly abundant at the Cagayan site possibly the egg parasites have more alternative hopper prey in the weedy areas than nymphaladult paisites

The seasonal dynamics of hopper egg parasites can be seen for upland rice in Batangas(Fig I 2) and for rainfed lowland lice in Pangasinan( 135)and Cagayan( 105) Levels of BPHWBPI-L and GLH egg parasitization were corn-parable in upland and irrigated lowland environments Except for some high rates of paraMsitization early in the season the parasitization rate tended to be steady in all envilonments However hopper nymphaladult parasi-tizution levels were more similar in the rainfed environments and lower than in irrigated rice

Overall parasites do not appear highly effective by themselves against upland or even lowland pests Among natural enemies parasites have attracted the most attention because they can be more readily assesscd

Predators and pathogens lend themselves better to management practices Predators the most important group of natural enemies in riccfields are difficult to quantify On the other hand pathogens appear less im-portant but can be readily cultured and disseminated

The natural enemy commulnity of upland rice is rich in species and differs significantly from that of lowland rice

These beneficial organisms must be conserved by applying insecticides judiciously particularly sprays Seed or soil placement of chemicals minimizes exposure to natural enemies The strategy to derive the greatest benefit from natural enemies isto allow the greatest number of beneficial species to thrive Then perhaps at least one species will be effective against each pest at any tine overcoming the variable abundance of each species within and between years

Programs to introduce exotic species or mass produce indigenous ones are ambitious and expensive because they require trained people on a sustained not ad hoc basis

Chemical control Insecticides offer rapid and efficient control of upland insect pests (190) but-should be used only after other control measures have been considered Insecticides are rarely used on upland rice because of cost (32) Upland rice yields are normally too low to justify the expenditure Also most upland rice is grown as a cash crop Spraying upland rice is more difficult than lowland rice because water is less accessible Government subsidy programs to provide pestishycides to tribal peoples for upland rice production have created problems of toxicity to humans because isolated people have had no experience to be able to handle pesticides safely

However ins-eticide use has beenjustified when a crop is

0Z

Table 8 Parasitization rates of lepidopterous pests in upland rainfed lowland and irrigated lowland environments Philippines 1976-84a

Parasitization (5)

Environment and site Year YSB GFB PSB Earcutting Swarming Skipper Greenborned RLF Brown larva larva caterpillar caterpillar caterpillar caterpillar larva semilooperEgg Larva larva larva larva larva larva

UplandBatangas 1977 - 0 (57) - 0 (63) - 0 (71) - 0 (106) 11(142) 2 (101)1978 43 (4697) 8(451) 5 (38) 3 (40) 6 (61) 2(58) 15 (57) 13 (129) 13 (1286) 4 (30)1979 59(4135) 12(320) 13 (11) 5(28) 9(45) 6(46) 7(23) 8(160) 9(1152) 8(26)

Rainfed lowland Iloilo 1976 - 0 (10) 0(10) 0 (7) 0 (19) 2 13) 15 (180) 5 (76)1978 0 (54) 0 (3) 0 (17) 9 (64) 4 (78) 3 (73) 0 (121) 0 (191)1979 64 (2350) 16 (84) 6 (17) 0(6) 18 (68) 25 (104)

1981 56 (1322) 18(304) 0(5) 12(51) 18(138)Pangasinan 1976 60 (7 7 )b 62(82) 0(111) 5 (39)

1978 58 (53)b 5(51) Cagayan 1979 59(713) 8 (101) 6 (34) 0(7)

1980 0(11) 4 (28) 15 (286)

62 ( 48(177) 61(140)1981 0(5) 12 (2) 13(45) 14 (92)

Irrigatedlowland Laguna 1974 62 8

1974-75 50 41 1975 43 72 1978 96 (623) 21(24) 33 (60)1979 83(2017) 18(110) 8(62) 11 (89) 22(400)1980 78 (1625) 20(75) 0 (16) 4 (72)1983 60

9 (80) 7 (270) 30

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R F Apostol unpublished Iloilo and Pangasnan 136 Cagayan B Canapi unpublished Laguna 175 P CPantua unpublished 118 234 143 Carino and Shepard unpublished YSB = yellow stem borer ScirpophagaincertulasGFB = gold fringed borer Chilo auricilius PSB = pink stem borerSesamia inferens earcutting caterpillar Alythimna separata swarming caterpillar Spcdoptera mauritia acronyctoides skipper Pelopidas mathias reen horned caterpillar Melanitls ledaismene RLF = rice leaffoldersCnaphalocrocismedinalis and Marasmiaspp brown semilooper Mocisfrugalis Figures in parentheses are sample sizes Egg masses

IRPS No 123 January 1987 35

Table 9 Effect of seeding rate and insecticide on plant density and Yield losses from insect pests in upland rice often areyield of Dagge upland rice Batangas Philippines 1976a comparable to those of lowland rainfed or irrigated rice

Seeding rate Insecticide Dosage Plant density Yield (0-Y30o) but usually from different pests The principal (kg aiha) seed treatment (kg aiha) (noIra-row) (tha) groups are soil pests feeding on sown seed and rootsat 14 DE followed by seedling pests These groups have largely gone

50 None - 35 a 20 a unnoticed leading researchers to conclude that stem borers50 Carbofuran F 10 38 ab 23 a and seed bugs are the most important pests (17) Yield loss 10 beCabofuranF100 None - 4551 c 23 a trials that pinpoint major pests need to includc seed

100 Dieldrin WP 10 52 cd 22 a treatments Granules applied in the furrow may not give150 None - 59 d 22 a adequate protection from many seed pests

aAv of 8 fields DE = days after crop emergence In a column means The floristic and pedological diversity of upland rice followed by a common letter are not significantly different at the environments is matched by the diversity of arthropods5level by D)MRT F = flowable Wi = wettable powder both pests and their natural enemies Irrigation has homoshy

genized lowland rice environments Flooded soils tend tothreatened by epidemics of locusts or armyworms New have the same properties and to eliminate soil pests The technology stressing efficiency has minimized insecticide spatial and temporal dominance of cultivated rice has cost benefited specialist pests

Seed treatment is normally inexpensive and can be Upland rice has no specialist arthropod species The economicallyjustified in -nanycases(66) Treating seed with monophagous pests on upland rice prefer the lowlands systemic insecticide (216) but not chlorinated hydrocarbons where rice evolved Rice in the uplands exists only because (247) is most effective in controlling seedling maggots of man and even though lowland rice predates upland riceFoliar sprays have to be repeated to obtain control and specialist species evolved Wheat maize and sorghum do often are washed off by frequent early season rains not have monophagous pests leading us to conclude that Granular insecticides are effective but required dosages are there is no evolutionary advantage for inssects to have too expensive to justify except under heavy infestation narrow host ranges in upland graminaceae(247) When directed against first-instar larvae banding Pests have adapted to the uplands not only by having granules in seed furrows (177) can be effective against white wider plant host ranges but also by having long life cyclesgrubs at low dosages (025 kg aii ha) (134) Seed treatments undergoing dormancy or being dispersive Each pest has a are not promising for white grub control (248) unique life history strategy suited to highly unpredictable

Seed treatment is also effective ainst ants and may be environments In agivenyear someinsect specics an better cheaper than increasing the seeding rate In Batangas exploit upland rice as a food source but no pest is a senious treating 50 kg seed ha was as effective as increasing the one every year on upland rice seeding rate to 100 kgha and using untreated seeds (Table The low yielding potential of upland rice means less 9) In Brazil seed treatments control termites and other soil quantity and lower quality of food for pests to exploit but insects including the lessercornstalk borer(66) Domiciano also less incentive for farmers to undertake control (58) however found seed treatments inconsistent in their The most effective control would be early and synchroshyeffect against high populations of lesser corn stalk borer nous planting of early-maturing upland rice varieties to

Baiting can be an inexpensive way of controlling ants prevent insect pest buildup Breeding for insect resistance seedling maggot flies mole crickets and field crickets Baits should initially focus on stem borers because they are verymade of locally available material and impregnated with difficult to control by other means Biocontrol efforts insecticide can be sparingly distributed taking adantage of should first stress conservation of natural enemies preshythe pests ability to disperse and encounter bait sites dators and pathogens rather than parasites Efficient

The new low-volume hand-held controlled-droplet chemical control includes seed treatment or baiting for seed sprayers may offer an advantage over high-volume knap- seedling and some root pests and using controlled dropletsack sprayes for controlling foliar pests Upland rice sprayers for spot treatments against foliar pests and seed canopies are more open than those in lowland rice and bugs when economic thresholds are reached better droplet penetration should result Controlled droplet applicators would be ideal for seedbugs and defoliators

CONCLUSION

Upland rice is attacked by a wider array of insect pests than is lowland rice mainly beca use of the addition of soil pests Generally population levels of lowland rice pests are lower in upland than lowland rice leading many people to conclude that insect pests are not important on upland rice

36 IRPS No 123 January 1987

REFERENCES CITED

1Abraham C C and K S R Mony 1978 Occurrence ofLeptocorisa acuta Fabr (Coreidae Hemiptera) as a pest of nutmeg trees J Bombay Nat ttistSoc 74553

2 Agyen-Sampong M 1975 HlierogrlphusdiaganensisKrauss (Orthop-tera Acrididae) a new pest of rice in Northern Ghana Ghana J Agric Sci 8249-253

3 Agyen-Samnpong M 1976 Varietal resistance and other inle-ratcd pest control components as factors of rice varietal improvement in West Africa Prcsented at the Second varietal improvement serinar 14-18 Sep 1976 Bouak Ivory Coast West Africa Rice Development Association Monrovia Liberia 18 p

4 Agyen-Sampong M 1977 Outbreak of Ihccrorrdnisorae in rice seedlings near Rokupr Sierra Leone lit Rice Res Newsl 2(6)16-17

5 Agyen-Sampong M 1980 Dirty panicles and rice yield reduction caused by hugs IntRice Res Newsl 5(l)11-12

6 Agyen-Sampong M 1982 The major pest problems of irrigated upland and mangrove swanip rice ecosystems ir the humid tropical Guinea savanna and Salel climate zones Pages 318-328 in Pro-ceedings training course cn concepts techniques and application of irtegrated pest management in rice inWest Africa 10-20 Jan 1982 Fendall Liberia West Africa Rice l)evelopioent Association Monrovia Liberia 505 p

7 Akibo-Betts V I and S A Raymundo 1978 Aphids as rice pests in Sierra Leone lit Rice Res Newsl 3(6)15-16

8 Akinsola F A 1975 Present status of different rice stemborers in parts of Nigeria iicL Entomo Newsl 328

9 Akinsola FA 1979 Ihe biology and ecology of rice stemborers in Nigeria Ph 1) thesis University ol lbadan Nigeria 217 p

10 Akinsola E A 1979 Guidelines of integrated management of insect pests of upland rice in West Africa Presented at the Seminar on integrated management of rice diseases and insect pests at Bobo-Dioulasso Upper Volta 17-22 Sep 1979 West Africa Rice Develop-ment Association Monrovia Liberia

II Akinsola IA 1984 Insect pests of upland rice in Africa Pages 301-305 in An overview of upland rice research Proceedings of the 1982 lIouak Iwsrv Coast upland rice workshop International Rice Research Institute 13O Box 933 Manila Philippines

12 Alam M S V TJohn and Kaung Zan 1985 Insect pests and diseases of rice in Africa Pages 67-82 in Rice improvement in Eastern Central and Southern Africa proceedings of the international rice workshop at Iusaka Zambia 9-19 April 1984

13 Alticri M A A van Schoonhovcn and J D Doll 1977 The ecological role of weeds in insect pest management systems a review illustiated with bean (Phaseohsvulgaris L)cropping systems PANS (Pest Artie News Summ) 23195-205

14 Anmante F 1967 The leaf-cutting ant Atia capiguaraa pest of pastures [in Portuguese] Biol6gico (Sao Paulo) 33113-120 (Rev Appl Entomol A56207)

15Andrade A C and D Puzzi 1951 Preliminary results of experi-merits on the control of Sculptcoris iastantus on sugarcane [in Portuguese] Bioulgico (Sao Paulo) 1744-49 (Rev Appl Entomol A41421422)

16 Appert J 1970 Maliarplia separaelha (rice stem borer) new observations of entomological problems of rice in Madagascar [in French] Agron Trop (Paris) 25329-334

17 Ariadeau Mand Z Harahap 1986 Relevant upland rice breeding objectives Pages 189-197 in Progress in upland rice research Proceedings of the 1985 Jakarta conference International Rice Research Institute 1O Box 933 Manila Philippines

18Asar S and Kisimoto 1975 Seasonal occurrence of the smaller brown planthopper in upland rice [in Japanese] Proc Kanto-Tosan Plant Prot Soc 2228

19 Bakker W 1974 Characterization and ecological aspects of rice yellow mottle virus in Kenya Meded Landbouwhogesh Wage-ningen 152 p

20 Balut F F 1970 White grubs attacking rice Oryza ativa L [in Portuguese] Biol6gico (So Paulo) 36321-322

21 Banerjee S N and L M Pramanik 1967 The lepidopterous stalk borers of rice and their life cycles in the tropics pages 103-124 in The major insect pests ofthe rice plant Proceedings ofa symposium at the International Rice Research Institute September 1964 The Johns Hopkins Press Baltimore Maryland USA

22 Barrion A T and J A Litsinger 1980 Ants a natural enemy of cliaiarac in dryland rice IntRice Res Newsl

5(4)22-23 23 Barrion A T and J A litsinger 1981 The spider fauna of

Philippine rice agroecosystem 1Dryland Philipp Entomnol 5139-166

24 Barrion A T and 1 A Iitsinger 1984 The spider fauna of Philippine rice agroccosystcms II Welland Philipp Eiitomol 611-37

25 Barrion A T1 and J A Litsinger 1984 lihanus (Diptera Tabanidae) eggs an alternative host of rice stei borer (SI3) egg parasite Telenoonis digns (Iliymenoptcra Scelionidae) lit Rice Res Newvsl 9(6)19

26 Barrion A T and J A Litsinger 1985 Identilication of rice leaffolders by wing markings Int Rice Res Newsl 10( 1)24

27 Barrion A T and J A Litsinger 1985 Chctiosspp (Coleoptera Carabidae) a leaffolder ILF)predator lit Rice Res Newsl 10(1)21

28 Bertels A 1970 Insect pests of rice and theircontrol [in Portuguese] Pelotas Brazil Instituto de Expcrimentacao Agropccuararias do Sul Circular 43 24 p

29 Bonzi S M 1982 Ctilo (flisilineus J de Joannis (Lepidoptera Pyralidae) stem borer of irrigated cereals in Upper Volta LioFrench] Agron Trop Riu Rizic Cult Vivrieres Trop 37207-209

30 Bourke T V T L Fenner J N LStihick G Baker EHassan D F OSullivan and C S li 1977 Insect pest survey for the year ending 30th June 1969 Papta New Guinea )epartment of Agrishyculture Stock and Fisheries Port Moresby 57 p

31 Bowden 1 1973 Migration of pests in the tropics Meded Rijksfac Landbouwwet Gent 38785-796

32 Breniere J 1973 The main rice parasites in West Africa and their control Pages 27-42 in Proceedings seminar on plant protection for the rice crop West Africa Rice Development Association Monrovia Liberia

33 Breniere J 1976 The principal insect pests of rice in West Africa and their control West Africa Rice Development Association Monrovia Liberia 52 p

34 Brown ES 1962 The African arinyworn Spoiolptera eXI)Ina (Walker) (Lepidoptera Noctuidae) a review of the literature Cornshymonwealth Institute of Entomology London UK57 p

35 Brown F B 1968 Upland rice in Latin America FAO-International Rice Commission 12th Session of the Working Party on Rice Production and Protection 2-14 Sept Peradeniya Sri lanka Food and Agriculture Organization Rome Italy 6 p

36 Buyckx F J 1 1962 Disease and insect pests of crops inCongo Rwanda and Burundi [iil French] Publications dc LInstilut National pour lestude Agrononiquedu Congo M Weissenbruck S A Brussels Belgium 708 p

37 Cariiio F 0 1981 Role of natural enentics inpopulation suppression and pest management of green rice lealhoppers Ph D thesis University ofthe Philippines at Los Bafitos PhilippinesLaguna 136 -

38 Carpente A J 1983 The development of rice cultivation and extension of vegetable productin in Zanzibar Final Report Vol II1 October 1983 Food and Agricultme Organization Ronte Italy

39 Carroll C P and S1 Risch 1983 Tropical annual cropping systems ant ecology Environ Manage 751-57

4(0 Cates R G 198 1 Host plant predictability and the feeding patterns of monophagous oligophagous and polyphagous insect herbivores Oecologia (Berl) 48319-326

IRPS No 123 January 1987 37

41 Cendcafia S M and F B Caora 1967 Insect pests of rice in the 62 Ene 1 C 1981 I)iiorldhol)ierusihilis Slater (licteroplera Philippines Pages 591-616 in The najor insect pests ofthe rice plant lygacidue Illissinac) a new rice pest front Nigeria lit Rice Res Proceedings of a symposium at the International Rice Research Newsl N3)15 Institute September 1964 The Johns Hopkins Press Baltimore 63 Fugenio T S 1965 rhe effect oi e-vCgetation on rice production Maryland USA 729 p and the possibility of overcoming it by leaf pruning MS thesis

42 Centro Ihternacional de Agricklittira Tropical 1984 Upland rice in University ol the Philippines at Los Baijos ILaguna Ihilippins 69 p Iatin Anerica Pages 93-119 hi An os ervies of uplaind rice research 04 Farrow R A 1975 The Alrican nmigratory locust ii is main Proceedings of the 1982 Boinakamp Ivory Coast upland rice workshop outbreak area of the middle Niger qlinlitatise studies of solitary International Rice Research Institute P0 Blox 933 Manila populations in relation to environmental factors Locusla No II Philippines 198 p

43 Chandra G 1979 Taxonomy ad hn riononlics ofthe imect parasites of 65 Fennal R G 1943 Food crop investigation Windward and rice Icalloppers and planthoppers in the Pihilippines and their Ieeward Islands First report Report onl work dunring the periodiiiportairce iii natural biol)gical cointrol Philipp Fntoltol 4119-139 April 1942-April 1943 Imperial College of Tropical Agriculture

44 Chang T T 1976 The originii colut ion enlIivation dissemination Trinidad 16 p (Rev of Appl Fintornol A3222-25) and diersificition of Asian and African rices -uphytica 25425-44 1 66 Ferreira F 1983 Integrated pest ritaageiret of upland rice [ill

45 Chatierjee P B 1979 Rice leaffolder attacks in India lit Rice Res Portuguese English summary] Pages 323-341 in Upland riceculttre Newsl 4(3)21 factors affecting production E Ferreira T Yamada and F-

46 Cheng Cbien-pan 19(4 Upland rice in Taiwan International Rice Malmolta eds Institutoda Potassae Foslato Institnto Internacional Cnmmission Wiorking Party oit Rice Production ard Protection I()th da Potassa Piracicaha Sao Paulo lraid Meeting 3-10 March 1964 Manila Philippines Food and Agricul- 67 Ferreira E arid F dl Silva Martins 1979 Occurrence of ture Organiaion Rome Italy 5 p Sogatohs orvziola (Muir) inupland rice in Goias Ilrail lit Rice

47 Chcrrett 1 M and 1) 1Peregrine 1976 A review of the status of Rcs Ncwssl 4(4)18 leaf-cutting ants and their control Ann Appl Biol 84124-128 68 Ferreira E and 1F da Silva Martiris 1984 Insect pestof rice ill

48 Chipeta F M 1974 Pests of rice in Malawi Pages 90-95 in Brazil and their control Jill Portugucsej FMIlRAtPA-CNPAF Proceedings Sith eastern Africa cereals research conference 10-15 Goiania Brmiil 67 p March 1974 DRB Mialda ed Ministry of Agriculture lilongwe 69 Ferreira FI F ca Silha Miartins S S Nero and 1 It P de Malawi Carvallio 1982 M tltiplc regression analysis todeterimine the effect of

49 Chu Yau-i 1979 The d atiaging period of the rice seedling fly insect populatiois ott upland rice yield [inIPortuguese Pesqui (Atlterigotiavxigu Stcin) (NItiscide Diptera) oil cori) at Fast Java Agropecu Bras 17671-675 Indonesia Plant Prot Bill 21403413 70 Ferreira F 1F da Silva Martins and F 1PZimmermann 1979

50 Conklin H C 1957 IHaninoo agriculture a report on aln integral Rice varietal resistance to tle lesser corn stalk borer [in Port uguese] systent of shi fti rig cult ivationi in the Pltilippincs Food and Agriculture Pesqui Agropccu Biras 14317-321 Organization Rome Italy 209 p 71 Fleming W F 1968 Biological control of lie hapanese beetle U S

51 Cramer H II 1967 Plant proection and world crop production Dep Agric RLs Sers [ech Bull 1383 78 p Pilaniensclhutz-Nachr 201-524 72 Food and Agriculture Organiation of the United Nations 1982

52 l)ani R C and Majunidar 1978 Prelininary observations on tie Country report surmimary Pages 27-37 in Report on the first incidence of root aphids on differeit rice cultnrcs Sci Cult 4488-89 FAOIUN)IPhail and regional training coiuirse in iniproved ctltural

53 Davidson R I 1979 lffects of root feeding on foliage yield Pages practices for upland rice 5-23 Oct 1981 Northern Region Agrculturd117-120 in T K Ci osb and R B Pottinger eds Proceedings of the Developririt Center Chiang Mai Food and Agricuilture Organiiashy2nd Australian conlerence on grassland invertebrate ecology Govt tion Rome Italy 37 n Printer Wellington New Zealand 73 Franco do Ariaral S 195 1 Residual action of insecticides in turfy

54 l)ean G 1W 1976 Rice insect pests inlaos Int Rice Res Newsl soil--the results of a field experiment [in Portuguese] Biol6gico (Sao 1(2)15 Paulo) 17209-214 (Rev Appl Entomol A41423)

55 I)ean G J W 1978 Insect pests of rice in Laos PANS 24(3)280-289 74 Freeman ) 1970 Report onithe Iban london School of -coroiics 56 )ingle H 1984 Behavior genes and lifc histories conplex adapta- Monographs on Social Anthropology No 41 Athlane Press

tions to uncertain environments Pages 169-194 in A new ecology London UK 317 p novel approaches to interactive systems P W Price C N Slobod- 75 Fuchs T W1 A Harding and 1W Smithlr 1979 Induction and chikoff and W S Gaud cdsl ohn Wiley New York USA termination oldiapause in the sugarcane borer Ann Flitonlol Soc

57 Dobelmann-JP and M Falais 1968 Upland rice cultivation ill tie Am 72271-274 North-western part of Madagascar in 1966-1967 [in French English 76 (hbriel B1) 1975 A review tf the major insect pests of some tpland summary] Trop Agron 2313-21 crops in tlc Philippines Pages 86-111 in Ad Htoc panel of experts on

58 Doniiciano N L 1979 Chemical control of tie lesser corn stalk pest disease arnd weed problems in some rainfed crops Bangkok (lolsonno~paloIgnowslhts) (Zeller 1848) (I-epid(ptera Ihycitid ae) Thailand Food aid Agriculture Organization Roire Italy

on rice in Ponta Gressa Parara 1977-1978 [in Portuguese] Pages 77 Garg D K and N K Shal 1983 White grub outbreak on rainfed 291-303 in Anais 1 Reuniao de tecnicos cmiirizicultura do estado de dryland rice in Uttar Pradeslh Int Rice Res Ncwsl 81(4)16-17 Sao Paulo-Campinas Cati Brazil 78 Garg I) K and IP Tandon 1982 Major insect pests of rice on hilly

59 Dunsmore 1 R 1970 Investigations on tlie varieties pests and tracts of Uttar Pradesh India lilt Rice Res Newsl 7(11)11-12diseases of upland rice (hill padi) in Sarawak Malaysia Int Rice 79 Garrity 1) P 1983 Upland rice research an international bibliograpy Comm Newsl 1929-35 1975-1982 Iternational Rice Research hnstitnte PO Box 933

60 Elias 1R1965 The new decline of rice seedlings is not caused by Manila Philippines 121 p parasites [in Portuguese] Biol6gico (Sao Paulo) 313-6 80 Ghauri M S K 1982 A new subspecies of Iinorh)rone bull

61 Empresa Brasiliera de Pesquisa Agropecuaria 1984 Upland rice itt cornlus Slater (Hemiptera Iygaeidae) front Papua New Guinea on Brazil Pages 121-134 it Av overview of upland rice research rice and carpet grass Ihll lEntoniol Res 72133-137 Proceedings ofthe 1982 Blouak Ivory Coast Upland rice workshop 81 Gonzales F 0 PArregoces R I ihernandef and OT Parada International Rice Research Institute PO Box 933 Manila 1983 Insect and imite pests of rice arid their control in Latin America Philippines [in Spanish] la Federacion Nacional de Arrocerts (Fedearroz)

Bogota Colombia 60 n

38 IRPS No 123 January 1987

82 Grist D H and R J A W Lever 1969 Pests of rice Longmans London UK520 p

83 Hale P R 1979 Two subterranean pests of upland rice in Papua New Guirnca IntRice Res Newsl 4(2)16-17

84 Hale P R and J T Hale 1975 Insect pests of rice in the New Guinea Islands Rice Entomol Newsl 34

85 Harris W V 1969 Termites as pests of crops and trees Common-wealth Institute of Entomology London UK 41 p

86 Hattori 1 1968 A method to distinguish between grass leaf roller (Ctaplha rocis medinalisGuenee) and the rice leaf roller (Susurtia exigua Butler) [in Japanese] Plant Prot (Japan) 23167-170

87 Henry I E and 1A Onsager 1984 Experimental control of the mormon cricket Iahrosshnph by NJsena locustae(Aficrospora Microsporidae) a protozoan parasite of grasshoppers (Ort Acri-didae) Entomophaga 27197-201

88 Hidaka 1 P Vungsilabulr and SKadkao 1974 Studies on ecology and control of the rice gall midge in Thailand Tech Bull 6 TropicalAgricultural Research Center Tokyo Japan 113 p

89 Hincklcy A 1) 1963 Ecology and control of rice planthoppers in Fiji Bull Entomrol Res 53467 481

90 Hirao J 1982 Ecology and chtInical control of the rice leaffolder Jpn Pestic Inf 4114-17

91 Ho DT 1984 Sten borer incifience in rice ecosystems in KenyaEast Africa Pages 307-320 in A overview of upland rice research Proceedings of the 1982 Bouac Ivory Coast upland rice workshop International Rice Research Institute PO Box 933 Manila Philippines

92 Ho )T J 1Njokah and J GKibuka 1983 Studies on rice sten borers in Kenya with emphasis on ifaliarpiaipiraellRag Insect Sci Appl 465-73

93 Ichihara I T Ito A NIatsuo and M Sawada 1971 Chemical control of the rice striped stem borer in upland rice grown in plastic mulch [iii Japanese] Proc Kanto-Tosan Plant Prot Soc 1882

94 IclhiharaIand A Matsuo 1969 Damage caused by rice striped stett borer to upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1686-87

95 Ichhara I atid A MIatsun11970 Conside ration oft lie extraordinary prevalence of rice striped stem borer in upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1778-79

96 Institut de Recherches Agrolnliqes Iropicale et des Cultures Vivrieres 1978 Insect resistance Pages 12-13 in Upland rice 1978 atnual report Bouak Ivory Coast 27 p

97 International Institute for Tropical Agriculture 1983 Aphids itt rice roots I ITA Res Briefs 4(1-3)4-5

98 International Institite fi Tropical Agriculture 1983 Hopperburn in rice caused by brown plathopper IITA Res Briefs 4(l-3)6-7

99 International Rice Research Institute 1975 Annual report for 1974 PO Box 933 Manila Philippines p 216

100 International Rice Restarch Institute 1976 Annual repiort for 1975 PO Box 933 Manila Philippines p 228

101 International Rice Research Institute 1979 Cropping systems program Component technology development and evaluation Page 404 i Annual report for 1978 11O Box 933 Manila Philippines

102 International Rice Research Institute Department of Entomology 1981) Insecticid evaluation 1979 Tables 73 74 PO Box 933Manila Philipes

103 International Rice Research Institute Department of Entonmology 1981 Insecticide evalomtion 1980 Table 81 POBox 933 Manila Philippines

104 International Rice Research Institute 1981 Report of a meeting of Asian rice-based cropping systems entomologists Eleventh cropping systems working group meeting 18-22 May 1981 PO Box 933 Manila Philippines p 132 133 144

105 International Rice Research Institute 1984 Cropping systems program Pages 368-373 in Annual report for 1983 PO Box 933 Manila Philippines

106 International Rice Research Institutc 1984 Upland rice in Asia Pages 45-68 in An overview of upland rice research Proceedings of the 1982 Bouakamp Ivory Coast upland rice workiiop loternational Rice Research Institute 11O Box 933 Manila Philippines

107 Irshad M 1977 Grasshoppers in rice in Pakistan lit Rice Res Newsl 2(3)8-9

108 Ito R and TOno 1964 Upland lice its culture and varieties in Japan lIternational Rice Cormmission loth Meeting of the Working Party on Rice Production and Protection 3-10 March 1964 Manila Philippines Food and Agriculture Organization Rome Italy 12 p

109 Jeffrey M I and V A Dyck 1983 Effect of moonlight on lightshytrap catches of brown planthopper lit Rice Res Newsl 8(6) 13

110 Jennings P R and T Alicia Pineda 1983 Management ofthe rice plnthoppers Sogatodsorivi(coIl in Latin America Presented at the meeting on stable plant resistance FAO and lIndocsian I)inectorate of Food Crops Protection Bali lndonesia 29 Novcniber-2 l)ccemshyber 9 p

IllJimenez Gomez J A 1979 Rice insect pests i lla ios ()riCntales lil Spanish] Pages 99-103 in Rice ciltivation Instituto (olotibiatto Agropecuario Compndio No 29

112 Jongeleen F JJ Soesanto and A Mockidjo 1979 Studies oti pathogens of two species of Scarabacidac froinIndonesia Z Angew Enlomnol 87412-417

113 Jouvenaz 1) PC S lofgren and W A Baiks 1981 Biological control of imported fire antsa review of cirrent knowledge Bull Entomol Soc Am 27203-208

114 Kainoh Y K Shiniiu S MarUani Y Taiaki 1980 lost finding behavior of tie rice bu lplotori+JchinvittjI)allas (liciniptera Corcidac) with special reference to diet pattern ol aggregation and feeding on the rice plant Appl Entomol Zool 15225-233

115 Kakkar K 0 P Sharma and P I Sharma 1977 Note on Ihteronyrchus liodresRedt (Coleopicra I)ynastidae) as asporadic pest of paddy and maie inHimachal Pradesh Corr Agric 156-57

116 Kalode M131 79 Major insect pest problems and their control in upland rice (Unpublishcd mineogriph CWI oJNaaiat lni Chandaratat (198 1)original not seen)

117 Kalshoven L i 98 I The pests of crops in I ldoniesia (translated by P A vatn der Laan) 1 T Ichtiar B3aru-vati Iloeve lakarta Indonesia 701 p

118 Kamal N Q 1981 Suppression of whitebacked planto ner Sogatella fircaifra (Horvath) aLId rice lealfolder (nhal -cis licdb lis (Guenee) populations by natural cnclmie hlI) tiesis Grcgorio Araneta University Foundation Victoncta Park Metto Manila Philippines 179 Ip

119 Karim A N M Reaul 1975 A species ofliealy bug Trioni i70s ceres new oii rice in the Philippines Rice Entoniol Newsl 247

120 Katanyukul Wand C Chandaratat 1981 Insect pestsof uplaid rice and their control Report firstFAOUNlFi Tliailand training course on improved cultural practices for tplad rice Chiang Mai Thailand 5-23 Oct 1981 Food and Agriculture Organiation Rome Italy 22 p

121 Kern F 1955 lhletwioa(I)uninemts) bhhit Biturm(Coleoptera) a new pest of sugarcane inVenezuela [inSpanish with Etnglish sunmary] Agron Trop (Maracay) 527-29 (Rev Appl Eitoniol A4572)

122 Khan K M and S Ghai 1974 White grubs ald their control in India Pesticides (1Bombay) 819-25

123 Kiritani K 1975 Integrated pest management of rice pests Pages 66-70 itt Proceedings platning workshop ott cooperative hIeld research in pest managementP SMotooka ed East-West Center Honololu Hawaii USA

124 Kisimoto R 1967 Ecology of insect vectors forecasting and chemical control Pages 243-255 in The virus diseases of tle rice plant Proceedings of a symposium atthe International Rice Research Institute April 1967 The Johns Hopkins Press altimore Maryshyland USA

125 Kisimoto R and V A Dyck 1976 Climate and rice insects Pages 367-391 in Climate and rice International Rice Research Institute PO Box 933 Manila Philippines

126 Kureha S S Kobayashi SAsari and T Ishihara 1977 Life history and control of the mole cricket Grrllotalpa africana Palisot de Beauvois [in Japanese English translation 7520 1RRI library] Proc Kanto-Tosan Plant Prot Soc 21192-197

127 Kushwaha K S 1981 The black beetle as a pest of rice in Haryana India Int Rice Res Newsl 6(4)14

128 Lawani S M 1982 A review of tile effects of various agronomic practices on cereal stem borers Trop Pest Manage 28266-276

129 Leuck D B 1966 Biology of the lesser cornstalk borer in South Georgia J Econ Entomol 59 797-801

130 lever R J A W 1955 Insect pests of the rice plant in Malaya Int Rice Comm Newsl 1612-21

131 Li Ching-Sing and Shin-Foon Chiu 1951 The study ofthe rice gall midge Iachydiplosis orvza Wood Mason [in Chinese with English summary] 1Taiwan Agric Res 21-13

132 Iitsinger J A 1984 Assesament of the need-based application for rice Proceedings TechnologyTransfer Workshop 15-16 March 1984 International Rice Research Institute 1PO Box 933 Manila Philippines 6 p

133 Litsinger J A 1984 Environmental diversity and insect pest abundance with reference to the Pacific Pages 285-307 in Proceedings sub-regional training course on methods of controlling diseases insects and other pests of plants in the South Pacific 4-20 Oct 1982 Vaini Kingdom of Tonga Nukualofa

134 Litsinger J A RF Apostol and M BObusan 1983 White grub Leucopholis irrorata (Coleoptera Scarabaeidae) pest status popula-tion dynamics and chemical conllol in arice-maize cropping pattern in the Philippines J Econ Entoinol 761133-1138

135 Litsinger J A C de la Cruz F Raymundo A T Barrion M D Lumaban J P Bandong M S Venugopal F Paragna and E Balete 1982 Insect pests and insecticide reponse of dry-seeded bunded rice Pages 179-193 in Report of workshop on cropping systems research in Asia International Rice Research Institute Los Bafos Philippine

136 Litsinger J A M 1) Luinaban J P Bandong A T Barrion W Katanyukul and N Panda 1978 Insect pest management in rainfcd lowland rice agroecosystms of tropical Asia International Rice Research Conference 17-21 April 1978 International Rice Research Institute Los Bafios Philippines

137 Litsinger J A and K Moody 1976 Integrated pest management in multiple cropping systems Pages 293-317 in R I Papendick P A Sanchez and G B Triplett eds Multiple cropping American Society of Agronomy Madison Wisconsin USA

138 Loevinsohn M E 1984 The ecology and control of rice pests in relation to the intensity and synchrony of cultivation Ph D thesis University of London 36) p

139 Loevinsohn M E J A Litsinger N Panda M A Austria R F Apostol C Andrion and J Bandong 1982 A basis for predicting rice insect pest abundance in cropping systems Paper presented at the 1982 International rice research conference on rice and cropping systems 19-23 April 1982 International Rice Research Institute Los Bafios Iaguna Philippines 25 p

140 Lopez A W 1930 Repot of the Entomology Department Pages 145-172 in Annual report of the Philippine Sugar Association 1929-30 Manila

141 Lopez AW 1931 Entomology Department Pages 227-266 in Annual report of the director or research 1930-31 Philippine Sugar Association Manila

142 Lopez A W 1932 Entomology Deparment Pages 252-279 in Annual report of the Philippine Sugar Association 1931-32 Manila

143 Lungibill P and G G Ainslee 1917 The lessercornstalk borer US Dep Agric Agric Res Serv Tech Bull 539 27 p

IRPS No 123 January 1987 39

144 McGuire J U Jr and BS Crandall 1967 Survey of insect pests and plant diseases ofselected food crops of Mexico Central America and Panama U SDepartment of Agriculture International Agriculshytural Development Service Agricultuial Researrh Service October 1967 11 p

145 McNeill Sand T R ESouthwood 1978 The role of nitrogen in the development of insectplant relations Pages 77-9 in lliochemical aspects of plant and animal coevolution J B Ilarbornu ed Academic Press London UK

146 Maebara H 1955 Spatial distribution of lastinstiarlanaeof purplish stem borer (Sesamia iqferens Walker) hibernating in an upland rice field Bull Kagoshima Univ Fac Agric 438-41

147 Mallick E H and N G Hajra 1977 Response of rice plant to defoliation treatments at different developmental stages Indian Agric 2151-56

148 Mansingh A 1971 Physiological classification of dormancies in insects Canl Entomol 103983-10)9

149 Martell C G 1974 Catalogue of plant feeding insects of Mexico [in Spanish] Fitofilo 27(69)1-176

150 Martinez 0 J 1982 Principal rice pests in Ilanos orientalis [in Spanish] P~ages 80-94 in Curso deact nalization de conocimientos en arroz Villaniencis Instituto (olonmbiano Agropecuario

151 Martins J F da Silva F Ferrcira A S Prabhu F 1 P Zimmermann 1980 Use of pesticides for the control of the main subterranean upland rice pests esqui Agropecu Bras 1553-u

152 Martins I F da Silva F Fe eira rod BdamSilveira Pinheiro I Q2 Damage simulation caused by defoliators of upland rice [in Portuguese with English summary] Pesqui Agropecu Bras 171113-1119

153 Martins 1F da Silva N(iyuen van Tan and 1Ida Silveira Pinheiro 1981 Stem borer resistance in upland rice associated with plant morphological characters [in Portuuce Iesqui Agropecu Bras 16187-192

154 Masaki S 1980 Sumnmer diapause Ann Rev Entomiol 251-25 155 Masuzawa T 11 Suwa and F Nakasoji 1983 l)ifkleences of

oviposition preferencc and survival rate of two skipper butterflies Parnara guttata and PIchlpidai iathiais (Iepidoptera HIesperiidac) on rice plant and cogon grass New Entomol 3247-56

156 Maurya D M and C P Vaish 1984 Upland rice in India Pages 69-92 in An overview oftpland rice research Proceedings ofthe 1982 Bouakamp Ivory Coast upland rice workshop International Rice Research Institute IO Box 933 Mnila Philippines

157 Menon M G R R N Katiyai S Kuar and S S Misra 1970 Termites attacking the paddy crop Sci Cult 36294-295

158 Miyahara Y and 1 Nishikubo 1967 Behavior of young nymphs of the red rice root aphid Iholahsipiun ri liabhdontinalisSasaki in relation to movement from the leaf to the roots of upland rice [in Japanese] Pioc Assoc Plant Prot Kyushu 1363-66

159 Mo ljoa Tjien 1952 White grub of upland rice (Holotrichiahelleri Brsk) [in Indonesian with English summary] Contrib Agric Res Stn Bogor No 13134 p

160 Mochida 0 and T Suryana 1979 The occurrence of the brown planthopper and ragged stunt on lowland and upland rice ill Central Java and logyakarta in tile late of January 1979 Entomnolgy Survey Report No 15 Central Research Institute of Agriculture (CRIA) Sukamandi Indonesia I February 1979 9 p

161Monet C 1979 Varietal resistance to upland rice pests [inI French] Pages 35-39 in Annual report 1977 Institut des Savanes Department des Cultures Vivrieres Bouak6 Ivory Coast

162 Monte 0 1942 A caterpillar ofrice-fields[in Portuguese] Biol6gico (Sao Paulo) 8161-163 (Rev Appl Entomol A31414)

163 Munroe I) D 1979 Pest management strategy for hill rice in four resettlement areas in Sarawak Malaysia Pages 89-102 i1 Guidelines for integrated control of rice insect pests Food and Agriculture Organization Plant Production and Protection Paper 14 Rome Italy 115 p

40 IRPS No 123 January 1987

164 Nakasuji F M Ishii and Jen-Zon Ito 1984 Rice skippers in Taiwan and their life histories ZZrC (Taipei) Tech Bull 82 10 p

165 Navas I) 1976 Fall armyworum in rice Pages 99-106 il Ecological animal control by habitat management Proc Tall Timbers Con-ference 28 Feb-I Mar 1974 Fall Timbers Res Stn Misc Publ 6 Tallahassee Horida USA

166 Nickel 1 1 1961 Biological control of rice stem borer A feasibility study Tcch Bull 2 International Rice Rcarch Institute PO Box 933 Manila Philippines IIl p

167 Nigan P M and R A Vernia 1985 Insect pests of uplaild rice in Uttar Pradesh Int Rice Res Newsl 10(6)22

168 Nilakhe S S 1985 -cological observations on spittle bugs with emphasis on their occirence in rice Pcsqtui Agropecu Bras 20407-414

169 Nixon G EJ 195 1 lie association of ats with aphids and coccids Commonwealth Iusti lcte of Entonology Iondon UK 36 p

170 Nurdin F an Ishak Manti 1985 Study oit seedling fly of upland rice Interlational Upland Rice Moitoring Tour Report 26 lebruary-2 MarLI Sukaranti West Sumatra

171 Oka 1 N 1979 Ciltural conttol of the brown planthopper Pages 357-369 in Brown planthopper threat to rice production in Asia International Rice Research Institutc 1p0 Box 933 Manila Philippines

172 Ohni M and A Vilnaii 1975 Rice insect pests of Italy biology of Sipha grcria (Kaltenbach) and its control [in Italian] Riso (Milan) 2459-71

173 Otanes F Q atnd 1 I Sison 194 1 Pests of rice Philipp J Agric 12211-259

174 Painter R I 1951 Insect icsistance in crop plants University Press of Kansits Lawrence Kansas USA 520 p

175 Pantua P C and 1A litsinger 1981 Arthropod abundance in continuous vs biannual rice cropping systeis Int Rice Res Ncwsl 6(2)20-21

176 Pathak M ) and V A l)yck 1975 Sttudics ott insect pests of upland rice Pages 186-191 in Major research in upland rice International Rice Research Institute PO Box 933 Ilanila Philippines

177 Patil B R B M IHasabe and 1) S Ajri 198 I Chemical control of white gritb lIditriihinhorraia Fabr intfesting groundntut paddy atnd sorghum Indian 1Entomol 43232-236

178 Pawar A 1) 1974 (ollection and identification of insect pests of rice rlerminal repirt lInternational Rice Research Institute PO Box 933 Manila Philippines 72 p (tnimco)

179 IerrajU A aid A S Reedy 1965 Observations ott diapausc itt tie steti borer ohrice Rice News Teller (lai-Mar)9-13

180 Pont A C 1972 A review of the Oriental species of theri ona Rondani (I)iptera Muscidac) of econtomic importance Pages 27-102 in Control of sorglium shoot fly M G lomtani and W R Young eds Oxford atd IH1 Ptthl Co New I)clhi India 324 p

181 Prasad S C 1PChand aid M 1PSingh 1983 Incidence of rice gall midge in the upland plateau region of Hihar Oryza 20235-236

182 Rai P S 198 0 Nisaga simpex damage to rice in the hill tracts of South India Int Rice kles News[ 5(I)12

183 Rathorc Y S 1969 Gonocplhahli dcprssunt Fabr as itpredator of (tilo zonwIlhis (Swinhoc) and other moths in field cage at Patnagar Uttar Pradesh Indian 1Entotnol 3127

184 Reed W 1983 Crop losses causcd by insect pists in the developing world Proceedings 10th Internatial Congress of Plant Protection lrighton UK 174-80

185 Reedy M V 1981 Ateactogasterinhens(Walker)a new root-grub pest of tipland paddy and inaize Curr Sci 501076

186 Reveno M A 1968 Catalogue of the insect pests of economic crops in Colombia [in Spaiish] Asociacion Latino Americana de Entomo-logia (AIAE) 129-33

187 Reyes G M 1957 Rice dwarf disease in the Philippines Food and Agriculture Organiiation Plant Prot Bull 617-19

188 Rice S E A A Grigarick and M 0 Way 1982 Effect of leaf and panicle feeding byarmyworm (Lepidoptera Noctuidae) larvae on rice grain yield Environ Entomol 7593-595

189 Risch S J 1) Andow and M A Altieri 1983 Agioccosystem diversity arnd pest control data tentative conclusions and nizw research dirctions Environ Entomol 12625-629

190 Rivera C T 1956 Three organic insecticides in thecontrol of insects affecting upland rice Philipp Agric 39465-472

191 Roffrey 1 1972 Locusta outbreaks in the Philippines Acrida 1177-188

192 Rojas R 1) 1941 The effect of pruning at varying ages upon the yield of Ramairio Phipp Agric 29851-860

193 Roonwal M i 1979 Ternite life and termite control in tropical South Asia Scientilic Pnbl odhpr India 139 p

194 Rosetto C1 F da Silva Martins N C Schmidt and L EAzzini 1978 Damage caused by pasturespittle bugs 4Dvoi)lavopictaandDJ schach) on rice lBragantia Nota 6 (Matcli)25-27

195 Rosztto C1- S S Neto I- Amantc I) Iink 1Garcia-Vieira D M de SoLza N V Bansa t and A M de Olivcira 1971 Rice pests of Brazil [in Portuguese] Pages 149-238 inrFA()-International Rice Commission 2nd meeting of the Rice Committee for the Americas 6-11 l)ecember 1971 Pelotas Rio Grande do Sul Brazil

196 Rothschild G H L 1969 Obseratiois on the armyworm Spodoptera matritia acronrctoiiles Gn (Lep Noctuidae) Bull Entomol Res 59143-160

197 Rothschild G 1- L 1974 Parasites of the armworm Spolopiera mauritiaacroncitoihes in Malaysia Fntornopliaga 19293-299

198 Sachan i N S K Gangwar and 1N Katujar 1980 Field crickets damaging cages in North Eastern IHill Region Indian J Entomol 42526-527

199 Sands 1) P A 1977 The biology and ecology of Lepocorisa (Hemiptera Alydidac) in Papua New Guinea Res Bull 18 Departshyment of Primary Industry Port Moresby Papua New Guinea 104 p

200 Sarawak 1968 titomology division Pages 97-106 in Report of the research branch of the Department of Altriculture 1966 Kuching (Rev AIppl Eintomol A56603)

201 Satpathy JM M S Das and K Naik 1977 Effect of multiple and mixed cropping on the incidence of scine important pestsI Entomol Res 178-85

202 Sauer IHF G 1939 Notes Ott Elasmpnlpualtsligniosellths Zeller (Lep Pyr) a serious pest ofcereals in Sao Paolo [in Portuguese] Arq Inst Biol 10199-206 (Rev Appl Entotnol A28378)

203 Satinders 1L A 1 SKing and C L Vargas S 1983 Crop pests of Central Anerica with t bibliography [in Spanish] Centro Agroshynotnico Tropical de Investigacion y Ensenanza Turrialba Costa Rica (CATIE) Tec Bol 9 90 p

204 Sauphanor B 1985 Some factors of upland rice tolerttce to stem borers in West Africa Inseu Sci Appl 6429434

205 Sen P atnd S Chakravorty 1976 Stentflics in Haritighata Nadia West Bengal India [it Rice Res Newsl 1(12) 15

206 Senapai B and 1M Satpathy 1983 The seedling fly Atherigona orrzae Malloch is a pest of rice in Orissa (India) J Entomol Res 777-79

207 Shhjahan M S Alatn M Ah nad and BN Islam 1983 A new scarabaeid beetle pest of dryland rice in Bangladesh Int Rice Res Ncwsl 8(I)16-17

208 Sharma G L S C Modgol and R C Gautam 1979 Effect of depth of submergence ott incidence of bacterial blight and white grub infestation in transplanted rice Int Rice Res Newsl 4(6)12-13

209 Sharma S 1985 Scarabaeid beetle outbreak itt upland rice Int Rice Res Newsl 1(1(l)24

210 Shiraki T 1917 The paddy borer Schoenobhits incertulas Walker [achaku Agric Exp Stn Formosa 256 p

211 Silva A de Brito and 11P Magalhaes 1981 Insect pests of rice [in Portuguese] Circular Tecnica No 22 EMBRAPA Centro de Pesquisa t ecuaria de Tropico Unido (CPAJU) Belem Brazil 14 p

212 Singh S A 1979 Emergence of rice leaf butterfly as a rice pest in Manipur India int Rice Res Newsl 4(4)16-17

213 SinghS A 1983 Changing trends of insect pestsof rice in Manipur Int Rice Res Newsl 8(f)12

214 Siwi S S and P H van Doesburg 1984 Lenovorisa Latreille in Indonesia (Heteroptera Coreidae Alydinae) Zool Meded (Leiden) 58117-129

215 Soejitno J 1977 Reaction of some upland rice varieties toseedling fly eltherikonaexigua Stein [in Indonesian] Siri Hana I Penyakit 630-35 Lembga I osat Fenclilian Pertanian Bogor Indonesia

216 Socnardi 1967 Insect pests of rice in Indonesia Pages675-683 in The major inse pests of thu rice plant Proceedings of a symposium at the Internatital Rice Rec Wi itt S pternber 1964 The Johns Hopkins Press Baltimore Maryland USA

217 Soto P E A T Petcz and I W Buddenhagen 1976 Survey of insect pests and diseases of rice in different ecological zones in Nigeria Rice Entomol Ncwsl 435-46

218 Soto 11E and Z Ziddiqi 1976 Screening for resistance to African rice insect pests Varietal improsetnent sCmi n r Bouak Ivory Coast 13-17 Sept 1976 West Africa Rice l)evelopment Association Monrovia Iiberia

219 Squire F A 1972 Entomological problems in Bolivia PANS (PestArtic News Sumr) 18249-268

220 Stapely 1 II 1978 The rice leaf roller in the Solomon Islands Int Rice Res Newsl 3(l)13

221 Suenaga H 1937 On aphids attacking the roots of rice Oyo-I)oubts Zasshi 9150-152 [in Japanese] (Rev Appl Entomol A25745)

222 Suenaga H 1952 On the seasonal abundance of three subterranean aphids on upland rice Bull Kyushu Agric Exp Sti 1249-262

223 Sutynso Yas and Dandi Soekaria 1964 Beberapa hana yang ttenperang tana man padi tanah kering [in Indonesian] Rapat kerja Padi Tanah Kering 6 sd 8 Juily 1964 Kaliurang 9 p

224 Tallamy I) W and R F Denno 1983 Alternative life history patterns in risky environments in example from lace bugs Pages 129-147 in Insect life history patterns R F l)cnio and If Dingle eds Spring-Verlag 225 p

225 Tamura 1 1939 Observations on some insect pests of field crops [in Japanese] 1 Plant Prot 26414-420 (Rev Appl EntomolA27-556)

226 Tamura and Suzuki 1964 Influences of the early-transplanting practice in rice culture upon development of the rice stem borer Chilo siql)ressalisWalker Bull Ilokuriku Natil Agric Exp Stn 761-94

227 Tan J P 1924 The rice root aphid Drowhi ihirsoia Philipp Agric 13277-288

228 Tanaka 1 1957 Taxonomy and distribution of some subterranean aphids injurious to upland rice in Japan with description of a new species [in Japanese English summary] Botyu-Kagaku 22168-176

229 Tanaka T 1961 The rice root aphids their ecology and control Spec [full College of Agriculture Utsunomiya University Japan F3 p

230 Tauber M IC A Tauber R Nechols and 1Obrycki 1983 Seasonal activity of parasitoids control by external internal and genetic factors Pages 87-108 itt I)iapause and life cycle strategies in insects V K Brown and I Ilodek eds )r W Junk The Hague Netherlands

231 Taylor W F 1972 Effects of artificial defoliation (simulating pest damage) on varieties of upland rice Exp Agric 879-83

232 feotia T P S and S Natud 1966 Biotoomics of the rice skipper Parnaraniathiis Fabricius (Lepidoptera Hesperiidac) Indian J Entomol 28181-186

233 Teran F 0 1971 Rice pests of Santa Cruz [in Spanish Ministerio de Agricultura Sita Cruz Bolivia Bull 45 8 p

234 Than Itun 1976 Population dynamics of the yellow rice borer Tryporza itcertulas (Walker) and its damage to the rice plant MS thesis University of the Philippines at Ios Bafios Laguna Philippines 92 p

235 Uichanco L B 1911 Pests Pages 55-74 it Rice Philipp Agric 11-200

IRPS No 123 January 1987 41

236 van der Laan P A 195 1 Life history of the rice seedling fly Contrib Gen Agric Res Stn Boor Indonesia 118 15 p

237 van der Laan 11A 1959 Correlation between rainfall in the dry season and the occurrence of the white rice borer (Scirpolhaga innotala WIk) in lava Entomol Fxp Appl 212-20

238 Van Driesche R G 1983 Meaning of percent parasitismin studies of insect parasitism Fnviron Entomol 121 I1- 122

239 Venugopal M S and A litsingcr 1984 Fffect ofcarbofuran on rice growth Prot Ecol 7313-317

240 Vietnam Directorate of Rural Affairs 1962 Annual work progress report on crop improvement program of rice sugarcane vegetable and field crops (lor the period from July 1961 to June 1962) (11) 299 p (Rev Appl Entomol A51526-527)

241 Wan M T K 1972 Observations on rice leaf- and planthoppers in Sarawak (Malaysian Borneo) Malay Agric 148308-335

242 West Africa Rice I)esclopnient Association 1978 Annual research report Vol 2 Statistics Monrovia Liberia

243 West Africa Rice Development Association 1979 Annual research report p47 Monrovia Liberia

244 Williams ) F and CS lofgren 1981 Fli I illy l-468 anew bait toxicant for control of the red imported lire at Fla Entomol 64472477

245 Williams I) 1 1970 The mealybugs OFlomoptera Coccoidea Iscudococcidac) of sugar-cane rice and sorghum Bull Entomol Res 0109-188

246 Williams D) 1 1982 Pulinaria icervi (Signoret) and its allies ott sugar cane and other grasses Btll Entoinol Res 72111-117

247 Wirjosuiardjo S 1976 Study ofchenmical cont rot of insect pests ott upland rice Imiu Pertanian (Jogjakarta) 1323-329

248 Wirjosuhardjo S and N Wirarta 1976 Chemical control of the white grub tIolorithiatlltriin upland rice in M uneng Irobolinggo Indonesia) [in Indonesian] Imut Pertanian (Jogjakarta) 1330-332

249 Yano K T Miyake and V F Eastop 1983 The biology and economic importance of rice aphids (Henuiptera Aphid idae) a review Bull Entomol Res 73539-566

250 Yelp D and P Rhodes 19(4 lhe success of drilled dryland rice iii the Fiji Islands International Rice Commission Working Party on Rice Production and Protection Manila Philippines 3-10 March1964 Food and Agriculture Organization Rome Italy 13 p

251 Young W R 1981 Fifty-five years of research on the sorghum shootfly Insect Sci Appi 23-9

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ISSN 05-3862Other papers in this series TITLES OF NUMBERS 1-45 ARE LISTED ON THE LAST PAGE OF NO46 THOSE OF NUMBERS 46-70 ARE ON THE LAST PAGE OF NO 71-80AND THOSE OF NUMBERS 71-1O0ARE ON

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No 101The economics of hybrid rice production in (hina No 113 Boron toxicity in rice No 102 Rice ratooning No 114 Energy analysis rice production systems and rice research No 103 Growth and development of the (Jeep water rice plant No 115 Production risk and optimal fertilizer rates an application of the

No 104 Faridpur acomputer-assisted instruction nmodel forrainfed lowland random coefficient model No 116 Consumer demand for rice grain quality in Thailand Indonesia andrice

No 105 A readingand listening comprehcnsion test inEnglish for non native the Philippines speakers applying for tiraining at IRIRI No 117 Morphological changes in rice panicle development a review of

No 106 Rice grassy stunt virus 2 a new strain of rice grassy stunt in tire literature

Philippines No 118 IRRI-Korca collaborative project for the development of cold-No 107 Physical losses and quality deterioration in rice plstprodnCtion tolerant lines through anthei culture

systemis No 119 Problen soils as potenial areas for adverse soils-tolerant rice No 108 (olptblication ofIRRI materials a survey of translators and varieties in South and Ssuthcasi Asia

publishers No 120 Changes isinall-farm iLtireliing techii)hgy in Thailand and [he No 109 Classification of Philippine rainfall patterns Philippines No 110 Contributions of modern rice varieties to rutrition in Asia No 121 Landforms and modern rice varieties No Ill Changes in rice breeding in 10 Asian countries 1965-84 No 122 Yield stability and modern rice technology No 112 Design parameters affecting the performance of the IRRI-designed

axial-flow pump

Page 26: UPLANDRICE. INSECTl--1i M T -E!R.EOLOU t RTANCE ,AND …delphacid.s3.amazonaws.com › 6958.pdf · The significance of abiotic factors in upland rice insect ecology will be apparent

26 IRPS No 123 January 1987

and leaffolde rs However because of the need to temper fertilizer use on upland rice o prevent blast fIrtilizer rates will be lower than is comonia r for lowlad rice causing relatixely less hopper and IaIfTfoide r population buildup However inupland rice areas without a history of blast such Is Batangas farmers apply high rates of N and conscqtrently the crop has high infcstation levels of WBPI]and RIF The fhrmrcrs leec insect buildup by splitting the fertili11r in0to two to three applicatiois bit do not applV fertilizer at the vegetati estage because this prodrces a tall crop which illeasily lodge during typhoons Greater crop fertility leads to greater insect survival (better nutrition) larger insects (to cat no re foliage) arid greater fec titd t (to lav nmre eggs) (128 145 171) Not all insect species respondequally hovr Increasing rates (if N l anrid K increased Dialraa s-acchralis and (iaeiurema flea beetle but reduced thrips (66) Zn rcdLuced deadheart density from the lesser cornIstalk borer

Fnldingi Although most Upland rice areas do nIot hawxcaccess to wa ter floodiig the fields is rRrrnriierdcd in areas of Brazil or root bug and white gib control ( 195)

Crop rotatioti Tillage before planting a crop after rc harvest will control stern boreis and other pests renain n the stubble arid will unearth soil pests Planting a non-gratriiaceous crop after rice is recommncrded for termite control (66) Crop rotation is less nttdcd in upland rice culture than in lowlands because atcrop-free period is assured even after harvest of crops following rice

t1ulchilt iryl plastic irli irig designed for weedcoitrol ii upland rice inJapan resulted in ntr highanri liy infestation of (ihilo sulytresvais (93 9-1 95) NIulched Upland rice grew more luxuriantlv and attracted more (Avipositing mothis than u lched rice Also the rice plant and stein borers imatoured 10rC quickly and a partialgeneratiori af vouing ista iNoverwintcred in the stubbleLotili--malitrt varieti Quickly maturing crops reduce the tmbCr of pCst generations that Calt build up Rapid crop establishment olan ca rl v-natu ring vaitvty if)pa 11icLa r will lessen white gib and stem borer damiage White grub larvae intle last iistar are highly destIruCtive so aii early-iiatri ng variety will escape serious root loss Stem borers build 1ip slowlyx in rice to ecormic abundant in a late-Maturing crop anrd caiuse whi tlieads Numibers of somespecies increase exponeitially with each succeeding genera-ti so a xarietyexhat atures I trowh earier thian another will hac less insect damage Early-naturing vaiicties however will not lessen seed pest daiage either ii the soil or oina standing crop

Strwhoi hAIJttam Inrscts tcd irig on thlie aerial portions of rice plats disperse from carly to late plantedfields AS iiiibCirs increase exponentially with eachinsect

gtscnrattm e eiglbrig f rmc~s1(uil upp wh rc generation pests build tIP where neighboring fihrTC e9r1sstagger their plantings II Batangas farmers plant uplandrice within I iro use 120-d s-arieties and plant at the start of the rainy season Ini Misarnis Oriental laricrs stagger their

plantings up to 3 mo apart and use a set of atieties that mature in 4-7 mo Insect pest danage particularly from stem borers is greater in Misarmis Oriental Also seedling maggots damage rice there because rainfall favors frequent planting of maic In Batangas rice escapes seedling maggots bCca usC of distinct drs season and synchron0us early planting with the onset of ains

Therefore combinations of cultural practices ea rlyplanting synchronous planing crop rotation and earlyshymaturing vaneties protect the rice crop against most insect pests

Ilmant resistance Beca tse upland rice has few specialized insect pests and many insect types attack the crop worldwide regional breeding and strong internatiornal cooperation are required

Sown-sced pests arc economically controlled by insectishycide seed treatment and should the relore receive lowxpriori ty inbreeding objectives

Root pests are normally cost ly to control with insecticides but finding resista t sonres mayvbe di fficult Tanaka (229) for exa -ple fi led to find varieties resistant to root aphids Selecting for large root bioniass perhaps should be the strategy for pests sLuch as Whlite grubs that renliove roots Fainter(174) howcvcr cites varietiesof crops tolerant ofor resistant t(i root pests Some sorghun varieties are high in cyanide so rice sorghun might incorporate broadshyspcctruri chemical resistance

Seedling vigor and drought tolerance are high prioritybreeding objectives so tupland rices should have large r root systernsand be able to tolerate ligherlevels of root loss Sonie root loss inolder plants is beneficial if it stimulates new root developnrut and therefore crihances intake of soil nutrients (53) The rice seedling inaggot rtiLotia spp may be

controlled thirough resistant aittics A hi rgc breeding effort on sorghniu has had fair success inideveloping arreties esrstait to Asoccata Rodani (251) A field trial comparing 10 upland ricc varieties showed differences aniong varieties va ring from II to 4(- damaged tilleis (215) Hovver resistarce to 1Iill-0r gtia species may rot cross ox-rti otier

Shiraki (210) reported that upland i-ice stens wcrc harder than lows-land rice stemsperhaps becaIusc (f a higher silicoricontent and thus were more resistant to yellow stern borer hc tortia Iow tillering (f uplaid iice xarneties prodtres a

higher percentage of infested tillers than would occor in higher tillering lowland iice under a siiilar egg density The corollary therefore is to select for high tilleriig upland rices

Stemii bor amcstheh main targes dobrecders inAfrica (571 204 )but nticl oifth e sc reen irig is dotie unde r lo wland co d t osrm s wih A h r~ o l pe i s n acorditions (11 96 218) As with lthrinrnwspecies mayhave to be dealt with independently however initial results show cross resistance between African and Asian species(92)

In Brazil screening Ior insect resistance focttses oil lou I pest problems Varieties tolerant of lasmolwalpus ligi-sellus and resistant to )iatraea saccharalis have been identified (70 153) Current work also emphasies 7iraca limativetntris and Thmialspis (=Iosis) flavouicta (66) These insect pests Ilt onlV citise IIigh yield losses tihey also ire difficult to control chemically ofr by other ilealls

ManV upland rice vaieties aic early maturing and are resistant to pests such as gall midge (I Xl ) Early mIturity is often a highly desirable trait to esuilpe pest buildup but it is not true resista nce bca usc ilthose va reties are plinted late they will be itsdamaged as ain early planted susc ible variety

P~erhaps tile cosiiiopolitall of all upland rice pestsimost are the sced pests (ripenling stagc) which pose a particular problem beta use scuds re ised IsIO for hitnit4ans Ayltoxic substatice introd tcld geticticllyilmay also be toxic to nal iniless it caln he placed old in thfe IdhuffSelectmi fo hairiness or lotng wns kill mIlct sthilrohleiiis dutilig threshing pa rtictfarl maitial threshinfl [hireshers ill complain of itcfimin atid itteascd (fust levels

Attempts itIRRI to fitid tcistatncc to the rice bug plorisa have ilotbeCnl prmitisilg lIIetIattonlids iorc

through the lenilna or palca hillhairs sedstVIor seeds wilh awns that deter bugs promote complaints from threshers Seed bugs liptocorisa and Stcnocoris enter the seed through the opening between tile lemna and palC but varieties with narrow openings are diifficult to mill

Breeding priorities should concentrate ott inotCI gruLtLIps that are most difficult to c nttrol by other meains Steil) horers appear to be the most widespread group that sigiiificantly dalag s rice aind is hard to manage by other Ilnca us

Biological control Very fe stutdic of mitatt~l eneties of rice pests in upland envirollnletits wee in tile wefouiid literature Therefore have not compiled a list of species Ve present a list of natural enemies iOtid in upland ricefields tuic latialln Batangas for some conitno iOldiar pest species ([itble 5)

Muich ofl thc lollowinle itfor ittiol cotlliCs rol Stutdies in the lowlatnds or roi crops otithcr thlin tice

Qu(ailtr o 11ll t11t( Nat trrtf lii hcoeits ight Ile used agaitist ile social insects tilt and termites Sollie predator specice live ill theirf nests n ilnic theirnllt ip-pearanIlCe aMRidbehavior IJtrouMdCing a mote aggressise ant species to displace pest ant codonies is a possibility (169) Patihogens as potential natural encnies have reccived little atteition (I13) Natural ncnuics asa group however hac not been toIl iporta it in Iegulating aintshown be and ternite numbers

The nocturnal behavior aid subteitmien habitat of lole crickets and field crickets protect them from many ntllral enemies There are only limited records of natural cnelics for these groups of pests some pathogens (87) but mainly

IRPS No 123 JattuIny I 271987

parasites - scelionids on eggsand sphecids on ilymphs and adults Sphecid wasps Larra carutmaria (SImitI hI L huzwuensis Rohwer and lsaltiuhta Willi ams speciali in mole crickets pa ralyzing them nd dragging the ninto thcir nests as food lot their voting ()tier sphCcid species Lins aurulenoa (Iabricius) lot maiac (Ashmead ) 1 lhshylssellaus (Smith) and Ml lal)oriosu (Snithf specialie in field clickets

White grUbsO the other hand hae Icti scriuns enuigh to have been extensively studied for cotttroi 1W nati al enemies( 112 140 141) Scoliid parasites C(ajsoMcispp have beetn intrtduced itto tilePhilippites for control of Ielxl ]udisiro la The rcsearchcts w Intttrdced themle during ill era claimed that the white gttionthreak as controlled (142) howevcr it isnormal for pest epidemics to subside and there was no diect evidence to indictte that tfe parasite was responsible A tmore promisitg titctllod might he to identify iso ltcttass prt~f tcartd apply hiactelaln fingal pathogens to tile soil Ihe lalattese beetle in North America has hcci coitrolled hv tite conmnleciall aailable hactcrial prcpa ration itacihlpu pav ofr milksdiscas (71) Spote prtyefott the soil itloted ilrtier reitiain actic fr Veas

Root aphid sardtnctly bugs readilslic tilib to predatots and parlsites once their tetndine ants are controlled The larvac and adtilts of a coccinellid beetle 1Scynms sp prey on llriadlouais in slash and burn upland rice Baits treated with insecticide may be used to control tending ants by killing their ourig (244)

Root-tkcditig blugs false wireoris wireorms and root wccils aic r ofsuch little ifilpoltatice that thenomttll role of liatural Cticniics hias tiot beei asscssed A possibility hoevrl tiglt he tit ItaIIitic niaCIMtodcs lintt1g11en Itese soil-itlhlfbitine mCtitodcs W5uhtfhave to be imiass Produiced

large miltibcr of niatltrll cnemics ircknown ft CltIoriS IC IpblCeii is thtt clttvortnlls ofonlie the crop soott alter land ptel)tritiOti a1tithe bcgit iniig of the raittv

Citsoln hfien tiatllral leniiiy populations aric lo Colltloil WOtuld iii ole mass prodtticing andirclcaslng key spccies

le egg stage is the fpirt (olthe Cedlitgt Ialot life cycle tI hich to f)ctis ftor tittlril elctit cottntrol hplraites o1 pleditors ttsit elleggs highly expoIsYCl I Sgg pailsites or predators stolld probahly e ittass andtbe produced released Littlh i toruialioll is a ailable ott what species to trV o ho to Iniss podLice thiCIi A etiloplhid wasp Icmihlaol sp effctiicly regushy

lates the leatf lcloalapolm2a spioanoiiitr heat andc rice antid should be collsctvcd

leaf beetles whCllose e am] eggs ate laid in the grutindhirO ould be motC difficutlt to control by niatulral enelV Mnll ipuhltiOn But species sUici IS c_gsthispid beetles ith

and larVle oilleascs ate vtilcratble to iparaites and pledators

Natural enemisc of tlirips otl rice are little known

28 IRPS No 123 January 1987

Table S Parasites recorded from upland rice insect pests in Tanauan Batangas Philippines 1977-84

Order Family Species Host stage

I lymttenoptera Braconidae Leaffolders Cnaphalocrocis Marasinia spy

Cardiochilesphilipplinensis Ashmead a Larva

Cotesia angustillasis(Gahan)Cotesia or cypris (Nixon)

Larva Larva

Cotesia or taeniaticornis(Nixon) Larva Cilonusinotakatae Munakata Chelonits SppAlacrocetrts philippinensis (Ashtmead)

Egg to larva Egg to larva Larva

Iclineumonidac Orgilts spp Isclutojppa hireator(Fabricius)

Larva Larva

Itoplectis naraigav(Ashmcad) Larva Tenelhtcha philippiensis (Ashmead) Larva Tenelchha stangli (Ashmead) a

irichomma cnaphalocrosis Uchida Larva Larva

Bethylidac Elasmidac

Goniozms or trianguli r Kieffer lasmtusalhopictus Crawford

Larva Larva

Diptera

Encyrtidac Chalcididac

Tachinidac

Elasmtus spp Copidosomopsisnacohiae (Fidy) Brach nteria evcarinata Gahan Biachyieria lasus (Walker) Ztgohothria ciliata (Wulp)

Larva Egg to larva Pupa Pupa Larva

Argyrophylx nigrotihialis(Blaranov) Larva

l-ymenoptera Braconidac Yellow stern borer Scirpopiaga incertulas

Braco chinensisSzeplegeti Larva CIthelotis spp Chelonus niunakatae Munakata Cotesi flaripes Cameron Stenoracon nicerilli (Bingham) a

Larva Egg to larva Larva Larva

Ichneumonidac Tropobracon schoenobii (Viereck) Amatroinorpiaaccepta metathoracica(Ashmiead)Frihorus sinicus (I lolmgren)

Larva Larva Larva

Ischnojoppaluteator (Fabricius) Larva Isotima nr dammeratiti Rohwer Larva Tenelucha philippinensis (Ashmead) a

Larva Tenehicha stangli (Ashmcad)Trichomma enaphalocrosis Uchida

Larva Lara

Chalcididac Eulophidac Trichogranmmatidae

Xanthopimnpla stemmator (Thunberg) Charops hrachYpterumn Gupta and Malteswary Brachtm eria sppTetrasticttus schoenohii Ferriere Trichorigranm chihunis Ishi

Larva to pupa Larva Pupa Egg Egg

Diptera

Scelionidac Pteromalidae Tachinidae

Tt ichograintnta aponiicuitt (Ashniead) Telenomits rowani (Gahan) a

Trichoinalopsis apattelctena (Crawford) Jeirhaea spp

Egg Fgg Egg pupa Larva

Z4gobothria ciliata (Wulp) Larva

Ilymenoptera lecneumonirlac Gol fringed borer (hilo auricilius

Trichomm enoptalocrosisUchida Larva

Ieioenoptera Braconidae

Aantholimlh steiniator (Thtinberg) a

Pink stet Ioret Sesamia inferens StoInacoil nicrilhi (Bingh am)

Loarva to pupa

larva

Diptera Ilyinenoptera

Armvtnyorms Mythimna separata Spodoptera mauritia acronyetoidesTachinidae Zvgohothriaatropiora( Robinea i -Desvoid y)Eulophidae It-phctrtiscitapadoe (Ashimead)

Larva Larva

(halcididae Braconidac

Brachpin ria spip (oesia spp

Larva larva

IHyntenoptera I)iptera

Rice skipper Ielop ida (halcididae Tachinidae

math ias aidgreen iortedci ciliarNIelan itis eda Brat lhincria sp iii matuqitatt Cameron Irgyrolhirlaxi trotlhiais(marumtv)a1

ismtne Larva Larva

Diptera Ilymenoptera

Tachnidae lelineutionidae

fuown se tilo per Mois frugalisIrgtrophrlax 1igrothiahiv 3 a rat ov) anthopimpa pmnctata (Iabricius)

Larva Larva to pupa

aMost dominant

IRPS No 123 January 1987 29

Normally predatory thrips prey on herbivorous thrips strepsiptera on both Spiders dominate tilenymphaladultNatural enemies of stem bugs --Scolinoplara and Tibraa predators Spittle bugs move to rice from pasture grasses - are in order of importance egg par~isites and predators and biocontrol would be successful only ifcarried out before fungal pathogens and nymphal adult parasites Egg tiley immigrate into ricefields parasites could either be introduced fron other areas or Biological control tactics for hemipterous seed bugs mass produced and released ould follow those outlined for stein bugs

Armyworms are normally lwld in hcck by the activities Coml)arison of environnewtns Natural enemies are of egg and larval parasites When these natural enemies fail perhaps faced with even greaterconstraints than insect pestsusually because of drought armyworns become epidemic to survive ind reproduce in upland environments TheyIt may be worthwhile therefore to release parasites during have adapted ani survived by having wide host ranges the the rainy months following a drought Virus diseases of ability to aestivate or hibernate over unfavorable seasonslarvae would beanotheravnue to exploreforarnyworms and the ability to disperse or by other mechanisms

Parasites have ben traditionallyconsidered for biolegical Comparing the most dominant and therefore the most control attempts against stein borers usually by rekasing adapted species of natural enemies collected in various exotic species( 166) This approach has not met with success environments in the Philippines we can understand how and new avenues should be explored The role of predators different rice environments are arid pathogens is little inderstood and deserves greater Anagqrms olptahiisa mymarid planthopper egg parasiteattention Egg parasitism rates are normally reasonably is cquallyadapted to all Philippine rice environments (Tablehigh but should be supplemented with effective egg 6) It is selective for planthoppers and even parasitizes eggspredators Orthopterans are potential egg predators of the n-iize planthopper Peregrinitsmaidis itt upland areas Metioche vittaticollis (Sta ) Anavipha nifiwnnis (Saus- Alavohis also parasitizes maize planthopper eggs but is sure) and Conocephahs hongipnni (de i-laan) feed on most prevalent on BPH in the irrigated wetlands (onashystem borer eggs etioche and Anaxil)ha specialize in eggs tocerus spp like A optalhilis on BPH and WBPI Iattacks with no hair such as Chilo spp The omnivorous (ono- the eggs of GLH in ell rie environments It is also found cephahsspecializes in eggs cov red with hairs it is a proven parasitizing the eggs ef the white rice licahopper (oimna egg predator of Scirlouphaga spp but has the discouraging spetra Oirosita naias a trichogrammatid egg parasitehabit of eating rice guin (65) attacks BIPH and WBPH in the wetlands and has no other

Grasshoppers have egg (scelionid) and nymphalad ult host A related species 0 aesopi specializes in G1-H in (nemestrinid tachinid and sarcophagid) v irasites Among wetland environments Dryinid strepsipt-an and pipunshypathogens protozoans have been recorded most frequently culid nymphal and adult parasites of planthoppers and Control of locust species by natural enemies could con- leafhoppers also specialize either in hopper species or centrate on their habitual breeding areas environments Among the dryinids Pseuhdgonatoipsflavi-

Leaffolders have rich complexes of natural enemies some feimur prefers the lowlands and BPH while P nuch has no adapted to upland rice (22 27) ranging from egg predators environmental prefirences but is adapted only to WBPH (gryllids coccinellids) to larval parasites (braconids ichncu- The main d ryinid species on GLH aptogonatop)usspp is monids) to larval prcdators (ants and carabids) to larval most dominant in rainfed lowlands and also attacks other pathogeiis (vira and fungal diseases) Efforts to augment leafhoppers within Amnrasca Cicadulinaand Bachaltuathese natural enemies should focus on the egg predators Strepsiptera are more dominant on plarithoppers than (rearing releasing)and viral or fungal pathogens (culturing lealhoppers The Elenchus species in the Philippines attack spraying) all species of planthoppers within the genera Sogawilla

Parasitization of the large conspicuous larvae of Sogatodels NilaparsataHarualia and Opiconsiva Elenshyskippers green horned caterpillars and other polyphagous cusrvasuntaisuiis adapted to the major rice environments Lepidoptera by tachinids and chalcids is normally low ELg but prefers BPH in rainfed arid WBPIH in wetland environshyparasites and predators are perhaps the key to their control ments Ilalictoph(utsmtnroeiand HVpectrus attack G LH

Gall midges are parasitized by pterormalids and platy- and COfimravcctra respectivelygasterids that aestivate within their larval host between Pipunculids are mainly parasitic oii lealhoppers Each seasons Shoot aphids are preyed upon by coccinellids species has a unique environmental preference Tomosshysyrphids and chrysopids varireIa oraetoraprefers Ne7 otetltix nigropictrs and

Planthoppers arid leafhoppers have egg parasites and Pilunwulhtsjavanensisattacks Dehoceplhahls spp which predators as well as nymphialad nlt parasitesand predators are more abundant in upland than irrigated lowland their rich complex of natural enemies includes fungal environments T suhvirescensattacks Nephotettix virescens pathogens The principal egg parasites are myrnarids and and prefers the lowlands Pilpuiulus nttillattls is an trichogramniatids Egg predators are norinally mirid plant upland rice parasite arid attacks only Nephotetix speciesbugs that suck out the yolk Nymphal ad nIt parasites are More than 17 recorded species of leaffolder parasites in dryinids on planthoppers pipunculids on lealhoppers and the Philippines attack species of C(naplwocrocis and

30 IRPS No 123 January 1987

Table 6 Dominant natural enemies of rice pests in upland rainfed lowland and irrigated lowland rice environments of the Philippines

Natural enemy

Host stage Species

Brown planthop-Parasite Egg

MymaridaeAnagrus optabilis(Perkins) Anagrusflaveolus Waterhouse

TrichogrammatidacOligosita naiasGirault

NymphAdult Dryinidac

Pseudogotatopuslaiifemttr Esaki et Hashimoto Elenchidae

Elenchus yasurnatsui Kifune and Hirashima

Environment

Upand Rainfed lowland Irrigated lowland

Ailaparata lugens

x x X x

x

x x

x X X Whitebacked planthoppers Sogatella furcifera and Sogatodes pusanusEgg

MymaridaeAnagrus optabilis(Perkins)

TrichogrammatidacOl~gositanaias Girault

NymphAdult Dryinidae

Pseudogonatopusnudus Perkins Elenchidae

Elenchus yasutatsui Kifune et Hirashima

Parasite Egg

MymaridaeGonatocerus spp

TrichogrammatidaeOligositaaesopi Girault

NymphAdult Pipunculidac

Tomosvaryella subviresccns Loew Tomosvaryella oryzaetora (Koizumi) Pipunclusmutillatus (de Meijere)

DryinidaeIlaplogonatopusspp

Egg Copidosomopsisnacoleiae (Eady)

Larva BraLonidae

Cardiochiles philipphnensisAshnead lchneumonidac

Trichiomnia cnaphalocrosisUchida Temelucha stangli (Ashmead)

Pupa Chalcididae

BrachymeriaexcarinataGahan Brachymerialasus (Walker) Brachymeria nr tachardiaegroup

Parasite Egg

Scelionidae Telenomus rowani (Gahan)

EulophidacTettasticusschoenobhi Ferriere

Larva Braconidae

Stenobraconnicevillei (Bingham) Iclhncurnonidac

Temelucha phiiippinensis(Ashinead)Temelucha stangli (Ashmead)

x x x

x x

X x X

x x Green leafhopper Nephotettix virescens

X x x

x x

X X X X

X

X Rice leaffolders CnaphalocrocisMarasmia

x X x

X

X X

x x x x x X x x x

Yellow stem borer Scirpophaga incertulas

X X x

x

x

x X XAinauromorphaaccepta metathoracica(Ashmead) x

continued on next page

IRPS No 123 January 1987 31

Table 6 cntinued

Natural enemy Environment

Host stage Species Upland Rainfed lowland Irrigated lowland

Larva-pupa Gold fringed borer Chilo attriciiuslchneumonidae

Xanthopimpla stemmator (Thunberg) x aPupa x

Pteromalidae Trichoinalopsis apanteloctena (Crawford) ax

Pink stem borerb Sesamnia inferensParasite

Larva Braconidae

Stenobracon nicevillei (Bingham) x b b Armyworms Mvthinna SpodopteraLarva

Tachinidac Zygobothriaarroptvora (Robineau-Desvoidy) x x

Braconidac Cotesia spp

X Skipper Pelopidas mathias and green horned caterpillar Melanitis leda ismeneF-gg

TrichogramrnatidacTrichogrannaspp x

Scelionidae Undetermined Scelionid

x Larva

Tachinidae Argyroph lax nitrotibialis (Baranov) X x

Pteromalidae Trichomnalopsis apanteloctena (Crawford)

X Predator

CoenagrionidaeAgriocnenis femina femina Brauer x xIschnura senegalensis tambur

Miridae CrtorhinsliidipennisReuter x x

Nabidac Stcnonabis tagalicus (Stal) x

VeliidaeAlicrovelia douglasi atrolineata (Bergroth) x

Coccinellidae Micraspis crocea (Mulsant) X x x

Vespidae Eumenes campaniformis (Fabricius) x

Formicidac Diaecamna spp xOdontoponera transversa (F Smith) x

Carabidae Chaenius spp OphioneaW- aJasciataSchmidt-Goebel

x

Ophionea ishii ishii Habu x x X

Gryllidav Anaxipta longipennis (Saussur-) x xMetioche ittaticollis (Sial) x x x

Tettigoniidae Conocephalus longipennis (de Haan) x x x

aNo rearing made on rainfed lowland bNo rearing made on rainfed lowland or Irrigated lowland

MrasmiaAmong the Laval parasites the braconid Cardio- lowlands and parasitizes oriental maize borer OsIriniachilesphilijpphensis is most adapted to an upland environ- furnacalis (Guenee) larvae The leaffolder pupal parasitesment It also parasitizes Hydelepta indicata a common not only occur equally in all environments but also haveleaffolder of legumes The ichneumonids Trichomma cna- wvide host ranges- parasitizing species of OsIrinia Marucaphalocrosis and Temehcha stangli occur more in the Hydelepta and Honiona - in maize and legumeslowlands T stangli is most adapted to irrigated lowlands The principal egg parasite of yeltow stem borer (YSB)and also parasitizes Chilo and Scirpophaga stem borer Telenomus rowani is widely adapted to all environmentslarvae T cnaphalocrosis is most prevalent in rainfed and has an unusual alternate host -Tabanus eggs (25)

32 IRPS No 123 January 1987

Tetrastictu schoenobii specializes only in YSB and occurs mainly in irrigated lowlands -- further proof that YSB is most adapted to the wetter environments

Larval parasites of YSB are highly environment-specific The braconid Stenohracon nicevillei occurs only in upland rice but also attacks the pink stem borer (PSB) Tieinchwa phiiiplPwnsis an ichneumonid occurs in upland rice and also parasitizes the larvae of (7hiu stqpresaisinatro-morpha accepta melaihoracica like 7 stangi is mnost adapted to the irrigated lowlands

The dark-headed stem borer(l)HSB) (ih pochit3sus was not collected in all rice environments and the gold fringed borer Chilo auricilius nd PSB were collected only in upland rice Xaantiholiplastenrnaloran ichneumonid larval parasite appears to be widely adapted whereas TrihomaslopisapanitI)cteiiaa pteromalid pupal parasite occurs mostly in wetlands and also parasitizes larvae of Chilo suppressais and Pelhuidas mathias

Armyworms Myit hinin and Spodujipera have two main larval parasites - a tachinid Zrgohothriaatropivora in rainfed environments and a braconid Cotesia spp in irrigated lowlands

The butterfly pests Pelopidas and 4elanitishave tricho-grammatid egg parasites found only in irrigated lowland environments

Two larval parasites show environmental preferences --Argrroph av nigrotihialis a tachinid in rainfed habitats and Trichonialopsisapantelkcnela in irrigated ricefields A limgrotibialisalsoattacks the sweet potato hornwormAgrius convolvtdi and T apanteloctena parasitizes DH S1B

Predators as a group show more distinct environmental preferences The aquatic damselflies Coenagrionidae are most abundant in the lowlands Cvriorhinus is also most adapted to the lowlands The nabid Stenonabis tagalicus occurs only in the uplands and isalso prevalent on legumes The coccinellid lady beetles wasps and ants are more prevalent in upland areas AlicrasIpis cit-ea larvae prey ona variety of aphids attacking legumes and maize Etunienes campanonia vespid mud wasp makes nests in trees and is therefore most adapted to the more botanically diverse uplands Soil-dwelling ants cannot tolerate flooding

Upland environments are habitats to arboreal carabid beetles Three species of Chlaeniusbeetles prey on leaffolder larvae and are perhaps more important than parasites (27)

The gryllids and tettigoniids are widely adapted egg predators not only on rice but also on maize and legumes They feed on eggs of most insects that are laid on leaves

The spidercommunity of upland rice environments isrich in species At one site (Batangas Philippines) 31 species have been recorded (23) Of the 176 spider species recorded in Philippine ricefields about one-half (82) occur in upland rice The spiderspecies of upland rice environments overlap more with those of rainfed lowland than with those of irrigated lowland rice (24) The same study showed that of the three environments irrigated rice has (ie greatest spider species diversity foiiowed by rainfed lowland and upland

However of the 10 most prevalent ricefield spiders 9 were abundant in upland rice showing wide adaptation All three spider guilds -- orb-web space-web and huting spiders shywere prevalent in upland ricefields There were however differences in environmental prelerences for some spider species

Of the orb-wcavers Tetragniathamanlihulatawas partishycila rlv abundant 7 japonica was low and Ltucatge dehcorata was absent in upland rice

As in other environments Aiypeiwnaifirtosana was the most dominant space-web spider however its relative numbers were lowerthan for lowland sites Among hunting spiders Ox|opes javanus and Lcosa leatcost fgla were more abundant and Lvcosa lpseudoanmnlata was less abundant in upland compared with lowland rice envishyroniments

Extensive rearing of rice insect pests in upland rainfed lowland and irrigated sites over an 8-yr period in the Philippines provided insight into the effectiveness of parasites as natural enemies (Table 7 8) Most of toe data are from large sample sizes taken over at least one crop which overcomes the pitfalls expressed by Van Driesche (238)

VSB egg parasitization was surprisingly similar across all environments Larval parasitization was highly dynamic among sites and even years within the same site but showed somewhat lowerlevels in uplandsthan in irrigated wetlands This would have been predicted for egg parasites as well since YSB is adapted to the lowlands and so must be its parasites This isevidence that Telenomus rowani is as good at dispersing as its main hosts Trowani clings onto the anal tufts of female YSB and parasitizes eggs as they are laid This phoresy may explain its wide adaptation The more specialized lowland YSB larval parasites therefore appear more effective than their upland counterparts with more alternate hosts

Low collection levels of PSB larvae especially in the wetlands make comparisons between environments difficult Low levels of larval parasifization were recorded in both rainfed environments Except in the Cagayan rainfed wetland site in 1980 low levels ef larval parasitization were evident at irrigated and rainfed sitesalike No differences by environment were evident Higher levels of parasitization occurred in Sarawak during a 1967 outbreak also from tachinids Rothschild (197) indicated that higher parasishytization occurred in upland rice areas

Skipper and green horned caterpillar larval parasitization rates were generally low and showed no environmental effect htighest activity occurred in Cagayan in 1980 (48 parasitization) Although larval parasitization levels of RLF were variable year to year activity appeared greater in the irrigated lowlands than in the uplands Again the highest levels were recorded in Cagayan in 1980 Cagayan has extensive grasslands and fallow around the rice areas and a relatively long rainfall period Rainfall in 1930 was good for crop growth and apparently for increase of larval parasites

IRPS No 123 January 1987 33

Table 7 Parasitization rates of rice planthoppers and green leafhopper in upland rainfed lowland and irrigated lowland environments Philipshypines 1976-840

Environment and site Year Egg parasitization () Nymphaladult parasitization ()

BPH W13P GLH BPII WBPH GLIH

UplandBatangas

Rainfed lowland

1976 1979 1980 1982

15 17 12

2(47)8(39)

4(51)

9(1800)14 (212)

12(304)

8 (101)6 (127)

9(116)

Iloilo

Cagayan

1976 1977 1978 1981 1980 1981 1982

0 (69)

11

44 48 40

-

-

46 55 44

0 (471)

11

52 56 49

0 (200)9 (214)

10 (1259)8(235)6 (900) 3(800)

-7 (462)

11 (270)3 (313)5 (900) 1 (800)

4 (600)10 (725)10 (362)12(870)5 (900) 3 (800)

Pangasinan

Irrigatedlowland

1984 1976 1978 1979 1982

13 (436) 13(379) 14 (457)

6(14)7 (197)

17(198) 11 (486) 18 (275)

3 (81)10 (274)3 (116)

16(871) 19(88)

28(211)9 (764)5 (647)7 (1204) 8(305)

Laguna 1977 11 7 21 1978 31 1979 14 4 4 28 1980 7 8 20 1981 1983

56 7(278)

26 16 (1650)

16 27(1921)

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R I Apostol unpublished 1lo1lo and Pangasinan 136 Cagayan BCanapi unpublished Liuna 175 I C Pantua unpublished 118 234 43 Carino and Shepard unpublished BPH = brown planthopper Nila parvara htgens W131Pi =whitebacked planthopper Sogatelia furcifera GLH = green leafhopper Nephotettrix virescens Figiires in parentheses are sample sizes

Tauher et al (230) reported that internal external and genetic factors influence the seasonal activity of parasitoids

Hopper egg parasites but not nymphaladult parasites were also parlicularly abundant at the Cagayan site possibly the egg parasites have more alternative hopper prey in the weedy areas than nymphaladult paisites

The seasonal dynamics of hopper egg parasites can be seen for upland rice in Batangas(Fig I 2) and for rainfed lowland lice in Pangasinan( 135)and Cagayan( 105) Levels of BPHWBPI-L and GLH egg parasitization were corn-parable in upland and irrigated lowland environments Except for some high rates of paraMsitization early in the season the parasitization rate tended to be steady in all envilonments However hopper nymphaladult parasi-tizution levels were more similar in the rainfed environments and lower than in irrigated rice

Overall parasites do not appear highly effective by themselves against upland or even lowland pests Among natural enemies parasites have attracted the most attention because they can be more readily assesscd

Predators and pathogens lend themselves better to management practices Predators the most important group of natural enemies in riccfields are difficult to quantify On the other hand pathogens appear less im-portant but can be readily cultured and disseminated

The natural enemy commulnity of upland rice is rich in species and differs significantly from that of lowland rice

These beneficial organisms must be conserved by applying insecticides judiciously particularly sprays Seed or soil placement of chemicals minimizes exposure to natural enemies The strategy to derive the greatest benefit from natural enemies isto allow the greatest number of beneficial species to thrive Then perhaps at least one species will be effective against each pest at any tine overcoming the variable abundance of each species within and between years

Programs to introduce exotic species or mass produce indigenous ones are ambitious and expensive because they require trained people on a sustained not ad hoc basis

Chemical control Insecticides offer rapid and efficient control of upland insect pests (190) but-should be used only after other control measures have been considered Insecticides are rarely used on upland rice because of cost (32) Upland rice yields are normally too low to justify the expenditure Also most upland rice is grown as a cash crop Spraying upland rice is more difficult than lowland rice because water is less accessible Government subsidy programs to provide pestishycides to tribal peoples for upland rice production have created problems of toxicity to humans because isolated people have had no experience to be able to handle pesticides safely

However ins-eticide use has beenjustified when a crop is

0Z

Table 8 Parasitization rates of lepidopterous pests in upland rainfed lowland and irrigated lowland environments Philippines 1976-84a

Parasitization (5)

Environment and site Year YSB GFB PSB Earcutting Swarming Skipper Greenborned RLF Brown larva larva caterpillar caterpillar caterpillar caterpillar larva semilooperEgg Larva larva larva larva larva larva

UplandBatangas 1977 - 0 (57) - 0 (63) - 0 (71) - 0 (106) 11(142) 2 (101)1978 43 (4697) 8(451) 5 (38) 3 (40) 6 (61) 2(58) 15 (57) 13 (129) 13 (1286) 4 (30)1979 59(4135) 12(320) 13 (11) 5(28) 9(45) 6(46) 7(23) 8(160) 9(1152) 8(26)

Rainfed lowland Iloilo 1976 - 0 (10) 0(10) 0 (7) 0 (19) 2 13) 15 (180) 5 (76)1978 0 (54) 0 (3) 0 (17) 9 (64) 4 (78) 3 (73) 0 (121) 0 (191)1979 64 (2350) 16 (84) 6 (17) 0(6) 18 (68) 25 (104)

1981 56 (1322) 18(304) 0(5) 12(51) 18(138)Pangasinan 1976 60 (7 7 )b 62(82) 0(111) 5 (39)

1978 58 (53)b 5(51) Cagayan 1979 59(713) 8 (101) 6 (34) 0(7)

1980 0(11) 4 (28) 15 (286)

62 ( 48(177) 61(140)1981 0(5) 12 (2) 13(45) 14 (92)

Irrigatedlowland Laguna 1974 62 8

1974-75 50 41 1975 43 72 1978 96 (623) 21(24) 33 (60)1979 83(2017) 18(110) 8(62) 11 (89) 22(400)1980 78 (1625) 20(75) 0 (16) 4 (72)1983 60

9 (80) 7 (270) 30

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R F Apostol unpublished Iloilo and Pangasnan 136 Cagayan B Canapi unpublished Laguna 175 P CPantua unpublished 118 234 143 Carino and Shepard unpublished YSB = yellow stem borer ScirpophagaincertulasGFB = gold fringed borer Chilo auricilius PSB = pink stem borerSesamia inferens earcutting caterpillar Alythimna separata swarming caterpillar Spcdoptera mauritia acronyctoides skipper Pelopidas mathias reen horned caterpillar Melanitls ledaismene RLF = rice leaffoldersCnaphalocrocismedinalis and Marasmiaspp brown semilooper Mocisfrugalis Figures in parentheses are sample sizes Egg masses

IRPS No 123 January 1987 35

Table 9 Effect of seeding rate and insecticide on plant density and Yield losses from insect pests in upland rice often areyield of Dagge upland rice Batangas Philippines 1976a comparable to those of lowland rainfed or irrigated rice

Seeding rate Insecticide Dosage Plant density Yield (0-Y30o) but usually from different pests The principal (kg aiha) seed treatment (kg aiha) (noIra-row) (tha) groups are soil pests feeding on sown seed and rootsat 14 DE followed by seedling pests These groups have largely gone

50 None - 35 a 20 a unnoticed leading researchers to conclude that stem borers50 Carbofuran F 10 38 ab 23 a and seed bugs are the most important pests (17) Yield loss 10 beCabofuranF100 None - 4551 c 23 a trials that pinpoint major pests need to includc seed

100 Dieldrin WP 10 52 cd 22 a treatments Granules applied in the furrow may not give150 None - 59 d 22 a adequate protection from many seed pests

aAv of 8 fields DE = days after crop emergence In a column means The floristic and pedological diversity of upland rice followed by a common letter are not significantly different at the environments is matched by the diversity of arthropods5level by D)MRT F = flowable Wi = wettable powder both pests and their natural enemies Irrigation has homoshy

genized lowland rice environments Flooded soils tend tothreatened by epidemics of locusts or armyworms New have the same properties and to eliminate soil pests The technology stressing efficiency has minimized insecticide spatial and temporal dominance of cultivated rice has cost benefited specialist pests

Seed treatment is normally inexpensive and can be Upland rice has no specialist arthropod species The economicallyjustified in -nanycases(66) Treating seed with monophagous pests on upland rice prefer the lowlands systemic insecticide (216) but not chlorinated hydrocarbons where rice evolved Rice in the uplands exists only because (247) is most effective in controlling seedling maggots of man and even though lowland rice predates upland riceFoliar sprays have to be repeated to obtain control and specialist species evolved Wheat maize and sorghum do often are washed off by frequent early season rains not have monophagous pests leading us to conclude that Granular insecticides are effective but required dosages are there is no evolutionary advantage for inssects to have too expensive to justify except under heavy infestation narrow host ranges in upland graminaceae(247) When directed against first-instar larvae banding Pests have adapted to the uplands not only by having granules in seed furrows (177) can be effective against white wider plant host ranges but also by having long life cyclesgrubs at low dosages (025 kg aii ha) (134) Seed treatments undergoing dormancy or being dispersive Each pest has a are not promising for white grub control (248) unique life history strategy suited to highly unpredictable

Seed treatment is also effective ainst ants and may be environments In agivenyear someinsect specics an better cheaper than increasing the seeding rate In Batangas exploit upland rice as a food source but no pest is a senious treating 50 kg seed ha was as effective as increasing the one every year on upland rice seeding rate to 100 kgha and using untreated seeds (Table The low yielding potential of upland rice means less 9) In Brazil seed treatments control termites and other soil quantity and lower quality of food for pests to exploit but insects including the lessercornstalk borer(66) Domiciano also less incentive for farmers to undertake control (58) however found seed treatments inconsistent in their The most effective control would be early and synchroshyeffect against high populations of lesser corn stalk borer nous planting of early-maturing upland rice varieties to

Baiting can be an inexpensive way of controlling ants prevent insect pest buildup Breeding for insect resistance seedling maggot flies mole crickets and field crickets Baits should initially focus on stem borers because they are verymade of locally available material and impregnated with difficult to control by other means Biocontrol efforts insecticide can be sparingly distributed taking adantage of should first stress conservation of natural enemies preshythe pests ability to disperse and encounter bait sites dators and pathogens rather than parasites Efficient

The new low-volume hand-held controlled-droplet chemical control includes seed treatment or baiting for seed sprayers may offer an advantage over high-volume knap- seedling and some root pests and using controlled dropletsack sprayes for controlling foliar pests Upland rice sprayers for spot treatments against foliar pests and seed canopies are more open than those in lowland rice and bugs when economic thresholds are reached better droplet penetration should result Controlled droplet applicators would be ideal for seedbugs and defoliators

CONCLUSION

Upland rice is attacked by a wider array of insect pests than is lowland rice mainly beca use of the addition of soil pests Generally population levels of lowland rice pests are lower in upland than lowland rice leading many people to conclude that insect pests are not important on upland rice

36 IRPS No 123 January 1987

REFERENCES CITED

1Abraham C C and K S R Mony 1978 Occurrence ofLeptocorisa acuta Fabr (Coreidae Hemiptera) as a pest of nutmeg trees J Bombay Nat ttistSoc 74553

2 Agyen-Sampong M 1975 HlierogrlphusdiaganensisKrauss (Orthop-tera Acrididae) a new pest of rice in Northern Ghana Ghana J Agric Sci 8249-253

3 Agyen-Samnpong M 1976 Varietal resistance and other inle-ratcd pest control components as factors of rice varietal improvement in West Africa Prcsented at the Second varietal improvement serinar 14-18 Sep 1976 Bouak Ivory Coast West Africa Rice Development Association Monrovia Liberia 18 p

4 Agyen-Sampong M 1977 Outbreak of Ihccrorrdnisorae in rice seedlings near Rokupr Sierra Leone lit Rice Res Newsl 2(6)16-17

5 Agyen-Sampong M 1980 Dirty panicles and rice yield reduction caused by hugs IntRice Res Newsl 5(l)11-12

6 Agyen-Sampong M 1982 The major pest problems of irrigated upland and mangrove swanip rice ecosystems ir the humid tropical Guinea savanna and Salel climate zones Pages 318-328 in Pro-ceedings training course cn concepts techniques and application of irtegrated pest management in rice inWest Africa 10-20 Jan 1982 Fendall Liberia West Africa Rice l)evelopioent Association Monrovia Liberia 505 p

7 Akibo-Betts V I and S A Raymundo 1978 Aphids as rice pests in Sierra Leone lit Rice Res Newsl 3(6)15-16

8 Akinsola F A 1975 Present status of different rice stemborers in parts of Nigeria iicL Entomo Newsl 328

9 Akinsola FA 1979 Ihe biology and ecology of rice stemborers in Nigeria Ph 1) thesis University ol lbadan Nigeria 217 p

10 Akinsola E A 1979 Guidelines of integrated management of insect pests of upland rice in West Africa Presented at the Seminar on integrated management of rice diseases and insect pests at Bobo-Dioulasso Upper Volta 17-22 Sep 1979 West Africa Rice Develop-ment Association Monrovia Liberia

II Akinsola IA 1984 Insect pests of upland rice in Africa Pages 301-305 in An overview of upland rice research Proceedings of the 1982 lIouak Iwsrv Coast upland rice workshop International Rice Research Institute 13O Box 933 Manila Philippines

12 Alam M S V TJohn and Kaung Zan 1985 Insect pests and diseases of rice in Africa Pages 67-82 in Rice improvement in Eastern Central and Southern Africa proceedings of the international rice workshop at Iusaka Zambia 9-19 April 1984

13 Alticri M A A van Schoonhovcn and J D Doll 1977 The ecological role of weeds in insect pest management systems a review illustiated with bean (Phaseohsvulgaris L)cropping systems PANS (Pest Artie News Summ) 23195-205

14 Anmante F 1967 The leaf-cutting ant Atia capiguaraa pest of pastures [in Portuguese] Biol6gico (Sao Paulo) 33113-120 (Rev Appl Entomol A56207)

15Andrade A C and D Puzzi 1951 Preliminary results of experi-merits on the control of Sculptcoris iastantus on sugarcane [in Portuguese] Bioulgico (Sao Paulo) 1744-49 (Rev Appl Entomol A41421422)

16 Appert J 1970 Maliarplia separaelha (rice stem borer) new observations of entomological problems of rice in Madagascar [in French] Agron Trop (Paris) 25329-334

17 Ariadeau Mand Z Harahap 1986 Relevant upland rice breeding objectives Pages 189-197 in Progress in upland rice research Proceedings of the 1985 Jakarta conference International Rice Research Institute 1O Box 933 Manila Philippines

18Asar S and Kisimoto 1975 Seasonal occurrence of the smaller brown planthopper in upland rice [in Japanese] Proc Kanto-Tosan Plant Prot Soc 2228

19 Bakker W 1974 Characterization and ecological aspects of rice yellow mottle virus in Kenya Meded Landbouwhogesh Wage-ningen 152 p

20 Balut F F 1970 White grubs attacking rice Oryza ativa L [in Portuguese] Biol6gico (So Paulo) 36321-322

21 Banerjee S N and L M Pramanik 1967 The lepidopterous stalk borers of rice and their life cycles in the tropics pages 103-124 in The major insect pests ofthe rice plant Proceedings ofa symposium at the International Rice Research Institute September 1964 The Johns Hopkins Press Baltimore Maryland USA

22 Barrion A T and J A Litsinger 1980 Ants a natural enemy of cliaiarac in dryland rice IntRice Res Newsl

5(4)22-23 23 Barrion A T and J A litsinger 1981 The spider fauna of

Philippine rice agroecosystem 1Dryland Philipp Entomnol 5139-166

24 Barrion A T and 1 A Iitsinger 1984 The spider fauna of Philippine rice agroccosystcms II Welland Philipp Eiitomol 611-37

25 Barrion A T1 and J A Litsinger 1984 lihanus (Diptera Tabanidae) eggs an alternative host of rice stei borer (SI3) egg parasite Telenoonis digns (Iliymenoptcra Scelionidae) lit Rice Res Newvsl 9(6)19

26 Barrion A T and J A Litsinger 1985 Identilication of rice leaffolders by wing markings Int Rice Res Newsl 10( 1)24

27 Barrion A T and J A Litsinger 1985 Chctiosspp (Coleoptera Carabidae) a leaffolder ILF)predator lit Rice Res Newsl 10(1)21

28 Bertels A 1970 Insect pests of rice and theircontrol [in Portuguese] Pelotas Brazil Instituto de Expcrimentacao Agropccuararias do Sul Circular 43 24 p

29 Bonzi S M 1982 Ctilo (flisilineus J de Joannis (Lepidoptera Pyralidae) stem borer of irrigated cereals in Upper Volta LioFrench] Agron Trop Riu Rizic Cult Vivrieres Trop 37207-209

30 Bourke T V T L Fenner J N LStihick G Baker EHassan D F OSullivan and C S li 1977 Insect pest survey for the year ending 30th June 1969 Papta New Guinea )epartment of Agrishyculture Stock and Fisheries Port Moresby 57 p

31 Bowden 1 1973 Migration of pests in the tropics Meded Rijksfac Landbouwwet Gent 38785-796

32 Breniere J 1973 The main rice parasites in West Africa and their control Pages 27-42 in Proceedings seminar on plant protection for the rice crop West Africa Rice Development Association Monrovia Liberia

33 Breniere J 1976 The principal insect pests of rice in West Africa and their control West Africa Rice Development Association Monrovia Liberia 52 p

34 Brown ES 1962 The African arinyworn Spoiolptera eXI)Ina (Walker) (Lepidoptera Noctuidae) a review of the literature Cornshymonwealth Institute of Entomology London UK57 p

35 Brown F B 1968 Upland rice in Latin America FAO-International Rice Commission 12th Session of the Working Party on Rice Production and Protection 2-14 Sept Peradeniya Sri lanka Food and Agriculture Organization Rome Italy 6 p

36 Buyckx F J 1 1962 Disease and insect pests of crops inCongo Rwanda and Burundi [iil French] Publications dc LInstilut National pour lestude Agrononiquedu Congo M Weissenbruck S A Brussels Belgium 708 p

37 Cariiio F 0 1981 Role of natural enentics inpopulation suppression and pest management of green rice lealhoppers Ph D thesis University ofthe Philippines at Los Bafitos PhilippinesLaguna 136 -

38 Carpente A J 1983 The development of rice cultivation and extension of vegetable productin in Zanzibar Final Report Vol II1 October 1983 Food and Agricultme Organization Ronte Italy

39 Carroll C P and S1 Risch 1983 Tropical annual cropping systems ant ecology Environ Manage 751-57

4(0 Cates R G 198 1 Host plant predictability and the feeding patterns of monophagous oligophagous and polyphagous insect herbivores Oecologia (Berl) 48319-326

IRPS No 123 January 1987 37

41 Cendcafia S M and F B Caora 1967 Insect pests of rice in the 62 Ene 1 C 1981 I)iiorldhol)ierusihilis Slater (licteroplera Philippines Pages 591-616 in The najor insect pests ofthe rice plant lygacidue Illissinac) a new rice pest front Nigeria lit Rice Res Proceedings of a symposium at the International Rice Research Newsl N3)15 Institute September 1964 The Johns Hopkins Press Baltimore 63 Fugenio T S 1965 rhe effect oi e-vCgetation on rice production Maryland USA 729 p and the possibility of overcoming it by leaf pruning MS thesis

42 Centro Ihternacional de Agricklittira Tropical 1984 Upland rice in University ol the Philippines at Los Baijos ILaguna Ihilippins 69 p Iatin Anerica Pages 93-119 hi An os ervies of uplaind rice research 04 Farrow R A 1975 The Alrican nmigratory locust ii is main Proceedings of the 1982 Boinakamp Ivory Coast upland rice workshop outbreak area of the middle Niger qlinlitatise studies of solitary International Rice Research Institute P0 Blox 933 Manila populations in relation to environmental factors Locusla No II Philippines 198 p

43 Chandra G 1979 Taxonomy ad hn riononlics ofthe imect parasites of 65 Fennal R G 1943 Food crop investigation Windward and rice Icalloppers and planthoppers in the Pihilippines and their Ieeward Islands First report Report onl work dunring the periodiiiportairce iii natural biol)gical cointrol Philipp Fntoltol 4119-139 April 1942-April 1943 Imperial College of Tropical Agriculture

44 Chang T T 1976 The originii colut ion enlIivation dissemination Trinidad 16 p (Rev of Appl Fintornol A3222-25) and diersificition of Asian and African rices -uphytica 25425-44 1 66 Ferreira F 1983 Integrated pest ritaageiret of upland rice [ill

45 Chatierjee P B 1979 Rice leaffolder attacks in India lit Rice Res Portuguese English summary] Pages 323-341 in Upland riceculttre Newsl 4(3)21 factors affecting production E Ferreira T Yamada and F-

46 Cheng Cbien-pan 19(4 Upland rice in Taiwan International Rice Malmolta eds Institutoda Potassae Foslato Institnto Internacional Cnmmission Wiorking Party oit Rice Production ard Protection I()th da Potassa Piracicaha Sao Paulo lraid Meeting 3-10 March 1964 Manila Philippines Food and Agricul- 67 Ferreira E arid F dl Silva Martins 1979 Occurrence of ture Organiaion Rome Italy 5 p Sogatohs orvziola (Muir) inupland rice in Goias Ilrail lit Rice

47 Chcrrett 1 M and 1) 1Peregrine 1976 A review of the status of Rcs Ncwssl 4(4)18 leaf-cutting ants and their control Ann Appl Biol 84124-128 68 Ferreira E and 1F da Silva Martiris 1984 Insect pestof rice ill

48 Chipeta F M 1974 Pests of rice in Malawi Pages 90-95 in Brazil and their control Jill Portugucsej FMIlRAtPA-CNPAF Proceedings Sith eastern Africa cereals research conference 10-15 Goiania Brmiil 67 p March 1974 DRB Mialda ed Ministry of Agriculture lilongwe 69 Ferreira FI F ca Silha Miartins S S Nero and 1 It P de Malawi Carvallio 1982 M tltiplc regression analysis todeterimine the effect of

49 Chu Yau-i 1979 The d atiaging period of the rice seedling fly insect populatiois ott upland rice yield [inIPortuguese Pesqui (Atlterigotiavxigu Stcin) (NItiscide Diptera) oil cori) at Fast Java Agropecu Bras 17671-675 Indonesia Plant Prot Bill 21403413 70 Ferreira F 1F da Silva Martins and F 1PZimmermann 1979

50 Conklin H C 1957 IHaninoo agriculture a report on aln integral Rice varietal resistance to tle lesser corn stalk borer [in Port uguese] systent of shi fti rig cult ivationi in the Pltilippincs Food and Agriculture Pesqui Agropccu Biras 14317-321 Organization Rome Italy 209 p 71 Fleming W F 1968 Biological control of lie hapanese beetle U S

51 Cramer H II 1967 Plant proection and world crop production Dep Agric RLs Sers [ech Bull 1383 78 p Pilaniensclhutz-Nachr 201-524 72 Food and Agriculture Organiation of the United Nations 1982

52 l)ani R C and Majunidar 1978 Prelininary observations on tie Country report surmimary Pages 27-37 in Report on the first incidence of root aphids on differeit rice cultnrcs Sci Cult 4488-89 FAOIUN)IPhail and regional training coiuirse in iniproved ctltural

53 Davidson R I 1979 lffects of root feeding on foliage yield Pages practices for upland rice 5-23 Oct 1981 Northern Region Agrculturd117-120 in T K Ci osb and R B Pottinger eds Proceedings of the Developririt Center Chiang Mai Food and Agricuilture Organiiashy2nd Australian conlerence on grassland invertebrate ecology Govt tion Rome Italy 37 n Printer Wellington New Zealand 73 Franco do Ariaral S 195 1 Residual action of insecticides in turfy

54 l)ean G 1W 1976 Rice insect pests inlaos Int Rice Res Newsl soil--the results of a field experiment [in Portuguese] Biol6gico (Sao 1(2)15 Paulo) 17209-214 (Rev Appl Entomol A41423)

55 I)ean G J W 1978 Insect pests of rice in Laos PANS 24(3)280-289 74 Freeman ) 1970 Report onithe Iban london School of -coroiics 56 )ingle H 1984 Behavior genes and lifc histories conplex adapta- Monographs on Social Anthropology No 41 Athlane Press

tions to uncertain environments Pages 169-194 in A new ecology London UK 317 p novel approaches to interactive systems P W Price C N Slobod- 75 Fuchs T W1 A Harding and 1W Smithlr 1979 Induction and chikoff and W S Gaud cdsl ohn Wiley New York USA termination oldiapause in the sugarcane borer Ann Flitonlol Soc

57 Dobelmann-JP and M Falais 1968 Upland rice cultivation ill tie Am 72271-274 North-western part of Madagascar in 1966-1967 [in French English 76 (hbriel B1) 1975 A review tf the major insect pests of some tpland summary] Trop Agron 2313-21 crops in tlc Philippines Pages 86-111 in Ad Htoc panel of experts on

58 Doniiciano N L 1979 Chemical control of tie lesser corn stalk pest disease arnd weed problems in some rainfed crops Bangkok (lolsonno~paloIgnowslhts) (Zeller 1848) (I-epid(ptera Ihycitid ae) Thailand Food aid Agriculture Organization Roire Italy

on rice in Ponta Gressa Parara 1977-1978 [in Portuguese] Pages 77 Garg D K and N K Shal 1983 White grub outbreak on rainfed 291-303 in Anais 1 Reuniao de tecnicos cmiirizicultura do estado de dryland rice in Uttar Pradeslh Int Rice Res Ncwsl 81(4)16-17 Sao Paulo-Campinas Cati Brazil 78 Garg I) K and IP Tandon 1982 Major insect pests of rice on hilly

59 Dunsmore 1 R 1970 Investigations on tlie varieties pests and tracts of Uttar Pradesh India lilt Rice Res Newsl 7(11)11-12diseases of upland rice (hill padi) in Sarawak Malaysia Int Rice 79 Garrity 1) P 1983 Upland rice research an international bibliograpy Comm Newsl 1929-35 1975-1982 Iternational Rice Research hnstitnte PO Box 933

60 Elias 1R1965 The new decline of rice seedlings is not caused by Manila Philippines 121 p parasites [in Portuguese] Biol6gico (Sao Paulo) 313-6 80 Ghauri M S K 1982 A new subspecies of Iinorh)rone bull

61 Empresa Brasiliera de Pesquisa Agropecuaria 1984 Upland rice itt cornlus Slater (Hemiptera Iygaeidae) front Papua New Guinea on Brazil Pages 121-134 it Av overview of upland rice research rice and carpet grass Ihll lEntoniol Res 72133-137 Proceedings ofthe 1982 Blouak Ivory Coast Upland rice workshop 81 Gonzales F 0 PArregoces R I ihernandef and OT Parada International Rice Research Institute PO Box 933 Manila 1983 Insect and imite pests of rice arid their control in Latin America Philippines [in Spanish] la Federacion Nacional de Arrocerts (Fedearroz)

Bogota Colombia 60 n

38 IRPS No 123 January 1987

82 Grist D H and R J A W Lever 1969 Pests of rice Longmans London UK520 p

83 Hale P R 1979 Two subterranean pests of upland rice in Papua New Guirnca IntRice Res Newsl 4(2)16-17

84 Hale P R and J T Hale 1975 Insect pests of rice in the New Guinea Islands Rice Entomol Newsl 34

85 Harris W V 1969 Termites as pests of crops and trees Common-wealth Institute of Entomology London UK 41 p

86 Hattori 1 1968 A method to distinguish between grass leaf roller (Ctaplha rocis medinalisGuenee) and the rice leaf roller (Susurtia exigua Butler) [in Japanese] Plant Prot (Japan) 23167-170

87 Henry I E and 1A Onsager 1984 Experimental control of the mormon cricket Iahrosshnph by NJsena locustae(Aficrospora Microsporidae) a protozoan parasite of grasshoppers (Ort Acri-didae) Entomophaga 27197-201

88 Hidaka 1 P Vungsilabulr and SKadkao 1974 Studies on ecology and control of the rice gall midge in Thailand Tech Bull 6 TropicalAgricultural Research Center Tokyo Japan 113 p

89 Hincklcy A 1) 1963 Ecology and control of rice planthoppers in Fiji Bull Entomrol Res 53467 481

90 Hirao J 1982 Ecology and chtInical control of the rice leaffolder Jpn Pestic Inf 4114-17

91 Ho DT 1984 Sten borer incifience in rice ecosystems in KenyaEast Africa Pages 307-320 in A overview of upland rice research Proceedings of the 1982 Bouac Ivory Coast upland rice workshop International Rice Research Institute PO Box 933 Manila Philippines

92 Ho )T J 1Njokah and J GKibuka 1983 Studies on rice sten borers in Kenya with emphasis on ifaliarpiaipiraellRag Insect Sci Appl 465-73

93 Ichihara I T Ito A NIatsuo and M Sawada 1971 Chemical control of the rice striped stem borer in upland rice grown in plastic mulch [iii Japanese] Proc Kanto-Tosan Plant Prot Soc 1882

94 IclhiharaIand A Matsuo 1969 Damage caused by rice striped stett borer to upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1686-87

95 Ichhara I atid A MIatsun11970 Conside ration oft lie extraordinary prevalence of rice striped stem borer in upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1778-79

96 Institut de Recherches Agrolnliqes Iropicale et des Cultures Vivrieres 1978 Insect resistance Pages 12-13 in Upland rice 1978 atnual report Bouak Ivory Coast 27 p

97 International Institute for Tropical Agriculture 1983 Aphids itt rice roots I ITA Res Briefs 4(1-3)4-5

98 International Institite fi Tropical Agriculture 1983 Hopperburn in rice caused by brown plathopper IITA Res Briefs 4(l-3)6-7

99 International Rice Research Institute 1975 Annual report for 1974 PO Box 933 Manila Philippines p 216

100 International Rice Restarch Institute 1976 Annual repiort for 1975 PO Box 933 Manila Philippines p 228

101 International Rice Research Institute 1979 Cropping systems program Component technology development and evaluation Page 404 i Annual report for 1978 11O Box 933 Manila Philippines

102 International Rice Research Institute Department of Entomology 1981) Insecticid evaluation 1979 Tables 73 74 PO Box 933Manila Philipes

103 International Rice Research Institute Department of Entonmology 1981 Insecticide evalomtion 1980 Table 81 POBox 933 Manila Philippines

104 International Rice Research Institute 1981 Report of a meeting of Asian rice-based cropping systems entomologists Eleventh cropping systems working group meeting 18-22 May 1981 PO Box 933 Manila Philippines p 132 133 144

105 International Rice Research Institute 1984 Cropping systems program Pages 368-373 in Annual report for 1983 PO Box 933 Manila Philippines

106 International Rice Research Institutc 1984 Upland rice in Asia Pages 45-68 in An overview of upland rice research Proceedings of the 1982 Bouakamp Ivory Coast upland rice workiiop loternational Rice Research Institute 11O Box 933 Manila Philippines

107 Irshad M 1977 Grasshoppers in rice in Pakistan lit Rice Res Newsl 2(3)8-9

108 Ito R and TOno 1964 Upland lice its culture and varieties in Japan lIternational Rice Cormmission loth Meeting of the Working Party on Rice Production and Protection 3-10 March 1964 Manila Philippines Food and Agriculture Organization Rome Italy 12 p

109 Jeffrey M I and V A Dyck 1983 Effect of moonlight on lightshytrap catches of brown planthopper lit Rice Res Newsl 8(6) 13

110 Jennings P R and T Alicia Pineda 1983 Management ofthe rice plnthoppers Sogatodsorivi(coIl in Latin America Presented at the meeting on stable plant resistance FAO and lIndocsian I)inectorate of Food Crops Protection Bali lndonesia 29 Novcniber-2 l)ccemshyber 9 p

IllJimenez Gomez J A 1979 Rice insect pests i lla ios ()riCntales lil Spanish] Pages 99-103 in Rice ciltivation Instituto (olotibiatto Agropecuario Compndio No 29

112 Jongeleen F JJ Soesanto and A Mockidjo 1979 Studies oti pathogens of two species of Scarabacidac froinIndonesia Z Angew Enlomnol 87412-417

113 Jouvenaz 1) PC S lofgren and W A Baiks 1981 Biological control of imported fire antsa review of cirrent knowledge Bull Entomol Soc Am 27203-208

114 Kainoh Y K Shiniiu S MarUani Y Taiaki 1980 lost finding behavior of tie rice bu lplotori+JchinvittjI)allas (liciniptera Corcidac) with special reference to diet pattern ol aggregation and feeding on the rice plant Appl Entomol Zool 15225-233

115 Kakkar K 0 P Sharma and P I Sharma 1977 Note on Ihteronyrchus liodresRedt (Coleopicra I)ynastidae) as asporadic pest of paddy and maie inHimachal Pradesh Corr Agric 156-57

116 Kalode M131 79 Major insect pest problems and their control in upland rice (Unpublishcd mineogriph CWI oJNaaiat lni Chandaratat (198 1)original not seen)

117 Kalshoven L i 98 I The pests of crops in I ldoniesia (translated by P A vatn der Laan) 1 T Ichtiar B3aru-vati Iloeve lakarta Indonesia 701 p

118 Kamal N Q 1981 Suppression of whitebacked planto ner Sogatella fircaifra (Horvath) aLId rice lealfolder (nhal -cis licdb lis (Guenee) populations by natural cnclmie hlI) tiesis Grcgorio Araneta University Foundation Victoncta Park Metto Manila Philippines 179 Ip

119 Karim A N M Reaul 1975 A species ofliealy bug Trioni i70s ceres new oii rice in the Philippines Rice Entoniol Newsl 247

120 Katanyukul Wand C Chandaratat 1981 Insect pestsof uplaid rice and their control Report firstFAOUNlFi Tliailand training course on improved cultural practices for tplad rice Chiang Mai Thailand 5-23 Oct 1981 Food and Agriculture Organiation Rome Italy 22 p

121 Kern F 1955 lhletwioa(I)uninemts) bhhit Biturm(Coleoptera) a new pest of sugarcane inVenezuela [inSpanish with Etnglish sunmary] Agron Trop (Maracay) 527-29 (Rev Appl Eitoniol A4572)

122 Khan K M and S Ghai 1974 White grubs ald their control in India Pesticides (1Bombay) 819-25

123 Kiritani K 1975 Integrated pest management of rice pests Pages 66-70 itt Proceedings platning workshop ott cooperative hIeld research in pest managementP SMotooka ed East-West Center Honololu Hawaii USA

124 Kisimoto R 1967 Ecology of insect vectors forecasting and chemical control Pages 243-255 in The virus diseases of tle rice plant Proceedings of a symposium atthe International Rice Research Institute April 1967 The Johns Hopkins Press altimore Maryshyland USA

125 Kisimoto R and V A Dyck 1976 Climate and rice insects Pages 367-391 in Climate and rice International Rice Research Institute PO Box 933 Manila Philippines

126 Kureha S S Kobayashi SAsari and T Ishihara 1977 Life history and control of the mole cricket Grrllotalpa africana Palisot de Beauvois [in Japanese English translation 7520 1RRI library] Proc Kanto-Tosan Plant Prot Soc 21192-197

127 Kushwaha K S 1981 The black beetle as a pest of rice in Haryana India Int Rice Res Newsl 6(4)14

128 Lawani S M 1982 A review of tile effects of various agronomic practices on cereal stem borers Trop Pest Manage 28266-276

129 Leuck D B 1966 Biology of the lesser cornstalk borer in South Georgia J Econ Entomol 59 797-801

130 lever R J A W 1955 Insect pests of the rice plant in Malaya Int Rice Comm Newsl 1612-21

131 Li Ching-Sing and Shin-Foon Chiu 1951 The study ofthe rice gall midge Iachydiplosis orvza Wood Mason [in Chinese with English summary] 1Taiwan Agric Res 21-13

132 Iitsinger J A 1984 Assesament of the need-based application for rice Proceedings TechnologyTransfer Workshop 15-16 March 1984 International Rice Research Institute 1PO Box 933 Manila Philippines 6 p

133 Litsinger J A 1984 Environmental diversity and insect pest abundance with reference to the Pacific Pages 285-307 in Proceedings sub-regional training course on methods of controlling diseases insects and other pests of plants in the South Pacific 4-20 Oct 1982 Vaini Kingdom of Tonga Nukualofa

134 Litsinger J A RF Apostol and M BObusan 1983 White grub Leucopholis irrorata (Coleoptera Scarabaeidae) pest status popula-tion dynamics and chemical conllol in arice-maize cropping pattern in the Philippines J Econ Entoinol 761133-1138

135 Litsinger J A C de la Cruz F Raymundo A T Barrion M D Lumaban J P Bandong M S Venugopal F Paragna and E Balete 1982 Insect pests and insecticide reponse of dry-seeded bunded rice Pages 179-193 in Report of workshop on cropping systems research in Asia International Rice Research Institute Los Bafos Philippine

136 Litsinger J A M 1) Luinaban J P Bandong A T Barrion W Katanyukul and N Panda 1978 Insect pest management in rainfcd lowland rice agroecosystms of tropical Asia International Rice Research Conference 17-21 April 1978 International Rice Research Institute Los Bafios Philippines

137 Litsinger J A and K Moody 1976 Integrated pest management in multiple cropping systems Pages 293-317 in R I Papendick P A Sanchez and G B Triplett eds Multiple cropping American Society of Agronomy Madison Wisconsin USA

138 Loevinsohn M E 1984 The ecology and control of rice pests in relation to the intensity and synchrony of cultivation Ph D thesis University of London 36) p

139 Loevinsohn M E J A Litsinger N Panda M A Austria R F Apostol C Andrion and J Bandong 1982 A basis for predicting rice insect pest abundance in cropping systems Paper presented at the 1982 International rice research conference on rice and cropping systems 19-23 April 1982 International Rice Research Institute Los Bafios Iaguna Philippines 25 p

140 Lopez A W 1930 Repot of the Entomology Department Pages 145-172 in Annual report of the Philippine Sugar Association 1929-30 Manila

141 Lopez AW 1931 Entomology Department Pages 227-266 in Annual report of the director or research 1930-31 Philippine Sugar Association Manila

142 Lopez A W 1932 Entomology Deparment Pages 252-279 in Annual report of the Philippine Sugar Association 1931-32 Manila

143 Lungibill P and G G Ainslee 1917 The lessercornstalk borer US Dep Agric Agric Res Serv Tech Bull 539 27 p

IRPS No 123 January 1987 39

144 McGuire J U Jr and BS Crandall 1967 Survey of insect pests and plant diseases ofselected food crops of Mexico Central America and Panama U SDepartment of Agriculture International Agriculshytural Development Service Agricultuial Researrh Service October 1967 11 p

145 McNeill Sand T R ESouthwood 1978 The role of nitrogen in the development of insectplant relations Pages 77-9 in lliochemical aspects of plant and animal coevolution J B Ilarbornu ed Academic Press London UK

146 Maebara H 1955 Spatial distribution of lastinstiarlanaeof purplish stem borer (Sesamia iqferens Walker) hibernating in an upland rice field Bull Kagoshima Univ Fac Agric 438-41

147 Mallick E H and N G Hajra 1977 Response of rice plant to defoliation treatments at different developmental stages Indian Agric 2151-56

148 Mansingh A 1971 Physiological classification of dormancies in insects Canl Entomol 103983-10)9

149 Martell C G 1974 Catalogue of plant feeding insects of Mexico [in Spanish] Fitofilo 27(69)1-176

150 Martinez 0 J 1982 Principal rice pests in Ilanos orientalis [in Spanish] P~ages 80-94 in Curso deact nalization de conocimientos en arroz Villaniencis Instituto (olonmbiano Agropecuario

151 Martins J F da Silva F Ferrcira A S Prabhu F 1 P Zimmermann 1980 Use of pesticides for the control of the main subterranean upland rice pests esqui Agropecu Bras 1553-u

152 Martins I F da Silva F Fe eira rod BdamSilveira Pinheiro I Q2 Damage simulation caused by defoliators of upland rice [in Portuguese with English summary] Pesqui Agropecu Bras 171113-1119

153 Martins 1F da Silva N(iyuen van Tan and 1Ida Silveira Pinheiro 1981 Stem borer resistance in upland rice associated with plant morphological characters [in Portuuce Iesqui Agropecu Bras 16187-192

154 Masaki S 1980 Sumnmer diapause Ann Rev Entomiol 251-25 155 Masuzawa T 11 Suwa and F Nakasoji 1983 l)ifkleences of

oviposition preferencc and survival rate of two skipper butterflies Parnara guttata and PIchlpidai iathiais (Iepidoptera HIesperiidac) on rice plant and cogon grass New Entomol 3247-56

156 Maurya D M and C P Vaish 1984 Upland rice in India Pages 69-92 in An overview oftpland rice research Proceedings ofthe 1982 Bouakamp Ivory Coast upland rice workshop International Rice Research Institute IO Box 933 Mnila Philippines

157 Menon M G R R N Katiyai S Kuar and S S Misra 1970 Termites attacking the paddy crop Sci Cult 36294-295

158 Miyahara Y and 1 Nishikubo 1967 Behavior of young nymphs of the red rice root aphid Iholahsipiun ri liabhdontinalisSasaki in relation to movement from the leaf to the roots of upland rice [in Japanese] Pioc Assoc Plant Prot Kyushu 1363-66

159 Mo ljoa Tjien 1952 White grub of upland rice (Holotrichiahelleri Brsk) [in Indonesian with English summary] Contrib Agric Res Stn Bogor No 13134 p

160 Mochida 0 and T Suryana 1979 The occurrence of the brown planthopper and ragged stunt on lowland and upland rice ill Central Java and logyakarta in tile late of January 1979 Entomnolgy Survey Report No 15 Central Research Institute of Agriculture (CRIA) Sukamandi Indonesia I February 1979 9 p

161Monet C 1979 Varietal resistance to upland rice pests [inI French] Pages 35-39 in Annual report 1977 Institut des Savanes Department des Cultures Vivrieres Bouak6 Ivory Coast

162 Monte 0 1942 A caterpillar ofrice-fields[in Portuguese] Biol6gico (Sao Paulo) 8161-163 (Rev Appl Entomol A31414)

163 Munroe I) D 1979 Pest management strategy for hill rice in four resettlement areas in Sarawak Malaysia Pages 89-102 i1 Guidelines for integrated control of rice insect pests Food and Agriculture Organization Plant Production and Protection Paper 14 Rome Italy 115 p

40 IRPS No 123 January 1987

164 Nakasuji F M Ishii and Jen-Zon Ito 1984 Rice skippers in Taiwan and their life histories ZZrC (Taipei) Tech Bull 82 10 p

165 Navas I) 1976 Fall armyworum in rice Pages 99-106 il Ecological animal control by habitat management Proc Tall Timbers Con-ference 28 Feb-I Mar 1974 Fall Timbers Res Stn Misc Publ 6 Tallahassee Horida USA

166 Nickel 1 1 1961 Biological control of rice stem borer A feasibility study Tcch Bull 2 International Rice Rcarch Institute PO Box 933 Manila Philippines IIl p

167 Nigan P M and R A Vernia 1985 Insect pests of uplaild rice in Uttar Pradesh Int Rice Res Newsl 10(6)22

168 Nilakhe S S 1985 -cological observations on spittle bugs with emphasis on their occirence in rice Pcsqtui Agropecu Bras 20407-414

169 Nixon G EJ 195 1 lie association of ats with aphids and coccids Commonwealth Iusti lcte of Entonology Iondon UK 36 p

170 Nurdin F an Ishak Manti 1985 Study oit seedling fly of upland rice Interlational Upland Rice Moitoring Tour Report 26 lebruary-2 MarLI Sukaranti West Sumatra

171 Oka 1 N 1979 Ciltural conttol of the brown planthopper Pages 357-369 in Brown planthopper threat to rice production in Asia International Rice Research Institutc 1p0 Box 933 Manila Philippines

172 Ohni M and A Vilnaii 1975 Rice insect pests of Italy biology of Sipha grcria (Kaltenbach) and its control [in Italian] Riso (Milan) 2459-71

173 Otanes F Q atnd 1 I Sison 194 1 Pests of rice Philipp J Agric 12211-259

174 Painter R I 1951 Insect icsistance in crop plants University Press of Kansits Lawrence Kansas USA 520 p

175 Pantua P C and 1A litsinger 1981 Arthropod abundance in continuous vs biannual rice cropping systeis Int Rice Res Ncwsl 6(2)20-21

176 Pathak M ) and V A l)yck 1975 Sttudics ott insect pests of upland rice Pages 186-191 in Major research in upland rice International Rice Research Institute PO Box 933 Ilanila Philippines

177 Patil B R B M IHasabe and 1) S Ajri 198 I Chemical control of white gritb lIditriihinhorraia Fabr intfesting groundntut paddy atnd sorghum Indian 1Entomol 43232-236

178 Pawar A 1) 1974 (ollection and identification of insect pests of rice rlerminal repirt lInternational Rice Research Institute PO Box 933 Manila Philippines 72 p (tnimco)

179 IerrajU A aid A S Reedy 1965 Observations ott diapausc itt tie steti borer ohrice Rice News Teller (lai-Mar)9-13

180 Pont A C 1972 A review of the Oriental species of theri ona Rondani (I)iptera Muscidac) of econtomic importance Pages 27-102 in Control of sorglium shoot fly M G lomtani and W R Young eds Oxford atd IH1 Ptthl Co New I)clhi India 324 p

181 Prasad S C 1PChand aid M 1PSingh 1983 Incidence of rice gall midge in the upland plateau region of Hihar Oryza 20235-236

182 Rai P S 198 0 Nisaga simpex damage to rice in the hill tracts of South India Int Rice kles News[ 5(I)12

183 Rathorc Y S 1969 Gonocplhahli dcprssunt Fabr as itpredator of (tilo zonwIlhis (Swinhoc) and other moths in field cage at Patnagar Uttar Pradesh Indian 1Entotnol 3127

184 Reed W 1983 Crop losses causcd by insect pists in the developing world Proceedings 10th Internatial Congress of Plant Protection lrighton UK 174-80

185 Reedy M V 1981 Ateactogasterinhens(Walker)a new root-grub pest of tipland paddy and inaize Curr Sci 501076

186 Reveno M A 1968 Catalogue of the insect pests of economic crops in Colombia [in Spaiish] Asociacion Latino Americana de Entomo-logia (AIAE) 129-33

187 Reyes G M 1957 Rice dwarf disease in the Philippines Food and Agriculture Organiiation Plant Prot Bull 617-19

188 Rice S E A A Grigarick and M 0 Way 1982 Effect of leaf and panicle feeding byarmyworm (Lepidoptera Noctuidae) larvae on rice grain yield Environ Entomol 7593-595

189 Risch S J 1) Andow and M A Altieri 1983 Agioccosystem diversity arnd pest control data tentative conclusions and nizw research dirctions Environ Entomol 12625-629

190 Rivera C T 1956 Three organic insecticides in thecontrol of insects affecting upland rice Philipp Agric 39465-472

191 Roffrey 1 1972 Locusta outbreaks in the Philippines Acrida 1177-188

192 Rojas R 1) 1941 The effect of pruning at varying ages upon the yield of Ramairio Phipp Agric 29851-860

193 Roonwal M i 1979 Ternite life and termite control in tropical South Asia Scientilic Pnbl odhpr India 139 p

194 Rosetto C1 F da Silva Martins N C Schmidt and L EAzzini 1978 Damage caused by pasturespittle bugs 4Dvoi)lavopictaandDJ schach) on rice lBragantia Nota 6 (Matcli)25-27

195 Rosztto C1- S S Neto I- Amantc I) Iink 1Garcia-Vieira D M de SoLza N V Bansa t and A M de Olivcira 1971 Rice pests of Brazil [in Portuguese] Pages 149-238 inrFA()-International Rice Commission 2nd meeting of the Rice Committee for the Americas 6-11 l)ecember 1971 Pelotas Rio Grande do Sul Brazil

196 Rothschild G H L 1969 Obseratiois on the armyworm Spodoptera matritia acronrctoiiles Gn (Lep Noctuidae) Bull Entomol Res 59143-160

197 Rothschild G 1- L 1974 Parasites of the armworm Spolopiera mauritiaacroncitoihes in Malaysia Fntornopliaga 19293-299

198 Sachan i N S K Gangwar and 1N Katujar 1980 Field crickets damaging cages in North Eastern IHill Region Indian J Entomol 42526-527

199 Sands 1) P A 1977 The biology and ecology of Lepocorisa (Hemiptera Alydidac) in Papua New Guinea Res Bull 18 Departshyment of Primary Industry Port Moresby Papua New Guinea 104 p

200 Sarawak 1968 titomology division Pages 97-106 in Report of the research branch of the Department of Altriculture 1966 Kuching (Rev AIppl Eintomol A56603)

201 Satpathy JM M S Das and K Naik 1977 Effect of multiple and mixed cropping on the incidence of scine important pestsI Entomol Res 178-85

202 Sauer IHF G 1939 Notes Ott Elasmpnlpualtsligniosellths Zeller (Lep Pyr) a serious pest ofcereals in Sao Paolo [in Portuguese] Arq Inst Biol 10199-206 (Rev Appl Entotnol A28378)

203 Satinders 1L A 1 SKing and C L Vargas S 1983 Crop pests of Central Anerica with t bibliography [in Spanish] Centro Agroshynotnico Tropical de Investigacion y Ensenanza Turrialba Costa Rica (CATIE) Tec Bol 9 90 p

204 Sauphanor B 1985 Some factors of upland rice tolerttce to stem borers in West Africa Inseu Sci Appl 6429434

205 Sen P atnd S Chakravorty 1976 Stentflics in Haritighata Nadia West Bengal India [it Rice Res Newsl 1(12) 15

206 Senapai B and 1M Satpathy 1983 The seedling fly Atherigona orrzae Malloch is a pest of rice in Orissa (India) J Entomol Res 777-79

207 Shhjahan M S Alatn M Ah nad and BN Islam 1983 A new scarabaeid beetle pest of dryland rice in Bangladesh Int Rice Res Ncwsl 8(I)16-17

208 Sharma G L S C Modgol and R C Gautam 1979 Effect of depth of submergence ott incidence of bacterial blight and white grub infestation in transplanted rice Int Rice Res Newsl 4(6)12-13

209 Sharma S 1985 Scarabaeid beetle outbreak itt upland rice Int Rice Res Newsl 1(1(l)24

210 Shiraki T 1917 The paddy borer Schoenobhits incertulas Walker [achaku Agric Exp Stn Formosa 256 p

211 Silva A de Brito and 11P Magalhaes 1981 Insect pests of rice [in Portuguese] Circular Tecnica No 22 EMBRAPA Centro de Pesquisa t ecuaria de Tropico Unido (CPAJU) Belem Brazil 14 p

212 Singh S A 1979 Emergence of rice leaf butterfly as a rice pest in Manipur India int Rice Res Newsl 4(4)16-17

213 SinghS A 1983 Changing trends of insect pestsof rice in Manipur Int Rice Res Newsl 8(f)12

214 Siwi S S and P H van Doesburg 1984 Lenovorisa Latreille in Indonesia (Heteroptera Coreidae Alydinae) Zool Meded (Leiden) 58117-129

215 Soejitno J 1977 Reaction of some upland rice varieties toseedling fly eltherikonaexigua Stein [in Indonesian] Siri Hana I Penyakit 630-35 Lembga I osat Fenclilian Pertanian Bogor Indonesia

216 Socnardi 1967 Insect pests of rice in Indonesia Pages675-683 in The major inse pests of thu rice plant Proceedings of a symposium at the Internatital Rice Rec Wi itt S pternber 1964 The Johns Hopkins Press Baltimore Maryland USA

217 Soto P E A T Petcz and I W Buddenhagen 1976 Survey of insect pests and diseases of rice in different ecological zones in Nigeria Rice Entomol Ncwsl 435-46

218 Soto 11E and Z Ziddiqi 1976 Screening for resistance to African rice insect pests Varietal improsetnent sCmi n r Bouak Ivory Coast 13-17 Sept 1976 West Africa Rice l)evelopment Association Monrovia Iiberia

219 Squire F A 1972 Entomological problems in Bolivia PANS (PestArtic News Sumr) 18249-268

220 Stapely 1 II 1978 The rice leaf roller in the Solomon Islands Int Rice Res Newsl 3(l)13

221 Suenaga H 1937 On aphids attacking the roots of rice Oyo-I)oubts Zasshi 9150-152 [in Japanese] (Rev Appl Entomol A25745)

222 Suenaga H 1952 On the seasonal abundance of three subterranean aphids on upland rice Bull Kyushu Agric Exp Sti 1249-262

223 Sutynso Yas and Dandi Soekaria 1964 Beberapa hana yang ttenperang tana man padi tanah kering [in Indonesian] Rapat kerja Padi Tanah Kering 6 sd 8 Juily 1964 Kaliurang 9 p

224 Tallamy I) W and R F Denno 1983 Alternative life history patterns in risky environments in example from lace bugs Pages 129-147 in Insect life history patterns R F l)cnio and If Dingle eds Spring-Verlag 225 p

225 Tamura 1 1939 Observations on some insect pests of field crops [in Japanese] 1 Plant Prot 26414-420 (Rev Appl EntomolA27-556)

226 Tamura and Suzuki 1964 Influences of the early-transplanting practice in rice culture upon development of the rice stem borer Chilo siql)ressalisWalker Bull Ilokuriku Natil Agric Exp Stn 761-94

227 Tan J P 1924 The rice root aphid Drowhi ihirsoia Philipp Agric 13277-288

228 Tanaka 1 1957 Taxonomy and distribution of some subterranean aphids injurious to upland rice in Japan with description of a new species [in Japanese English summary] Botyu-Kagaku 22168-176

229 Tanaka T 1961 The rice root aphids their ecology and control Spec [full College of Agriculture Utsunomiya University Japan F3 p

230 Tauber M IC A Tauber R Nechols and 1Obrycki 1983 Seasonal activity of parasitoids control by external internal and genetic factors Pages 87-108 itt I)iapause and life cycle strategies in insects V K Brown and I Ilodek eds )r W Junk The Hague Netherlands

231 Taylor W F 1972 Effects of artificial defoliation (simulating pest damage) on varieties of upland rice Exp Agric 879-83

232 feotia T P S and S Natud 1966 Biotoomics of the rice skipper Parnaraniathiis Fabricius (Lepidoptera Hesperiidac) Indian J Entomol 28181-186

233 Teran F 0 1971 Rice pests of Santa Cruz [in Spanish Ministerio de Agricultura Sita Cruz Bolivia Bull 45 8 p

234 Than Itun 1976 Population dynamics of the yellow rice borer Tryporza itcertulas (Walker) and its damage to the rice plant MS thesis University of the Philippines at Ios Bafios Laguna Philippines 92 p

235 Uichanco L B 1911 Pests Pages 55-74 it Rice Philipp Agric 11-200

IRPS No 123 January 1987 41

236 van der Laan P A 195 1 Life history of the rice seedling fly Contrib Gen Agric Res Stn Boor Indonesia 118 15 p

237 van der Laan 11A 1959 Correlation between rainfall in the dry season and the occurrence of the white rice borer (Scirpolhaga innotala WIk) in lava Entomol Fxp Appl 212-20

238 Van Driesche R G 1983 Meaning of percent parasitismin studies of insect parasitism Fnviron Entomol 121 I1- 122

239 Venugopal M S and A litsingcr 1984 Fffect ofcarbofuran on rice growth Prot Ecol 7313-317

240 Vietnam Directorate of Rural Affairs 1962 Annual work progress report on crop improvement program of rice sugarcane vegetable and field crops (lor the period from July 1961 to June 1962) (11) 299 p (Rev Appl Entomol A51526-527)

241 Wan M T K 1972 Observations on rice leaf- and planthoppers in Sarawak (Malaysian Borneo) Malay Agric 148308-335

242 West Africa Rice I)esclopnient Association 1978 Annual research report Vol 2 Statistics Monrovia Liberia

243 West Africa Rice Development Association 1979 Annual research report p47 Monrovia Liberia

244 Williams ) F and CS lofgren 1981 Fli I illy l-468 anew bait toxicant for control of the red imported lire at Fla Entomol 64472477

245 Williams I) 1 1970 The mealybugs OFlomoptera Coccoidea Iscudococcidac) of sugar-cane rice and sorghum Bull Entomol Res 0109-188

246 Williams D) 1 1982 Pulinaria icervi (Signoret) and its allies ott sugar cane and other grasses Btll Entoinol Res 72111-117

247 Wirjosuiardjo S 1976 Study ofchenmical cont rot of insect pests ott upland rice Imiu Pertanian (Jogjakarta) 1323-329

248 Wirjosuhardjo S and N Wirarta 1976 Chemical control of the white grub tIolorithiatlltriin upland rice in M uneng Irobolinggo Indonesia) [in Indonesian] Imut Pertanian (Jogjakarta) 1330-332

249 Yano K T Miyake and V F Eastop 1983 The biology and economic importance of rice aphids (Henuiptera Aphid idae) a review Bull Entomol Res 73539-566

250 Yelp D and P Rhodes 19(4 lhe success of drilled dryland rice iii the Fiji Islands International Rice Commission Working Party on Rice Production and Protection Manila Philippines 3-10 March1964 Food and Agriculture Organization Rome Italy 13 p

251 Young W R 1981 Fifty-five years of research on the sorghum shootfly Insect Sci Appi 23-9

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No 101The economics of hybrid rice production in (hina No 113 Boron toxicity in rice No 102 Rice ratooning No 114 Energy analysis rice production systems and rice research No 103 Growth and development of the (Jeep water rice plant No 115 Production risk and optimal fertilizer rates an application of the

No 104 Faridpur acomputer-assisted instruction nmodel forrainfed lowland random coefficient model No 116 Consumer demand for rice grain quality in Thailand Indonesia andrice

No 105 A readingand listening comprehcnsion test inEnglish for non native the Philippines speakers applying for tiraining at IRIRI No 117 Morphological changes in rice panicle development a review of

No 106 Rice grassy stunt virus 2 a new strain of rice grassy stunt in tire literature

Philippines No 118 IRRI-Korca collaborative project for the development of cold-No 107 Physical losses and quality deterioration in rice plstprodnCtion tolerant lines through anthei culture

systemis No 119 Problen soils as potenial areas for adverse soils-tolerant rice No 108 (olptblication ofIRRI materials a survey of translators and varieties in South and Ssuthcasi Asia

publishers No 120 Changes isinall-farm iLtireliing techii)hgy in Thailand and [he No 109 Classification of Philippine rainfall patterns Philippines No 110 Contributions of modern rice varieties to rutrition in Asia No 121 Landforms and modern rice varieties No Ill Changes in rice breeding in 10 Asian countries 1965-84 No 122 Yield stability and modern rice technology No 112 Design parameters affecting the performance of the IRRI-designed

axial-flow pump

Page 27: UPLANDRICE. INSECTl--1i M T -E!R.EOLOU t RTANCE ,AND …delphacid.s3.amazonaws.com › 6958.pdf · The significance of abiotic factors in upland rice insect ecology will be apparent

In Brazil screening Ior insect resistance focttses oil lou I pest problems Varieties tolerant of lasmolwalpus ligi-sellus and resistant to )iatraea saccharalis have been identified (70 153) Current work also emphasies 7iraca limativetntris and Thmialspis (=Iosis) flavouicta (66) These insect pests Ilt onlV citise IIigh yield losses tihey also ire difficult to control chemically ofr by other ilealls

ManV upland rice vaieties aic early maturing and are resistant to pests such as gall midge (I Xl ) Early mIturity is often a highly desirable trait to esuilpe pest buildup but it is not true resista nce bca usc ilthose va reties are plinted late they will be itsdamaged as ain early planted susc ible variety

P~erhaps tile cosiiiopolitall of all upland rice pestsimost are the sced pests (ripenling stagc) which pose a particular problem beta use scuds re ised IsIO for hitnit4ans Ayltoxic substatice introd tcld geticticllyilmay also be toxic to nal iniless it caln he placed old in thfe IdhuffSelectmi fo hairiness or lotng wns kill mIlct sthilrohleiiis dutilig threshing pa rtictfarl maitial threshinfl [hireshers ill complain of itcfimin atid itteascd (fust levels

Attempts itIRRI to fitid tcistatncc to the rice bug plorisa have ilotbeCnl prmitisilg lIIetIattonlids iorc

through the lenilna or palca hillhairs sedstVIor seeds wilh awns that deter bugs promote complaints from threshers Seed bugs liptocorisa and Stcnocoris enter the seed through the opening between tile lemna and palC but varieties with narrow openings are diifficult to mill

Breeding priorities should concentrate ott inotCI gruLtLIps that are most difficult to c nttrol by other meains Steil) horers appear to be the most widespread group that sigiiificantly dalag s rice aind is hard to manage by other Ilnca us

Biological control Very fe stutdic of mitatt~l eneties of rice pests in upland envirollnletits wee in tile wefouiid literature Therefore have not compiled a list of species Ve present a list of natural enemies iOtid in upland ricefields tuic latialln Batangas for some conitno iOldiar pest species ([itble 5)

Muich ofl thc lollowinle itfor ittiol cotlliCs rol Stutdies in the lowlatnds or roi crops otithcr thlin tice

Qu(ailtr o 11ll t11t( Nat trrtf lii hcoeits ight Ile used agaitist ile social insects tilt and termites Sollie predator specice live ill theirf nests n ilnic theirnllt ip-pearanIlCe aMRidbehavior IJtrouMdCing a mote aggressise ant species to displace pest ant codonies is a possibility (169) Patihogens as potential natural encnies have reccived little atteition (I13) Natural ncnuics asa group however hac not been toIl iporta it in Iegulating aintshown be and ternite numbers

The nocturnal behavior aid subteitmien habitat of lole crickets and field crickets protect them from many ntllral enemies There are only limited records of natural cnelics for these groups of pests some pathogens (87) but mainly

IRPS No 123 JattuIny I 271987

parasites - scelionids on eggsand sphecids on ilymphs and adults Sphecid wasps Larra carutmaria (SImitI hI L huzwuensis Rohwer and lsaltiuhta Willi ams speciali in mole crickets pa ralyzing them nd dragging the ninto thcir nests as food lot their voting ()tier sphCcid species Lins aurulenoa (Iabricius) lot maiac (Ashmead ) 1 lhshylssellaus (Smith) and Ml lal)oriosu (Snithf specialie in field clickets

White grUbsO the other hand hae Icti scriuns enuigh to have been extensively studied for cotttroi 1W nati al enemies( 112 140 141) Scoliid parasites C(ajsoMcispp have beetn intrtduced itto tilePhilippites for control of Ielxl ]udisiro la The rcsearchcts w Intttrdced themle during ill era claimed that the white gttionthreak as controlled (142) howevcr it isnormal for pest epidemics to subside and there was no diect evidence to indictte that tfe parasite was responsible A tmore promisitg titctllod might he to identify iso ltcttass prt~f tcartd apply hiactelaln fingal pathogens to tile soil Ihe lalattese beetle in North America has hcci coitrolled hv tite conmnleciall aailable hactcrial prcpa ration itacihlpu pav ofr milksdiscas (71) Spote prtyefott the soil itloted ilrtier reitiain actic fr Veas

Root aphid sardtnctly bugs readilslic tilib to predatots and parlsites once their tetndine ants are controlled The larvac and adtilts of a coccinellid beetle 1Scynms sp prey on llriadlouais in slash and burn upland rice Baits treated with insecticide may be used to control tending ants by killing their ourig (244)

Root-tkcditig blugs false wireoris wireorms and root wccils aic r ofsuch little ifilpoltatice that thenomttll role of liatural Cticniics hias tiot beei asscssed A possibility hoevrl tiglt he tit ItaIIitic niaCIMtodcs lintt1g11en Itese soil-itlhlfbitine mCtitodcs W5uhtfhave to be imiass Produiced

large miltibcr of niatltrll cnemics ircknown ft CltIoriS IC IpblCeii is thtt clttvortnlls ofonlie the crop soott alter land ptel)tritiOti a1tithe bcgit iniig of the raittv

Citsoln hfien tiatllral leniiiy populations aric lo Colltloil WOtuld iii ole mass prodtticing andirclcaslng key spccies

le egg stage is the fpirt (olthe Cedlitgt Ialot life cycle tI hich to f)ctis ftor tittlril elctit cottntrol hplraites o1 pleditors ttsit elleggs highly expoIsYCl I Sgg pailsites or predators stolld probahly e ittass andtbe produced released Littlh i toruialioll is a ailable ott what species to trV o ho to Iniss podLice thiCIi A etiloplhid wasp Icmihlaol sp effctiicly regushy

lates the leatf lcloalapolm2a spioanoiiitr heat andc rice antid should be collsctvcd

leaf beetles whCllose e am] eggs ate laid in the grutindhirO ould be motC difficutlt to control by niatulral enelV Mnll ipuhltiOn But species sUici IS c_gsthispid beetles ith

and larVle oilleascs ate vtilcratble to iparaites and pledators

Natural enemisc of tlirips otl rice are little known

28 IRPS No 123 January 1987

Table S Parasites recorded from upland rice insect pests in Tanauan Batangas Philippines 1977-84

Order Family Species Host stage

I lymttenoptera Braconidae Leaffolders Cnaphalocrocis Marasinia spy

Cardiochilesphilipplinensis Ashmead a Larva

Cotesia angustillasis(Gahan)Cotesia or cypris (Nixon)

Larva Larva

Cotesia or taeniaticornis(Nixon) Larva Cilonusinotakatae Munakata Chelonits SppAlacrocetrts philippinensis (Ashtmead)

Egg to larva Egg to larva Larva

Iclineumonidac Orgilts spp Isclutojppa hireator(Fabricius)

Larva Larva

Itoplectis naraigav(Ashmcad) Larva Tenelhtcha philippiensis (Ashmead) Larva Tenelchha stangli (Ashmead) a

irichomma cnaphalocrosis Uchida Larva Larva

Bethylidac Elasmidac

Goniozms or trianguli r Kieffer lasmtusalhopictus Crawford

Larva Larva

Diptera

Encyrtidac Chalcididac

Tachinidac

Elasmtus spp Copidosomopsisnacohiae (Fidy) Brach nteria evcarinata Gahan Biachyieria lasus (Walker) Ztgohothria ciliata (Wulp)

Larva Egg to larva Pupa Pupa Larva

Argyrophylx nigrotihialis(Blaranov) Larva

l-ymenoptera Braconidac Yellow stern borer Scirpopiaga incertulas

Braco chinensisSzeplegeti Larva CIthelotis spp Chelonus niunakatae Munakata Cotesi flaripes Cameron Stenoracon nicerilli (Bingham) a

Larva Egg to larva Larva Larva

Ichneumonidac Tropobracon schoenobii (Viereck) Amatroinorpiaaccepta metathoracica(Ashmiead)Frihorus sinicus (I lolmgren)

Larva Larva Larva

Ischnojoppaluteator (Fabricius) Larva Isotima nr dammeratiti Rohwer Larva Tenelucha philippinensis (Ashmead) a

Larva Tenehicha stangli (Ashmcad)Trichomma enaphalocrosis Uchida

Larva Lara

Chalcididac Eulophidac Trichogranmmatidae

Xanthopimnpla stemmator (Thunberg) Charops hrachYpterumn Gupta and Malteswary Brachtm eria sppTetrasticttus schoenohii Ferriere Trichorigranm chihunis Ishi

Larva to pupa Larva Pupa Egg Egg

Diptera

Scelionidac Pteromalidae Tachinidae

Tt ichograintnta aponiicuitt (Ashniead) Telenomits rowani (Gahan) a

Trichoinalopsis apattelctena (Crawford) Jeirhaea spp

Egg Fgg Egg pupa Larva

Z4gobothria ciliata (Wulp) Larva

Ilymenoptera lecneumonirlac Gol fringed borer (hilo auricilius

Trichomm enoptalocrosisUchida Larva

Ieioenoptera Braconidae

Aantholimlh steiniator (Thtinberg) a

Pink stet Ioret Sesamia inferens StoInacoil nicrilhi (Bingh am)

Loarva to pupa

larva

Diptera Ilyinenoptera

Armvtnyorms Mythimna separata Spodoptera mauritia acronyetoidesTachinidae Zvgohothriaatropiora( Robinea i -Desvoid y)Eulophidae It-phctrtiscitapadoe (Ashimead)

Larva Larva

(halcididae Braconidac

Brachpin ria spip (oesia spp

Larva larva

IHyntenoptera I)iptera

Rice skipper Ielop ida (halcididae Tachinidae

math ias aidgreen iortedci ciliarNIelan itis eda Brat lhincria sp iii matuqitatt Cameron Irgyrolhirlaxi trotlhiais(marumtv)a1

ismtne Larva Larva

Diptera Ilymenoptera

Tachnidae lelineutionidae

fuown se tilo per Mois frugalisIrgtrophrlax 1igrothiahiv 3 a rat ov) anthopimpa pmnctata (Iabricius)

Larva Larva to pupa

aMost dominant

IRPS No 123 January 1987 29

Normally predatory thrips prey on herbivorous thrips strepsiptera on both Spiders dominate tilenymphaladultNatural enemies of stem bugs --Scolinoplara and Tibraa predators Spittle bugs move to rice from pasture grasses - are in order of importance egg par~isites and predators and biocontrol would be successful only ifcarried out before fungal pathogens and nymphal adult parasites Egg tiley immigrate into ricefields parasites could either be introduced fron other areas or Biological control tactics for hemipterous seed bugs mass produced and released ould follow those outlined for stein bugs

Armyworms are normally lwld in hcck by the activities Coml)arison of environnewtns Natural enemies are of egg and larval parasites When these natural enemies fail perhaps faced with even greaterconstraints than insect pestsusually because of drought armyworns become epidemic to survive ind reproduce in upland environments TheyIt may be worthwhile therefore to release parasites during have adapted ani survived by having wide host ranges the the rainy months following a drought Virus diseases of ability to aestivate or hibernate over unfavorable seasonslarvae would beanotheravnue to exploreforarnyworms and the ability to disperse or by other mechanisms

Parasites have ben traditionallyconsidered for biolegical Comparing the most dominant and therefore the most control attempts against stein borers usually by rekasing adapted species of natural enemies collected in various exotic species( 166) This approach has not met with success environments in the Philippines we can understand how and new avenues should be explored The role of predators different rice environments are arid pathogens is little inderstood and deserves greater Anagqrms olptahiisa mymarid planthopper egg parasiteattention Egg parasitism rates are normally reasonably is cquallyadapted to all Philippine rice environments (Tablehigh but should be supplemented with effective egg 6) It is selective for planthoppers and even parasitizes eggspredators Orthopterans are potential egg predators of the n-iize planthopper Peregrinitsmaidis itt upland areas Metioche vittaticollis (Sta ) Anavipha nifiwnnis (Saus- Alavohis also parasitizes maize planthopper eggs but is sure) and Conocephahs hongipnni (de i-laan) feed on most prevalent on BPH in the irrigated wetlands (onashystem borer eggs etioche and Anaxil)ha specialize in eggs tocerus spp like A optalhilis on BPH and WBPI Iattacks with no hair such as Chilo spp The omnivorous (ono- the eggs of GLH in ell rie environments It is also found cephahsspecializes in eggs cov red with hairs it is a proven parasitizing the eggs ef the white rice licahopper (oimna egg predator of Scirlouphaga spp but has the discouraging spetra Oirosita naias a trichogrammatid egg parasitehabit of eating rice guin (65) attacks BIPH and WBPH in the wetlands and has no other

Grasshoppers have egg (scelionid) and nymphalad ult host A related species 0 aesopi specializes in G1-H in (nemestrinid tachinid and sarcophagid) v irasites Among wetland environments Dryinid strepsipt-an and pipunshypathogens protozoans have been recorded most frequently culid nymphal and adult parasites of planthoppers and Control of locust species by natural enemies could con- leafhoppers also specialize either in hopper species or centrate on their habitual breeding areas environments Among the dryinids Pseuhdgonatoipsflavi-

Leaffolders have rich complexes of natural enemies some feimur prefers the lowlands and BPH while P nuch has no adapted to upland rice (22 27) ranging from egg predators environmental prefirences but is adapted only to WBPH (gryllids coccinellids) to larval parasites (braconids ichncu- The main d ryinid species on GLH aptogonatop)usspp is monids) to larval prcdators (ants and carabids) to larval most dominant in rainfed lowlands and also attacks other pathogeiis (vira and fungal diseases) Efforts to augment leafhoppers within Amnrasca Cicadulinaand Bachaltuathese natural enemies should focus on the egg predators Strepsiptera are more dominant on plarithoppers than (rearing releasing)and viral or fungal pathogens (culturing lealhoppers The Elenchus species in the Philippines attack spraying) all species of planthoppers within the genera Sogawilla

Parasitization of the large conspicuous larvae of Sogatodels NilaparsataHarualia and Opiconsiva Elenshyskippers green horned caterpillars and other polyphagous cusrvasuntaisuiis adapted to the major rice environments Lepidoptera by tachinids and chalcids is normally low ELg but prefers BPH in rainfed arid WBPIH in wetland environshyparasites and predators are perhaps the key to their control ments Ilalictoph(utsmtnroeiand HVpectrus attack G LH

Gall midges are parasitized by pterormalids and platy- and COfimravcctra respectivelygasterids that aestivate within their larval host between Pipunculids are mainly parasitic oii lealhoppers Each seasons Shoot aphids are preyed upon by coccinellids species has a unique environmental preference Tomosshysyrphids and chrysopids varireIa oraetoraprefers Ne7 otetltix nigropictrs and

Planthoppers arid leafhoppers have egg parasites and Pilunwulhtsjavanensisattacks Dehoceplhahls spp which predators as well as nymphialad nlt parasitesand predators are more abundant in upland than irrigated lowland their rich complex of natural enemies includes fungal environments T suhvirescensattacks Nephotettix virescens pathogens The principal egg parasites are myrnarids and and prefers the lowlands Pilpuiulus nttillattls is an trichogramniatids Egg predators are norinally mirid plant upland rice parasite arid attacks only Nephotetix speciesbugs that suck out the yolk Nymphal ad nIt parasites are More than 17 recorded species of leaffolder parasites in dryinids on planthoppers pipunculids on lealhoppers and the Philippines attack species of C(naplwocrocis and

30 IRPS No 123 January 1987

Table 6 Dominant natural enemies of rice pests in upland rainfed lowland and irrigated lowland rice environments of the Philippines

Natural enemy

Host stage Species

Brown planthop-Parasite Egg

MymaridaeAnagrus optabilis(Perkins) Anagrusflaveolus Waterhouse

TrichogrammatidacOligosita naiasGirault

NymphAdult Dryinidac

Pseudogotatopuslaiifemttr Esaki et Hashimoto Elenchidae

Elenchus yasurnatsui Kifune and Hirashima

Environment

Upand Rainfed lowland Irrigated lowland

Ailaparata lugens

x x X x

x

x x

x X X Whitebacked planthoppers Sogatella furcifera and Sogatodes pusanusEgg

MymaridaeAnagrus optabilis(Perkins)

TrichogrammatidacOl~gositanaias Girault

NymphAdult Dryinidae

Pseudogonatopusnudus Perkins Elenchidae

Elenchus yasutatsui Kifune et Hirashima

Parasite Egg

MymaridaeGonatocerus spp

TrichogrammatidaeOligositaaesopi Girault

NymphAdult Pipunculidac

Tomosvaryella subviresccns Loew Tomosvaryella oryzaetora (Koizumi) Pipunclusmutillatus (de Meijere)

DryinidaeIlaplogonatopusspp

Egg Copidosomopsisnacoleiae (Eady)

Larva BraLonidae

Cardiochiles philipphnensisAshnead lchneumonidac

Trichiomnia cnaphalocrosisUchida Temelucha stangli (Ashmead)

Pupa Chalcididae

BrachymeriaexcarinataGahan Brachymerialasus (Walker) Brachymeria nr tachardiaegroup

Parasite Egg

Scelionidae Telenomus rowani (Gahan)

EulophidacTettasticusschoenobhi Ferriere

Larva Braconidae

Stenobraconnicevillei (Bingham) Iclhncurnonidac

Temelucha phiiippinensis(Ashinead)Temelucha stangli (Ashmead)

x x x

x x

X x X

x x Green leafhopper Nephotettix virescens

X x x

x x

X X X X

X

X Rice leaffolders CnaphalocrocisMarasmia

x X x

X

X X

x x x x x X x x x

Yellow stem borer Scirpophaga incertulas

X X x

x

x

x X XAinauromorphaaccepta metathoracica(Ashmead) x

continued on next page

IRPS No 123 January 1987 31

Table 6 cntinued

Natural enemy Environment

Host stage Species Upland Rainfed lowland Irrigated lowland

Larva-pupa Gold fringed borer Chilo attriciiuslchneumonidae

Xanthopimpla stemmator (Thunberg) x aPupa x

Pteromalidae Trichoinalopsis apanteloctena (Crawford) ax

Pink stem borerb Sesamnia inferensParasite

Larva Braconidae

Stenobracon nicevillei (Bingham) x b b Armyworms Mvthinna SpodopteraLarva

Tachinidac Zygobothriaarroptvora (Robineau-Desvoidy) x x

Braconidac Cotesia spp

X Skipper Pelopidas mathias and green horned caterpillar Melanitis leda ismeneF-gg

TrichogramrnatidacTrichogrannaspp x

Scelionidae Undetermined Scelionid

x Larva

Tachinidae Argyroph lax nitrotibialis (Baranov) X x

Pteromalidae Trichomnalopsis apanteloctena (Crawford)

X Predator

CoenagrionidaeAgriocnenis femina femina Brauer x xIschnura senegalensis tambur

Miridae CrtorhinsliidipennisReuter x x

Nabidac Stcnonabis tagalicus (Stal) x

VeliidaeAlicrovelia douglasi atrolineata (Bergroth) x

Coccinellidae Micraspis crocea (Mulsant) X x x

Vespidae Eumenes campaniformis (Fabricius) x

Formicidac Diaecamna spp xOdontoponera transversa (F Smith) x

Carabidae Chaenius spp OphioneaW- aJasciataSchmidt-Goebel

x

Ophionea ishii ishii Habu x x X

Gryllidav Anaxipta longipennis (Saussur-) x xMetioche ittaticollis (Sial) x x x

Tettigoniidae Conocephalus longipennis (de Haan) x x x

aNo rearing made on rainfed lowland bNo rearing made on rainfed lowland or Irrigated lowland

MrasmiaAmong the Laval parasites the braconid Cardio- lowlands and parasitizes oriental maize borer OsIriniachilesphilijpphensis is most adapted to an upland environ- furnacalis (Guenee) larvae The leaffolder pupal parasitesment It also parasitizes Hydelepta indicata a common not only occur equally in all environments but also haveleaffolder of legumes The ichneumonids Trichomma cna- wvide host ranges- parasitizing species of OsIrinia Marucaphalocrosis and Temehcha stangli occur more in the Hydelepta and Honiona - in maize and legumeslowlands T stangli is most adapted to irrigated lowlands The principal egg parasite of yeltow stem borer (YSB)and also parasitizes Chilo and Scirpophaga stem borer Telenomus rowani is widely adapted to all environmentslarvae T cnaphalocrosis is most prevalent in rainfed and has an unusual alternate host -Tabanus eggs (25)

32 IRPS No 123 January 1987

Tetrastictu schoenobii specializes only in YSB and occurs mainly in irrigated lowlands -- further proof that YSB is most adapted to the wetter environments

Larval parasites of YSB are highly environment-specific The braconid Stenohracon nicevillei occurs only in upland rice but also attacks the pink stem borer (PSB) Tieinchwa phiiiplPwnsis an ichneumonid occurs in upland rice and also parasitizes the larvae of (7hiu stqpresaisinatro-morpha accepta melaihoracica like 7 stangi is mnost adapted to the irrigated lowlands

The dark-headed stem borer(l)HSB) (ih pochit3sus was not collected in all rice environments and the gold fringed borer Chilo auricilius nd PSB were collected only in upland rice Xaantiholiplastenrnaloran ichneumonid larval parasite appears to be widely adapted whereas TrihomaslopisapanitI)cteiiaa pteromalid pupal parasite occurs mostly in wetlands and also parasitizes larvae of Chilo suppressais and Pelhuidas mathias

Armyworms Myit hinin and Spodujipera have two main larval parasites - a tachinid Zrgohothriaatropivora in rainfed environments and a braconid Cotesia spp in irrigated lowlands

The butterfly pests Pelopidas and 4elanitishave tricho-grammatid egg parasites found only in irrigated lowland environments

Two larval parasites show environmental preferences --Argrroph av nigrotihialis a tachinid in rainfed habitats and Trichonialopsisapantelkcnela in irrigated ricefields A limgrotibialisalsoattacks the sweet potato hornwormAgrius convolvtdi and T apanteloctena parasitizes DH S1B

Predators as a group show more distinct environmental preferences The aquatic damselflies Coenagrionidae are most abundant in the lowlands Cvriorhinus is also most adapted to the lowlands The nabid Stenonabis tagalicus occurs only in the uplands and isalso prevalent on legumes The coccinellid lady beetles wasps and ants are more prevalent in upland areas AlicrasIpis cit-ea larvae prey ona variety of aphids attacking legumes and maize Etunienes campanonia vespid mud wasp makes nests in trees and is therefore most adapted to the more botanically diverse uplands Soil-dwelling ants cannot tolerate flooding

Upland environments are habitats to arboreal carabid beetles Three species of Chlaeniusbeetles prey on leaffolder larvae and are perhaps more important than parasites (27)

The gryllids and tettigoniids are widely adapted egg predators not only on rice but also on maize and legumes They feed on eggs of most insects that are laid on leaves

The spidercommunity of upland rice environments isrich in species At one site (Batangas Philippines) 31 species have been recorded (23) Of the 176 spider species recorded in Philippine ricefields about one-half (82) occur in upland rice The spiderspecies of upland rice environments overlap more with those of rainfed lowland than with those of irrigated lowland rice (24) The same study showed that of the three environments irrigated rice has (ie greatest spider species diversity foiiowed by rainfed lowland and upland

However of the 10 most prevalent ricefield spiders 9 were abundant in upland rice showing wide adaptation All three spider guilds -- orb-web space-web and huting spiders shywere prevalent in upland ricefields There were however differences in environmental prelerences for some spider species

Of the orb-wcavers Tetragniathamanlihulatawas partishycila rlv abundant 7 japonica was low and Ltucatge dehcorata was absent in upland rice

As in other environments Aiypeiwnaifirtosana was the most dominant space-web spider however its relative numbers were lowerthan for lowland sites Among hunting spiders Ox|opes javanus and Lcosa leatcost fgla were more abundant and Lvcosa lpseudoanmnlata was less abundant in upland compared with lowland rice envishyroniments

Extensive rearing of rice insect pests in upland rainfed lowland and irrigated sites over an 8-yr period in the Philippines provided insight into the effectiveness of parasites as natural enemies (Table 7 8) Most of toe data are from large sample sizes taken over at least one crop which overcomes the pitfalls expressed by Van Driesche (238)

VSB egg parasitization was surprisingly similar across all environments Larval parasitization was highly dynamic among sites and even years within the same site but showed somewhat lowerlevels in uplandsthan in irrigated wetlands This would have been predicted for egg parasites as well since YSB is adapted to the lowlands and so must be its parasites This isevidence that Telenomus rowani is as good at dispersing as its main hosts Trowani clings onto the anal tufts of female YSB and parasitizes eggs as they are laid This phoresy may explain its wide adaptation The more specialized lowland YSB larval parasites therefore appear more effective than their upland counterparts with more alternate hosts

Low collection levels of PSB larvae especially in the wetlands make comparisons between environments difficult Low levels of larval parasifization were recorded in both rainfed environments Except in the Cagayan rainfed wetland site in 1980 low levels ef larval parasitization were evident at irrigated and rainfed sitesalike No differences by environment were evident Higher levels of parasitization occurred in Sarawak during a 1967 outbreak also from tachinids Rothschild (197) indicated that higher parasishytization occurred in upland rice areas

Skipper and green horned caterpillar larval parasitization rates were generally low and showed no environmental effect htighest activity occurred in Cagayan in 1980 (48 parasitization) Although larval parasitization levels of RLF were variable year to year activity appeared greater in the irrigated lowlands than in the uplands Again the highest levels were recorded in Cagayan in 1980 Cagayan has extensive grasslands and fallow around the rice areas and a relatively long rainfall period Rainfall in 1930 was good for crop growth and apparently for increase of larval parasites

IRPS No 123 January 1987 33

Table 7 Parasitization rates of rice planthoppers and green leafhopper in upland rainfed lowland and irrigated lowland environments Philipshypines 1976-840

Environment and site Year Egg parasitization () Nymphaladult parasitization ()

BPH W13P GLH BPII WBPH GLIH

UplandBatangas

Rainfed lowland

1976 1979 1980 1982

15 17 12

2(47)8(39)

4(51)

9(1800)14 (212)

12(304)

8 (101)6 (127)

9(116)

Iloilo

Cagayan

1976 1977 1978 1981 1980 1981 1982

0 (69)

11

44 48 40

-

-

46 55 44

0 (471)

11

52 56 49

0 (200)9 (214)

10 (1259)8(235)6 (900) 3(800)

-7 (462)

11 (270)3 (313)5 (900) 1 (800)

4 (600)10 (725)10 (362)12(870)5 (900) 3 (800)

Pangasinan

Irrigatedlowland

1984 1976 1978 1979 1982

13 (436) 13(379) 14 (457)

6(14)7 (197)

17(198) 11 (486) 18 (275)

3 (81)10 (274)3 (116)

16(871) 19(88)

28(211)9 (764)5 (647)7 (1204) 8(305)

Laguna 1977 11 7 21 1978 31 1979 14 4 4 28 1980 7 8 20 1981 1983

56 7(278)

26 16 (1650)

16 27(1921)

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R I Apostol unpublished 1lo1lo and Pangasinan 136 Cagayan BCanapi unpublished Liuna 175 I C Pantua unpublished 118 234 43 Carino and Shepard unpublished BPH = brown planthopper Nila parvara htgens W131Pi =whitebacked planthopper Sogatelia furcifera GLH = green leafhopper Nephotettrix virescens Figiires in parentheses are sample sizes

Tauher et al (230) reported that internal external and genetic factors influence the seasonal activity of parasitoids

Hopper egg parasites but not nymphaladult parasites were also parlicularly abundant at the Cagayan site possibly the egg parasites have more alternative hopper prey in the weedy areas than nymphaladult paisites

The seasonal dynamics of hopper egg parasites can be seen for upland rice in Batangas(Fig I 2) and for rainfed lowland lice in Pangasinan( 135)and Cagayan( 105) Levels of BPHWBPI-L and GLH egg parasitization were corn-parable in upland and irrigated lowland environments Except for some high rates of paraMsitization early in the season the parasitization rate tended to be steady in all envilonments However hopper nymphaladult parasi-tizution levels were more similar in the rainfed environments and lower than in irrigated rice

Overall parasites do not appear highly effective by themselves against upland or even lowland pests Among natural enemies parasites have attracted the most attention because they can be more readily assesscd

Predators and pathogens lend themselves better to management practices Predators the most important group of natural enemies in riccfields are difficult to quantify On the other hand pathogens appear less im-portant but can be readily cultured and disseminated

The natural enemy commulnity of upland rice is rich in species and differs significantly from that of lowland rice

These beneficial organisms must be conserved by applying insecticides judiciously particularly sprays Seed or soil placement of chemicals minimizes exposure to natural enemies The strategy to derive the greatest benefit from natural enemies isto allow the greatest number of beneficial species to thrive Then perhaps at least one species will be effective against each pest at any tine overcoming the variable abundance of each species within and between years

Programs to introduce exotic species or mass produce indigenous ones are ambitious and expensive because they require trained people on a sustained not ad hoc basis

Chemical control Insecticides offer rapid and efficient control of upland insect pests (190) but-should be used only after other control measures have been considered Insecticides are rarely used on upland rice because of cost (32) Upland rice yields are normally too low to justify the expenditure Also most upland rice is grown as a cash crop Spraying upland rice is more difficult than lowland rice because water is less accessible Government subsidy programs to provide pestishycides to tribal peoples for upland rice production have created problems of toxicity to humans because isolated people have had no experience to be able to handle pesticides safely

However ins-eticide use has beenjustified when a crop is

0Z

Table 8 Parasitization rates of lepidopterous pests in upland rainfed lowland and irrigated lowland environments Philippines 1976-84a

Parasitization (5)

Environment and site Year YSB GFB PSB Earcutting Swarming Skipper Greenborned RLF Brown larva larva caterpillar caterpillar caterpillar caterpillar larva semilooperEgg Larva larva larva larva larva larva

UplandBatangas 1977 - 0 (57) - 0 (63) - 0 (71) - 0 (106) 11(142) 2 (101)1978 43 (4697) 8(451) 5 (38) 3 (40) 6 (61) 2(58) 15 (57) 13 (129) 13 (1286) 4 (30)1979 59(4135) 12(320) 13 (11) 5(28) 9(45) 6(46) 7(23) 8(160) 9(1152) 8(26)

Rainfed lowland Iloilo 1976 - 0 (10) 0(10) 0 (7) 0 (19) 2 13) 15 (180) 5 (76)1978 0 (54) 0 (3) 0 (17) 9 (64) 4 (78) 3 (73) 0 (121) 0 (191)1979 64 (2350) 16 (84) 6 (17) 0(6) 18 (68) 25 (104)

1981 56 (1322) 18(304) 0(5) 12(51) 18(138)Pangasinan 1976 60 (7 7 )b 62(82) 0(111) 5 (39)

1978 58 (53)b 5(51) Cagayan 1979 59(713) 8 (101) 6 (34) 0(7)

1980 0(11) 4 (28) 15 (286)

62 ( 48(177) 61(140)1981 0(5) 12 (2) 13(45) 14 (92)

Irrigatedlowland Laguna 1974 62 8

1974-75 50 41 1975 43 72 1978 96 (623) 21(24) 33 (60)1979 83(2017) 18(110) 8(62) 11 (89) 22(400)1980 78 (1625) 20(75) 0 (16) 4 (72)1983 60

9 (80) 7 (270) 30

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R F Apostol unpublished Iloilo and Pangasnan 136 Cagayan B Canapi unpublished Laguna 175 P CPantua unpublished 118 234 143 Carino and Shepard unpublished YSB = yellow stem borer ScirpophagaincertulasGFB = gold fringed borer Chilo auricilius PSB = pink stem borerSesamia inferens earcutting caterpillar Alythimna separata swarming caterpillar Spcdoptera mauritia acronyctoides skipper Pelopidas mathias reen horned caterpillar Melanitls ledaismene RLF = rice leaffoldersCnaphalocrocismedinalis and Marasmiaspp brown semilooper Mocisfrugalis Figures in parentheses are sample sizes Egg masses

IRPS No 123 January 1987 35

Table 9 Effect of seeding rate and insecticide on plant density and Yield losses from insect pests in upland rice often areyield of Dagge upland rice Batangas Philippines 1976a comparable to those of lowland rainfed or irrigated rice

Seeding rate Insecticide Dosage Plant density Yield (0-Y30o) but usually from different pests The principal (kg aiha) seed treatment (kg aiha) (noIra-row) (tha) groups are soil pests feeding on sown seed and rootsat 14 DE followed by seedling pests These groups have largely gone

50 None - 35 a 20 a unnoticed leading researchers to conclude that stem borers50 Carbofuran F 10 38 ab 23 a and seed bugs are the most important pests (17) Yield loss 10 beCabofuranF100 None - 4551 c 23 a trials that pinpoint major pests need to includc seed

100 Dieldrin WP 10 52 cd 22 a treatments Granules applied in the furrow may not give150 None - 59 d 22 a adequate protection from many seed pests

aAv of 8 fields DE = days after crop emergence In a column means The floristic and pedological diversity of upland rice followed by a common letter are not significantly different at the environments is matched by the diversity of arthropods5level by D)MRT F = flowable Wi = wettable powder both pests and their natural enemies Irrigation has homoshy

genized lowland rice environments Flooded soils tend tothreatened by epidemics of locusts or armyworms New have the same properties and to eliminate soil pests The technology stressing efficiency has minimized insecticide spatial and temporal dominance of cultivated rice has cost benefited specialist pests

Seed treatment is normally inexpensive and can be Upland rice has no specialist arthropod species The economicallyjustified in -nanycases(66) Treating seed with monophagous pests on upland rice prefer the lowlands systemic insecticide (216) but not chlorinated hydrocarbons where rice evolved Rice in the uplands exists only because (247) is most effective in controlling seedling maggots of man and even though lowland rice predates upland riceFoliar sprays have to be repeated to obtain control and specialist species evolved Wheat maize and sorghum do often are washed off by frequent early season rains not have monophagous pests leading us to conclude that Granular insecticides are effective but required dosages are there is no evolutionary advantage for inssects to have too expensive to justify except under heavy infestation narrow host ranges in upland graminaceae(247) When directed against first-instar larvae banding Pests have adapted to the uplands not only by having granules in seed furrows (177) can be effective against white wider plant host ranges but also by having long life cyclesgrubs at low dosages (025 kg aii ha) (134) Seed treatments undergoing dormancy or being dispersive Each pest has a are not promising for white grub control (248) unique life history strategy suited to highly unpredictable

Seed treatment is also effective ainst ants and may be environments In agivenyear someinsect specics an better cheaper than increasing the seeding rate In Batangas exploit upland rice as a food source but no pest is a senious treating 50 kg seed ha was as effective as increasing the one every year on upland rice seeding rate to 100 kgha and using untreated seeds (Table The low yielding potential of upland rice means less 9) In Brazil seed treatments control termites and other soil quantity and lower quality of food for pests to exploit but insects including the lessercornstalk borer(66) Domiciano also less incentive for farmers to undertake control (58) however found seed treatments inconsistent in their The most effective control would be early and synchroshyeffect against high populations of lesser corn stalk borer nous planting of early-maturing upland rice varieties to

Baiting can be an inexpensive way of controlling ants prevent insect pest buildup Breeding for insect resistance seedling maggot flies mole crickets and field crickets Baits should initially focus on stem borers because they are verymade of locally available material and impregnated with difficult to control by other means Biocontrol efforts insecticide can be sparingly distributed taking adantage of should first stress conservation of natural enemies preshythe pests ability to disperse and encounter bait sites dators and pathogens rather than parasites Efficient

The new low-volume hand-held controlled-droplet chemical control includes seed treatment or baiting for seed sprayers may offer an advantage over high-volume knap- seedling and some root pests and using controlled dropletsack sprayes for controlling foliar pests Upland rice sprayers for spot treatments against foliar pests and seed canopies are more open than those in lowland rice and bugs when economic thresholds are reached better droplet penetration should result Controlled droplet applicators would be ideal for seedbugs and defoliators

CONCLUSION

Upland rice is attacked by a wider array of insect pests than is lowland rice mainly beca use of the addition of soil pests Generally population levels of lowland rice pests are lower in upland than lowland rice leading many people to conclude that insect pests are not important on upland rice

36 IRPS No 123 January 1987

REFERENCES CITED

1Abraham C C and K S R Mony 1978 Occurrence ofLeptocorisa acuta Fabr (Coreidae Hemiptera) as a pest of nutmeg trees J Bombay Nat ttistSoc 74553

2 Agyen-Sampong M 1975 HlierogrlphusdiaganensisKrauss (Orthop-tera Acrididae) a new pest of rice in Northern Ghana Ghana J Agric Sci 8249-253

3 Agyen-Samnpong M 1976 Varietal resistance and other inle-ratcd pest control components as factors of rice varietal improvement in West Africa Prcsented at the Second varietal improvement serinar 14-18 Sep 1976 Bouak Ivory Coast West Africa Rice Development Association Monrovia Liberia 18 p

4 Agyen-Sampong M 1977 Outbreak of Ihccrorrdnisorae in rice seedlings near Rokupr Sierra Leone lit Rice Res Newsl 2(6)16-17

5 Agyen-Sampong M 1980 Dirty panicles and rice yield reduction caused by hugs IntRice Res Newsl 5(l)11-12

6 Agyen-Sampong M 1982 The major pest problems of irrigated upland and mangrove swanip rice ecosystems ir the humid tropical Guinea savanna and Salel climate zones Pages 318-328 in Pro-ceedings training course cn concepts techniques and application of irtegrated pest management in rice inWest Africa 10-20 Jan 1982 Fendall Liberia West Africa Rice l)evelopioent Association Monrovia Liberia 505 p

7 Akibo-Betts V I and S A Raymundo 1978 Aphids as rice pests in Sierra Leone lit Rice Res Newsl 3(6)15-16

8 Akinsola F A 1975 Present status of different rice stemborers in parts of Nigeria iicL Entomo Newsl 328

9 Akinsola FA 1979 Ihe biology and ecology of rice stemborers in Nigeria Ph 1) thesis University ol lbadan Nigeria 217 p

10 Akinsola E A 1979 Guidelines of integrated management of insect pests of upland rice in West Africa Presented at the Seminar on integrated management of rice diseases and insect pests at Bobo-Dioulasso Upper Volta 17-22 Sep 1979 West Africa Rice Develop-ment Association Monrovia Liberia

II Akinsola IA 1984 Insect pests of upland rice in Africa Pages 301-305 in An overview of upland rice research Proceedings of the 1982 lIouak Iwsrv Coast upland rice workshop International Rice Research Institute 13O Box 933 Manila Philippines

12 Alam M S V TJohn and Kaung Zan 1985 Insect pests and diseases of rice in Africa Pages 67-82 in Rice improvement in Eastern Central and Southern Africa proceedings of the international rice workshop at Iusaka Zambia 9-19 April 1984

13 Alticri M A A van Schoonhovcn and J D Doll 1977 The ecological role of weeds in insect pest management systems a review illustiated with bean (Phaseohsvulgaris L)cropping systems PANS (Pest Artie News Summ) 23195-205

14 Anmante F 1967 The leaf-cutting ant Atia capiguaraa pest of pastures [in Portuguese] Biol6gico (Sao Paulo) 33113-120 (Rev Appl Entomol A56207)

15Andrade A C and D Puzzi 1951 Preliminary results of experi-merits on the control of Sculptcoris iastantus on sugarcane [in Portuguese] Bioulgico (Sao Paulo) 1744-49 (Rev Appl Entomol A41421422)

16 Appert J 1970 Maliarplia separaelha (rice stem borer) new observations of entomological problems of rice in Madagascar [in French] Agron Trop (Paris) 25329-334

17 Ariadeau Mand Z Harahap 1986 Relevant upland rice breeding objectives Pages 189-197 in Progress in upland rice research Proceedings of the 1985 Jakarta conference International Rice Research Institute 1O Box 933 Manila Philippines

18Asar S and Kisimoto 1975 Seasonal occurrence of the smaller brown planthopper in upland rice [in Japanese] Proc Kanto-Tosan Plant Prot Soc 2228

19 Bakker W 1974 Characterization and ecological aspects of rice yellow mottle virus in Kenya Meded Landbouwhogesh Wage-ningen 152 p

20 Balut F F 1970 White grubs attacking rice Oryza ativa L [in Portuguese] Biol6gico (So Paulo) 36321-322

21 Banerjee S N and L M Pramanik 1967 The lepidopterous stalk borers of rice and their life cycles in the tropics pages 103-124 in The major insect pests ofthe rice plant Proceedings ofa symposium at the International Rice Research Institute September 1964 The Johns Hopkins Press Baltimore Maryland USA

22 Barrion A T and J A Litsinger 1980 Ants a natural enemy of cliaiarac in dryland rice IntRice Res Newsl

5(4)22-23 23 Barrion A T and J A litsinger 1981 The spider fauna of

Philippine rice agroecosystem 1Dryland Philipp Entomnol 5139-166

24 Barrion A T and 1 A Iitsinger 1984 The spider fauna of Philippine rice agroccosystcms II Welland Philipp Eiitomol 611-37

25 Barrion A T1 and J A Litsinger 1984 lihanus (Diptera Tabanidae) eggs an alternative host of rice stei borer (SI3) egg parasite Telenoonis digns (Iliymenoptcra Scelionidae) lit Rice Res Newvsl 9(6)19

26 Barrion A T and J A Litsinger 1985 Identilication of rice leaffolders by wing markings Int Rice Res Newsl 10( 1)24

27 Barrion A T and J A Litsinger 1985 Chctiosspp (Coleoptera Carabidae) a leaffolder ILF)predator lit Rice Res Newsl 10(1)21

28 Bertels A 1970 Insect pests of rice and theircontrol [in Portuguese] Pelotas Brazil Instituto de Expcrimentacao Agropccuararias do Sul Circular 43 24 p

29 Bonzi S M 1982 Ctilo (flisilineus J de Joannis (Lepidoptera Pyralidae) stem borer of irrigated cereals in Upper Volta LioFrench] Agron Trop Riu Rizic Cult Vivrieres Trop 37207-209

30 Bourke T V T L Fenner J N LStihick G Baker EHassan D F OSullivan and C S li 1977 Insect pest survey for the year ending 30th June 1969 Papta New Guinea )epartment of Agrishyculture Stock and Fisheries Port Moresby 57 p

31 Bowden 1 1973 Migration of pests in the tropics Meded Rijksfac Landbouwwet Gent 38785-796

32 Breniere J 1973 The main rice parasites in West Africa and their control Pages 27-42 in Proceedings seminar on plant protection for the rice crop West Africa Rice Development Association Monrovia Liberia

33 Breniere J 1976 The principal insect pests of rice in West Africa and their control West Africa Rice Development Association Monrovia Liberia 52 p

34 Brown ES 1962 The African arinyworn Spoiolptera eXI)Ina (Walker) (Lepidoptera Noctuidae) a review of the literature Cornshymonwealth Institute of Entomology London UK57 p

35 Brown F B 1968 Upland rice in Latin America FAO-International Rice Commission 12th Session of the Working Party on Rice Production and Protection 2-14 Sept Peradeniya Sri lanka Food and Agriculture Organization Rome Italy 6 p

36 Buyckx F J 1 1962 Disease and insect pests of crops inCongo Rwanda and Burundi [iil French] Publications dc LInstilut National pour lestude Agrononiquedu Congo M Weissenbruck S A Brussels Belgium 708 p

37 Cariiio F 0 1981 Role of natural enentics inpopulation suppression and pest management of green rice lealhoppers Ph D thesis University ofthe Philippines at Los Bafitos PhilippinesLaguna 136 -

38 Carpente A J 1983 The development of rice cultivation and extension of vegetable productin in Zanzibar Final Report Vol II1 October 1983 Food and Agricultme Organization Ronte Italy

39 Carroll C P and S1 Risch 1983 Tropical annual cropping systems ant ecology Environ Manage 751-57

4(0 Cates R G 198 1 Host plant predictability and the feeding patterns of monophagous oligophagous and polyphagous insect herbivores Oecologia (Berl) 48319-326

IRPS No 123 January 1987 37

41 Cendcafia S M and F B Caora 1967 Insect pests of rice in the 62 Ene 1 C 1981 I)iiorldhol)ierusihilis Slater (licteroplera Philippines Pages 591-616 in The najor insect pests ofthe rice plant lygacidue Illissinac) a new rice pest front Nigeria lit Rice Res Proceedings of a symposium at the International Rice Research Newsl N3)15 Institute September 1964 The Johns Hopkins Press Baltimore 63 Fugenio T S 1965 rhe effect oi e-vCgetation on rice production Maryland USA 729 p and the possibility of overcoming it by leaf pruning MS thesis

42 Centro Ihternacional de Agricklittira Tropical 1984 Upland rice in University ol the Philippines at Los Baijos ILaguna Ihilippins 69 p Iatin Anerica Pages 93-119 hi An os ervies of uplaind rice research 04 Farrow R A 1975 The Alrican nmigratory locust ii is main Proceedings of the 1982 Boinakamp Ivory Coast upland rice workshop outbreak area of the middle Niger qlinlitatise studies of solitary International Rice Research Institute P0 Blox 933 Manila populations in relation to environmental factors Locusla No II Philippines 198 p

43 Chandra G 1979 Taxonomy ad hn riononlics ofthe imect parasites of 65 Fennal R G 1943 Food crop investigation Windward and rice Icalloppers and planthoppers in the Pihilippines and their Ieeward Islands First report Report onl work dunring the periodiiiportairce iii natural biol)gical cointrol Philipp Fntoltol 4119-139 April 1942-April 1943 Imperial College of Tropical Agriculture

44 Chang T T 1976 The originii colut ion enlIivation dissemination Trinidad 16 p (Rev of Appl Fintornol A3222-25) and diersificition of Asian and African rices -uphytica 25425-44 1 66 Ferreira F 1983 Integrated pest ritaageiret of upland rice [ill

45 Chatierjee P B 1979 Rice leaffolder attacks in India lit Rice Res Portuguese English summary] Pages 323-341 in Upland riceculttre Newsl 4(3)21 factors affecting production E Ferreira T Yamada and F-

46 Cheng Cbien-pan 19(4 Upland rice in Taiwan International Rice Malmolta eds Institutoda Potassae Foslato Institnto Internacional Cnmmission Wiorking Party oit Rice Production ard Protection I()th da Potassa Piracicaha Sao Paulo lraid Meeting 3-10 March 1964 Manila Philippines Food and Agricul- 67 Ferreira E arid F dl Silva Martins 1979 Occurrence of ture Organiaion Rome Italy 5 p Sogatohs orvziola (Muir) inupland rice in Goias Ilrail lit Rice

47 Chcrrett 1 M and 1) 1Peregrine 1976 A review of the status of Rcs Ncwssl 4(4)18 leaf-cutting ants and their control Ann Appl Biol 84124-128 68 Ferreira E and 1F da Silva Martiris 1984 Insect pestof rice ill

48 Chipeta F M 1974 Pests of rice in Malawi Pages 90-95 in Brazil and their control Jill Portugucsej FMIlRAtPA-CNPAF Proceedings Sith eastern Africa cereals research conference 10-15 Goiania Brmiil 67 p March 1974 DRB Mialda ed Ministry of Agriculture lilongwe 69 Ferreira FI F ca Silha Miartins S S Nero and 1 It P de Malawi Carvallio 1982 M tltiplc regression analysis todeterimine the effect of

49 Chu Yau-i 1979 The d atiaging period of the rice seedling fly insect populatiois ott upland rice yield [inIPortuguese Pesqui (Atlterigotiavxigu Stcin) (NItiscide Diptera) oil cori) at Fast Java Agropecu Bras 17671-675 Indonesia Plant Prot Bill 21403413 70 Ferreira F 1F da Silva Martins and F 1PZimmermann 1979

50 Conklin H C 1957 IHaninoo agriculture a report on aln integral Rice varietal resistance to tle lesser corn stalk borer [in Port uguese] systent of shi fti rig cult ivationi in the Pltilippincs Food and Agriculture Pesqui Agropccu Biras 14317-321 Organization Rome Italy 209 p 71 Fleming W F 1968 Biological control of lie hapanese beetle U S

51 Cramer H II 1967 Plant proection and world crop production Dep Agric RLs Sers [ech Bull 1383 78 p Pilaniensclhutz-Nachr 201-524 72 Food and Agriculture Organiation of the United Nations 1982

52 l)ani R C and Majunidar 1978 Prelininary observations on tie Country report surmimary Pages 27-37 in Report on the first incidence of root aphids on differeit rice cultnrcs Sci Cult 4488-89 FAOIUN)IPhail and regional training coiuirse in iniproved ctltural

53 Davidson R I 1979 lffects of root feeding on foliage yield Pages practices for upland rice 5-23 Oct 1981 Northern Region Agrculturd117-120 in T K Ci osb and R B Pottinger eds Proceedings of the Developririt Center Chiang Mai Food and Agricuilture Organiiashy2nd Australian conlerence on grassland invertebrate ecology Govt tion Rome Italy 37 n Printer Wellington New Zealand 73 Franco do Ariaral S 195 1 Residual action of insecticides in turfy

54 l)ean G 1W 1976 Rice insect pests inlaos Int Rice Res Newsl soil--the results of a field experiment [in Portuguese] Biol6gico (Sao 1(2)15 Paulo) 17209-214 (Rev Appl Entomol A41423)

55 I)ean G J W 1978 Insect pests of rice in Laos PANS 24(3)280-289 74 Freeman ) 1970 Report onithe Iban london School of -coroiics 56 )ingle H 1984 Behavior genes and lifc histories conplex adapta- Monographs on Social Anthropology No 41 Athlane Press

tions to uncertain environments Pages 169-194 in A new ecology London UK 317 p novel approaches to interactive systems P W Price C N Slobod- 75 Fuchs T W1 A Harding and 1W Smithlr 1979 Induction and chikoff and W S Gaud cdsl ohn Wiley New York USA termination oldiapause in the sugarcane borer Ann Flitonlol Soc

57 Dobelmann-JP and M Falais 1968 Upland rice cultivation ill tie Am 72271-274 North-western part of Madagascar in 1966-1967 [in French English 76 (hbriel B1) 1975 A review tf the major insect pests of some tpland summary] Trop Agron 2313-21 crops in tlc Philippines Pages 86-111 in Ad Htoc panel of experts on

58 Doniiciano N L 1979 Chemical control of tie lesser corn stalk pest disease arnd weed problems in some rainfed crops Bangkok (lolsonno~paloIgnowslhts) (Zeller 1848) (I-epid(ptera Ihycitid ae) Thailand Food aid Agriculture Organization Roire Italy

on rice in Ponta Gressa Parara 1977-1978 [in Portuguese] Pages 77 Garg D K and N K Shal 1983 White grub outbreak on rainfed 291-303 in Anais 1 Reuniao de tecnicos cmiirizicultura do estado de dryland rice in Uttar Pradeslh Int Rice Res Ncwsl 81(4)16-17 Sao Paulo-Campinas Cati Brazil 78 Garg I) K and IP Tandon 1982 Major insect pests of rice on hilly

59 Dunsmore 1 R 1970 Investigations on tlie varieties pests and tracts of Uttar Pradesh India lilt Rice Res Newsl 7(11)11-12diseases of upland rice (hill padi) in Sarawak Malaysia Int Rice 79 Garrity 1) P 1983 Upland rice research an international bibliograpy Comm Newsl 1929-35 1975-1982 Iternational Rice Research hnstitnte PO Box 933

60 Elias 1R1965 The new decline of rice seedlings is not caused by Manila Philippines 121 p parasites [in Portuguese] Biol6gico (Sao Paulo) 313-6 80 Ghauri M S K 1982 A new subspecies of Iinorh)rone bull

61 Empresa Brasiliera de Pesquisa Agropecuaria 1984 Upland rice itt cornlus Slater (Hemiptera Iygaeidae) front Papua New Guinea on Brazil Pages 121-134 it Av overview of upland rice research rice and carpet grass Ihll lEntoniol Res 72133-137 Proceedings ofthe 1982 Blouak Ivory Coast Upland rice workshop 81 Gonzales F 0 PArregoces R I ihernandef and OT Parada International Rice Research Institute PO Box 933 Manila 1983 Insect and imite pests of rice arid their control in Latin America Philippines [in Spanish] la Federacion Nacional de Arrocerts (Fedearroz)

Bogota Colombia 60 n

38 IRPS No 123 January 1987

82 Grist D H and R J A W Lever 1969 Pests of rice Longmans London UK520 p

83 Hale P R 1979 Two subterranean pests of upland rice in Papua New Guirnca IntRice Res Newsl 4(2)16-17

84 Hale P R and J T Hale 1975 Insect pests of rice in the New Guinea Islands Rice Entomol Newsl 34

85 Harris W V 1969 Termites as pests of crops and trees Common-wealth Institute of Entomology London UK 41 p

86 Hattori 1 1968 A method to distinguish between grass leaf roller (Ctaplha rocis medinalisGuenee) and the rice leaf roller (Susurtia exigua Butler) [in Japanese] Plant Prot (Japan) 23167-170

87 Henry I E and 1A Onsager 1984 Experimental control of the mormon cricket Iahrosshnph by NJsena locustae(Aficrospora Microsporidae) a protozoan parasite of grasshoppers (Ort Acri-didae) Entomophaga 27197-201

88 Hidaka 1 P Vungsilabulr and SKadkao 1974 Studies on ecology and control of the rice gall midge in Thailand Tech Bull 6 TropicalAgricultural Research Center Tokyo Japan 113 p

89 Hincklcy A 1) 1963 Ecology and control of rice planthoppers in Fiji Bull Entomrol Res 53467 481

90 Hirao J 1982 Ecology and chtInical control of the rice leaffolder Jpn Pestic Inf 4114-17

91 Ho DT 1984 Sten borer incifience in rice ecosystems in KenyaEast Africa Pages 307-320 in A overview of upland rice research Proceedings of the 1982 Bouac Ivory Coast upland rice workshop International Rice Research Institute PO Box 933 Manila Philippines

92 Ho )T J 1Njokah and J GKibuka 1983 Studies on rice sten borers in Kenya with emphasis on ifaliarpiaipiraellRag Insect Sci Appl 465-73

93 Ichihara I T Ito A NIatsuo and M Sawada 1971 Chemical control of the rice striped stem borer in upland rice grown in plastic mulch [iii Japanese] Proc Kanto-Tosan Plant Prot Soc 1882

94 IclhiharaIand A Matsuo 1969 Damage caused by rice striped stett borer to upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1686-87

95 Ichhara I atid A MIatsun11970 Conside ration oft lie extraordinary prevalence of rice striped stem borer in upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1778-79

96 Institut de Recherches Agrolnliqes Iropicale et des Cultures Vivrieres 1978 Insect resistance Pages 12-13 in Upland rice 1978 atnual report Bouak Ivory Coast 27 p

97 International Institute for Tropical Agriculture 1983 Aphids itt rice roots I ITA Res Briefs 4(1-3)4-5

98 International Institite fi Tropical Agriculture 1983 Hopperburn in rice caused by brown plathopper IITA Res Briefs 4(l-3)6-7

99 International Rice Research Institute 1975 Annual report for 1974 PO Box 933 Manila Philippines p 216

100 International Rice Restarch Institute 1976 Annual repiort for 1975 PO Box 933 Manila Philippines p 228

101 International Rice Research Institute 1979 Cropping systems program Component technology development and evaluation Page 404 i Annual report for 1978 11O Box 933 Manila Philippines

102 International Rice Research Institute Department of Entomology 1981) Insecticid evaluation 1979 Tables 73 74 PO Box 933Manila Philipes

103 International Rice Research Institute Department of Entonmology 1981 Insecticide evalomtion 1980 Table 81 POBox 933 Manila Philippines

104 International Rice Research Institute 1981 Report of a meeting of Asian rice-based cropping systems entomologists Eleventh cropping systems working group meeting 18-22 May 1981 PO Box 933 Manila Philippines p 132 133 144

105 International Rice Research Institute 1984 Cropping systems program Pages 368-373 in Annual report for 1983 PO Box 933 Manila Philippines

106 International Rice Research Institutc 1984 Upland rice in Asia Pages 45-68 in An overview of upland rice research Proceedings of the 1982 Bouakamp Ivory Coast upland rice workiiop loternational Rice Research Institute 11O Box 933 Manila Philippines

107 Irshad M 1977 Grasshoppers in rice in Pakistan lit Rice Res Newsl 2(3)8-9

108 Ito R and TOno 1964 Upland lice its culture and varieties in Japan lIternational Rice Cormmission loth Meeting of the Working Party on Rice Production and Protection 3-10 March 1964 Manila Philippines Food and Agriculture Organization Rome Italy 12 p

109 Jeffrey M I and V A Dyck 1983 Effect of moonlight on lightshytrap catches of brown planthopper lit Rice Res Newsl 8(6) 13

110 Jennings P R and T Alicia Pineda 1983 Management ofthe rice plnthoppers Sogatodsorivi(coIl in Latin America Presented at the meeting on stable plant resistance FAO and lIndocsian I)inectorate of Food Crops Protection Bali lndonesia 29 Novcniber-2 l)ccemshyber 9 p

IllJimenez Gomez J A 1979 Rice insect pests i lla ios ()riCntales lil Spanish] Pages 99-103 in Rice ciltivation Instituto (olotibiatto Agropecuario Compndio No 29

112 Jongeleen F JJ Soesanto and A Mockidjo 1979 Studies oti pathogens of two species of Scarabacidac froinIndonesia Z Angew Enlomnol 87412-417

113 Jouvenaz 1) PC S lofgren and W A Baiks 1981 Biological control of imported fire antsa review of cirrent knowledge Bull Entomol Soc Am 27203-208

114 Kainoh Y K Shiniiu S MarUani Y Taiaki 1980 lost finding behavior of tie rice bu lplotori+JchinvittjI)allas (liciniptera Corcidac) with special reference to diet pattern ol aggregation and feeding on the rice plant Appl Entomol Zool 15225-233

115 Kakkar K 0 P Sharma and P I Sharma 1977 Note on Ihteronyrchus liodresRedt (Coleopicra I)ynastidae) as asporadic pest of paddy and maie inHimachal Pradesh Corr Agric 156-57

116 Kalode M131 79 Major insect pest problems and their control in upland rice (Unpublishcd mineogriph CWI oJNaaiat lni Chandaratat (198 1)original not seen)

117 Kalshoven L i 98 I The pests of crops in I ldoniesia (translated by P A vatn der Laan) 1 T Ichtiar B3aru-vati Iloeve lakarta Indonesia 701 p

118 Kamal N Q 1981 Suppression of whitebacked planto ner Sogatella fircaifra (Horvath) aLId rice lealfolder (nhal -cis licdb lis (Guenee) populations by natural cnclmie hlI) tiesis Grcgorio Araneta University Foundation Victoncta Park Metto Manila Philippines 179 Ip

119 Karim A N M Reaul 1975 A species ofliealy bug Trioni i70s ceres new oii rice in the Philippines Rice Entoniol Newsl 247

120 Katanyukul Wand C Chandaratat 1981 Insect pestsof uplaid rice and their control Report firstFAOUNlFi Tliailand training course on improved cultural practices for tplad rice Chiang Mai Thailand 5-23 Oct 1981 Food and Agriculture Organiation Rome Italy 22 p

121 Kern F 1955 lhletwioa(I)uninemts) bhhit Biturm(Coleoptera) a new pest of sugarcane inVenezuela [inSpanish with Etnglish sunmary] Agron Trop (Maracay) 527-29 (Rev Appl Eitoniol A4572)

122 Khan K M and S Ghai 1974 White grubs ald their control in India Pesticides (1Bombay) 819-25

123 Kiritani K 1975 Integrated pest management of rice pests Pages 66-70 itt Proceedings platning workshop ott cooperative hIeld research in pest managementP SMotooka ed East-West Center Honololu Hawaii USA

124 Kisimoto R 1967 Ecology of insect vectors forecasting and chemical control Pages 243-255 in The virus diseases of tle rice plant Proceedings of a symposium atthe International Rice Research Institute April 1967 The Johns Hopkins Press altimore Maryshyland USA

125 Kisimoto R and V A Dyck 1976 Climate and rice insects Pages 367-391 in Climate and rice International Rice Research Institute PO Box 933 Manila Philippines

126 Kureha S S Kobayashi SAsari and T Ishihara 1977 Life history and control of the mole cricket Grrllotalpa africana Palisot de Beauvois [in Japanese English translation 7520 1RRI library] Proc Kanto-Tosan Plant Prot Soc 21192-197

127 Kushwaha K S 1981 The black beetle as a pest of rice in Haryana India Int Rice Res Newsl 6(4)14

128 Lawani S M 1982 A review of tile effects of various agronomic practices on cereal stem borers Trop Pest Manage 28266-276

129 Leuck D B 1966 Biology of the lesser cornstalk borer in South Georgia J Econ Entomol 59 797-801

130 lever R J A W 1955 Insect pests of the rice plant in Malaya Int Rice Comm Newsl 1612-21

131 Li Ching-Sing and Shin-Foon Chiu 1951 The study ofthe rice gall midge Iachydiplosis orvza Wood Mason [in Chinese with English summary] 1Taiwan Agric Res 21-13

132 Iitsinger J A 1984 Assesament of the need-based application for rice Proceedings TechnologyTransfer Workshop 15-16 March 1984 International Rice Research Institute 1PO Box 933 Manila Philippines 6 p

133 Litsinger J A 1984 Environmental diversity and insect pest abundance with reference to the Pacific Pages 285-307 in Proceedings sub-regional training course on methods of controlling diseases insects and other pests of plants in the South Pacific 4-20 Oct 1982 Vaini Kingdom of Tonga Nukualofa

134 Litsinger J A RF Apostol and M BObusan 1983 White grub Leucopholis irrorata (Coleoptera Scarabaeidae) pest status popula-tion dynamics and chemical conllol in arice-maize cropping pattern in the Philippines J Econ Entoinol 761133-1138

135 Litsinger J A C de la Cruz F Raymundo A T Barrion M D Lumaban J P Bandong M S Venugopal F Paragna and E Balete 1982 Insect pests and insecticide reponse of dry-seeded bunded rice Pages 179-193 in Report of workshop on cropping systems research in Asia International Rice Research Institute Los Bafos Philippine

136 Litsinger J A M 1) Luinaban J P Bandong A T Barrion W Katanyukul and N Panda 1978 Insect pest management in rainfcd lowland rice agroecosystms of tropical Asia International Rice Research Conference 17-21 April 1978 International Rice Research Institute Los Bafios Philippines

137 Litsinger J A and K Moody 1976 Integrated pest management in multiple cropping systems Pages 293-317 in R I Papendick P A Sanchez and G B Triplett eds Multiple cropping American Society of Agronomy Madison Wisconsin USA

138 Loevinsohn M E 1984 The ecology and control of rice pests in relation to the intensity and synchrony of cultivation Ph D thesis University of London 36) p

139 Loevinsohn M E J A Litsinger N Panda M A Austria R F Apostol C Andrion and J Bandong 1982 A basis for predicting rice insect pest abundance in cropping systems Paper presented at the 1982 International rice research conference on rice and cropping systems 19-23 April 1982 International Rice Research Institute Los Bafios Iaguna Philippines 25 p

140 Lopez A W 1930 Repot of the Entomology Department Pages 145-172 in Annual report of the Philippine Sugar Association 1929-30 Manila

141 Lopez AW 1931 Entomology Department Pages 227-266 in Annual report of the director or research 1930-31 Philippine Sugar Association Manila

142 Lopez A W 1932 Entomology Deparment Pages 252-279 in Annual report of the Philippine Sugar Association 1931-32 Manila

143 Lungibill P and G G Ainslee 1917 The lessercornstalk borer US Dep Agric Agric Res Serv Tech Bull 539 27 p

IRPS No 123 January 1987 39

144 McGuire J U Jr and BS Crandall 1967 Survey of insect pests and plant diseases ofselected food crops of Mexico Central America and Panama U SDepartment of Agriculture International Agriculshytural Development Service Agricultuial Researrh Service October 1967 11 p

145 McNeill Sand T R ESouthwood 1978 The role of nitrogen in the development of insectplant relations Pages 77-9 in lliochemical aspects of plant and animal coevolution J B Ilarbornu ed Academic Press London UK

146 Maebara H 1955 Spatial distribution of lastinstiarlanaeof purplish stem borer (Sesamia iqferens Walker) hibernating in an upland rice field Bull Kagoshima Univ Fac Agric 438-41

147 Mallick E H and N G Hajra 1977 Response of rice plant to defoliation treatments at different developmental stages Indian Agric 2151-56

148 Mansingh A 1971 Physiological classification of dormancies in insects Canl Entomol 103983-10)9

149 Martell C G 1974 Catalogue of plant feeding insects of Mexico [in Spanish] Fitofilo 27(69)1-176

150 Martinez 0 J 1982 Principal rice pests in Ilanos orientalis [in Spanish] P~ages 80-94 in Curso deact nalization de conocimientos en arroz Villaniencis Instituto (olonmbiano Agropecuario

151 Martins J F da Silva F Ferrcira A S Prabhu F 1 P Zimmermann 1980 Use of pesticides for the control of the main subterranean upland rice pests esqui Agropecu Bras 1553-u

152 Martins I F da Silva F Fe eira rod BdamSilveira Pinheiro I Q2 Damage simulation caused by defoliators of upland rice [in Portuguese with English summary] Pesqui Agropecu Bras 171113-1119

153 Martins 1F da Silva N(iyuen van Tan and 1Ida Silveira Pinheiro 1981 Stem borer resistance in upland rice associated with plant morphological characters [in Portuuce Iesqui Agropecu Bras 16187-192

154 Masaki S 1980 Sumnmer diapause Ann Rev Entomiol 251-25 155 Masuzawa T 11 Suwa and F Nakasoji 1983 l)ifkleences of

oviposition preferencc and survival rate of two skipper butterflies Parnara guttata and PIchlpidai iathiais (Iepidoptera HIesperiidac) on rice plant and cogon grass New Entomol 3247-56

156 Maurya D M and C P Vaish 1984 Upland rice in India Pages 69-92 in An overview oftpland rice research Proceedings ofthe 1982 Bouakamp Ivory Coast upland rice workshop International Rice Research Institute IO Box 933 Mnila Philippines

157 Menon M G R R N Katiyai S Kuar and S S Misra 1970 Termites attacking the paddy crop Sci Cult 36294-295

158 Miyahara Y and 1 Nishikubo 1967 Behavior of young nymphs of the red rice root aphid Iholahsipiun ri liabhdontinalisSasaki in relation to movement from the leaf to the roots of upland rice [in Japanese] Pioc Assoc Plant Prot Kyushu 1363-66

159 Mo ljoa Tjien 1952 White grub of upland rice (Holotrichiahelleri Brsk) [in Indonesian with English summary] Contrib Agric Res Stn Bogor No 13134 p

160 Mochida 0 and T Suryana 1979 The occurrence of the brown planthopper and ragged stunt on lowland and upland rice ill Central Java and logyakarta in tile late of January 1979 Entomnolgy Survey Report No 15 Central Research Institute of Agriculture (CRIA) Sukamandi Indonesia I February 1979 9 p

161Monet C 1979 Varietal resistance to upland rice pests [inI French] Pages 35-39 in Annual report 1977 Institut des Savanes Department des Cultures Vivrieres Bouak6 Ivory Coast

162 Monte 0 1942 A caterpillar ofrice-fields[in Portuguese] Biol6gico (Sao Paulo) 8161-163 (Rev Appl Entomol A31414)

163 Munroe I) D 1979 Pest management strategy for hill rice in four resettlement areas in Sarawak Malaysia Pages 89-102 i1 Guidelines for integrated control of rice insect pests Food and Agriculture Organization Plant Production and Protection Paper 14 Rome Italy 115 p

40 IRPS No 123 January 1987

164 Nakasuji F M Ishii and Jen-Zon Ito 1984 Rice skippers in Taiwan and their life histories ZZrC (Taipei) Tech Bull 82 10 p

165 Navas I) 1976 Fall armyworum in rice Pages 99-106 il Ecological animal control by habitat management Proc Tall Timbers Con-ference 28 Feb-I Mar 1974 Fall Timbers Res Stn Misc Publ 6 Tallahassee Horida USA

166 Nickel 1 1 1961 Biological control of rice stem borer A feasibility study Tcch Bull 2 International Rice Rcarch Institute PO Box 933 Manila Philippines IIl p

167 Nigan P M and R A Vernia 1985 Insect pests of uplaild rice in Uttar Pradesh Int Rice Res Newsl 10(6)22

168 Nilakhe S S 1985 -cological observations on spittle bugs with emphasis on their occirence in rice Pcsqtui Agropecu Bras 20407-414

169 Nixon G EJ 195 1 lie association of ats with aphids and coccids Commonwealth Iusti lcte of Entonology Iondon UK 36 p

170 Nurdin F an Ishak Manti 1985 Study oit seedling fly of upland rice Interlational Upland Rice Moitoring Tour Report 26 lebruary-2 MarLI Sukaranti West Sumatra

171 Oka 1 N 1979 Ciltural conttol of the brown planthopper Pages 357-369 in Brown planthopper threat to rice production in Asia International Rice Research Institutc 1p0 Box 933 Manila Philippines

172 Ohni M and A Vilnaii 1975 Rice insect pests of Italy biology of Sipha grcria (Kaltenbach) and its control [in Italian] Riso (Milan) 2459-71

173 Otanes F Q atnd 1 I Sison 194 1 Pests of rice Philipp J Agric 12211-259

174 Painter R I 1951 Insect icsistance in crop plants University Press of Kansits Lawrence Kansas USA 520 p

175 Pantua P C and 1A litsinger 1981 Arthropod abundance in continuous vs biannual rice cropping systeis Int Rice Res Ncwsl 6(2)20-21

176 Pathak M ) and V A l)yck 1975 Sttudics ott insect pests of upland rice Pages 186-191 in Major research in upland rice International Rice Research Institute PO Box 933 Ilanila Philippines

177 Patil B R B M IHasabe and 1) S Ajri 198 I Chemical control of white gritb lIditriihinhorraia Fabr intfesting groundntut paddy atnd sorghum Indian 1Entomol 43232-236

178 Pawar A 1) 1974 (ollection and identification of insect pests of rice rlerminal repirt lInternational Rice Research Institute PO Box 933 Manila Philippines 72 p (tnimco)

179 IerrajU A aid A S Reedy 1965 Observations ott diapausc itt tie steti borer ohrice Rice News Teller (lai-Mar)9-13

180 Pont A C 1972 A review of the Oriental species of theri ona Rondani (I)iptera Muscidac) of econtomic importance Pages 27-102 in Control of sorglium shoot fly M G lomtani and W R Young eds Oxford atd IH1 Ptthl Co New I)clhi India 324 p

181 Prasad S C 1PChand aid M 1PSingh 1983 Incidence of rice gall midge in the upland plateau region of Hihar Oryza 20235-236

182 Rai P S 198 0 Nisaga simpex damage to rice in the hill tracts of South India Int Rice kles News[ 5(I)12

183 Rathorc Y S 1969 Gonocplhahli dcprssunt Fabr as itpredator of (tilo zonwIlhis (Swinhoc) and other moths in field cage at Patnagar Uttar Pradesh Indian 1Entotnol 3127

184 Reed W 1983 Crop losses causcd by insect pists in the developing world Proceedings 10th Internatial Congress of Plant Protection lrighton UK 174-80

185 Reedy M V 1981 Ateactogasterinhens(Walker)a new root-grub pest of tipland paddy and inaize Curr Sci 501076

186 Reveno M A 1968 Catalogue of the insect pests of economic crops in Colombia [in Spaiish] Asociacion Latino Americana de Entomo-logia (AIAE) 129-33

187 Reyes G M 1957 Rice dwarf disease in the Philippines Food and Agriculture Organiiation Plant Prot Bull 617-19

188 Rice S E A A Grigarick and M 0 Way 1982 Effect of leaf and panicle feeding byarmyworm (Lepidoptera Noctuidae) larvae on rice grain yield Environ Entomol 7593-595

189 Risch S J 1) Andow and M A Altieri 1983 Agioccosystem diversity arnd pest control data tentative conclusions and nizw research dirctions Environ Entomol 12625-629

190 Rivera C T 1956 Three organic insecticides in thecontrol of insects affecting upland rice Philipp Agric 39465-472

191 Roffrey 1 1972 Locusta outbreaks in the Philippines Acrida 1177-188

192 Rojas R 1) 1941 The effect of pruning at varying ages upon the yield of Ramairio Phipp Agric 29851-860

193 Roonwal M i 1979 Ternite life and termite control in tropical South Asia Scientilic Pnbl odhpr India 139 p

194 Rosetto C1 F da Silva Martins N C Schmidt and L EAzzini 1978 Damage caused by pasturespittle bugs 4Dvoi)lavopictaandDJ schach) on rice lBragantia Nota 6 (Matcli)25-27

195 Rosztto C1- S S Neto I- Amantc I) Iink 1Garcia-Vieira D M de SoLza N V Bansa t and A M de Olivcira 1971 Rice pests of Brazil [in Portuguese] Pages 149-238 inrFA()-International Rice Commission 2nd meeting of the Rice Committee for the Americas 6-11 l)ecember 1971 Pelotas Rio Grande do Sul Brazil

196 Rothschild G H L 1969 Obseratiois on the armyworm Spodoptera matritia acronrctoiiles Gn (Lep Noctuidae) Bull Entomol Res 59143-160

197 Rothschild G 1- L 1974 Parasites of the armworm Spolopiera mauritiaacroncitoihes in Malaysia Fntornopliaga 19293-299

198 Sachan i N S K Gangwar and 1N Katujar 1980 Field crickets damaging cages in North Eastern IHill Region Indian J Entomol 42526-527

199 Sands 1) P A 1977 The biology and ecology of Lepocorisa (Hemiptera Alydidac) in Papua New Guinea Res Bull 18 Departshyment of Primary Industry Port Moresby Papua New Guinea 104 p

200 Sarawak 1968 titomology division Pages 97-106 in Report of the research branch of the Department of Altriculture 1966 Kuching (Rev AIppl Eintomol A56603)

201 Satpathy JM M S Das and K Naik 1977 Effect of multiple and mixed cropping on the incidence of scine important pestsI Entomol Res 178-85

202 Sauer IHF G 1939 Notes Ott Elasmpnlpualtsligniosellths Zeller (Lep Pyr) a serious pest ofcereals in Sao Paolo [in Portuguese] Arq Inst Biol 10199-206 (Rev Appl Entotnol A28378)

203 Satinders 1L A 1 SKing and C L Vargas S 1983 Crop pests of Central Anerica with t bibliography [in Spanish] Centro Agroshynotnico Tropical de Investigacion y Ensenanza Turrialba Costa Rica (CATIE) Tec Bol 9 90 p

204 Sauphanor B 1985 Some factors of upland rice tolerttce to stem borers in West Africa Inseu Sci Appl 6429434

205 Sen P atnd S Chakravorty 1976 Stentflics in Haritighata Nadia West Bengal India [it Rice Res Newsl 1(12) 15

206 Senapai B and 1M Satpathy 1983 The seedling fly Atherigona orrzae Malloch is a pest of rice in Orissa (India) J Entomol Res 777-79

207 Shhjahan M S Alatn M Ah nad and BN Islam 1983 A new scarabaeid beetle pest of dryland rice in Bangladesh Int Rice Res Ncwsl 8(I)16-17

208 Sharma G L S C Modgol and R C Gautam 1979 Effect of depth of submergence ott incidence of bacterial blight and white grub infestation in transplanted rice Int Rice Res Newsl 4(6)12-13

209 Sharma S 1985 Scarabaeid beetle outbreak itt upland rice Int Rice Res Newsl 1(1(l)24

210 Shiraki T 1917 The paddy borer Schoenobhits incertulas Walker [achaku Agric Exp Stn Formosa 256 p

211 Silva A de Brito and 11P Magalhaes 1981 Insect pests of rice [in Portuguese] Circular Tecnica No 22 EMBRAPA Centro de Pesquisa t ecuaria de Tropico Unido (CPAJU) Belem Brazil 14 p

212 Singh S A 1979 Emergence of rice leaf butterfly as a rice pest in Manipur India int Rice Res Newsl 4(4)16-17

213 SinghS A 1983 Changing trends of insect pestsof rice in Manipur Int Rice Res Newsl 8(f)12

214 Siwi S S and P H van Doesburg 1984 Lenovorisa Latreille in Indonesia (Heteroptera Coreidae Alydinae) Zool Meded (Leiden) 58117-129

215 Soejitno J 1977 Reaction of some upland rice varieties toseedling fly eltherikonaexigua Stein [in Indonesian] Siri Hana I Penyakit 630-35 Lembga I osat Fenclilian Pertanian Bogor Indonesia

216 Socnardi 1967 Insect pests of rice in Indonesia Pages675-683 in The major inse pests of thu rice plant Proceedings of a symposium at the Internatital Rice Rec Wi itt S pternber 1964 The Johns Hopkins Press Baltimore Maryland USA

217 Soto P E A T Petcz and I W Buddenhagen 1976 Survey of insect pests and diseases of rice in different ecological zones in Nigeria Rice Entomol Ncwsl 435-46

218 Soto 11E and Z Ziddiqi 1976 Screening for resistance to African rice insect pests Varietal improsetnent sCmi n r Bouak Ivory Coast 13-17 Sept 1976 West Africa Rice l)evelopment Association Monrovia Iiberia

219 Squire F A 1972 Entomological problems in Bolivia PANS (PestArtic News Sumr) 18249-268

220 Stapely 1 II 1978 The rice leaf roller in the Solomon Islands Int Rice Res Newsl 3(l)13

221 Suenaga H 1937 On aphids attacking the roots of rice Oyo-I)oubts Zasshi 9150-152 [in Japanese] (Rev Appl Entomol A25745)

222 Suenaga H 1952 On the seasonal abundance of three subterranean aphids on upland rice Bull Kyushu Agric Exp Sti 1249-262

223 Sutynso Yas and Dandi Soekaria 1964 Beberapa hana yang ttenperang tana man padi tanah kering [in Indonesian] Rapat kerja Padi Tanah Kering 6 sd 8 Juily 1964 Kaliurang 9 p

224 Tallamy I) W and R F Denno 1983 Alternative life history patterns in risky environments in example from lace bugs Pages 129-147 in Insect life history patterns R F l)cnio and If Dingle eds Spring-Verlag 225 p

225 Tamura 1 1939 Observations on some insect pests of field crops [in Japanese] 1 Plant Prot 26414-420 (Rev Appl EntomolA27-556)

226 Tamura and Suzuki 1964 Influences of the early-transplanting practice in rice culture upon development of the rice stem borer Chilo siql)ressalisWalker Bull Ilokuriku Natil Agric Exp Stn 761-94

227 Tan J P 1924 The rice root aphid Drowhi ihirsoia Philipp Agric 13277-288

228 Tanaka 1 1957 Taxonomy and distribution of some subterranean aphids injurious to upland rice in Japan with description of a new species [in Japanese English summary] Botyu-Kagaku 22168-176

229 Tanaka T 1961 The rice root aphids their ecology and control Spec [full College of Agriculture Utsunomiya University Japan F3 p

230 Tauber M IC A Tauber R Nechols and 1Obrycki 1983 Seasonal activity of parasitoids control by external internal and genetic factors Pages 87-108 itt I)iapause and life cycle strategies in insects V K Brown and I Ilodek eds )r W Junk The Hague Netherlands

231 Taylor W F 1972 Effects of artificial defoliation (simulating pest damage) on varieties of upland rice Exp Agric 879-83

232 feotia T P S and S Natud 1966 Biotoomics of the rice skipper Parnaraniathiis Fabricius (Lepidoptera Hesperiidac) Indian J Entomol 28181-186

233 Teran F 0 1971 Rice pests of Santa Cruz [in Spanish Ministerio de Agricultura Sita Cruz Bolivia Bull 45 8 p

234 Than Itun 1976 Population dynamics of the yellow rice borer Tryporza itcertulas (Walker) and its damage to the rice plant MS thesis University of the Philippines at Ios Bafios Laguna Philippines 92 p

235 Uichanco L B 1911 Pests Pages 55-74 it Rice Philipp Agric 11-200

IRPS No 123 January 1987 41

236 van der Laan P A 195 1 Life history of the rice seedling fly Contrib Gen Agric Res Stn Boor Indonesia 118 15 p

237 van der Laan 11A 1959 Correlation between rainfall in the dry season and the occurrence of the white rice borer (Scirpolhaga innotala WIk) in lava Entomol Fxp Appl 212-20

238 Van Driesche R G 1983 Meaning of percent parasitismin studies of insect parasitism Fnviron Entomol 121 I1- 122

239 Venugopal M S and A litsingcr 1984 Fffect ofcarbofuran on rice growth Prot Ecol 7313-317

240 Vietnam Directorate of Rural Affairs 1962 Annual work progress report on crop improvement program of rice sugarcane vegetable and field crops (lor the period from July 1961 to June 1962) (11) 299 p (Rev Appl Entomol A51526-527)

241 Wan M T K 1972 Observations on rice leaf- and planthoppers in Sarawak (Malaysian Borneo) Malay Agric 148308-335

242 West Africa Rice I)esclopnient Association 1978 Annual research report Vol 2 Statistics Monrovia Liberia

243 West Africa Rice Development Association 1979 Annual research report p47 Monrovia Liberia

244 Williams ) F and CS lofgren 1981 Fli I illy l-468 anew bait toxicant for control of the red imported lire at Fla Entomol 64472477

245 Williams I) 1 1970 The mealybugs OFlomoptera Coccoidea Iscudococcidac) of sugar-cane rice and sorghum Bull Entomol Res 0109-188

246 Williams D) 1 1982 Pulinaria icervi (Signoret) and its allies ott sugar cane and other grasses Btll Entoinol Res 72111-117

247 Wirjosuiardjo S 1976 Study ofchenmical cont rot of insect pests ott upland rice Imiu Pertanian (Jogjakarta) 1323-329

248 Wirjosuhardjo S and N Wirarta 1976 Chemical control of the white grub tIolorithiatlltriin upland rice in M uneng Irobolinggo Indonesia) [in Indonesian] Imut Pertanian (Jogjakarta) 1330-332

249 Yano K T Miyake and V F Eastop 1983 The biology and economic importance of rice aphids (Henuiptera Aphid idae) a review Bull Entomol Res 73539-566

250 Yelp D and P Rhodes 19(4 lhe success of drilled dryland rice iii the Fiji Islands International Rice Commission Working Party on Rice Production and Protection Manila Philippines 3-10 March1964 Food and Agriculture Organization Rome Italy 13 p

251 Young W R 1981 Fifty-five years of research on the sorghum shootfly Insect Sci Appi 23-9

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No 104 Faridpur acomputer-assisted instruction nmodel forrainfed lowland random coefficient model No 116 Consumer demand for rice grain quality in Thailand Indonesia andrice

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No 106 Rice grassy stunt virus 2 a new strain of rice grassy stunt in tire literature

Philippines No 118 IRRI-Korca collaborative project for the development of cold-No 107 Physical losses and quality deterioration in rice plstprodnCtion tolerant lines through anthei culture

systemis No 119 Problen soils as potenial areas for adverse soils-tolerant rice No 108 (olptblication ofIRRI materials a survey of translators and varieties in South and Ssuthcasi Asia

publishers No 120 Changes isinall-farm iLtireliing techii)hgy in Thailand and [he No 109 Classification of Philippine rainfall patterns Philippines No 110 Contributions of modern rice varieties to rutrition in Asia No 121 Landforms and modern rice varieties No Ill Changes in rice breeding in 10 Asian countries 1965-84 No 122 Yield stability and modern rice technology No 112 Design parameters affecting the performance of the IRRI-designed

axial-flow pump

Page 28: UPLANDRICE. INSECTl--1i M T -E!R.EOLOU t RTANCE ,AND …delphacid.s3.amazonaws.com › 6958.pdf · The significance of abiotic factors in upland rice insect ecology will be apparent

28 IRPS No 123 January 1987

Table S Parasites recorded from upland rice insect pests in Tanauan Batangas Philippines 1977-84

Order Family Species Host stage

I lymttenoptera Braconidae Leaffolders Cnaphalocrocis Marasinia spy

Cardiochilesphilipplinensis Ashmead a Larva

Cotesia angustillasis(Gahan)Cotesia or cypris (Nixon)

Larva Larva

Cotesia or taeniaticornis(Nixon) Larva Cilonusinotakatae Munakata Chelonits SppAlacrocetrts philippinensis (Ashtmead)

Egg to larva Egg to larva Larva

Iclineumonidac Orgilts spp Isclutojppa hireator(Fabricius)

Larva Larva

Itoplectis naraigav(Ashmcad) Larva Tenelhtcha philippiensis (Ashmead) Larva Tenelchha stangli (Ashmead) a

irichomma cnaphalocrosis Uchida Larva Larva

Bethylidac Elasmidac

Goniozms or trianguli r Kieffer lasmtusalhopictus Crawford

Larva Larva

Diptera

Encyrtidac Chalcididac

Tachinidac

Elasmtus spp Copidosomopsisnacohiae (Fidy) Brach nteria evcarinata Gahan Biachyieria lasus (Walker) Ztgohothria ciliata (Wulp)

Larva Egg to larva Pupa Pupa Larva

Argyrophylx nigrotihialis(Blaranov) Larva

l-ymenoptera Braconidac Yellow stern borer Scirpopiaga incertulas

Braco chinensisSzeplegeti Larva CIthelotis spp Chelonus niunakatae Munakata Cotesi flaripes Cameron Stenoracon nicerilli (Bingham) a

Larva Egg to larva Larva Larva

Ichneumonidac Tropobracon schoenobii (Viereck) Amatroinorpiaaccepta metathoracica(Ashmiead)Frihorus sinicus (I lolmgren)

Larva Larva Larva

Ischnojoppaluteator (Fabricius) Larva Isotima nr dammeratiti Rohwer Larva Tenelucha philippinensis (Ashmead) a

Larva Tenehicha stangli (Ashmcad)Trichomma enaphalocrosis Uchida

Larva Lara

Chalcididac Eulophidac Trichogranmmatidae

Xanthopimnpla stemmator (Thunberg) Charops hrachYpterumn Gupta and Malteswary Brachtm eria sppTetrasticttus schoenohii Ferriere Trichorigranm chihunis Ishi

Larva to pupa Larva Pupa Egg Egg

Diptera

Scelionidac Pteromalidae Tachinidae

Tt ichograintnta aponiicuitt (Ashniead) Telenomits rowani (Gahan) a

Trichoinalopsis apattelctena (Crawford) Jeirhaea spp

Egg Fgg Egg pupa Larva

Z4gobothria ciliata (Wulp) Larva

Ilymenoptera lecneumonirlac Gol fringed borer (hilo auricilius

Trichomm enoptalocrosisUchida Larva

Ieioenoptera Braconidae

Aantholimlh steiniator (Thtinberg) a

Pink stet Ioret Sesamia inferens StoInacoil nicrilhi (Bingh am)

Loarva to pupa

larva

Diptera Ilyinenoptera

Armvtnyorms Mythimna separata Spodoptera mauritia acronyetoidesTachinidae Zvgohothriaatropiora( Robinea i -Desvoid y)Eulophidae It-phctrtiscitapadoe (Ashimead)

Larva Larva

(halcididae Braconidac

Brachpin ria spip (oesia spp

Larva larva

IHyntenoptera I)iptera

Rice skipper Ielop ida (halcididae Tachinidae

math ias aidgreen iortedci ciliarNIelan itis eda Brat lhincria sp iii matuqitatt Cameron Irgyrolhirlaxi trotlhiais(marumtv)a1

ismtne Larva Larva

Diptera Ilymenoptera

Tachnidae lelineutionidae

fuown se tilo per Mois frugalisIrgtrophrlax 1igrothiahiv 3 a rat ov) anthopimpa pmnctata (Iabricius)

Larva Larva to pupa

aMost dominant

IRPS No 123 January 1987 29

Normally predatory thrips prey on herbivorous thrips strepsiptera on both Spiders dominate tilenymphaladultNatural enemies of stem bugs --Scolinoplara and Tibraa predators Spittle bugs move to rice from pasture grasses - are in order of importance egg par~isites and predators and biocontrol would be successful only ifcarried out before fungal pathogens and nymphal adult parasites Egg tiley immigrate into ricefields parasites could either be introduced fron other areas or Biological control tactics for hemipterous seed bugs mass produced and released ould follow those outlined for stein bugs

Armyworms are normally lwld in hcck by the activities Coml)arison of environnewtns Natural enemies are of egg and larval parasites When these natural enemies fail perhaps faced with even greaterconstraints than insect pestsusually because of drought armyworns become epidemic to survive ind reproduce in upland environments TheyIt may be worthwhile therefore to release parasites during have adapted ani survived by having wide host ranges the the rainy months following a drought Virus diseases of ability to aestivate or hibernate over unfavorable seasonslarvae would beanotheravnue to exploreforarnyworms and the ability to disperse or by other mechanisms

Parasites have ben traditionallyconsidered for biolegical Comparing the most dominant and therefore the most control attempts against stein borers usually by rekasing adapted species of natural enemies collected in various exotic species( 166) This approach has not met with success environments in the Philippines we can understand how and new avenues should be explored The role of predators different rice environments are arid pathogens is little inderstood and deserves greater Anagqrms olptahiisa mymarid planthopper egg parasiteattention Egg parasitism rates are normally reasonably is cquallyadapted to all Philippine rice environments (Tablehigh but should be supplemented with effective egg 6) It is selective for planthoppers and even parasitizes eggspredators Orthopterans are potential egg predators of the n-iize planthopper Peregrinitsmaidis itt upland areas Metioche vittaticollis (Sta ) Anavipha nifiwnnis (Saus- Alavohis also parasitizes maize planthopper eggs but is sure) and Conocephahs hongipnni (de i-laan) feed on most prevalent on BPH in the irrigated wetlands (onashystem borer eggs etioche and Anaxil)ha specialize in eggs tocerus spp like A optalhilis on BPH and WBPI Iattacks with no hair such as Chilo spp The omnivorous (ono- the eggs of GLH in ell rie environments It is also found cephahsspecializes in eggs cov red with hairs it is a proven parasitizing the eggs ef the white rice licahopper (oimna egg predator of Scirlouphaga spp but has the discouraging spetra Oirosita naias a trichogrammatid egg parasitehabit of eating rice guin (65) attacks BIPH and WBPH in the wetlands and has no other

Grasshoppers have egg (scelionid) and nymphalad ult host A related species 0 aesopi specializes in G1-H in (nemestrinid tachinid and sarcophagid) v irasites Among wetland environments Dryinid strepsipt-an and pipunshypathogens protozoans have been recorded most frequently culid nymphal and adult parasites of planthoppers and Control of locust species by natural enemies could con- leafhoppers also specialize either in hopper species or centrate on their habitual breeding areas environments Among the dryinids Pseuhdgonatoipsflavi-

Leaffolders have rich complexes of natural enemies some feimur prefers the lowlands and BPH while P nuch has no adapted to upland rice (22 27) ranging from egg predators environmental prefirences but is adapted only to WBPH (gryllids coccinellids) to larval parasites (braconids ichncu- The main d ryinid species on GLH aptogonatop)usspp is monids) to larval prcdators (ants and carabids) to larval most dominant in rainfed lowlands and also attacks other pathogeiis (vira and fungal diseases) Efforts to augment leafhoppers within Amnrasca Cicadulinaand Bachaltuathese natural enemies should focus on the egg predators Strepsiptera are more dominant on plarithoppers than (rearing releasing)and viral or fungal pathogens (culturing lealhoppers The Elenchus species in the Philippines attack spraying) all species of planthoppers within the genera Sogawilla

Parasitization of the large conspicuous larvae of Sogatodels NilaparsataHarualia and Opiconsiva Elenshyskippers green horned caterpillars and other polyphagous cusrvasuntaisuiis adapted to the major rice environments Lepidoptera by tachinids and chalcids is normally low ELg but prefers BPH in rainfed arid WBPIH in wetland environshyparasites and predators are perhaps the key to their control ments Ilalictoph(utsmtnroeiand HVpectrus attack G LH

Gall midges are parasitized by pterormalids and platy- and COfimravcctra respectivelygasterids that aestivate within their larval host between Pipunculids are mainly parasitic oii lealhoppers Each seasons Shoot aphids are preyed upon by coccinellids species has a unique environmental preference Tomosshysyrphids and chrysopids varireIa oraetoraprefers Ne7 otetltix nigropictrs and

Planthoppers arid leafhoppers have egg parasites and Pilunwulhtsjavanensisattacks Dehoceplhahls spp which predators as well as nymphialad nlt parasitesand predators are more abundant in upland than irrigated lowland their rich complex of natural enemies includes fungal environments T suhvirescensattacks Nephotettix virescens pathogens The principal egg parasites are myrnarids and and prefers the lowlands Pilpuiulus nttillattls is an trichogramniatids Egg predators are norinally mirid plant upland rice parasite arid attacks only Nephotetix speciesbugs that suck out the yolk Nymphal ad nIt parasites are More than 17 recorded species of leaffolder parasites in dryinids on planthoppers pipunculids on lealhoppers and the Philippines attack species of C(naplwocrocis and

30 IRPS No 123 January 1987

Table 6 Dominant natural enemies of rice pests in upland rainfed lowland and irrigated lowland rice environments of the Philippines

Natural enemy

Host stage Species

Brown planthop-Parasite Egg

MymaridaeAnagrus optabilis(Perkins) Anagrusflaveolus Waterhouse

TrichogrammatidacOligosita naiasGirault

NymphAdult Dryinidac

Pseudogotatopuslaiifemttr Esaki et Hashimoto Elenchidae

Elenchus yasurnatsui Kifune and Hirashima

Environment

Upand Rainfed lowland Irrigated lowland

Ailaparata lugens

x x X x

x

x x

x X X Whitebacked planthoppers Sogatella furcifera and Sogatodes pusanusEgg

MymaridaeAnagrus optabilis(Perkins)

TrichogrammatidacOl~gositanaias Girault

NymphAdult Dryinidae

Pseudogonatopusnudus Perkins Elenchidae

Elenchus yasutatsui Kifune et Hirashima

Parasite Egg

MymaridaeGonatocerus spp

TrichogrammatidaeOligositaaesopi Girault

NymphAdult Pipunculidac

Tomosvaryella subviresccns Loew Tomosvaryella oryzaetora (Koizumi) Pipunclusmutillatus (de Meijere)

DryinidaeIlaplogonatopusspp

Egg Copidosomopsisnacoleiae (Eady)

Larva BraLonidae

Cardiochiles philipphnensisAshnead lchneumonidac

Trichiomnia cnaphalocrosisUchida Temelucha stangli (Ashmead)

Pupa Chalcididae

BrachymeriaexcarinataGahan Brachymerialasus (Walker) Brachymeria nr tachardiaegroup

Parasite Egg

Scelionidae Telenomus rowani (Gahan)

EulophidacTettasticusschoenobhi Ferriere

Larva Braconidae

Stenobraconnicevillei (Bingham) Iclhncurnonidac

Temelucha phiiippinensis(Ashinead)Temelucha stangli (Ashmead)

x x x

x x

X x X

x x Green leafhopper Nephotettix virescens

X x x

x x

X X X X

X

X Rice leaffolders CnaphalocrocisMarasmia

x X x

X

X X

x x x x x X x x x

Yellow stem borer Scirpophaga incertulas

X X x

x

x

x X XAinauromorphaaccepta metathoracica(Ashmead) x

continued on next page

IRPS No 123 January 1987 31

Table 6 cntinued

Natural enemy Environment

Host stage Species Upland Rainfed lowland Irrigated lowland

Larva-pupa Gold fringed borer Chilo attriciiuslchneumonidae

Xanthopimpla stemmator (Thunberg) x aPupa x

Pteromalidae Trichoinalopsis apanteloctena (Crawford) ax

Pink stem borerb Sesamnia inferensParasite

Larva Braconidae

Stenobracon nicevillei (Bingham) x b b Armyworms Mvthinna SpodopteraLarva

Tachinidac Zygobothriaarroptvora (Robineau-Desvoidy) x x

Braconidac Cotesia spp

X Skipper Pelopidas mathias and green horned caterpillar Melanitis leda ismeneF-gg

TrichogramrnatidacTrichogrannaspp x

Scelionidae Undetermined Scelionid

x Larva

Tachinidae Argyroph lax nitrotibialis (Baranov) X x

Pteromalidae Trichomnalopsis apanteloctena (Crawford)

X Predator

CoenagrionidaeAgriocnenis femina femina Brauer x xIschnura senegalensis tambur

Miridae CrtorhinsliidipennisReuter x x

Nabidac Stcnonabis tagalicus (Stal) x

VeliidaeAlicrovelia douglasi atrolineata (Bergroth) x

Coccinellidae Micraspis crocea (Mulsant) X x x

Vespidae Eumenes campaniformis (Fabricius) x

Formicidac Diaecamna spp xOdontoponera transversa (F Smith) x

Carabidae Chaenius spp OphioneaW- aJasciataSchmidt-Goebel

x

Ophionea ishii ishii Habu x x X

Gryllidav Anaxipta longipennis (Saussur-) x xMetioche ittaticollis (Sial) x x x

Tettigoniidae Conocephalus longipennis (de Haan) x x x

aNo rearing made on rainfed lowland bNo rearing made on rainfed lowland or Irrigated lowland

MrasmiaAmong the Laval parasites the braconid Cardio- lowlands and parasitizes oriental maize borer OsIriniachilesphilijpphensis is most adapted to an upland environ- furnacalis (Guenee) larvae The leaffolder pupal parasitesment It also parasitizes Hydelepta indicata a common not only occur equally in all environments but also haveleaffolder of legumes The ichneumonids Trichomma cna- wvide host ranges- parasitizing species of OsIrinia Marucaphalocrosis and Temehcha stangli occur more in the Hydelepta and Honiona - in maize and legumeslowlands T stangli is most adapted to irrigated lowlands The principal egg parasite of yeltow stem borer (YSB)and also parasitizes Chilo and Scirpophaga stem borer Telenomus rowani is widely adapted to all environmentslarvae T cnaphalocrosis is most prevalent in rainfed and has an unusual alternate host -Tabanus eggs (25)

32 IRPS No 123 January 1987

Tetrastictu schoenobii specializes only in YSB and occurs mainly in irrigated lowlands -- further proof that YSB is most adapted to the wetter environments

Larval parasites of YSB are highly environment-specific The braconid Stenohracon nicevillei occurs only in upland rice but also attacks the pink stem borer (PSB) Tieinchwa phiiiplPwnsis an ichneumonid occurs in upland rice and also parasitizes the larvae of (7hiu stqpresaisinatro-morpha accepta melaihoracica like 7 stangi is mnost adapted to the irrigated lowlands

The dark-headed stem borer(l)HSB) (ih pochit3sus was not collected in all rice environments and the gold fringed borer Chilo auricilius nd PSB were collected only in upland rice Xaantiholiplastenrnaloran ichneumonid larval parasite appears to be widely adapted whereas TrihomaslopisapanitI)cteiiaa pteromalid pupal parasite occurs mostly in wetlands and also parasitizes larvae of Chilo suppressais and Pelhuidas mathias

Armyworms Myit hinin and Spodujipera have two main larval parasites - a tachinid Zrgohothriaatropivora in rainfed environments and a braconid Cotesia spp in irrigated lowlands

The butterfly pests Pelopidas and 4elanitishave tricho-grammatid egg parasites found only in irrigated lowland environments

Two larval parasites show environmental preferences --Argrroph av nigrotihialis a tachinid in rainfed habitats and Trichonialopsisapantelkcnela in irrigated ricefields A limgrotibialisalsoattacks the sweet potato hornwormAgrius convolvtdi and T apanteloctena parasitizes DH S1B

Predators as a group show more distinct environmental preferences The aquatic damselflies Coenagrionidae are most abundant in the lowlands Cvriorhinus is also most adapted to the lowlands The nabid Stenonabis tagalicus occurs only in the uplands and isalso prevalent on legumes The coccinellid lady beetles wasps and ants are more prevalent in upland areas AlicrasIpis cit-ea larvae prey ona variety of aphids attacking legumes and maize Etunienes campanonia vespid mud wasp makes nests in trees and is therefore most adapted to the more botanically diverse uplands Soil-dwelling ants cannot tolerate flooding

Upland environments are habitats to arboreal carabid beetles Three species of Chlaeniusbeetles prey on leaffolder larvae and are perhaps more important than parasites (27)

The gryllids and tettigoniids are widely adapted egg predators not only on rice but also on maize and legumes They feed on eggs of most insects that are laid on leaves

The spidercommunity of upland rice environments isrich in species At one site (Batangas Philippines) 31 species have been recorded (23) Of the 176 spider species recorded in Philippine ricefields about one-half (82) occur in upland rice The spiderspecies of upland rice environments overlap more with those of rainfed lowland than with those of irrigated lowland rice (24) The same study showed that of the three environments irrigated rice has (ie greatest spider species diversity foiiowed by rainfed lowland and upland

However of the 10 most prevalent ricefield spiders 9 were abundant in upland rice showing wide adaptation All three spider guilds -- orb-web space-web and huting spiders shywere prevalent in upland ricefields There were however differences in environmental prelerences for some spider species

Of the orb-wcavers Tetragniathamanlihulatawas partishycila rlv abundant 7 japonica was low and Ltucatge dehcorata was absent in upland rice

As in other environments Aiypeiwnaifirtosana was the most dominant space-web spider however its relative numbers were lowerthan for lowland sites Among hunting spiders Ox|opes javanus and Lcosa leatcost fgla were more abundant and Lvcosa lpseudoanmnlata was less abundant in upland compared with lowland rice envishyroniments

Extensive rearing of rice insect pests in upland rainfed lowland and irrigated sites over an 8-yr period in the Philippines provided insight into the effectiveness of parasites as natural enemies (Table 7 8) Most of toe data are from large sample sizes taken over at least one crop which overcomes the pitfalls expressed by Van Driesche (238)

VSB egg parasitization was surprisingly similar across all environments Larval parasitization was highly dynamic among sites and even years within the same site but showed somewhat lowerlevels in uplandsthan in irrigated wetlands This would have been predicted for egg parasites as well since YSB is adapted to the lowlands and so must be its parasites This isevidence that Telenomus rowani is as good at dispersing as its main hosts Trowani clings onto the anal tufts of female YSB and parasitizes eggs as they are laid This phoresy may explain its wide adaptation The more specialized lowland YSB larval parasites therefore appear more effective than their upland counterparts with more alternate hosts

Low collection levels of PSB larvae especially in the wetlands make comparisons between environments difficult Low levels of larval parasifization were recorded in both rainfed environments Except in the Cagayan rainfed wetland site in 1980 low levels ef larval parasitization were evident at irrigated and rainfed sitesalike No differences by environment were evident Higher levels of parasitization occurred in Sarawak during a 1967 outbreak also from tachinids Rothschild (197) indicated that higher parasishytization occurred in upland rice areas

Skipper and green horned caterpillar larval parasitization rates were generally low and showed no environmental effect htighest activity occurred in Cagayan in 1980 (48 parasitization) Although larval parasitization levels of RLF were variable year to year activity appeared greater in the irrigated lowlands than in the uplands Again the highest levels were recorded in Cagayan in 1980 Cagayan has extensive grasslands and fallow around the rice areas and a relatively long rainfall period Rainfall in 1930 was good for crop growth and apparently for increase of larval parasites

IRPS No 123 January 1987 33

Table 7 Parasitization rates of rice planthoppers and green leafhopper in upland rainfed lowland and irrigated lowland environments Philipshypines 1976-840

Environment and site Year Egg parasitization () Nymphaladult parasitization ()

BPH W13P GLH BPII WBPH GLIH

UplandBatangas

Rainfed lowland

1976 1979 1980 1982

15 17 12

2(47)8(39)

4(51)

9(1800)14 (212)

12(304)

8 (101)6 (127)

9(116)

Iloilo

Cagayan

1976 1977 1978 1981 1980 1981 1982

0 (69)

11

44 48 40

-

-

46 55 44

0 (471)

11

52 56 49

0 (200)9 (214)

10 (1259)8(235)6 (900) 3(800)

-7 (462)

11 (270)3 (313)5 (900) 1 (800)

4 (600)10 (725)10 (362)12(870)5 (900) 3 (800)

Pangasinan

Irrigatedlowland

1984 1976 1978 1979 1982

13 (436) 13(379) 14 (457)

6(14)7 (197)

17(198) 11 (486) 18 (275)

3 (81)10 (274)3 (116)

16(871) 19(88)

28(211)9 (764)5 (647)7 (1204) 8(305)

Laguna 1977 11 7 21 1978 31 1979 14 4 4 28 1980 7 8 20 1981 1983

56 7(278)

26 16 (1650)

16 27(1921)

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R I Apostol unpublished 1lo1lo and Pangasinan 136 Cagayan BCanapi unpublished Liuna 175 I C Pantua unpublished 118 234 43 Carino and Shepard unpublished BPH = brown planthopper Nila parvara htgens W131Pi =whitebacked planthopper Sogatelia furcifera GLH = green leafhopper Nephotettrix virescens Figiires in parentheses are sample sizes

Tauher et al (230) reported that internal external and genetic factors influence the seasonal activity of parasitoids

Hopper egg parasites but not nymphaladult parasites were also parlicularly abundant at the Cagayan site possibly the egg parasites have more alternative hopper prey in the weedy areas than nymphaladult paisites

The seasonal dynamics of hopper egg parasites can be seen for upland rice in Batangas(Fig I 2) and for rainfed lowland lice in Pangasinan( 135)and Cagayan( 105) Levels of BPHWBPI-L and GLH egg parasitization were corn-parable in upland and irrigated lowland environments Except for some high rates of paraMsitization early in the season the parasitization rate tended to be steady in all envilonments However hopper nymphaladult parasi-tizution levels were more similar in the rainfed environments and lower than in irrigated rice

Overall parasites do not appear highly effective by themselves against upland or even lowland pests Among natural enemies parasites have attracted the most attention because they can be more readily assesscd

Predators and pathogens lend themselves better to management practices Predators the most important group of natural enemies in riccfields are difficult to quantify On the other hand pathogens appear less im-portant but can be readily cultured and disseminated

The natural enemy commulnity of upland rice is rich in species and differs significantly from that of lowland rice

These beneficial organisms must be conserved by applying insecticides judiciously particularly sprays Seed or soil placement of chemicals minimizes exposure to natural enemies The strategy to derive the greatest benefit from natural enemies isto allow the greatest number of beneficial species to thrive Then perhaps at least one species will be effective against each pest at any tine overcoming the variable abundance of each species within and between years

Programs to introduce exotic species or mass produce indigenous ones are ambitious and expensive because they require trained people on a sustained not ad hoc basis

Chemical control Insecticides offer rapid and efficient control of upland insect pests (190) but-should be used only after other control measures have been considered Insecticides are rarely used on upland rice because of cost (32) Upland rice yields are normally too low to justify the expenditure Also most upland rice is grown as a cash crop Spraying upland rice is more difficult than lowland rice because water is less accessible Government subsidy programs to provide pestishycides to tribal peoples for upland rice production have created problems of toxicity to humans because isolated people have had no experience to be able to handle pesticides safely

However ins-eticide use has beenjustified when a crop is

0Z

Table 8 Parasitization rates of lepidopterous pests in upland rainfed lowland and irrigated lowland environments Philippines 1976-84a

Parasitization (5)

Environment and site Year YSB GFB PSB Earcutting Swarming Skipper Greenborned RLF Brown larva larva caterpillar caterpillar caterpillar caterpillar larva semilooperEgg Larva larva larva larva larva larva

UplandBatangas 1977 - 0 (57) - 0 (63) - 0 (71) - 0 (106) 11(142) 2 (101)1978 43 (4697) 8(451) 5 (38) 3 (40) 6 (61) 2(58) 15 (57) 13 (129) 13 (1286) 4 (30)1979 59(4135) 12(320) 13 (11) 5(28) 9(45) 6(46) 7(23) 8(160) 9(1152) 8(26)

Rainfed lowland Iloilo 1976 - 0 (10) 0(10) 0 (7) 0 (19) 2 13) 15 (180) 5 (76)1978 0 (54) 0 (3) 0 (17) 9 (64) 4 (78) 3 (73) 0 (121) 0 (191)1979 64 (2350) 16 (84) 6 (17) 0(6) 18 (68) 25 (104)

1981 56 (1322) 18(304) 0(5) 12(51) 18(138)Pangasinan 1976 60 (7 7 )b 62(82) 0(111) 5 (39)

1978 58 (53)b 5(51) Cagayan 1979 59(713) 8 (101) 6 (34) 0(7)

1980 0(11) 4 (28) 15 (286)

62 ( 48(177) 61(140)1981 0(5) 12 (2) 13(45) 14 (92)

Irrigatedlowland Laguna 1974 62 8

1974-75 50 41 1975 43 72 1978 96 (623) 21(24) 33 (60)1979 83(2017) 18(110) 8(62) 11 (89) 22(400)1980 78 (1625) 20(75) 0 (16) 4 (72)1983 60

9 (80) 7 (270) 30

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R F Apostol unpublished Iloilo and Pangasnan 136 Cagayan B Canapi unpublished Laguna 175 P CPantua unpublished 118 234 143 Carino and Shepard unpublished YSB = yellow stem borer ScirpophagaincertulasGFB = gold fringed borer Chilo auricilius PSB = pink stem borerSesamia inferens earcutting caterpillar Alythimna separata swarming caterpillar Spcdoptera mauritia acronyctoides skipper Pelopidas mathias reen horned caterpillar Melanitls ledaismene RLF = rice leaffoldersCnaphalocrocismedinalis and Marasmiaspp brown semilooper Mocisfrugalis Figures in parentheses are sample sizes Egg masses

IRPS No 123 January 1987 35

Table 9 Effect of seeding rate and insecticide on plant density and Yield losses from insect pests in upland rice often areyield of Dagge upland rice Batangas Philippines 1976a comparable to those of lowland rainfed or irrigated rice

Seeding rate Insecticide Dosage Plant density Yield (0-Y30o) but usually from different pests The principal (kg aiha) seed treatment (kg aiha) (noIra-row) (tha) groups are soil pests feeding on sown seed and rootsat 14 DE followed by seedling pests These groups have largely gone

50 None - 35 a 20 a unnoticed leading researchers to conclude that stem borers50 Carbofuran F 10 38 ab 23 a and seed bugs are the most important pests (17) Yield loss 10 beCabofuranF100 None - 4551 c 23 a trials that pinpoint major pests need to includc seed

100 Dieldrin WP 10 52 cd 22 a treatments Granules applied in the furrow may not give150 None - 59 d 22 a adequate protection from many seed pests

aAv of 8 fields DE = days after crop emergence In a column means The floristic and pedological diversity of upland rice followed by a common letter are not significantly different at the environments is matched by the diversity of arthropods5level by D)MRT F = flowable Wi = wettable powder both pests and their natural enemies Irrigation has homoshy

genized lowland rice environments Flooded soils tend tothreatened by epidemics of locusts or armyworms New have the same properties and to eliminate soil pests The technology stressing efficiency has minimized insecticide spatial and temporal dominance of cultivated rice has cost benefited specialist pests

Seed treatment is normally inexpensive and can be Upland rice has no specialist arthropod species The economicallyjustified in -nanycases(66) Treating seed with monophagous pests on upland rice prefer the lowlands systemic insecticide (216) but not chlorinated hydrocarbons where rice evolved Rice in the uplands exists only because (247) is most effective in controlling seedling maggots of man and even though lowland rice predates upland riceFoliar sprays have to be repeated to obtain control and specialist species evolved Wheat maize and sorghum do often are washed off by frequent early season rains not have monophagous pests leading us to conclude that Granular insecticides are effective but required dosages are there is no evolutionary advantage for inssects to have too expensive to justify except under heavy infestation narrow host ranges in upland graminaceae(247) When directed against first-instar larvae banding Pests have adapted to the uplands not only by having granules in seed furrows (177) can be effective against white wider plant host ranges but also by having long life cyclesgrubs at low dosages (025 kg aii ha) (134) Seed treatments undergoing dormancy or being dispersive Each pest has a are not promising for white grub control (248) unique life history strategy suited to highly unpredictable

Seed treatment is also effective ainst ants and may be environments In agivenyear someinsect specics an better cheaper than increasing the seeding rate In Batangas exploit upland rice as a food source but no pest is a senious treating 50 kg seed ha was as effective as increasing the one every year on upland rice seeding rate to 100 kgha and using untreated seeds (Table The low yielding potential of upland rice means less 9) In Brazil seed treatments control termites and other soil quantity and lower quality of food for pests to exploit but insects including the lessercornstalk borer(66) Domiciano also less incentive for farmers to undertake control (58) however found seed treatments inconsistent in their The most effective control would be early and synchroshyeffect against high populations of lesser corn stalk borer nous planting of early-maturing upland rice varieties to

Baiting can be an inexpensive way of controlling ants prevent insect pest buildup Breeding for insect resistance seedling maggot flies mole crickets and field crickets Baits should initially focus on stem borers because they are verymade of locally available material and impregnated with difficult to control by other means Biocontrol efforts insecticide can be sparingly distributed taking adantage of should first stress conservation of natural enemies preshythe pests ability to disperse and encounter bait sites dators and pathogens rather than parasites Efficient

The new low-volume hand-held controlled-droplet chemical control includes seed treatment or baiting for seed sprayers may offer an advantage over high-volume knap- seedling and some root pests and using controlled dropletsack sprayes for controlling foliar pests Upland rice sprayers for spot treatments against foliar pests and seed canopies are more open than those in lowland rice and bugs when economic thresholds are reached better droplet penetration should result Controlled droplet applicators would be ideal for seedbugs and defoliators

CONCLUSION

Upland rice is attacked by a wider array of insect pests than is lowland rice mainly beca use of the addition of soil pests Generally population levels of lowland rice pests are lower in upland than lowland rice leading many people to conclude that insect pests are not important on upland rice

36 IRPS No 123 January 1987

REFERENCES CITED

1Abraham C C and K S R Mony 1978 Occurrence ofLeptocorisa acuta Fabr (Coreidae Hemiptera) as a pest of nutmeg trees J Bombay Nat ttistSoc 74553

2 Agyen-Sampong M 1975 HlierogrlphusdiaganensisKrauss (Orthop-tera Acrididae) a new pest of rice in Northern Ghana Ghana J Agric Sci 8249-253

3 Agyen-Samnpong M 1976 Varietal resistance and other inle-ratcd pest control components as factors of rice varietal improvement in West Africa Prcsented at the Second varietal improvement serinar 14-18 Sep 1976 Bouak Ivory Coast West Africa Rice Development Association Monrovia Liberia 18 p

4 Agyen-Sampong M 1977 Outbreak of Ihccrorrdnisorae in rice seedlings near Rokupr Sierra Leone lit Rice Res Newsl 2(6)16-17

5 Agyen-Sampong M 1980 Dirty panicles and rice yield reduction caused by hugs IntRice Res Newsl 5(l)11-12

6 Agyen-Sampong M 1982 The major pest problems of irrigated upland and mangrove swanip rice ecosystems ir the humid tropical Guinea savanna and Salel climate zones Pages 318-328 in Pro-ceedings training course cn concepts techniques and application of irtegrated pest management in rice inWest Africa 10-20 Jan 1982 Fendall Liberia West Africa Rice l)evelopioent Association Monrovia Liberia 505 p

7 Akibo-Betts V I and S A Raymundo 1978 Aphids as rice pests in Sierra Leone lit Rice Res Newsl 3(6)15-16

8 Akinsola F A 1975 Present status of different rice stemborers in parts of Nigeria iicL Entomo Newsl 328

9 Akinsola FA 1979 Ihe biology and ecology of rice stemborers in Nigeria Ph 1) thesis University ol lbadan Nigeria 217 p

10 Akinsola E A 1979 Guidelines of integrated management of insect pests of upland rice in West Africa Presented at the Seminar on integrated management of rice diseases and insect pests at Bobo-Dioulasso Upper Volta 17-22 Sep 1979 West Africa Rice Develop-ment Association Monrovia Liberia

II Akinsola IA 1984 Insect pests of upland rice in Africa Pages 301-305 in An overview of upland rice research Proceedings of the 1982 lIouak Iwsrv Coast upland rice workshop International Rice Research Institute 13O Box 933 Manila Philippines

12 Alam M S V TJohn and Kaung Zan 1985 Insect pests and diseases of rice in Africa Pages 67-82 in Rice improvement in Eastern Central and Southern Africa proceedings of the international rice workshop at Iusaka Zambia 9-19 April 1984

13 Alticri M A A van Schoonhovcn and J D Doll 1977 The ecological role of weeds in insect pest management systems a review illustiated with bean (Phaseohsvulgaris L)cropping systems PANS (Pest Artie News Summ) 23195-205

14 Anmante F 1967 The leaf-cutting ant Atia capiguaraa pest of pastures [in Portuguese] Biol6gico (Sao Paulo) 33113-120 (Rev Appl Entomol A56207)

15Andrade A C and D Puzzi 1951 Preliminary results of experi-merits on the control of Sculptcoris iastantus on sugarcane [in Portuguese] Bioulgico (Sao Paulo) 1744-49 (Rev Appl Entomol A41421422)

16 Appert J 1970 Maliarplia separaelha (rice stem borer) new observations of entomological problems of rice in Madagascar [in French] Agron Trop (Paris) 25329-334

17 Ariadeau Mand Z Harahap 1986 Relevant upland rice breeding objectives Pages 189-197 in Progress in upland rice research Proceedings of the 1985 Jakarta conference International Rice Research Institute 1O Box 933 Manila Philippines

18Asar S and Kisimoto 1975 Seasonal occurrence of the smaller brown planthopper in upland rice [in Japanese] Proc Kanto-Tosan Plant Prot Soc 2228

19 Bakker W 1974 Characterization and ecological aspects of rice yellow mottle virus in Kenya Meded Landbouwhogesh Wage-ningen 152 p

20 Balut F F 1970 White grubs attacking rice Oryza ativa L [in Portuguese] Biol6gico (So Paulo) 36321-322

21 Banerjee S N and L M Pramanik 1967 The lepidopterous stalk borers of rice and their life cycles in the tropics pages 103-124 in The major insect pests ofthe rice plant Proceedings ofa symposium at the International Rice Research Institute September 1964 The Johns Hopkins Press Baltimore Maryland USA

22 Barrion A T and J A Litsinger 1980 Ants a natural enemy of cliaiarac in dryland rice IntRice Res Newsl

5(4)22-23 23 Barrion A T and J A litsinger 1981 The spider fauna of

Philippine rice agroecosystem 1Dryland Philipp Entomnol 5139-166

24 Barrion A T and 1 A Iitsinger 1984 The spider fauna of Philippine rice agroccosystcms II Welland Philipp Eiitomol 611-37

25 Barrion A T1 and J A Litsinger 1984 lihanus (Diptera Tabanidae) eggs an alternative host of rice stei borer (SI3) egg parasite Telenoonis digns (Iliymenoptcra Scelionidae) lit Rice Res Newvsl 9(6)19

26 Barrion A T and J A Litsinger 1985 Identilication of rice leaffolders by wing markings Int Rice Res Newsl 10( 1)24

27 Barrion A T and J A Litsinger 1985 Chctiosspp (Coleoptera Carabidae) a leaffolder ILF)predator lit Rice Res Newsl 10(1)21

28 Bertels A 1970 Insect pests of rice and theircontrol [in Portuguese] Pelotas Brazil Instituto de Expcrimentacao Agropccuararias do Sul Circular 43 24 p

29 Bonzi S M 1982 Ctilo (flisilineus J de Joannis (Lepidoptera Pyralidae) stem borer of irrigated cereals in Upper Volta LioFrench] Agron Trop Riu Rizic Cult Vivrieres Trop 37207-209

30 Bourke T V T L Fenner J N LStihick G Baker EHassan D F OSullivan and C S li 1977 Insect pest survey for the year ending 30th June 1969 Papta New Guinea )epartment of Agrishyculture Stock and Fisheries Port Moresby 57 p

31 Bowden 1 1973 Migration of pests in the tropics Meded Rijksfac Landbouwwet Gent 38785-796

32 Breniere J 1973 The main rice parasites in West Africa and their control Pages 27-42 in Proceedings seminar on plant protection for the rice crop West Africa Rice Development Association Monrovia Liberia

33 Breniere J 1976 The principal insect pests of rice in West Africa and their control West Africa Rice Development Association Monrovia Liberia 52 p

34 Brown ES 1962 The African arinyworn Spoiolptera eXI)Ina (Walker) (Lepidoptera Noctuidae) a review of the literature Cornshymonwealth Institute of Entomology London UK57 p

35 Brown F B 1968 Upland rice in Latin America FAO-International Rice Commission 12th Session of the Working Party on Rice Production and Protection 2-14 Sept Peradeniya Sri lanka Food and Agriculture Organization Rome Italy 6 p

36 Buyckx F J 1 1962 Disease and insect pests of crops inCongo Rwanda and Burundi [iil French] Publications dc LInstilut National pour lestude Agrononiquedu Congo M Weissenbruck S A Brussels Belgium 708 p

37 Cariiio F 0 1981 Role of natural enentics inpopulation suppression and pest management of green rice lealhoppers Ph D thesis University ofthe Philippines at Los Bafitos PhilippinesLaguna 136 -

38 Carpente A J 1983 The development of rice cultivation and extension of vegetable productin in Zanzibar Final Report Vol II1 October 1983 Food and Agricultme Organization Ronte Italy

39 Carroll C P and S1 Risch 1983 Tropical annual cropping systems ant ecology Environ Manage 751-57

4(0 Cates R G 198 1 Host plant predictability and the feeding patterns of monophagous oligophagous and polyphagous insect herbivores Oecologia (Berl) 48319-326

IRPS No 123 January 1987 37

41 Cendcafia S M and F B Caora 1967 Insect pests of rice in the 62 Ene 1 C 1981 I)iiorldhol)ierusihilis Slater (licteroplera Philippines Pages 591-616 in The najor insect pests ofthe rice plant lygacidue Illissinac) a new rice pest front Nigeria lit Rice Res Proceedings of a symposium at the International Rice Research Newsl N3)15 Institute September 1964 The Johns Hopkins Press Baltimore 63 Fugenio T S 1965 rhe effect oi e-vCgetation on rice production Maryland USA 729 p and the possibility of overcoming it by leaf pruning MS thesis

42 Centro Ihternacional de Agricklittira Tropical 1984 Upland rice in University ol the Philippines at Los Baijos ILaguna Ihilippins 69 p Iatin Anerica Pages 93-119 hi An os ervies of uplaind rice research 04 Farrow R A 1975 The Alrican nmigratory locust ii is main Proceedings of the 1982 Boinakamp Ivory Coast upland rice workshop outbreak area of the middle Niger qlinlitatise studies of solitary International Rice Research Institute P0 Blox 933 Manila populations in relation to environmental factors Locusla No II Philippines 198 p

43 Chandra G 1979 Taxonomy ad hn riononlics ofthe imect parasites of 65 Fennal R G 1943 Food crop investigation Windward and rice Icalloppers and planthoppers in the Pihilippines and their Ieeward Islands First report Report onl work dunring the periodiiiportairce iii natural biol)gical cointrol Philipp Fntoltol 4119-139 April 1942-April 1943 Imperial College of Tropical Agriculture

44 Chang T T 1976 The originii colut ion enlIivation dissemination Trinidad 16 p (Rev of Appl Fintornol A3222-25) and diersificition of Asian and African rices -uphytica 25425-44 1 66 Ferreira F 1983 Integrated pest ritaageiret of upland rice [ill

45 Chatierjee P B 1979 Rice leaffolder attacks in India lit Rice Res Portuguese English summary] Pages 323-341 in Upland riceculttre Newsl 4(3)21 factors affecting production E Ferreira T Yamada and F-

46 Cheng Cbien-pan 19(4 Upland rice in Taiwan International Rice Malmolta eds Institutoda Potassae Foslato Institnto Internacional Cnmmission Wiorking Party oit Rice Production ard Protection I()th da Potassa Piracicaha Sao Paulo lraid Meeting 3-10 March 1964 Manila Philippines Food and Agricul- 67 Ferreira E arid F dl Silva Martins 1979 Occurrence of ture Organiaion Rome Italy 5 p Sogatohs orvziola (Muir) inupland rice in Goias Ilrail lit Rice

47 Chcrrett 1 M and 1) 1Peregrine 1976 A review of the status of Rcs Ncwssl 4(4)18 leaf-cutting ants and their control Ann Appl Biol 84124-128 68 Ferreira E and 1F da Silva Martiris 1984 Insect pestof rice ill

48 Chipeta F M 1974 Pests of rice in Malawi Pages 90-95 in Brazil and their control Jill Portugucsej FMIlRAtPA-CNPAF Proceedings Sith eastern Africa cereals research conference 10-15 Goiania Brmiil 67 p March 1974 DRB Mialda ed Ministry of Agriculture lilongwe 69 Ferreira FI F ca Silha Miartins S S Nero and 1 It P de Malawi Carvallio 1982 M tltiplc regression analysis todeterimine the effect of

49 Chu Yau-i 1979 The d atiaging period of the rice seedling fly insect populatiois ott upland rice yield [inIPortuguese Pesqui (Atlterigotiavxigu Stcin) (NItiscide Diptera) oil cori) at Fast Java Agropecu Bras 17671-675 Indonesia Plant Prot Bill 21403413 70 Ferreira F 1F da Silva Martins and F 1PZimmermann 1979

50 Conklin H C 1957 IHaninoo agriculture a report on aln integral Rice varietal resistance to tle lesser corn stalk borer [in Port uguese] systent of shi fti rig cult ivationi in the Pltilippincs Food and Agriculture Pesqui Agropccu Biras 14317-321 Organization Rome Italy 209 p 71 Fleming W F 1968 Biological control of lie hapanese beetle U S

51 Cramer H II 1967 Plant proection and world crop production Dep Agric RLs Sers [ech Bull 1383 78 p Pilaniensclhutz-Nachr 201-524 72 Food and Agriculture Organiation of the United Nations 1982

52 l)ani R C and Majunidar 1978 Prelininary observations on tie Country report surmimary Pages 27-37 in Report on the first incidence of root aphids on differeit rice cultnrcs Sci Cult 4488-89 FAOIUN)IPhail and regional training coiuirse in iniproved ctltural

53 Davidson R I 1979 lffects of root feeding on foliage yield Pages practices for upland rice 5-23 Oct 1981 Northern Region Agrculturd117-120 in T K Ci osb and R B Pottinger eds Proceedings of the Developririt Center Chiang Mai Food and Agricuilture Organiiashy2nd Australian conlerence on grassland invertebrate ecology Govt tion Rome Italy 37 n Printer Wellington New Zealand 73 Franco do Ariaral S 195 1 Residual action of insecticides in turfy

54 l)ean G 1W 1976 Rice insect pests inlaos Int Rice Res Newsl soil--the results of a field experiment [in Portuguese] Biol6gico (Sao 1(2)15 Paulo) 17209-214 (Rev Appl Entomol A41423)

55 I)ean G J W 1978 Insect pests of rice in Laos PANS 24(3)280-289 74 Freeman ) 1970 Report onithe Iban london School of -coroiics 56 )ingle H 1984 Behavior genes and lifc histories conplex adapta- Monographs on Social Anthropology No 41 Athlane Press

tions to uncertain environments Pages 169-194 in A new ecology London UK 317 p novel approaches to interactive systems P W Price C N Slobod- 75 Fuchs T W1 A Harding and 1W Smithlr 1979 Induction and chikoff and W S Gaud cdsl ohn Wiley New York USA termination oldiapause in the sugarcane borer Ann Flitonlol Soc

57 Dobelmann-JP and M Falais 1968 Upland rice cultivation ill tie Am 72271-274 North-western part of Madagascar in 1966-1967 [in French English 76 (hbriel B1) 1975 A review tf the major insect pests of some tpland summary] Trop Agron 2313-21 crops in tlc Philippines Pages 86-111 in Ad Htoc panel of experts on

58 Doniiciano N L 1979 Chemical control of tie lesser corn stalk pest disease arnd weed problems in some rainfed crops Bangkok (lolsonno~paloIgnowslhts) (Zeller 1848) (I-epid(ptera Ihycitid ae) Thailand Food aid Agriculture Organization Roire Italy

on rice in Ponta Gressa Parara 1977-1978 [in Portuguese] Pages 77 Garg D K and N K Shal 1983 White grub outbreak on rainfed 291-303 in Anais 1 Reuniao de tecnicos cmiirizicultura do estado de dryland rice in Uttar Pradeslh Int Rice Res Ncwsl 81(4)16-17 Sao Paulo-Campinas Cati Brazil 78 Garg I) K and IP Tandon 1982 Major insect pests of rice on hilly

59 Dunsmore 1 R 1970 Investigations on tlie varieties pests and tracts of Uttar Pradesh India lilt Rice Res Newsl 7(11)11-12diseases of upland rice (hill padi) in Sarawak Malaysia Int Rice 79 Garrity 1) P 1983 Upland rice research an international bibliograpy Comm Newsl 1929-35 1975-1982 Iternational Rice Research hnstitnte PO Box 933

60 Elias 1R1965 The new decline of rice seedlings is not caused by Manila Philippines 121 p parasites [in Portuguese] Biol6gico (Sao Paulo) 313-6 80 Ghauri M S K 1982 A new subspecies of Iinorh)rone bull

61 Empresa Brasiliera de Pesquisa Agropecuaria 1984 Upland rice itt cornlus Slater (Hemiptera Iygaeidae) front Papua New Guinea on Brazil Pages 121-134 it Av overview of upland rice research rice and carpet grass Ihll lEntoniol Res 72133-137 Proceedings ofthe 1982 Blouak Ivory Coast Upland rice workshop 81 Gonzales F 0 PArregoces R I ihernandef and OT Parada International Rice Research Institute PO Box 933 Manila 1983 Insect and imite pests of rice arid their control in Latin America Philippines [in Spanish] la Federacion Nacional de Arrocerts (Fedearroz)

Bogota Colombia 60 n

38 IRPS No 123 January 1987

82 Grist D H and R J A W Lever 1969 Pests of rice Longmans London UK520 p

83 Hale P R 1979 Two subterranean pests of upland rice in Papua New Guirnca IntRice Res Newsl 4(2)16-17

84 Hale P R and J T Hale 1975 Insect pests of rice in the New Guinea Islands Rice Entomol Newsl 34

85 Harris W V 1969 Termites as pests of crops and trees Common-wealth Institute of Entomology London UK 41 p

86 Hattori 1 1968 A method to distinguish between grass leaf roller (Ctaplha rocis medinalisGuenee) and the rice leaf roller (Susurtia exigua Butler) [in Japanese] Plant Prot (Japan) 23167-170

87 Henry I E and 1A Onsager 1984 Experimental control of the mormon cricket Iahrosshnph by NJsena locustae(Aficrospora Microsporidae) a protozoan parasite of grasshoppers (Ort Acri-didae) Entomophaga 27197-201

88 Hidaka 1 P Vungsilabulr and SKadkao 1974 Studies on ecology and control of the rice gall midge in Thailand Tech Bull 6 TropicalAgricultural Research Center Tokyo Japan 113 p

89 Hincklcy A 1) 1963 Ecology and control of rice planthoppers in Fiji Bull Entomrol Res 53467 481

90 Hirao J 1982 Ecology and chtInical control of the rice leaffolder Jpn Pestic Inf 4114-17

91 Ho DT 1984 Sten borer incifience in rice ecosystems in KenyaEast Africa Pages 307-320 in A overview of upland rice research Proceedings of the 1982 Bouac Ivory Coast upland rice workshop International Rice Research Institute PO Box 933 Manila Philippines

92 Ho )T J 1Njokah and J GKibuka 1983 Studies on rice sten borers in Kenya with emphasis on ifaliarpiaipiraellRag Insect Sci Appl 465-73

93 Ichihara I T Ito A NIatsuo and M Sawada 1971 Chemical control of the rice striped stem borer in upland rice grown in plastic mulch [iii Japanese] Proc Kanto-Tosan Plant Prot Soc 1882

94 IclhiharaIand A Matsuo 1969 Damage caused by rice striped stett borer to upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1686-87

95 Ichhara I atid A MIatsun11970 Conside ration oft lie extraordinary prevalence of rice striped stem borer in upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1778-79

96 Institut de Recherches Agrolnliqes Iropicale et des Cultures Vivrieres 1978 Insect resistance Pages 12-13 in Upland rice 1978 atnual report Bouak Ivory Coast 27 p

97 International Institute for Tropical Agriculture 1983 Aphids itt rice roots I ITA Res Briefs 4(1-3)4-5

98 International Institite fi Tropical Agriculture 1983 Hopperburn in rice caused by brown plathopper IITA Res Briefs 4(l-3)6-7

99 International Rice Research Institute 1975 Annual report for 1974 PO Box 933 Manila Philippines p 216

100 International Rice Restarch Institute 1976 Annual repiort for 1975 PO Box 933 Manila Philippines p 228

101 International Rice Research Institute 1979 Cropping systems program Component technology development and evaluation Page 404 i Annual report for 1978 11O Box 933 Manila Philippines

102 International Rice Research Institute Department of Entomology 1981) Insecticid evaluation 1979 Tables 73 74 PO Box 933Manila Philipes

103 International Rice Research Institute Department of Entonmology 1981 Insecticide evalomtion 1980 Table 81 POBox 933 Manila Philippines

104 International Rice Research Institute 1981 Report of a meeting of Asian rice-based cropping systems entomologists Eleventh cropping systems working group meeting 18-22 May 1981 PO Box 933 Manila Philippines p 132 133 144

105 International Rice Research Institute 1984 Cropping systems program Pages 368-373 in Annual report for 1983 PO Box 933 Manila Philippines

106 International Rice Research Institutc 1984 Upland rice in Asia Pages 45-68 in An overview of upland rice research Proceedings of the 1982 Bouakamp Ivory Coast upland rice workiiop loternational Rice Research Institute 11O Box 933 Manila Philippines

107 Irshad M 1977 Grasshoppers in rice in Pakistan lit Rice Res Newsl 2(3)8-9

108 Ito R and TOno 1964 Upland lice its culture and varieties in Japan lIternational Rice Cormmission loth Meeting of the Working Party on Rice Production and Protection 3-10 March 1964 Manila Philippines Food and Agriculture Organization Rome Italy 12 p

109 Jeffrey M I and V A Dyck 1983 Effect of moonlight on lightshytrap catches of brown planthopper lit Rice Res Newsl 8(6) 13

110 Jennings P R and T Alicia Pineda 1983 Management ofthe rice plnthoppers Sogatodsorivi(coIl in Latin America Presented at the meeting on stable plant resistance FAO and lIndocsian I)inectorate of Food Crops Protection Bali lndonesia 29 Novcniber-2 l)ccemshyber 9 p

IllJimenez Gomez J A 1979 Rice insect pests i lla ios ()riCntales lil Spanish] Pages 99-103 in Rice ciltivation Instituto (olotibiatto Agropecuario Compndio No 29

112 Jongeleen F JJ Soesanto and A Mockidjo 1979 Studies oti pathogens of two species of Scarabacidac froinIndonesia Z Angew Enlomnol 87412-417

113 Jouvenaz 1) PC S lofgren and W A Baiks 1981 Biological control of imported fire antsa review of cirrent knowledge Bull Entomol Soc Am 27203-208

114 Kainoh Y K Shiniiu S MarUani Y Taiaki 1980 lost finding behavior of tie rice bu lplotori+JchinvittjI)allas (liciniptera Corcidac) with special reference to diet pattern ol aggregation and feeding on the rice plant Appl Entomol Zool 15225-233

115 Kakkar K 0 P Sharma and P I Sharma 1977 Note on Ihteronyrchus liodresRedt (Coleopicra I)ynastidae) as asporadic pest of paddy and maie inHimachal Pradesh Corr Agric 156-57

116 Kalode M131 79 Major insect pest problems and their control in upland rice (Unpublishcd mineogriph CWI oJNaaiat lni Chandaratat (198 1)original not seen)

117 Kalshoven L i 98 I The pests of crops in I ldoniesia (translated by P A vatn der Laan) 1 T Ichtiar B3aru-vati Iloeve lakarta Indonesia 701 p

118 Kamal N Q 1981 Suppression of whitebacked planto ner Sogatella fircaifra (Horvath) aLId rice lealfolder (nhal -cis licdb lis (Guenee) populations by natural cnclmie hlI) tiesis Grcgorio Araneta University Foundation Victoncta Park Metto Manila Philippines 179 Ip

119 Karim A N M Reaul 1975 A species ofliealy bug Trioni i70s ceres new oii rice in the Philippines Rice Entoniol Newsl 247

120 Katanyukul Wand C Chandaratat 1981 Insect pestsof uplaid rice and their control Report firstFAOUNlFi Tliailand training course on improved cultural practices for tplad rice Chiang Mai Thailand 5-23 Oct 1981 Food and Agriculture Organiation Rome Italy 22 p

121 Kern F 1955 lhletwioa(I)uninemts) bhhit Biturm(Coleoptera) a new pest of sugarcane inVenezuela [inSpanish with Etnglish sunmary] Agron Trop (Maracay) 527-29 (Rev Appl Eitoniol A4572)

122 Khan K M and S Ghai 1974 White grubs ald their control in India Pesticides (1Bombay) 819-25

123 Kiritani K 1975 Integrated pest management of rice pests Pages 66-70 itt Proceedings platning workshop ott cooperative hIeld research in pest managementP SMotooka ed East-West Center Honololu Hawaii USA

124 Kisimoto R 1967 Ecology of insect vectors forecasting and chemical control Pages 243-255 in The virus diseases of tle rice plant Proceedings of a symposium atthe International Rice Research Institute April 1967 The Johns Hopkins Press altimore Maryshyland USA

125 Kisimoto R and V A Dyck 1976 Climate and rice insects Pages 367-391 in Climate and rice International Rice Research Institute PO Box 933 Manila Philippines

126 Kureha S S Kobayashi SAsari and T Ishihara 1977 Life history and control of the mole cricket Grrllotalpa africana Palisot de Beauvois [in Japanese English translation 7520 1RRI library] Proc Kanto-Tosan Plant Prot Soc 21192-197

127 Kushwaha K S 1981 The black beetle as a pest of rice in Haryana India Int Rice Res Newsl 6(4)14

128 Lawani S M 1982 A review of tile effects of various agronomic practices on cereal stem borers Trop Pest Manage 28266-276

129 Leuck D B 1966 Biology of the lesser cornstalk borer in South Georgia J Econ Entomol 59 797-801

130 lever R J A W 1955 Insect pests of the rice plant in Malaya Int Rice Comm Newsl 1612-21

131 Li Ching-Sing and Shin-Foon Chiu 1951 The study ofthe rice gall midge Iachydiplosis orvza Wood Mason [in Chinese with English summary] 1Taiwan Agric Res 21-13

132 Iitsinger J A 1984 Assesament of the need-based application for rice Proceedings TechnologyTransfer Workshop 15-16 March 1984 International Rice Research Institute 1PO Box 933 Manila Philippines 6 p

133 Litsinger J A 1984 Environmental diversity and insect pest abundance with reference to the Pacific Pages 285-307 in Proceedings sub-regional training course on methods of controlling diseases insects and other pests of plants in the South Pacific 4-20 Oct 1982 Vaini Kingdom of Tonga Nukualofa

134 Litsinger J A RF Apostol and M BObusan 1983 White grub Leucopholis irrorata (Coleoptera Scarabaeidae) pest status popula-tion dynamics and chemical conllol in arice-maize cropping pattern in the Philippines J Econ Entoinol 761133-1138

135 Litsinger J A C de la Cruz F Raymundo A T Barrion M D Lumaban J P Bandong M S Venugopal F Paragna and E Balete 1982 Insect pests and insecticide reponse of dry-seeded bunded rice Pages 179-193 in Report of workshop on cropping systems research in Asia International Rice Research Institute Los Bafos Philippine

136 Litsinger J A M 1) Luinaban J P Bandong A T Barrion W Katanyukul and N Panda 1978 Insect pest management in rainfcd lowland rice agroecosystms of tropical Asia International Rice Research Conference 17-21 April 1978 International Rice Research Institute Los Bafios Philippines

137 Litsinger J A and K Moody 1976 Integrated pest management in multiple cropping systems Pages 293-317 in R I Papendick P A Sanchez and G B Triplett eds Multiple cropping American Society of Agronomy Madison Wisconsin USA

138 Loevinsohn M E 1984 The ecology and control of rice pests in relation to the intensity and synchrony of cultivation Ph D thesis University of London 36) p

139 Loevinsohn M E J A Litsinger N Panda M A Austria R F Apostol C Andrion and J Bandong 1982 A basis for predicting rice insect pest abundance in cropping systems Paper presented at the 1982 International rice research conference on rice and cropping systems 19-23 April 1982 International Rice Research Institute Los Bafios Iaguna Philippines 25 p

140 Lopez A W 1930 Repot of the Entomology Department Pages 145-172 in Annual report of the Philippine Sugar Association 1929-30 Manila

141 Lopez AW 1931 Entomology Department Pages 227-266 in Annual report of the director or research 1930-31 Philippine Sugar Association Manila

142 Lopez A W 1932 Entomology Deparment Pages 252-279 in Annual report of the Philippine Sugar Association 1931-32 Manila

143 Lungibill P and G G Ainslee 1917 The lessercornstalk borer US Dep Agric Agric Res Serv Tech Bull 539 27 p

IRPS No 123 January 1987 39

144 McGuire J U Jr and BS Crandall 1967 Survey of insect pests and plant diseases ofselected food crops of Mexico Central America and Panama U SDepartment of Agriculture International Agriculshytural Development Service Agricultuial Researrh Service October 1967 11 p

145 McNeill Sand T R ESouthwood 1978 The role of nitrogen in the development of insectplant relations Pages 77-9 in lliochemical aspects of plant and animal coevolution J B Ilarbornu ed Academic Press London UK

146 Maebara H 1955 Spatial distribution of lastinstiarlanaeof purplish stem borer (Sesamia iqferens Walker) hibernating in an upland rice field Bull Kagoshima Univ Fac Agric 438-41

147 Mallick E H and N G Hajra 1977 Response of rice plant to defoliation treatments at different developmental stages Indian Agric 2151-56

148 Mansingh A 1971 Physiological classification of dormancies in insects Canl Entomol 103983-10)9

149 Martell C G 1974 Catalogue of plant feeding insects of Mexico [in Spanish] Fitofilo 27(69)1-176

150 Martinez 0 J 1982 Principal rice pests in Ilanos orientalis [in Spanish] P~ages 80-94 in Curso deact nalization de conocimientos en arroz Villaniencis Instituto (olonmbiano Agropecuario

151 Martins J F da Silva F Ferrcira A S Prabhu F 1 P Zimmermann 1980 Use of pesticides for the control of the main subterranean upland rice pests esqui Agropecu Bras 1553-u

152 Martins I F da Silva F Fe eira rod BdamSilveira Pinheiro I Q2 Damage simulation caused by defoliators of upland rice [in Portuguese with English summary] Pesqui Agropecu Bras 171113-1119

153 Martins 1F da Silva N(iyuen van Tan and 1Ida Silveira Pinheiro 1981 Stem borer resistance in upland rice associated with plant morphological characters [in Portuuce Iesqui Agropecu Bras 16187-192

154 Masaki S 1980 Sumnmer diapause Ann Rev Entomiol 251-25 155 Masuzawa T 11 Suwa and F Nakasoji 1983 l)ifkleences of

oviposition preferencc and survival rate of two skipper butterflies Parnara guttata and PIchlpidai iathiais (Iepidoptera HIesperiidac) on rice plant and cogon grass New Entomol 3247-56

156 Maurya D M and C P Vaish 1984 Upland rice in India Pages 69-92 in An overview oftpland rice research Proceedings ofthe 1982 Bouakamp Ivory Coast upland rice workshop International Rice Research Institute IO Box 933 Mnila Philippines

157 Menon M G R R N Katiyai S Kuar and S S Misra 1970 Termites attacking the paddy crop Sci Cult 36294-295

158 Miyahara Y and 1 Nishikubo 1967 Behavior of young nymphs of the red rice root aphid Iholahsipiun ri liabhdontinalisSasaki in relation to movement from the leaf to the roots of upland rice [in Japanese] Pioc Assoc Plant Prot Kyushu 1363-66

159 Mo ljoa Tjien 1952 White grub of upland rice (Holotrichiahelleri Brsk) [in Indonesian with English summary] Contrib Agric Res Stn Bogor No 13134 p

160 Mochida 0 and T Suryana 1979 The occurrence of the brown planthopper and ragged stunt on lowland and upland rice ill Central Java and logyakarta in tile late of January 1979 Entomnolgy Survey Report No 15 Central Research Institute of Agriculture (CRIA) Sukamandi Indonesia I February 1979 9 p

161Monet C 1979 Varietal resistance to upland rice pests [inI French] Pages 35-39 in Annual report 1977 Institut des Savanes Department des Cultures Vivrieres Bouak6 Ivory Coast

162 Monte 0 1942 A caterpillar ofrice-fields[in Portuguese] Biol6gico (Sao Paulo) 8161-163 (Rev Appl Entomol A31414)

163 Munroe I) D 1979 Pest management strategy for hill rice in four resettlement areas in Sarawak Malaysia Pages 89-102 i1 Guidelines for integrated control of rice insect pests Food and Agriculture Organization Plant Production and Protection Paper 14 Rome Italy 115 p

40 IRPS No 123 January 1987

164 Nakasuji F M Ishii and Jen-Zon Ito 1984 Rice skippers in Taiwan and their life histories ZZrC (Taipei) Tech Bull 82 10 p

165 Navas I) 1976 Fall armyworum in rice Pages 99-106 il Ecological animal control by habitat management Proc Tall Timbers Con-ference 28 Feb-I Mar 1974 Fall Timbers Res Stn Misc Publ 6 Tallahassee Horida USA

166 Nickel 1 1 1961 Biological control of rice stem borer A feasibility study Tcch Bull 2 International Rice Rcarch Institute PO Box 933 Manila Philippines IIl p

167 Nigan P M and R A Vernia 1985 Insect pests of uplaild rice in Uttar Pradesh Int Rice Res Newsl 10(6)22

168 Nilakhe S S 1985 -cological observations on spittle bugs with emphasis on their occirence in rice Pcsqtui Agropecu Bras 20407-414

169 Nixon G EJ 195 1 lie association of ats with aphids and coccids Commonwealth Iusti lcte of Entonology Iondon UK 36 p

170 Nurdin F an Ishak Manti 1985 Study oit seedling fly of upland rice Interlational Upland Rice Moitoring Tour Report 26 lebruary-2 MarLI Sukaranti West Sumatra

171 Oka 1 N 1979 Ciltural conttol of the brown planthopper Pages 357-369 in Brown planthopper threat to rice production in Asia International Rice Research Institutc 1p0 Box 933 Manila Philippines

172 Ohni M and A Vilnaii 1975 Rice insect pests of Italy biology of Sipha grcria (Kaltenbach) and its control [in Italian] Riso (Milan) 2459-71

173 Otanes F Q atnd 1 I Sison 194 1 Pests of rice Philipp J Agric 12211-259

174 Painter R I 1951 Insect icsistance in crop plants University Press of Kansits Lawrence Kansas USA 520 p

175 Pantua P C and 1A litsinger 1981 Arthropod abundance in continuous vs biannual rice cropping systeis Int Rice Res Ncwsl 6(2)20-21

176 Pathak M ) and V A l)yck 1975 Sttudics ott insect pests of upland rice Pages 186-191 in Major research in upland rice International Rice Research Institute PO Box 933 Ilanila Philippines

177 Patil B R B M IHasabe and 1) S Ajri 198 I Chemical control of white gritb lIditriihinhorraia Fabr intfesting groundntut paddy atnd sorghum Indian 1Entomol 43232-236

178 Pawar A 1) 1974 (ollection and identification of insect pests of rice rlerminal repirt lInternational Rice Research Institute PO Box 933 Manila Philippines 72 p (tnimco)

179 IerrajU A aid A S Reedy 1965 Observations ott diapausc itt tie steti borer ohrice Rice News Teller (lai-Mar)9-13

180 Pont A C 1972 A review of the Oriental species of theri ona Rondani (I)iptera Muscidac) of econtomic importance Pages 27-102 in Control of sorglium shoot fly M G lomtani and W R Young eds Oxford atd IH1 Ptthl Co New I)clhi India 324 p

181 Prasad S C 1PChand aid M 1PSingh 1983 Incidence of rice gall midge in the upland plateau region of Hihar Oryza 20235-236

182 Rai P S 198 0 Nisaga simpex damage to rice in the hill tracts of South India Int Rice kles News[ 5(I)12

183 Rathorc Y S 1969 Gonocplhahli dcprssunt Fabr as itpredator of (tilo zonwIlhis (Swinhoc) and other moths in field cage at Patnagar Uttar Pradesh Indian 1Entotnol 3127

184 Reed W 1983 Crop losses causcd by insect pists in the developing world Proceedings 10th Internatial Congress of Plant Protection lrighton UK 174-80

185 Reedy M V 1981 Ateactogasterinhens(Walker)a new root-grub pest of tipland paddy and inaize Curr Sci 501076

186 Reveno M A 1968 Catalogue of the insect pests of economic crops in Colombia [in Spaiish] Asociacion Latino Americana de Entomo-logia (AIAE) 129-33

187 Reyes G M 1957 Rice dwarf disease in the Philippines Food and Agriculture Organiiation Plant Prot Bull 617-19

188 Rice S E A A Grigarick and M 0 Way 1982 Effect of leaf and panicle feeding byarmyworm (Lepidoptera Noctuidae) larvae on rice grain yield Environ Entomol 7593-595

189 Risch S J 1) Andow and M A Altieri 1983 Agioccosystem diversity arnd pest control data tentative conclusions and nizw research dirctions Environ Entomol 12625-629

190 Rivera C T 1956 Three organic insecticides in thecontrol of insects affecting upland rice Philipp Agric 39465-472

191 Roffrey 1 1972 Locusta outbreaks in the Philippines Acrida 1177-188

192 Rojas R 1) 1941 The effect of pruning at varying ages upon the yield of Ramairio Phipp Agric 29851-860

193 Roonwal M i 1979 Ternite life and termite control in tropical South Asia Scientilic Pnbl odhpr India 139 p

194 Rosetto C1 F da Silva Martins N C Schmidt and L EAzzini 1978 Damage caused by pasturespittle bugs 4Dvoi)lavopictaandDJ schach) on rice lBragantia Nota 6 (Matcli)25-27

195 Rosztto C1- S S Neto I- Amantc I) Iink 1Garcia-Vieira D M de SoLza N V Bansa t and A M de Olivcira 1971 Rice pests of Brazil [in Portuguese] Pages 149-238 inrFA()-International Rice Commission 2nd meeting of the Rice Committee for the Americas 6-11 l)ecember 1971 Pelotas Rio Grande do Sul Brazil

196 Rothschild G H L 1969 Obseratiois on the armyworm Spodoptera matritia acronrctoiiles Gn (Lep Noctuidae) Bull Entomol Res 59143-160

197 Rothschild G 1- L 1974 Parasites of the armworm Spolopiera mauritiaacroncitoihes in Malaysia Fntornopliaga 19293-299

198 Sachan i N S K Gangwar and 1N Katujar 1980 Field crickets damaging cages in North Eastern IHill Region Indian J Entomol 42526-527

199 Sands 1) P A 1977 The biology and ecology of Lepocorisa (Hemiptera Alydidac) in Papua New Guinea Res Bull 18 Departshyment of Primary Industry Port Moresby Papua New Guinea 104 p

200 Sarawak 1968 titomology division Pages 97-106 in Report of the research branch of the Department of Altriculture 1966 Kuching (Rev AIppl Eintomol A56603)

201 Satpathy JM M S Das and K Naik 1977 Effect of multiple and mixed cropping on the incidence of scine important pestsI Entomol Res 178-85

202 Sauer IHF G 1939 Notes Ott Elasmpnlpualtsligniosellths Zeller (Lep Pyr) a serious pest ofcereals in Sao Paolo [in Portuguese] Arq Inst Biol 10199-206 (Rev Appl Entotnol A28378)

203 Satinders 1L A 1 SKing and C L Vargas S 1983 Crop pests of Central Anerica with t bibliography [in Spanish] Centro Agroshynotnico Tropical de Investigacion y Ensenanza Turrialba Costa Rica (CATIE) Tec Bol 9 90 p

204 Sauphanor B 1985 Some factors of upland rice tolerttce to stem borers in West Africa Inseu Sci Appl 6429434

205 Sen P atnd S Chakravorty 1976 Stentflics in Haritighata Nadia West Bengal India [it Rice Res Newsl 1(12) 15

206 Senapai B and 1M Satpathy 1983 The seedling fly Atherigona orrzae Malloch is a pest of rice in Orissa (India) J Entomol Res 777-79

207 Shhjahan M S Alatn M Ah nad and BN Islam 1983 A new scarabaeid beetle pest of dryland rice in Bangladesh Int Rice Res Ncwsl 8(I)16-17

208 Sharma G L S C Modgol and R C Gautam 1979 Effect of depth of submergence ott incidence of bacterial blight and white grub infestation in transplanted rice Int Rice Res Newsl 4(6)12-13

209 Sharma S 1985 Scarabaeid beetle outbreak itt upland rice Int Rice Res Newsl 1(1(l)24

210 Shiraki T 1917 The paddy borer Schoenobhits incertulas Walker [achaku Agric Exp Stn Formosa 256 p

211 Silva A de Brito and 11P Magalhaes 1981 Insect pests of rice [in Portuguese] Circular Tecnica No 22 EMBRAPA Centro de Pesquisa t ecuaria de Tropico Unido (CPAJU) Belem Brazil 14 p

212 Singh S A 1979 Emergence of rice leaf butterfly as a rice pest in Manipur India int Rice Res Newsl 4(4)16-17

213 SinghS A 1983 Changing trends of insect pestsof rice in Manipur Int Rice Res Newsl 8(f)12

214 Siwi S S and P H van Doesburg 1984 Lenovorisa Latreille in Indonesia (Heteroptera Coreidae Alydinae) Zool Meded (Leiden) 58117-129

215 Soejitno J 1977 Reaction of some upland rice varieties toseedling fly eltherikonaexigua Stein [in Indonesian] Siri Hana I Penyakit 630-35 Lembga I osat Fenclilian Pertanian Bogor Indonesia

216 Socnardi 1967 Insect pests of rice in Indonesia Pages675-683 in The major inse pests of thu rice plant Proceedings of a symposium at the Internatital Rice Rec Wi itt S pternber 1964 The Johns Hopkins Press Baltimore Maryland USA

217 Soto P E A T Petcz and I W Buddenhagen 1976 Survey of insect pests and diseases of rice in different ecological zones in Nigeria Rice Entomol Ncwsl 435-46

218 Soto 11E and Z Ziddiqi 1976 Screening for resistance to African rice insect pests Varietal improsetnent sCmi n r Bouak Ivory Coast 13-17 Sept 1976 West Africa Rice l)evelopment Association Monrovia Iiberia

219 Squire F A 1972 Entomological problems in Bolivia PANS (PestArtic News Sumr) 18249-268

220 Stapely 1 II 1978 The rice leaf roller in the Solomon Islands Int Rice Res Newsl 3(l)13

221 Suenaga H 1937 On aphids attacking the roots of rice Oyo-I)oubts Zasshi 9150-152 [in Japanese] (Rev Appl Entomol A25745)

222 Suenaga H 1952 On the seasonal abundance of three subterranean aphids on upland rice Bull Kyushu Agric Exp Sti 1249-262

223 Sutynso Yas and Dandi Soekaria 1964 Beberapa hana yang ttenperang tana man padi tanah kering [in Indonesian] Rapat kerja Padi Tanah Kering 6 sd 8 Juily 1964 Kaliurang 9 p

224 Tallamy I) W and R F Denno 1983 Alternative life history patterns in risky environments in example from lace bugs Pages 129-147 in Insect life history patterns R F l)cnio and If Dingle eds Spring-Verlag 225 p

225 Tamura 1 1939 Observations on some insect pests of field crops [in Japanese] 1 Plant Prot 26414-420 (Rev Appl EntomolA27-556)

226 Tamura and Suzuki 1964 Influences of the early-transplanting practice in rice culture upon development of the rice stem borer Chilo siql)ressalisWalker Bull Ilokuriku Natil Agric Exp Stn 761-94

227 Tan J P 1924 The rice root aphid Drowhi ihirsoia Philipp Agric 13277-288

228 Tanaka 1 1957 Taxonomy and distribution of some subterranean aphids injurious to upland rice in Japan with description of a new species [in Japanese English summary] Botyu-Kagaku 22168-176

229 Tanaka T 1961 The rice root aphids their ecology and control Spec [full College of Agriculture Utsunomiya University Japan F3 p

230 Tauber M IC A Tauber R Nechols and 1Obrycki 1983 Seasonal activity of parasitoids control by external internal and genetic factors Pages 87-108 itt I)iapause and life cycle strategies in insects V K Brown and I Ilodek eds )r W Junk The Hague Netherlands

231 Taylor W F 1972 Effects of artificial defoliation (simulating pest damage) on varieties of upland rice Exp Agric 879-83

232 feotia T P S and S Natud 1966 Biotoomics of the rice skipper Parnaraniathiis Fabricius (Lepidoptera Hesperiidac) Indian J Entomol 28181-186

233 Teran F 0 1971 Rice pests of Santa Cruz [in Spanish Ministerio de Agricultura Sita Cruz Bolivia Bull 45 8 p

234 Than Itun 1976 Population dynamics of the yellow rice borer Tryporza itcertulas (Walker) and its damage to the rice plant MS thesis University of the Philippines at Ios Bafios Laguna Philippines 92 p

235 Uichanco L B 1911 Pests Pages 55-74 it Rice Philipp Agric 11-200

IRPS No 123 January 1987 41

236 van der Laan P A 195 1 Life history of the rice seedling fly Contrib Gen Agric Res Stn Boor Indonesia 118 15 p

237 van der Laan 11A 1959 Correlation between rainfall in the dry season and the occurrence of the white rice borer (Scirpolhaga innotala WIk) in lava Entomol Fxp Appl 212-20

238 Van Driesche R G 1983 Meaning of percent parasitismin studies of insect parasitism Fnviron Entomol 121 I1- 122

239 Venugopal M S and A litsingcr 1984 Fffect ofcarbofuran on rice growth Prot Ecol 7313-317

240 Vietnam Directorate of Rural Affairs 1962 Annual work progress report on crop improvement program of rice sugarcane vegetable and field crops (lor the period from July 1961 to June 1962) (11) 299 p (Rev Appl Entomol A51526-527)

241 Wan M T K 1972 Observations on rice leaf- and planthoppers in Sarawak (Malaysian Borneo) Malay Agric 148308-335

242 West Africa Rice I)esclopnient Association 1978 Annual research report Vol 2 Statistics Monrovia Liberia

243 West Africa Rice Development Association 1979 Annual research report p47 Monrovia Liberia

244 Williams ) F and CS lofgren 1981 Fli I illy l-468 anew bait toxicant for control of the red imported lire at Fla Entomol 64472477

245 Williams I) 1 1970 The mealybugs OFlomoptera Coccoidea Iscudococcidac) of sugar-cane rice and sorghum Bull Entomol Res 0109-188

246 Williams D) 1 1982 Pulinaria icervi (Signoret) and its allies ott sugar cane and other grasses Btll Entoinol Res 72111-117

247 Wirjosuiardjo S 1976 Study ofchenmical cont rot of insect pests ott upland rice Imiu Pertanian (Jogjakarta) 1323-329

248 Wirjosuhardjo S and N Wirarta 1976 Chemical control of the white grub tIolorithiatlltriin upland rice in M uneng Irobolinggo Indonesia) [in Indonesian] Imut Pertanian (Jogjakarta) 1330-332

249 Yano K T Miyake and V F Eastop 1983 The biology and economic importance of rice aphids (Henuiptera Aphid idae) a review Bull Entomol Res 73539-566

250 Yelp D and P Rhodes 19(4 lhe success of drilled dryland rice iii the Fiji Islands International Rice Commission Working Party on Rice Production and Protection Manila Philippines 3-10 March1964 Food and Agriculture Organization Rome Italy 13 p

251 Young W R 1981 Fifty-five years of research on the sorghum shootfly Insect Sci Appi 23-9

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No 101The economics of hybrid rice production in (hina No 113 Boron toxicity in rice No 102 Rice ratooning No 114 Energy analysis rice production systems and rice research No 103 Growth and development of the (Jeep water rice plant No 115 Production risk and optimal fertilizer rates an application of the

No 104 Faridpur acomputer-assisted instruction nmodel forrainfed lowland random coefficient model No 116 Consumer demand for rice grain quality in Thailand Indonesia andrice

No 105 A readingand listening comprehcnsion test inEnglish for non native the Philippines speakers applying for tiraining at IRIRI No 117 Morphological changes in rice panicle development a review of

No 106 Rice grassy stunt virus 2 a new strain of rice grassy stunt in tire literature

Philippines No 118 IRRI-Korca collaborative project for the development of cold-No 107 Physical losses and quality deterioration in rice plstprodnCtion tolerant lines through anthei culture

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publishers No 120 Changes isinall-farm iLtireliing techii)hgy in Thailand and [he No 109 Classification of Philippine rainfall patterns Philippines No 110 Contributions of modern rice varieties to rutrition in Asia No 121 Landforms and modern rice varieties No Ill Changes in rice breeding in 10 Asian countries 1965-84 No 122 Yield stability and modern rice technology No 112 Design parameters affecting the performance of the IRRI-designed

axial-flow pump

Page 29: UPLANDRICE. INSECTl--1i M T -E!R.EOLOU t RTANCE ,AND …delphacid.s3.amazonaws.com › 6958.pdf · The significance of abiotic factors in upland rice insect ecology will be apparent

IRPS No 123 January 1987 29

Normally predatory thrips prey on herbivorous thrips strepsiptera on both Spiders dominate tilenymphaladultNatural enemies of stem bugs --Scolinoplara and Tibraa predators Spittle bugs move to rice from pasture grasses - are in order of importance egg par~isites and predators and biocontrol would be successful only ifcarried out before fungal pathogens and nymphal adult parasites Egg tiley immigrate into ricefields parasites could either be introduced fron other areas or Biological control tactics for hemipterous seed bugs mass produced and released ould follow those outlined for stein bugs

Armyworms are normally lwld in hcck by the activities Coml)arison of environnewtns Natural enemies are of egg and larval parasites When these natural enemies fail perhaps faced with even greaterconstraints than insect pestsusually because of drought armyworns become epidemic to survive ind reproduce in upland environments TheyIt may be worthwhile therefore to release parasites during have adapted ani survived by having wide host ranges the the rainy months following a drought Virus diseases of ability to aestivate or hibernate over unfavorable seasonslarvae would beanotheravnue to exploreforarnyworms and the ability to disperse or by other mechanisms

Parasites have ben traditionallyconsidered for biolegical Comparing the most dominant and therefore the most control attempts against stein borers usually by rekasing adapted species of natural enemies collected in various exotic species( 166) This approach has not met with success environments in the Philippines we can understand how and new avenues should be explored The role of predators different rice environments are arid pathogens is little inderstood and deserves greater Anagqrms olptahiisa mymarid planthopper egg parasiteattention Egg parasitism rates are normally reasonably is cquallyadapted to all Philippine rice environments (Tablehigh but should be supplemented with effective egg 6) It is selective for planthoppers and even parasitizes eggspredators Orthopterans are potential egg predators of the n-iize planthopper Peregrinitsmaidis itt upland areas Metioche vittaticollis (Sta ) Anavipha nifiwnnis (Saus- Alavohis also parasitizes maize planthopper eggs but is sure) and Conocephahs hongipnni (de i-laan) feed on most prevalent on BPH in the irrigated wetlands (onashystem borer eggs etioche and Anaxil)ha specialize in eggs tocerus spp like A optalhilis on BPH and WBPI Iattacks with no hair such as Chilo spp The omnivorous (ono- the eggs of GLH in ell rie environments It is also found cephahsspecializes in eggs cov red with hairs it is a proven parasitizing the eggs ef the white rice licahopper (oimna egg predator of Scirlouphaga spp but has the discouraging spetra Oirosita naias a trichogrammatid egg parasitehabit of eating rice guin (65) attacks BIPH and WBPH in the wetlands and has no other

Grasshoppers have egg (scelionid) and nymphalad ult host A related species 0 aesopi specializes in G1-H in (nemestrinid tachinid and sarcophagid) v irasites Among wetland environments Dryinid strepsipt-an and pipunshypathogens protozoans have been recorded most frequently culid nymphal and adult parasites of planthoppers and Control of locust species by natural enemies could con- leafhoppers also specialize either in hopper species or centrate on their habitual breeding areas environments Among the dryinids Pseuhdgonatoipsflavi-

Leaffolders have rich complexes of natural enemies some feimur prefers the lowlands and BPH while P nuch has no adapted to upland rice (22 27) ranging from egg predators environmental prefirences but is adapted only to WBPH (gryllids coccinellids) to larval parasites (braconids ichncu- The main d ryinid species on GLH aptogonatop)usspp is monids) to larval prcdators (ants and carabids) to larval most dominant in rainfed lowlands and also attacks other pathogeiis (vira and fungal diseases) Efforts to augment leafhoppers within Amnrasca Cicadulinaand Bachaltuathese natural enemies should focus on the egg predators Strepsiptera are more dominant on plarithoppers than (rearing releasing)and viral or fungal pathogens (culturing lealhoppers The Elenchus species in the Philippines attack spraying) all species of planthoppers within the genera Sogawilla

Parasitization of the large conspicuous larvae of Sogatodels NilaparsataHarualia and Opiconsiva Elenshyskippers green horned caterpillars and other polyphagous cusrvasuntaisuiis adapted to the major rice environments Lepidoptera by tachinids and chalcids is normally low ELg but prefers BPH in rainfed arid WBPIH in wetland environshyparasites and predators are perhaps the key to their control ments Ilalictoph(utsmtnroeiand HVpectrus attack G LH

Gall midges are parasitized by pterormalids and platy- and COfimravcctra respectivelygasterids that aestivate within their larval host between Pipunculids are mainly parasitic oii lealhoppers Each seasons Shoot aphids are preyed upon by coccinellids species has a unique environmental preference Tomosshysyrphids and chrysopids varireIa oraetoraprefers Ne7 otetltix nigropictrs and

Planthoppers arid leafhoppers have egg parasites and Pilunwulhtsjavanensisattacks Dehoceplhahls spp which predators as well as nymphialad nlt parasitesand predators are more abundant in upland than irrigated lowland their rich complex of natural enemies includes fungal environments T suhvirescensattacks Nephotettix virescens pathogens The principal egg parasites are myrnarids and and prefers the lowlands Pilpuiulus nttillattls is an trichogramniatids Egg predators are norinally mirid plant upland rice parasite arid attacks only Nephotetix speciesbugs that suck out the yolk Nymphal ad nIt parasites are More than 17 recorded species of leaffolder parasites in dryinids on planthoppers pipunculids on lealhoppers and the Philippines attack species of C(naplwocrocis and

30 IRPS No 123 January 1987

Table 6 Dominant natural enemies of rice pests in upland rainfed lowland and irrigated lowland rice environments of the Philippines

Natural enemy

Host stage Species

Brown planthop-Parasite Egg

MymaridaeAnagrus optabilis(Perkins) Anagrusflaveolus Waterhouse

TrichogrammatidacOligosita naiasGirault

NymphAdult Dryinidac

Pseudogotatopuslaiifemttr Esaki et Hashimoto Elenchidae

Elenchus yasurnatsui Kifune and Hirashima

Environment

Upand Rainfed lowland Irrigated lowland

Ailaparata lugens

x x X x

x

x x

x X X Whitebacked planthoppers Sogatella furcifera and Sogatodes pusanusEgg

MymaridaeAnagrus optabilis(Perkins)

TrichogrammatidacOl~gositanaias Girault

NymphAdult Dryinidae

Pseudogonatopusnudus Perkins Elenchidae

Elenchus yasutatsui Kifune et Hirashima

Parasite Egg

MymaridaeGonatocerus spp

TrichogrammatidaeOligositaaesopi Girault

NymphAdult Pipunculidac

Tomosvaryella subviresccns Loew Tomosvaryella oryzaetora (Koizumi) Pipunclusmutillatus (de Meijere)

DryinidaeIlaplogonatopusspp

Egg Copidosomopsisnacoleiae (Eady)

Larva BraLonidae

Cardiochiles philipphnensisAshnead lchneumonidac

Trichiomnia cnaphalocrosisUchida Temelucha stangli (Ashmead)

Pupa Chalcididae

BrachymeriaexcarinataGahan Brachymerialasus (Walker) Brachymeria nr tachardiaegroup

Parasite Egg

Scelionidae Telenomus rowani (Gahan)

EulophidacTettasticusschoenobhi Ferriere

Larva Braconidae

Stenobraconnicevillei (Bingham) Iclhncurnonidac

Temelucha phiiippinensis(Ashinead)Temelucha stangli (Ashmead)

x x x

x x

X x X

x x Green leafhopper Nephotettix virescens

X x x

x x

X X X X

X

X Rice leaffolders CnaphalocrocisMarasmia

x X x

X

X X

x x x x x X x x x

Yellow stem borer Scirpophaga incertulas

X X x

x

x

x X XAinauromorphaaccepta metathoracica(Ashmead) x

continued on next page

IRPS No 123 January 1987 31

Table 6 cntinued

Natural enemy Environment

Host stage Species Upland Rainfed lowland Irrigated lowland

Larva-pupa Gold fringed borer Chilo attriciiuslchneumonidae

Xanthopimpla stemmator (Thunberg) x aPupa x

Pteromalidae Trichoinalopsis apanteloctena (Crawford) ax

Pink stem borerb Sesamnia inferensParasite

Larva Braconidae

Stenobracon nicevillei (Bingham) x b b Armyworms Mvthinna SpodopteraLarva

Tachinidac Zygobothriaarroptvora (Robineau-Desvoidy) x x

Braconidac Cotesia spp

X Skipper Pelopidas mathias and green horned caterpillar Melanitis leda ismeneF-gg

TrichogramrnatidacTrichogrannaspp x

Scelionidae Undetermined Scelionid

x Larva

Tachinidae Argyroph lax nitrotibialis (Baranov) X x

Pteromalidae Trichomnalopsis apanteloctena (Crawford)

X Predator

CoenagrionidaeAgriocnenis femina femina Brauer x xIschnura senegalensis tambur

Miridae CrtorhinsliidipennisReuter x x

Nabidac Stcnonabis tagalicus (Stal) x

VeliidaeAlicrovelia douglasi atrolineata (Bergroth) x

Coccinellidae Micraspis crocea (Mulsant) X x x

Vespidae Eumenes campaniformis (Fabricius) x

Formicidac Diaecamna spp xOdontoponera transversa (F Smith) x

Carabidae Chaenius spp OphioneaW- aJasciataSchmidt-Goebel

x

Ophionea ishii ishii Habu x x X

Gryllidav Anaxipta longipennis (Saussur-) x xMetioche ittaticollis (Sial) x x x

Tettigoniidae Conocephalus longipennis (de Haan) x x x

aNo rearing made on rainfed lowland bNo rearing made on rainfed lowland or Irrigated lowland

MrasmiaAmong the Laval parasites the braconid Cardio- lowlands and parasitizes oriental maize borer OsIriniachilesphilijpphensis is most adapted to an upland environ- furnacalis (Guenee) larvae The leaffolder pupal parasitesment It also parasitizes Hydelepta indicata a common not only occur equally in all environments but also haveleaffolder of legumes The ichneumonids Trichomma cna- wvide host ranges- parasitizing species of OsIrinia Marucaphalocrosis and Temehcha stangli occur more in the Hydelepta and Honiona - in maize and legumeslowlands T stangli is most adapted to irrigated lowlands The principal egg parasite of yeltow stem borer (YSB)and also parasitizes Chilo and Scirpophaga stem borer Telenomus rowani is widely adapted to all environmentslarvae T cnaphalocrosis is most prevalent in rainfed and has an unusual alternate host -Tabanus eggs (25)

32 IRPS No 123 January 1987

Tetrastictu schoenobii specializes only in YSB and occurs mainly in irrigated lowlands -- further proof that YSB is most adapted to the wetter environments

Larval parasites of YSB are highly environment-specific The braconid Stenohracon nicevillei occurs only in upland rice but also attacks the pink stem borer (PSB) Tieinchwa phiiiplPwnsis an ichneumonid occurs in upland rice and also parasitizes the larvae of (7hiu stqpresaisinatro-morpha accepta melaihoracica like 7 stangi is mnost adapted to the irrigated lowlands

The dark-headed stem borer(l)HSB) (ih pochit3sus was not collected in all rice environments and the gold fringed borer Chilo auricilius nd PSB were collected only in upland rice Xaantiholiplastenrnaloran ichneumonid larval parasite appears to be widely adapted whereas TrihomaslopisapanitI)cteiiaa pteromalid pupal parasite occurs mostly in wetlands and also parasitizes larvae of Chilo suppressais and Pelhuidas mathias

Armyworms Myit hinin and Spodujipera have two main larval parasites - a tachinid Zrgohothriaatropivora in rainfed environments and a braconid Cotesia spp in irrigated lowlands

The butterfly pests Pelopidas and 4elanitishave tricho-grammatid egg parasites found only in irrigated lowland environments

Two larval parasites show environmental preferences --Argrroph av nigrotihialis a tachinid in rainfed habitats and Trichonialopsisapantelkcnela in irrigated ricefields A limgrotibialisalsoattacks the sweet potato hornwormAgrius convolvtdi and T apanteloctena parasitizes DH S1B

Predators as a group show more distinct environmental preferences The aquatic damselflies Coenagrionidae are most abundant in the lowlands Cvriorhinus is also most adapted to the lowlands The nabid Stenonabis tagalicus occurs only in the uplands and isalso prevalent on legumes The coccinellid lady beetles wasps and ants are more prevalent in upland areas AlicrasIpis cit-ea larvae prey ona variety of aphids attacking legumes and maize Etunienes campanonia vespid mud wasp makes nests in trees and is therefore most adapted to the more botanically diverse uplands Soil-dwelling ants cannot tolerate flooding

Upland environments are habitats to arboreal carabid beetles Three species of Chlaeniusbeetles prey on leaffolder larvae and are perhaps more important than parasites (27)

The gryllids and tettigoniids are widely adapted egg predators not only on rice but also on maize and legumes They feed on eggs of most insects that are laid on leaves

The spidercommunity of upland rice environments isrich in species At one site (Batangas Philippines) 31 species have been recorded (23) Of the 176 spider species recorded in Philippine ricefields about one-half (82) occur in upland rice The spiderspecies of upland rice environments overlap more with those of rainfed lowland than with those of irrigated lowland rice (24) The same study showed that of the three environments irrigated rice has (ie greatest spider species diversity foiiowed by rainfed lowland and upland

However of the 10 most prevalent ricefield spiders 9 were abundant in upland rice showing wide adaptation All three spider guilds -- orb-web space-web and huting spiders shywere prevalent in upland ricefields There were however differences in environmental prelerences for some spider species

Of the orb-wcavers Tetragniathamanlihulatawas partishycila rlv abundant 7 japonica was low and Ltucatge dehcorata was absent in upland rice

As in other environments Aiypeiwnaifirtosana was the most dominant space-web spider however its relative numbers were lowerthan for lowland sites Among hunting spiders Ox|opes javanus and Lcosa leatcost fgla were more abundant and Lvcosa lpseudoanmnlata was less abundant in upland compared with lowland rice envishyroniments

Extensive rearing of rice insect pests in upland rainfed lowland and irrigated sites over an 8-yr period in the Philippines provided insight into the effectiveness of parasites as natural enemies (Table 7 8) Most of toe data are from large sample sizes taken over at least one crop which overcomes the pitfalls expressed by Van Driesche (238)

VSB egg parasitization was surprisingly similar across all environments Larval parasitization was highly dynamic among sites and even years within the same site but showed somewhat lowerlevels in uplandsthan in irrigated wetlands This would have been predicted for egg parasites as well since YSB is adapted to the lowlands and so must be its parasites This isevidence that Telenomus rowani is as good at dispersing as its main hosts Trowani clings onto the anal tufts of female YSB and parasitizes eggs as they are laid This phoresy may explain its wide adaptation The more specialized lowland YSB larval parasites therefore appear more effective than their upland counterparts with more alternate hosts

Low collection levels of PSB larvae especially in the wetlands make comparisons between environments difficult Low levels of larval parasifization were recorded in both rainfed environments Except in the Cagayan rainfed wetland site in 1980 low levels ef larval parasitization were evident at irrigated and rainfed sitesalike No differences by environment were evident Higher levels of parasitization occurred in Sarawak during a 1967 outbreak also from tachinids Rothschild (197) indicated that higher parasishytization occurred in upland rice areas

Skipper and green horned caterpillar larval parasitization rates were generally low and showed no environmental effect htighest activity occurred in Cagayan in 1980 (48 parasitization) Although larval parasitization levels of RLF were variable year to year activity appeared greater in the irrigated lowlands than in the uplands Again the highest levels were recorded in Cagayan in 1980 Cagayan has extensive grasslands and fallow around the rice areas and a relatively long rainfall period Rainfall in 1930 was good for crop growth and apparently for increase of larval parasites

IRPS No 123 January 1987 33

Table 7 Parasitization rates of rice planthoppers and green leafhopper in upland rainfed lowland and irrigated lowland environments Philipshypines 1976-840

Environment and site Year Egg parasitization () Nymphaladult parasitization ()

BPH W13P GLH BPII WBPH GLIH

UplandBatangas

Rainfed lowland

1976 1979 1980 1982

15 17 12

2(47)8(39)

4(51)

9(1800)14 (212)

12(304)

8 (101)6 (127)

9(116)

Iloilo

Cagayan

1976 1977 1978 1981 1980 1981 1982

0 (69)

11

44 48 40

-

-

46 55 44

0 (471)

11

52 56 49

0 (200)9 (214)

10 (1259)8(235)6 (900) 3(800)

-7 (462)

11 (270)3 (313)5 (900) 1 (800)

4 (600)10 (725)10 (362)12(870)5 (900) 3 (800)

Pangasinan

Irrigatedlowland

1984 1976 1978 1979 1982

13 (436) 13(379) 14 (457)

6(14)7 (197)

17(198) 11 (486) 18 (275)

3 (81)10 (274)3 (116)

16(871) 19(88)

28(211)9 (764)5 (647)7 (1204) 8(305)

Laguna 1977 11 7 21 1978 31 1979 14 4 4 28 1980 7 8 20 1981 1983

56 7(278)

26 16 (1650)

16 27(1921)

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R I Apostol unpublished 1lo1lo and Pangasinan 136 Cagayan BCanapi unpublished Liuna 175 I C Pantua unpublished 118 234 43 Carino and Shepard unpublished BPH = brown planthopper Nila parvara htgens W131Pi =whitebacked planthopper Sogatelia furcifera GLH = green leafhopper Nephotettrix virescens Figiires in parentheses are sample sizes

Tauher et al (230) reported that internal external and genetic factors influence the seasonal activity of parasitoids

Hopper egg parasites but not nymphaladult parasites were also parlicularly abundant at the Cagayan site possibly the egg parasites have more alternative hopper prey in the weedy areas than nymphaladult paisites

The seasonal dynamics of hopper egg parasites can be seen for upland rice in Batangas(Fig I 2) and for rainfed lowland lice in Pangasinan( 135)and Cagayan( 105) Levels of BPHWBPI-L and GLH egg parasitization were corn-parable in upland and irrigated lowland environments Except for some high rates of paraMsitization early in the season the parasitization rate tended to be steady in all envilonments However hopper nymphaladult parasi-tizution levels were more similar in the rainfed environments and lower than in irrigated rice

Overall parasites do not appear highly effective by themselves against upland or even lowland pests Among natural enemies parasites have attracted the most attention because they can be more readily assesscd

Predators and pathogens lend themselves better to management practices Predators the most important group of natural enemies in riccfields are difficult to quantify On the other hand pathogens appear less im-portant but can be readily cultured and disseminated

The natural enemy commulnity of upland rice is rich in species and differs significantly from that of lowland rice

These beneficial organisms must be conserved by applying insecticides judiciously particularly sprays Seed or soil placement of chemicals minimizes exposure to natural enemies The strategy to derive the greatest benefit from natural enemies isto allow the greatest number of beneficial species to thrive Then perhaps at least one species will be effective against each pest at any tine overcoming the variable abundance of each species within and between years

Programs to introduce exotic species or mass produce indigenous ones are ambitious and expensive because they require trained people on a sustained not ad hoc basis

Chemical control Insecticides offer rapid and efficient control of upland insect pests (190) but-should be used only after other control measures have been considered Insecticides are rarely used on upland rice because of cost (32) Upland rice yields are normally too low to justify the expenditure Also most upland rice is grown as a cash crop Spraying upland rice is more difficult than lowland rice because water is less accessible Government subsidy programs to provide pestishycides to tribal peoples for upland rice production have created problems of toxicity to humans because isolated people have had no experience to be able to handle pesticides safely

However ins-eticide use has beenjustified when a crop is

0Z

Table 8 Parasitization rates of lepidopterous pests in upland rainfed lowland and irrigated lowland environments Philippines 1976-84a

Parasitization (5)

Environment and site Year YSB GFB PSB Earcutting Swarming Skipper Greenborned RLF Brown larva larva caterpillar caterpillar caterpillar caterpillar larva semilooperEgg Larva larva larva larva larva larva

UplandBatangas 1977 - 0 (57) - 0 (63) - 0 (71) - 0 (106) 11(142) 2 (101)1978 43 (4697) 8(451) 5 (38) 3 (40) 6 (61) 2(58) 15 (57) 13 (129) 13 (1286) 4 (30)1979 59(4135) 12(320) 13 (11) 5(28) 9(45) 6(46) 7(23) 8(160) 9(1152) 8(26)

Rainfed lowland Iloilo 1976 - 0 (10) 0(10) 0 (7) 0 (19) 2 13) 15 (180) 5 (76)1978 0 (54) 0 (3) 0 (17) 9 (64) 4 (78) 3 (73) 0 (121) 0 (191)1979 64 (2350) 16 (84) 6 (17) 0(6) 18 (68) 25 (104)

1981 56 (1322) 18(304) 0(5) 12(51) 18(138)Pangasinan 1976 60 (7 7 )b 62(82) 0(111) 5 (39)

1978 58 (53)b 5(51) Cagayan 1979 59(713) 8 (101) 6 (34) 0(7)

1980 0(11) 4 (28) 15 (286)

62 ( 48(177) 61(140)1981 0(5) 12 (2) 13(45) 14 (92)

Irrigatedlowland Laguna 1974 62 8

1974-75 50 41 1975 43 72 1978 96 (623) 21(24) 33 (60)1979 83(2017) 18(110) 8(62) 11 (89) 22(400)1980 78 (1625) 20(75) 0 (16) 4 (72)1983 60

9 (80) 7 (270) 30

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R F Apostol unpublished Iloilo and Pangasnan 136 Cagayan B Canapi unpublished Laguna 175 P CPantua unpublished 118 234 143 Carino and Shepard unpublished YSB = yellow stem borer ScirpophagaincertulasGFB = gold fringed borer Chilo auricilius PSB = pink stem borerSesamia inferens earcutting caterpillar Alythimna separata swarming caterpillar Spcdoptera mauritia acronyctoides skipper Pelopidas mathias reen horned caterpillar Melanitls ledaismene RLF = rice leaffoldersCnaphalocrocismedinalis and Marasmiaspp brown semilooper Mocisfrugalis Figures in parentheses are sample sizes Egg masses

IRPS No 123 January 1987 35

Table 9 Effect of seeding rate and insecticide on plant density and Yield losses from insect pests in upland rice often areyield of Dagge upland rice Batangas Philippines 1976a comparable to those of lowland rainfed or irrigated rice

Seeding rate Insecticide Dosage Plant density Yield (0-Y30o) but usually from different pests The principal (kg aiha) seed treatment (kg aiha) (noIra-row) (tha) groups are soil pests feeding on sown seed and rootsat 14 DE followed by seedling pests These groups have largely gone

50 None - 35 a 20 a unnoticed leading researchers to conclude that stem borers50 Carbofuran F 10 38 ab 23 a and seed bugs are the most important pests (17) Yield loss 10 beCabofuranF100 None - 4551 c 23 a trials that pinpoint major pests need to includc seed

100 Dieldrin WP 10 52 cd 22 a treatments Granules applied in the furrow may not give150 None - 59 d 22 a adequate protection from many seed pests

aAv of 8 fields DE = days after crop emergence In a column means The floristic and pedological diversity of upland rice followed by a common letter are not significantly different at the environments is matched by the diversity of arthropods5level by D)MRT F = flowable Wi = wettable powder both pests and their natural enemies Irrigation has homoshy

genized lowland rice environments Flooded soils tend tothreatened by epidemics of locusts or armyworms New have the same properties and to eliminate soil pests The technology stressing efficiency has minimized insecticide spatial and temporal dominance of cultivated rice has cost benefited specialist pests

Seed treatment is normally inexpensive and can be Upland rice has no specialist arthropod species The economicallyjustified in -nanycases(66) Treating seed with monophagous pests on upland rice prefer the lowlands systemic insecticide (216) but not chlorinated hydrocarbons where rice evolved Rice in the uplands exists only because (247) is most effective in controlling seedling maggots of man and even though lowland rice predates upland riceFoliar sprays have to be repeated to obtain control and specialist species evolved Wheat maize and sorghum do often are washed off by frequent early season rains not have monophagous pests leading us to conclude that Granular insecticides are effective but required dosages are there is no evolutionary advantage for inssects to have too expensive to justify except under heavy infestation narrow host ranges in upland graminaceae(247) When directed against first-instar larvae banding Pests have adapted to the uplands not only by having granules in seed furrows (177) can be effective against white wider plant host ranges but also by having long life cyclesgrubs at low dosages (025 kg aii ha) (134) Seed treatments undergoing dormancy or being dispersive Each pest has a are not promising for white grub control (248) unique life history strategy suited to highly unpredictable

Seed treatment is also effective ainst ants and may be environments In agivenyear someinsect specics an better cheaper than increasing the seeding rate In Batangas exploit upland rice as a food source but no pest is a senious treating 50 kg seed ha was as effective as increasing the one every year on upland rice seeding rate to 100 kgha and using untreated seeds (Table The low yielding potential of upland rice means less 9) In Brazil seed treatments control termites and other soil quantity and lower quality of food for pests to exploit but insects including the lessercornstalk borer(66) Domiciano also less incentive for farmers to undertake control (58) however found seed treatments inconsistent in their The most effective control would be early and synchroshyeffect against high populations of lesser corn stalk borer nous planting of early-maturing upland rice varieties to

Baiting can be an inexpensive way of controlling ants prevent insect pest buildup Breeding for insect resistance seedling maggot flies mole crickets and field crickets Baits should initially focus on stem borers because they are verymade of locally available material and impregnated with difficult to control by other means Biocontrol efforts insecticide can be sparingly distributed taking adantage of should first stress conservation of natural enemies preshythe pests ability to disperse and encounter bait sites dators and pathogens rather than parasites Efficient

The new low-volume hand-held controlled-droplet chemical control includes seed treatment or baiting for seed sprayers may offer an advantage over high-volume knap- seedling and some root pests and using controlled dropletsack sprayes for controlling foliar pests Upland rice sprayers for spot treatments against foliar pests and seed canopies are more open than those in lowland rice and bugs when economic thresholds are reached better droplet penetration should result Controlled droplet applicators would be ideal for seedbugs and defoliators

CONCLUSION

Upland rice is attacked by a wider array of insect pests than is lowland rice mainly beca use of the addition of soil pests Generally population levels of lowland rice pests are lower in upland than lowland rice leading many people to conclude that insect pests are not important on upland rice

36 IRPS No 123 January 1987

REFERENCES CITED

1Abraham C C and K S R Mony 1978 Occurrence ofLeptocorisa acuta Fabr (Coreidae Hemiptera) as a pest of nutmeg trees J Bombay Nat ttistSoc 74553

2 Agyen-Sampong M 1975 HlierogrlphusdiaganensisKrauss (Orthop-tera Acrididae) a new pest of rice in Northern Ghana Ghana J Agric Sci 8249-253

3 Agyen-Samnpong M 1976 Varietal resistance and other inle-ratcd pest control components as factors of rice varietal improvement in West Africa Prcsented at the Second varietal improvement serinar 14-18 Sep 1976 Bouak Ivory Coast West Africa Rice Development Association Monrovia Liberia 18 p

4 Agyen-Sampong M 1977 Outbreak of Ihccrorrdnisorae in rice seedlings near Rokupr Sierra Leone lit Rice Res Newsl 2(6)16-17

5 Agyen-Sampong M 1980 Dirty panicles and rice yield reduction caused by hugs IntRice Res Newsl 5(l)11-12

6 Agyen-Sampong M 1982 The major pest problems of irrigated upland and mangrove swanip rice ecosystems ir the humid tropical Guinea savanna and Salel climate zones Pages 318-328 in Pro-ceedings training course cn concepts techniques and application of irtegrated pest management in rice inWest Africa 10-20 Jan 1982 Fendall Liberia West Africa Rice l)evelopioent Association Monrovia Liberia 505 p

7 Akibo-Betts V I and S A Raymundo 1978 Aphids as rice pests in Sierra Leone lit Rice Res Newsl 3(6)15-16

8 Akinsola F A 1975 Present status of different rice stemborers in parts of Nigeria iicL Entomo Newsl 328

9 Akinsola FA 1979 Ihe biology and ecology of rice stemborers in Nigeria Ph 1) thesis University ol lbadan Nigeria 217 p

10 Akinsola E A 1979 Guidelines of integrated management of insect pests of upland rice in West Africa Presented at the Seminar on integrated management of rice diseases and insect pests at Bobo-Dioulasso Upper Volta 17-22 Sep 1979 West Africa Rice Develop-ment Association Monrovia Liberia

II Akinsola IA 1984 Insect pests of upland rice in Africa Pages 301-305 in An overview of upland rice research Proceedings of the 1982 lIouak Iwsrv Coast upland rice workshop International Rice Research Institute 13O Box 933 Manila Philippines

12 Alam M S V TJohn and Kaung Zan 1985 Insect pests and diseases of rice in Africa Pages 67-82 in Rice improvement in Eastern Central and Southern Africa proceedings of the international rice workshop at Iusaka Zambia 9-19 April 1984

13 Alticri M A A van Schoonhovcn and J D Doll 1977 The ecological role of weeds in insect pest management systems a review illustiated with bean (Phaseohsvulgaris L)cropping systems PANS (Pest Artie News Summ) 23195-205

14 Anmante F 1967 The leaf-cutting ant Atia capiguaraa pest of pastures [in Portuguese] Biol6gico (Sao Paulo) 33113-120 (Rev Appl Entomol A56207)

15Andrade A C and D Puzzi 1951 Preliminary results of experi-merits on the control of Sculptcoris iastantus on sugarcane [in Portuguese] Bioulgico (Sao Paulo) 1744-49 (Rev Appl Entomol A41421422)

16 Appert J 1970 Maliarplia separaelha (rice stem borer) new observations of entomological problems of rice in Madagascar [in French] Agron Trop (Paris) 25329-334

17 Ariadeau Mand Z Harahap 1986 Relevant upland rice breeding objectives Pages 189-197 in Progress in upland rice research Proceedings of the 1985 Jakarta conference International Rice Research Institute 1O Box 933 Manila Philippines

18Asar S and Kisimoto 1975 Seasonal occurrence of the smaller brown planthopper in upland rice [in Japanese] Proc Kanto-Tosan Plant Prot Soc 2228

19 Bakker W 1974 Characterization and ecological aspects of rice yellow mottle virus in Kenya Meded Landbouwhogesh Wage-ningen 152 p

20 Balut F F 1970 White grubs attacking rice Oryza ativa L [in Portuguese] Biol6gico (So Paulo) 36321-322

21 Banerjee S N and L M Pramanik 1967 The lepidopterous stalk borers of rice and their life cycles in the tropics pages 103-124 in The major insect pests ofthe rice plant Proceedings ofa symposium at the International Rice Research Institute September 1964 The Johns Hopkins Press Baltimore Maryland USA

22 Barrion A T and J A Litsinger 1980 Ants a natural enemy of cliaiarac in dryland rice IntRice Res Newsl

5(4)22-23 23 Barrion A T and J A litsinger 1981 The spider fauna of

Philippine rice agroecosystem 1Dryland Philipp Entomnol 5139-166

24 Barrion A T and 1 A Iitsinger 1984 The spider fauna of Philippine rice agroccosystcms II Welland Philipp Eiitomol 611-37

25 Barrion A T1 and J A Litsinger 1984 lihanus (Diptera Tabanidae) eggs an alternative host of rice stei borer (SI3) egg parasite Telenoonis digns (Iliymenoptcra Scelionidae) lit Rice Res Newvsl 9(6)19

26 Barrion A T and J A Litsinger 1985 Identilication of rice leaffolders by wing markings Int Rice Res Newsl 10( 1)24

27 Barrion A T and J A Litsinger 1985 Chctiosspp (Coleoptera Carabidae) a leaffolder ILF)predator lit Rice Res Newsl 10(1)21

28 Bertels A 1970 Insect pests of rice and theircontrol [in Portuguese] Pelotas Brazil Instituto de Expcrimentacao Agropccuararias do Sul Circular 43 24 p

29 Bonzi S M 1982 Ctilo (flisilineus J de Joannis (Lepidoptera Pyralidae) stem borer of irrigated cereals in Upper Volta LioFrench] Agron Trop Riu Rizic Cult Vivrieres Trop 37207-209

30 Bourke T V T L Fenner J N LStihick G Baker EHassan D F OSullivan and C S li 1977 Insect pest survey for the year ending 30th June 1969 Papta New Guinea )epartment of Agrishyculture Stock and Fisheries Port Moresby 57 p

31 Bowden 1 1973 Migration of pests in the tropics Meded Rijksfac Landbouwwet Gent 38785-796

32 Breniere J 1973 The main rice parasites in West Africa and their control Pages 27-42 in Proceedings seminar on plant protection for the rice crop West Africa Rice Development Association Monrovia Liberia

33 Breniere J 1976 The principal insect pests of rice in West Africa and their control West Africa Rice Development Association Monrovia Liberia 52 p

34 Brown ES 1962 The African arinyworn Spoiolptera eXI)Ina (Walker) (Lepidoptera Noctuidae) a review of the literature Cornshymonwealth Institute of Entomology London UK57 p

35 Brown F B 1968 Upland rice in Latin America FAO-International Rice Commission 12th Session of the Working Party on Rice Production and Protection 2-14 Sept Peradeniya Sri lanka Food and Agriculture Organization Rome Italy 6 p

36 Buyckx F J 1 1962 Disease and insect pests of crops inCongo Rwanda and Burundi [iil French] Publications dc LInstilut National pour lestude Agrononiquedu Congo M Weissenbruck S A Brussels Belgium 708 p

37 Cariiio F 0 1981 Role of natural enentics inpopulation suppression and pest management of green rice lealhoppers Ph D thesis University ofthe Philippines at Los Bafitos PhilippinesLaguna 136 -

38 Carpente A J 1983 The development of rice cultivation and extension of vegetable productin in Zanzibar Final Report Vol II1 October 1983 Food and Agricultme Organization Ronte Italy

39 Carroll C P and S1 Risch 1983 Tropical annual cropping systems ant ecology Environ Manage 751-57

4(0 Cates R G 198 1 Host plant predictability and the feeding patterns of monophagous oligophagous and polyphagous insect herbivores Oecologia (Berl) 48319-326

IRPS No 123 January 1987 37

41 Cendcafia S M and F B Caora 1967 Insect pests of rice in the 62 Ene 1 C 1981 I)iiorldhol)ierusihilis Slater (licteroplera Philippines Pages 591-616 in The najor insect pests ofthe rice plant lygacidue Illissinac) a new rice pest front Nigeria lit Rice Res Proceedings of a symposium at the International Rice Research Newsl N3)15 Institute September 1964 The Johns Hopkins Press Baltimore 63 Fugenio T S 1965 rhe effect oi e-vCgetation on rice production Maryland USA 729 p and the possibility of overcoming it by leaf pruning MS thesis

42 Centro Ihternacional de Agricklittira Tropical 1984 Upland rice in University ol the Philippines at Los Baijos ILaguna Ihilippins 69 p Iatin Anerica Pages 93-119 hi An os ervies of uplaind rice research 04 Farrow R A 1975 The Alrican nmigratory locust ii is main Proceedings of the 1982 Boinakamp Ivory Coast upland rice workshop outbreak area of the middle Niger qlinlitatise studies of solitary International Rice Research Institute P0 Blox 933 Manila populations in relation to environmental factors Locusla No II Philippines 198 p

43 Chandra G 1979 Taxonomy ad hn riononlics ofthe imect parasites of 65 Fennal R G 1943 Food crop investigation Windward and rice Icalloppers and planthoppers in the Pihilippines and their Ieeward Islands First report Report onl work dunring the periodiiiportairce iii natural biol)gical cointrol Philipp Fntoltol 4119-139 April 1942-April 1943 Imperial College of Tropical Agriculture

44 Chang T T 1976 The originii colut ion enlIivation dissemination Trinidad 16 p (Rev of Appl Fintornol A3222-25) and diersificition of Asian and African rices -uphytica 25425-44 1 66 Ferreira F 1983 Integrated pest ritaageiret of upland rice [ill

45 Chatierjee P B 1979 Rice leaffolder attacks in India lit Rice Res Portuguese English summary] Pages 323-341 in Upland riceculttre Newsl 4(3)21 factors affecting production E Ferreira T Yamada and F-

46 Cheng Cbien-pan 19(4 Upland rice in Taiwan International Rice Malmolta eds Institutoda Potassae Foslato Institnto Internacional Cnmmission Wiorking Party oit Rice Production ard Protection I()th da Potassa Piracicaha Sao Paulo lraid Meeting 3-10 March 1964 Manila Philippines Food and Agricul- 67 Ferreira E arid F dl Silva Martins 1979 Occurrence of ture Organiaion Rome Italy 5 p Sogatohs orvziola (Muir) inupland rice in Goias Ilrail lit Rice

47 Chcrrett 1 M and 1) 1Peregrine 1976 A review of the status of Rcs Ncwssl 4(4)18 leaf-cutting ants and their control Ann Appl Biol 84124-128 68 Ferreira E and 1F da Silva Martiris 1984 Insect pestof rice ill

48 Chipeta F M 1974 Pests of rice in Malawi Pages 90-95 in Brazil and their control Jill Portugucsej FMIlRAtPA-CNPAF Proceedings Sith eastern Africa cereals research conference 10-15 Goiania Brmiil 67 p March 1974 DRB Mialda ed Ministry of Agriculture lilongwe 69 Ferreira FI F ca Silha Miartins S S Nero and 1 It P de Malawi Carvallio 1982 M tltiplc regression analysis todeterimine the effect of

49 Chu Yau-i 1979 The d atiaging period of the rice seedling fly insect populatiois ott upland rice yield [inIPortuguese Pesqui (Atlterigotiavxigu Stcin) (NItiscide Diptera) oil cori) at Fast Java Agropecu Bras 17671-675 Indonesia Plant Prot Bill 21403413 70 Ferreira F 1F da Silva Martins and F 1PZimmermann 1979

50 Conklin H C 1957 IHaninoo agriculture a report on aln integral Rice varietal resistance to tle lesser corn stalk borer [in Port uguese] systent of shi fti rig cult ivationi in the Pltilippincs Food and Agriculture Pesqui Agropccu Biras 14317-321 Organization Rome Italy 209 p 71 Fleming W F 1968 Biological control of lie hapanese beetle U S

51 Cramer H II 1967 Plant proection and world crop production Dep Agric RLs Sers [ech Bull 1383 78 p Pilaniensclhutz-Nachr 201-524 72 Food and Agriculture Organiation of the United Nations 1982

52 l)ani R C and Majunidar 1978 Prelininary observations on tie Country report surmimary Pages 27-37 in Report on the first incidence of root aphids on differeit rice cultnrcs Sci Cult 4488-89 FAOIUN)IPhail and regional training coiuirse in iniproved ctltural

53 Davidson R I 1979 lffects of root feeding on foliage yield Pages practices for upland rice 5-23 Oct 1981 Northern Region Agrculturd117-120 in T K Ci osb and R B Pottinger eds Proceedings of the Developririt Center Chiang Mai Food and Agricuilture Organiiashy2nd Australian conlerence on grassland invertebrate ecology Govt tion Rome Italy 37 n Printer Wellington New Zealand 73 Franco do Ariaral S 195 1 Residual action of insecticides in turfy

54 l)ean G 1W 1976 Rice insect pests inlaos Int Rice Res Newsl soil--the results of a field experiment [in Portuguese] Biol6gico (Sao 1(2)15 Paulo) 17209-214 (Rev Appl Entomol A41423)

55 I)ean G J W 1978 Insect pests of rice in Laos PANS 24(3)280-289 74 Freeman ) 1970 Report onithe Iban london School of -coroiics 56 )ingle H 1984 Behavior genes and lifc histories conplex adapta- Monographs on Social Anthropology No 41 Athlane Press

tions to uncertain environments Pages 169-194 in A new ecology London UK 317 p novel approaches to interactive systems P W Price C N Slobod- 75 Fuchs T W1 A Harding and 1W Smithlr 1979 Induction and chikoff and W S Gaud cdsl ohn Wiley New York USA termination oldiapause in the sugarcane borer Ann Flitonlol Soc

57 Dobelmann-JP and M Falais 1968 Upland rice cultivation ill tie Am 72271-274 North-western part of Madagascar in 1966-1967 [in French English 76 (hbriel B1) 1975 A review tf the major insect pests of some tpland summary] Trop Agron 2313-21 crops in tlc Philippines Pages 86-111 in Ad Htoc panel of experts on

58 Doniiciano N L 1979 Chemical control of tie lesser corn stalk pest disease arnd weed problems in some rainfed crops Bangkok (lolsonno~paloIgnowslhts) (Zeller 1848) (I-epid(ptera Ihycitid ae) Thailand Food aid Agriculture Organization Roire Italy

on rice in Ponta Gressa Parara 1977-1978 [in Portuguese] Pages 77 Garg D K and N K Shal 1983 White grub outbreak on rainfed 291-303 in Anais 1 Reuniao de tecnicos cmiirizicultura do estado de dryland rice in Uttar Pradeslh Int Rice Res Ncwsl 81(4)16-17 Sao Paulo-Campinas Cati Brazil 78 Garg I) K and IP Tandon 1982 Major insect pests of rice on hilly

59 Dunsmore 1 R 1970 Investigations on tlie varieties pests and tracts of Uttar Pradesh India lilt Rice Res Newsl 7(11)11-12diseases of upland rice (hill padi) in Sarawak Malaysia Int Rice 79 Garrity 1) P 1983 Upland rice research an international bibliograpy Comm Newsl 1929-35 1975-1982 Iternational Rice Research hnstitnte PO Box 933

60 Elias 1R1965 The new decline of rice seedlings is not caused by Manila Philippines 121 p parasites [in Portuguese] Biol6gico (Sao Paulo) 313-6 80 Ghauri M S K 1982 A new subspecies of Iinorh)rone bull

61 Empresa Brasiliera de Pesquisa Agropecuaria 1984 Upland rice itt cornlus Slater (Hemiptera Iygaeidae) front Papua New Guinea on Brazil Pages 121-134 it Av overview of upland rice research rice and carpet grass Ihll lEntoniol Res 72133-137 Proceedings ofthe 1982 Blouak Ivory Coast Upland rice workshop 81 Gonzales F 0 PArregoces R I ihernandef and OT Parada International Rice Research Institute PO Box 933 Manila 1983 Insect and imite pests of rice arid their control in Latin America Philippines [in Spanish] la Federacion Nacional de Arrocerts (Fedearroz)

Bogota Colombia 60 n

38 IRPS No 123 January 1987

82 Grist D H and R J A W Lever 1969 Pests of rice Longmans London UK520 p

83 Hale P R 1979 Two subterranean pests of upland rice in Papua New Guirnca IntRice Res Newsl 4(2)16-17

84 Hale P R and J T Hale 1975 Insect pests of rice in the New Guinea Islands Rice Entomol Newsl 34

85 Harris W V 1969 Termites as pests of crops and trees Common-wealth Institute of Entomology London UK 41 p

86 Hattori 1 1968 A method to distinguish between grass leaf roller (Ctaplha rocis medinalisGuenee) and the rice leaf roller (Susurtia exigua Butler) [in Japanese] Plant Prot (Japan) 23167-170

87 Henry I E and 1A Onsager 1984 Experimental control of the mormon cricket Iahrosshnph by NJsena locustae(Aficrospora Microsporidae) a protozoan parasite of grasshoppers (Ort Acri-didae) Entomophaga 27197-201

88 Hidaka 1 P Vungsilabulr and SKadkao 1974 Studies on ecology and control of the rice gall midge in Thailand Tech Bull 6 TropicalAgricultural Research Center Tokyo Japan 113 p

89 Hincklcy A 1) 1963 Ecology and control of rice planthoppers in Fiji Bull Entomrol Res 53467 481

90 Hirao J 1982 Ecology and chtInical control of the rice leaffolder Jpn Pestic Inf 4114-17

91 Ho DT 1984 Sten borer incifience in rice ecosystems in KenyaEast Africa Pages 307-320 in A overview of upland rice research Proceedings of the 1982 Bouac Ivory Coast upland rice workshop International Rice Research Institute PO Box 933 Manila Philippines

92 Ho )T J 1Njokah and J GKibuka 1983 Studies on rice sten borers in Kenya with emphasis on ifaliarpiaipiraellRag Insect Sci Appl 465-73

93 Ichihara I T Ito A NIatsuo and M Sawada 1971 Chemical control of the rice striped stem borer in upland rice grown in plastic mulch [iii Japanese] Proc Kanto-Tosan Plant Prot Soc 1882

94 IclhiharaIand A Matsuo 1969 Damage caused by rice striped stett borer to upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1686-87

95 Ichhara I atid A MIatsun11970 Conside ration oft lie extraordinary prevalence of rice striped stem borer in upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1778-79

96 Institut de Recherches Agrolnliqes Iropicale et des Cultures Vivrieres 1978 Insect resistance Pages 12-13 in Upland rice 1978 atnual report Bouak Ivory Coast 27 p

97 International Institute for Tropical Agriculture 1983 Aphids itt rice roots I ITA Res Briefs 4(1-3)4-5

98 International Institite fi Tropical Agriculture 1983 Hopperburn in rice caused by brown plathopper IITA Res Briefs 4(l-3)6-7

99 International Rice Research Institute 1975 Annual report for 1974 PO Box 933 Manila Philippines p 216

100 International Rice Restarch Institute 1976 Annual repiort for 1975 PO Box 933 Manila Philippines p 228

101 International Rice Research Institute 1979 Cropping systems program Component technology development and evaluation Page 404 i Annual report for 1978 11O Box 933 Manila Philippines

102 International Rice Research Institute Department of Entomology 1981) Insecticid evaluation 1979 Tables 73 74 PO Box 933Manila Philipes

103 International Rice Research Institute Department of Entonmology 1981 Insecticide evalomtion 1980 Table 81 POBox 933 Manila Philippines

104 International Rice Research Institute 1981 Report of a meeting of Asian rice-based cropping systems entomologists Eleventh cropping systems working group meeting 18-22 May 1981 PO Box 933 Manila Philippines p 132 133 144

105 International Rice Research Institute 1984 Cropping systems program Pages 368-373 in Annual report for 1983 PO Box 933 Manila Philippines

106 International Rice Research Institutc 1984 Upland rice in Asia Pages 45-68 in An overview of upland rice research Proceedings of the 1982 Bouakamp Ivory Coast upland rice workiiop loternational Rice Research Institute 11O Box 933 Manila Philippines

107 Irshad M 1977 Grasshoppers in rice in Pakistan lit Rice Res Newsl 2(3)8-9

108 Ito R and TOno 1964 Upland lice its culture and varieties in Japan lIternational Rice Cormmission loth Meeting of the Working Party on Rice Production and Protection 3-10 March 1964 Manila Philippines Food and Agriculture Organization Rome Italy 12 p

109 Jeffrey M I and V A Dyck 1983 Effect of moonlight on lightshytrap catches of brown planthopper lit Rice Res Newsl 8(6) 13

110 Jennings P R and T Alicia Pineda 1983 Management ofthe rice plnthoppers Sogatodsorivi(coIl in Latin America Presented at the meeting on stable plant resistance FAO and lIndocsian I)inectorate of Food Crops Protection Bali lndonesia 29 Novcniber-2 l)ccemshyber 9 p

IllJimenez Gomez J A 1979 Rice insect pests i lla ios ()riCntales lil Spanish] Pages 99-103 in Rice ciltivation Instituto (olotibiatto Agropecuario Compndio No 29

112 Jongeleen F JJ Soesanto and A Mockidjo 1979 Studies oti pathogens of two species of Scarabacidac froinIndonesia Z Angew Enlomnol 87412-417

113 Jouvenaz 1) PC S lofgren and W A Baiks 1981 Biological control of imported fire antsa review of cirrent knowledge Bull Entomol Soc Am 27203-208

114 Kainoh Y K Shiniiu S MarUani Y Taiaki 1980 lost finding behavior of tie rice bu lplotori+JchinvittjI)allas (liciniptera Corcidac) with special reference to diet pattern ol aggregation and feeding on the rice plant Appl Entomol Zool 15225-233

115 Kakkar K 0 P Sharma and P I Sharma 1977 Note on Ihteronyrchus liodresRedt (Coleopicra I)ynastidae) as asporadic pest of paddy and maie inHimachal Pradesh Corr Agric 156-57

116 Kalode M131 79 Major insect pest problems and their control in upland rice (Unpublishcd mineogriph CWI oJNaaiat lni Chandaratat (198 1)original not seen)

117 Kalshoven L i 98 I The pests of crops in I ldoniesia (translated by P A vatn der Laan) 1 T Ichtiar B3aru-vati Iloeve lakarta Indonesia 701 p

118 Kamal N Q 1981 Suppression of whitebacked planto ner Sogatella fircaifra (Horvath) aLId rice lealfolder (nhal -cis licdb lis (Guenee) populations by natural cnclmie hlI) tiesis Grcgorio Araneta University Foundation Victoncta Park Metto Manila Philippines 179 Ip

119 Karim A N M Reaul 1975 A species ofliealy bug Trioni i70s ceres new oii rice in the Philippines Rice Entoniol Newsl 247

120 Katanyukul Wand C Chandaratat 1981 Insect pestsof uplaid rice and their control Report firstFAOUNlFi Tliailand training course on improved cultural practices for tplad rice Chiang Mai Thailand 5-23 Oct 1981 Food and Agriculture Organiation Rome Italy 22 p

121 Kern F 1955 lhletwioa(I)uninemts) bhhit Biturm(Coleoptera) a new pest of sugarcane inVenezuela [inSpanish with Etnglish sunmary] Agron Trop (Maracay) 527-29 (Rev Appl Eitoniol A4572)

122 Khan K M and S Ghai 1974 White grubs ald their control in India Pesticides (1Bombay) 819-25

123 Kiritani K 1975 Integrated pest management of rice pests Pages 66-70 itt Proceedings platning workshop ott cooperative hIeld research in pest managementP SMotooka ed East-West Center Honololu Hawaii USA

124 Kisimoto R 1967 Ecology of insect vectors forecasting and chemical control Pages 243-255 in The virus diseases of tle rice plant Proceedings of a symposium atthe International Rice Research Institute April 1967 The Johns Hopkins Press altimore Maryshyland USA

125 Kisimoto R and V A Dyck 1976 Climate and rice insects Pages 367-391 in Climate and rice International Rice Research Institute PO Box 933 Manila Philippines

126 Kureha S S Kobayashi SAsari and T Ishihara 1977 Life history and control of the mole cricket Grrllotalpa africana Palisot de Beauvois [in Japanese English translation 7520 1RRI library] Proc Kanto-Tosan Plant Prot Soc 21192-197

127 Kushwaha K S 1981 The black beetle as a pest of rice in Haryana India Int Rice Res Newsl 6(4)14

128 Lawani S M 1982 A review of tile effects of various agronomic practices on cereal stem borers Trop Pest Manage 28266-276

129 Leuck D B 1966 Biology of the lesser cornstalk borer in South Georgia J Econ Entomol 59 797-801

130 lever R J A W 1955 Insect pests of the rice plant in Malaya Int Rice Comm Newsl 1612-21

131 Li Ching-Sing and Shin-Foon Chiu 1951 The study ofthe rice gall midge Iachydiplosis orvza Wood Mason [in Chinese with English summary] 1Taiwan Agric Res 21-13

132 Iitsinger J A 1984 Assesament of the need-based application for rice Proceedings TechnologyTransfer Workshop 15-16 March 1984 International Rice Research Institute 1PO Box 933 Manila Philippines 6 p

133 Litsinger J A 1984 Environmental diversity and insect pest abundance with reference to the Pacific Pages 285-307 in Proceedings sub-regional training course on methods of controlling diseases insects and other pests of plants in the South Pacific 4-20 Oct 1982 Vaini Kingdom of Tonga Nukualofa

134 Litsinger J A RF Apostol and M BObusan 1983 White grub Leucopholis irrorata (Coleoptera Scarabaeidae) pest status popula-tion dynamics and chemical conllol in arice-maize cropping pattern in the Philippines J Econ Entoinol 761133-1138

135 Litsinger J A C de la Cruz F Raymundo A T Barrion M D Lumaban J P Bandong M S Venugopal F Paragna and E Balete 1982 Insect pests and insecticide reponse of dry-seeded bunded rice Pages 179-193 in Report of workshop on cropping systems research in Asia International Rice Research Institute Los Bafos Philippine

136 Litsinger J A M 1) Luinaban J P Bandong A T Barrion W Katanyukul and N Panda 1978 Insect pest management in rainfcd lowland rice agroecosystms of tropical Asia International Rice Research Conference 17-21 April 1978 International Rice Research Institute Los Bafios Philippines

137 Litsinger J A and K Moody 1976 Integrated pest management in multiple cropping systems Pages 293-317 in R I Papendick P A Sanchez and G B Triplett eds Multiple cropping American Society of Agronomy Madison Wisconsin USA

138 Loevinsohn M E 1984 The ecology and control of rice pests in relation to the intensity and synchrony of cultivation Ph D thesis University of London 36) p

139 Loevinsohn M E J A Litsinger N Panda M A Austria R F Apostol C Andrion and J Bandong 1982 A basis for predicting rice insect pest abundance in cropping systems Paper presented at the 1982 International rice research conference on rice and cropping systems 19-23 April 1982 International Rice Research Institute Los Bafios Iaguna Philippines 25 p

140 Lopez A W 1930 Repot of the Entomology Department Pages 145-172 in Annual report of the Philippine Sugar Association 1929-30 Manila

141 Lopez AW 1931 Entomology Department Pages 227-266 in Annual report of the director or research 1930-31 Philippine Sugar Association Manila

142 Lopez A W 1932 Entomology Deparment Pages 252-279 in Annual report of the Philippine Sugar Association 1931-32 Manila

143 Lungibill P and G G Ainslee 1917 The lessercornstalk borer US Dep Agric Agric Res Serv Tech Bull 539 27 p

IRPS No 123 January 1987 39

144 McGuire J U Jr and BS Crandall 1967 Survey of insect pests and plant diseases ofselected food crops of Mexico Central America and Panama U SDepartment of Agriculture International Agriculshytural Development Service Agricultuial Researrh Service October 1967 11 p

145 McNeill Sand T R ESouthwood 1978 The role of nitrogen in the development of insectplant relations Pages 77-9 in lliochemical aspects of plant and animal coevolution J B Ilarbornu ed Academic Press London UK

146 Maebara H 1955 Spatial distribution of lastinstiarlanaeof purplish stem borer (Sesamia iqferens Walker) hibernating in an upland rice field Bull Kagoshima Univ Fac Agric 438-41

147 Mallick E H and N G Hajra 1977 Response of rice plant to defoliation treatments at different developmental stages Indian Agric 2151-56

148 Mansingh A 1971 Physiological classification of dormancies in insects Canl Entomol 103983-10)9

149 Martell C G 1974 Catalogue of plant feeding insects of Mexico [in Spanish] Fitofilo 27(69)1-176

150 Martinez 0 J 1982 Principal rice pests in Ilanos orientalis [in Spanish] P~ages 80-94 in Curso deact nalization de conocimientos en arroz Villaniencis Instituto (olonmbiano Agropecuario

151 Martins J F da Silva F Ferrcira A S Prabhu F 1 P Zimmermann 1980 Use of pesticides for the control of the main subterranean upland rice pests esqui Agropecu Bras 1553-u

152 Martins I F da Silva F Fe eira rod BdamSilveira Pinheiro I Q2 Damage simulation caused by defoliators of upland rice [in Portuguese with English summary] Pesqui Agropecu Bras 171113-1119

153 Martins 1F da Silva N(iyuen van Tan and 1Ida Silveira Pinheiro 1981 Stem borer resistance in upland rice associated with plant morphological characters [in Portuuce Iesqui Agropecu Bras 16187-192

154 Masaki S 1980 Sumnmer diapause Ann Rev Entomiol 251-25 155 Masuzawa T 11 Suwa and F Nakasoji 1983 l)ifkleences of

oviposition preferencc and survival rate of two skipper butterflies Parnara guttata and PIchlpidai iathiais (Iepidoptera HIesperiidac) on rice plant and cogon grass New Entomol 3247-56

156 Maurya D M and C P Vaish 1984 Upland rice in India Pages 69-92 in An overview oftpland rice research Proceedings ofthe 1982 Bouakamp Ivory Coast upland rice workshop International Rice Research Institute IO Box 933 Mnila Philippines

157 Menon M G R R N Katiyai S Kuar and S S Misra 1970 Termites attacking the paddy crop Sci Cult 36294-295

158 Miyahara Y and 1 Nishikubo 1967 Behavior of young nymphs of the red rice root aphid Iholahsipiun ri liabhdontinalisSasaki in relation to movement from the leaf to the roots of upland rice [in Japanese] Pioc Assoc Plant Prot Kyushu 1363-66

159 Mo ljoa Tjien 1952 White grub of upland rice (Holotrichiahelleri Brsk) [in Indonesian with English summary] Contrib Agric Res Stn Bogor No 13134 p

160 Mochida 0 and T Suryana 1979 The occurrence of the brown planthopper and ragged stunt on lowland and upland rice ill Central Java and logyakarta in tile late of January 1979 Entomnolgy Survey Report No 15 Central Research Institute of Agriculture (CRIA) Sukamandi Indonesia I February 1979 9 p

161Monet C 1979 Varietal resistance to upland rice pests [inI French] Pages 35-39 in Annual report 1977 Institut des Savanes Department des Cultures Vivrieres Bouak6 Ivory Coast

162 Monte 0 1942 A caterpillar ofrice-fields[in Portuguese] Biol6gico (Sao Paulo) 8161-163 (Rev Appl Entomol A31414)

163 Munroe I) D 1979 Pest management strategy for hill rice in four resettlement areas in Sarawak Malaysia Pages 89-102 i1 Guidelines for integrated control of rice insect pests Food and Agriculture Organization Plant Production and Protection Paper 14 Rome Italy 115 p

40 IRPS No 123 January 1987

164 Nakasuji F M Ishii and Jen-Zon Ito 1984 Rice skippers in Taiwan and their life histories ZZrC (Taipei) Tech Bull 82 10 p

165 Navas I) 1976 Fall armyworum in rice Pages 99-106 il Ecological animal control by habitat management Proc Tall Timbers Con-ference 28 Feb-I Mar 1974 Fall Timbers Res Stn Misc Publ 6 Tallahassee Horida USA

166 Nickel 1 1 1961 Biological control of rice stem borer A feasibility study Tcch Bull 2 International Rice Rcarch Institute PO Box 933 Manila Philippines IIl p

167 Nigan P M and R A Vernia 1985 Insect pests of uplaild rice in Uttar Pradesh Int Rice Res Newsl 10(6)22

168 Nilakhe S S 1985 -cological observations on spittle bugs with emphasis on their occirence in rice Pcsqtui Agropecu Bras 20407-414

169 Nixon G EJ 195 1 lie association of ats with aphids and coccids Commonwealth Iusti lcte of Entonology Iondon UK 36 p

170 Nurdin F an Ishak Manti 1985 Study oit seedling fly of upland rice Interlational Upland Rice Moitoring Tour Report 26 lebruary-2 MarLI Sukaranti West Sumatra

171 Oka 1 N 1979 Ciltural conttol of the brown planthopper Pages 357-369 in Brown planthopper threat to rice production in Asia International Rice Research Institutc 1p0 Box 933 Manila Philippines

172 Ohni M and A Vilnaii 1975 Rice insect pests of Italy biology of Sipha grcria (Kaltenbach) and its control [in Italian] Riso (Milan) 2459-71

173 Otanes F Q atnd 1 I Sison 194 1 Pests of rice Philipp J Agric 12211-259

174 Painter R I 1951 Insect icsistance in crop plants University Press of Kansits Lawrence Kansas USA 520 p

175 Pantua P C and 1A litsinger 1981 Arthropod abundance in continuous vs biannual rice cropping systeis Int Rice Res Ncwsl 6(2)20-21

176 Pathak M ) and V A l)yck 1975 Sttudics ott insect pests of upland rice Pages 186-191 in Major research in upland rice International Rice Research Institute PO Box 933 Ilanila Philippines

177 Patil B R B M IHasabe and 1) S Ajri 198 I Chemical control of white gritb lIditriihinhorraia Fabr intfesting groundntut paddy atnd sorghum Indian 1Entomol 43232-236

178 Pawar A 1) 1974 (ollection and identification of insect pests of rice rlerminal repirt lInternational Rice Research Institute PO Box 933 Manila Philippines 72 p (tnimco)

179 IerrajU A aid A S Reedy 1965 Observations ott diapausc itt tie steti borer ohrice Rice News Teller (lai-Mar)9-13

180 Pont A C 1972 A review of the Oriental species of theri ona Rondani (I)iptera Muscidac) of econtomic importance Pages 27-102 in Control of sorglium shoot fly M G lomtani and W R Young eds Oxford atd IH1 Ptthl Co New I)clhi India 324 p

181 Prasad S C 1PChand aid M 1PSingh 1983 Incidence of rice gall midge in the upland plateau region of Hihar Oryza 20235-236

182 Rai P S 198 0 Nisaga simpex damage to rice in the hill tracts of South India Int Rice kles News[ 5(I)12

183 Rathorc Y S 1969 Gonocplhahli dcprssunt Fabr as itpredator of (tilo zonwIlhis (Swinhoc) and other moths in field cage at Patnagar Uttar Pradesh Indian 1Entotnol 3127

184 Reed W 1983 Crop losses causcd by insect pists in the developing world Proceedings 10th Internatial Congress of Plant Protection lrighton UK 174-80

185 Reedy M V 1981 Ateactogasterinhens(Walker)a new root-grub pest of tipland paddy and inaize Curr Sci 501076

186 Reveno M A 1968 Catalogue of the insect pests of economic crops in Colombia [in Spaiish] Asociacion Latino Americana de Entomo-logia (AIAE) 129-33

187 Reyes G M 1957 Rice dwarf disease in the Philippines Food and Agriculture Organiiation Plant Prot Bull 617-19

188 Rice S E A A Grigarick and M 0 Way 1982 Effect of leaf and panicle feeding byarmyworm (Lepidoptera Noctuidae) larvae on rice grain yield Environ Entomol 7593-595

189 Risch S J 1) Andow and M A Altieri 1983 Agioccosystem diversity arnd pest control data tentative conclusions and nizw research dirctions Environ Entomol 12625-629

190 Rivera C T 1956 Three organic insecticides in thecontrol of insects affecting upland rice Philipp Agric 39465-472

191 Roffrey 1 1972 Locusta outbreaks in the Philippines Acrida 1177-188

192 Rojas R 1) 1941 The effect of pruning at varying ages upon the yield of Ramairio Phipp Agric 29851-860

193 Roonwal M i 1979 Ternite life and termite control in tropical South Asia Scientilic Pnbl odhpr India 139 p

194 Rosetto C1 F da Silva Martins N C Schmidt and L EAzzini 1978 Damage caused by pasturespittle bugs 4Dvoi)lavopictaandDJ schach) on rice lBragantia Nota 6 (Matcli)25-27

195 Rosztto C1- S S Neto I- Amantc I) Iink 1Garcia-Vieira D M de SoLza N V Bansa t and A M de Olivcira 1971 Rice pests of Brazil [in Portuguese] Pages 149-238 inrFA()-International Rice Commission 2nd meeting of the Rice Committee for the Americas 6-11 l)ecember 1971 Pelotas Rio Grande do Sul Brazil

196 Rothschild G H L 1969 Obseratiois on the armyworm Spodoptera matritia acronrctoiiles Gn (Lep Noctuidae) Bull Entomol Res 59143-160

197 Rothschild G 1- L 1974 Parasites of the armworm Spolopiera mauritiaacroncitoihes in Malaysia Fntornopliaga 19293-299

198 Sachan i N S K Gangwar and 1N Katujar 1980 Field crickets damaging cages in North Eastern IHill Region Indian J Entomol 42526-527

199 Sands 1) P A 1977 The biology and ecology of Lepocorisa (Hemiptera Alydidac) in Papua New Guinea Res Bull 18 Departshyment of Primary Industry Port Moresby Papua New Guinea 104 p

200 Sarawak 1968 titomology division Pages 97-106 in Report of the research branch of the Department of Altriculture 1966 Kuching (Rev AIppl Eintomol A56603)

201 Satpathy JM M S Das and K Naik 1977 Effect of multiple and mixed cropping on the incidence of scine important pestsI Entomol Res 178-85

202 Sauer IHF G 1939 Notes Ott Elasmpnlpualtsligniosellths Zeller (Lep Pyr) a serious pest ofcereals in Sao Paolo [in Portuguese] Arq Inst Biol 10199-206 (Rev Appl Entotnol A28378)

203 Satinders 1L A 1 SKing and C L Vargas S 1983 Crop pests of Central Anerica with t bibliography [in Spanish] Centro Agroshynotnico Tropical de Investigacion y Ensenanza Turrialba Costa Rica (CATIE) Tec Bol 9 90 p

204 Sauphanor B 1985 Some factors of upland rice tolerttce to stem borers in West Africa Inseu Sci Appl 6429434

205 Sen P atnd S Chakravorty 1976 Stentflics in Haritighata Nadia West Bengal India [it Rice Res Newsl 1(12) 15

206 Senapai B and 1M Satpathy 1983 The seedling fly Atherigona orrzae Malloch is a pest of rice in Orissa (India) J Entomol Res 777-79

207 Shhjahan M S Alatn M Ah nad and BN Islam 1983 A new scarabaeid beetle pest of dryland rice in Bangladesh Int Rice Res Ncwsl 8(I)16-17

208 Sharma G L S C Modgol and R C Gautam 1979 Effect of depth of submergence ott incidence of bacterial blight and white grub infestation in transplanted rice Int Rice Res Newsl 4(6)12-13

209 Sharma S 1985 Scarabaeid beetle outbreak itt upland rice Int Rice Res Newsl 1(1(l)24

210 Shiraki T 1917 The paddy borer Schoenobhits incertulas Walker [achaku Agric Exp Stn Formosa 256 p

211 Silva A de Brito and 11P Magalhaes 1981 Insect pests of rice [in Portuguese] Circular Tecnica No 22 EMBRAPA Centro de Pesquisa t ecuaria de Tropico Unido (CPAJU) Belem Brazil 14 p

212 Singh S A 1979 Emergence of rice leaf butterfly as a rice pest in Manipur India int Rice Res Newsl 4(4)16-17

213 SinghS A 1983 Changing trends of insect pestsof rice in Manipur Int Rice Res Newsl 8(f)12

214 Siwi S S and P H van Doesburg 1984 Lenovorisa Latreille in Indonesia (Heteroptera Coreidae Alydinae) Zool Meded (Leiden) 58117-129

215 Soejitno J 1977 Reaction of some upland rice varieties toseedling fly eltherikonaexigua Stein [in Indonesian] Siri Hana I Penyakit 630-35 Lembga I osat Fenclilian Pertanian Bogor Indonesia

216 Socnardi 1967 Insect pests of rice in Indonesia Pages675-683 in The major inse pests of thu rice plant Proceedings of a symposium at the Internatital Rice Rec Wi itt S pternber 1964 The Johns Hopkins Press Baltimore Maryland USA

217 Soto P E A T Petcz and I W Buddenhagen 1976 Survey of insect pests and diseases of rice in different ecological zones in Nigeria Rice Entomol Ncwsl 435-46

218 Soto 11E and Z Ziddiqi 1976 Screening for resistance to African rice insect pests Varietal improsetnent sCmi n r Bouak Ivory Coast 13-17 Sept 1976 West Africa Rice l)evelopment Association Monrovia Iiberia

219 Squire F A 1972 Entomological problems in Bolivia PANS (PestArtic News Sumr) 18249-268

220 Stapely 1 II 1978 The rice leaf roller in the Solomon Islands Int Rice Res Newsl 3(l)13

221 Suenaga H 1937 On aphids attacking the roots of rice Oyo-I)oubts Zasshi 9150-152 [in Japanese] (Rev Appl Entomol A25745)

222 Suenaga H 1952 On the seasonal abundance of three subterranean aphids on upland rice Bull Kyushu Agric Exp Sti 1249-262

223 Sutynso Yas and Dandi Soekaria 1964 Beberapa hana yang ttenperang tana man padi tanah kering [in Indonesian] Rapat kerja Padi Tanah Kering 6 sd 8 Juily 1964 Kaliurang 9 p

224 Tallamy I) W and R F Denno 1983 Alternative life history patterns in risky environments in example from lace bugs Pages 129-147 in Insect life history patterns R F l)cnio and If Dingle eds Spring-Verlag 225 p

225 Tamura 1 1939 Observations on some insect pests of field crops [in Japanese] 1 Plant Prot 26414-420 (Rev Appl EntomolA27-556)

226 Tamura and Suzuki 1964 Influences of the early-transplanting practice in rice culture upon development of the rice stem borer Chilo siql)ressalisWalker Bull Ilokuriku Natil Agric Exp Stn 761-94

227 Tan J P 1924 The rice root aphid Drowhi ihirsoia Philipp Agric 13277-288

228 Tanaka 1 1957 Taxonomy and distribution of some subterranean aphids injurious to upland rice in Japan with description of a new species [in Japanese English summary] Botyu-Kagaku 22168-176

229 Tanaka T 1961 The rice root aphids their ecology and control Spec [full College of Agriculture Utsunomiya University Japan F3 p

230 Tauber M IC A Tauber R Nechols and 1Obrycki 1983 Seasonal activity of parasitoids control by external internal and genetic factors Pages 87-108 itt I)iapause and life cycle strategies in insects V K Brown and I Ilodek eds )r W Junk The Hague Netherlands

231 Taylor W F 1972 Effects of artificial defoliation (simulating pest damage) on varieties of upland rice Exp Agric 879-83

232 feotia T P S and S Natud 1966 Biotoomics of the rice skipper Parnaraniathiis Fabricius (Lepidoptera Hesperiidac) Indian J Entomol 28181-186

233 Teran F 0 1971 Rice pests of Santa Cruz [in Spanish Ministerio de Agricultura Sita Cruz Bolivia Bull 45 8 p

234 Than Itun 1976 Population dynamics of the yellow rice borer Tryporza itcertulas (Walker) and its damage to the rice plant MS thesis University of the Philippines at Ios Bafios Laguna Philippines 92 p

235 Uichanco L B 1911 Pests Pages 55-74 it Rice Philipp Agric 11-200

IRPS No 123 January 1987 41

236 van der Laan P A 195 1 Life history of the rice seedling fly Contrib Gen Agric Res Stn Boor Indonesia 118 15 p

237 van der Laan 11A 1959 Correlation between rainfall in the dry season and the occurrence of the white rice borer (Scirpolhaga innotala WIk) in lava Entomol Fxp Appl 212-20

238 Van Driesche R G 1983 Meaning of percent parasitismin studies of insect parasitism Fnviron Entomol 121 I1- 122

239 Venugopal M S and A litsingcr 1984 Fffect ofcarbofuran on rice growth Prot Ecol 7313-317

240 Vietnam Directorate of Rural Affairs 1962 Annual work progress report on crop improvement program of rice sugarcane vegetable and field crops (lor the period from July 1961 to June 1962) (11) 299 p (Rev Appl Entomol A51526-527)

241 Wan M T K 1972 Observations on rice leaf- and planthoppers in Sarawak (Malaysian Borneo) Malay Agric 148308-335

242 West Africa Rice I)esclopnient Association 1978 Annual research report Vol 2 Statistics Monrovia Liberia

243 West Africa Rice Development Association 1979 Annual research report p47 Monrovia Liberia

244 Williams ) F and CS lofgren 1981 Fli I illy l-468 anew bait toxicant for control of the red imported lire at Fla Entomol 64472477

245 Williams I) 1 1970 The mealybugs OFlomoptera Coccoidea Iscudococcidac) of sugar-cane rice and sorghum Bull Entomol Res 0109-188

246 Williams D) 1 1982 Pulinaria icervi (Signoret) and its allies ott sugar cane and other grasses Btll Entoinol Res 72111-117

247 Wirjosuiardjo S 1976 Study ofchenmical cont rot of insect pests ott upland rice Imiu Pertanian (Jogjakarta) 1323-329

248 Wirjosuhardjo S and N Wirarta 1976 Chemical control of the white grub tIolorithiatlltriin upland rice in M uneng Irobolinggo Indonesia) [in Indonesian] Imut Pertanian (Jogjakarta) 1330-332

249 Yano K T Miyake and V F Eastop 1983 The biology and economic importance of rice aphids (Henuiptera Aphid idae) a review Bull Entomol Res 73539-566

250 Yelp D and P Rhodes 19(4 lhe success of drilled dryland rice iii the Fiji Islands International Rice Commission Working Party on Rice Production and Protection Manila Philippines 3-10 March1964 Food and Agriculture Organization Rome Italy 13 p

251 Young W R 1981 Fifty-five years of research on the sorghum shootfly Insect Sci Appi 23-9

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No 101The economics of hybrid rice production in (hina No 113 Boron toxicity in rice No 102 Rice ratooning No 114 Energy analysis rice production systems and rice research No 103 Growth and development of the (Jeep water rice plant No 115 Production risk and optimal fertilizer rates an application of the

No 104 Faridpur acomputer-assisted instruction nmodel forrainfed lowland random coefficient model No 116 Consumer demand for rice grain quality in Thailand Indonesia andrice

No 105 A readingand listening comprehcnsion test inEnglish for non native the Philippines speakers applying for tiraining at IRIRI No 117 Morphological changes in rice panicle development a review of

No 106 Rice grassy stunt virus 2 a new strain of rice grassy stunt in tire literature

Philippines No 118 IRRI-Korca collaborative project for the development of cold-No 107 Physical losses and quality deterioration in rice plstprodnCtion tolerant lines through anthei culture

systemis No 119 Problen soils as potenial areas for adverse soils-tolerant rice No 108 (olptblication ofIRRI materials a survey of translators and varieties in South and Ssuthcasi Asia

publishers No 120 Changes isinall-farm iLtireliing techii)hgy in Thailand and [he No 109 Classification of Philippine rainfall patterns Philippines No 110 Contributions of modern rice varieties to rutrition in Asia No 121 Landforms and modern rice varieties No Ill Changes in rice breeding in 10 Asian countries 1965-84 No 122 Yield stability and modern rice technology No 112 Design parameters affecting the performance of the IRRI-designed

axial-flow pump

Page 30: UPLANDRICE. INSECTl--1i M T -E!R.EOLOU t RTANCE ,AND …delphacid.s3.amazonaws.com › 6958.pdf · The significance of abiotic factors in upland rice insect ecology will be apparent

30 IRPS No 123 January 1987

Table 6 Dominant natural enemies of rice pests in upland rainfed lowland and irrigated lowland rice environments of the Philippines

Natural enemy

Host stage Species

Brown planthop-Parasite Egg

MymaridaeAnagrus optabilis(Perkins) Anagrusflaveolus Waterhouse

TrichogrammatidacOligosita naiasGirault

NymphAdult Dryinidac

Pseudogotatopuslaiifemttr Esaki et Hashimoto Elenchidae

Elenchus yasurnatsui Kifune and Hirashima

Environment

Upand Rainfed lowland Irrigated lowland

Ailaparata lugens

x x X x

x

x x

x X X Whitebacked planthoppers Sogatella furcifera and Sogatodes pusanusEgg

MymaridaeAnagrus optabilis(Perkins)

TrichogrammatidacOl~gositanaias Girault

NymphAdult Dryinidae

Pseudogonatopusnudus Perkins Elenchidae

Elenchus yasutatsui Kifune et Hirashima

Parasite Egg

MymaridaeGonatocerus spp

TrichogrammatidaeOligositaaesopi Girault

NymphAdult Pipunculidac

Tomosvaryella subviresccns Loew Tomosvaryella oryzaetora (Koizumi) Pipunclusmutillatus (de Meijere)

DryinidaeIlaplogonatopusspp

Egg Copidosomopsisnacoleiae (Eady)

Larva BraLonidae

Cardiochiles philipphnensisAshnead lchneumonidac

Trichiomnia cnaphalocrosisUchida Temelucha stangli (Ashmead)

Pupa Chalcididae

BrachymeriaexcarinataGahan Brachymerialasus (Walker) Brachymeria nr tachardiaegroup

Parasite Egg

Scelionidae Telenomus rowani (Gahan)

EulophidacTettasticusschoenobhi Ferriere

Larva Braconidae

Stenobraconnicevillei (Bingham) Iclhncurnonidac

Temelucha phiiippinensis(Ashinead)Temelucha stangli (Ashmead)

x x x

x x

X x X

x x Green leafhopper Nephotettix virescens

X x x

x x

X X X X

X

X Rice leaffolders CnaphalocrocisMarasmia

x X x

X

X X

x x x x x X x x x

Yellow stem borer Scirpophaga incertulas

X X x

x

x

x X XAinauromorphaaccepta metathoracica(Ashmead) x

continued on next page

IRPS No 123 January 1987 31

Table 6 cntinued

Natural enemy Environment

Host stage Species Upland Rainfed lowland Irrigated lowland

Larva-pupa Gold fringed borer Chilo attriciiuslchneumonidae

Xanthopimpla stemmator (Thunberg) x aPupa x

Pteromalidae Trichoinalopsis apanteloctena (Crawford) ax

Pink stem borerb Sesamnia inferensParasite

Larva Braconidae

Stenobracon nicevillei (Bingham) x b b Armyworms Mvthinna SpodopteraLarva

Tachinidac Zygobothriaarroptvora (Robineau-Desvoidy) x x

Braconidac Cotesia spp

X Skipper Pelopidas mathias and green horned caterpillar Melanitis leda ismeneF-gg

TrichogramrnatidacTrichogrannaspp x

Scelionidae Undetermined Scelionid

x Larva

Tachinidae Argyroph lax nitrotibialis (Baranov) X x

Pteromalidae Trichomnalopsis apanteloctena (Crawford)

X Predator

CoenagrionidaeAgriocnenis femina femina Brauer x xIschnura senegalensis tambur

Miridae CrtorhinsliidipennisReuter x x

Nabidac Stcnonabis tagalicus (Stal) x

VeliidaeAlicrovelia douglasi atrolineata (Bergroth) x

Coccinellidae Micraspis crocea (Mulsant) X x x

Vespidae Eumenes campaniformis (Fabricius) x

Formicidac Diaecamna spp xOdontoponera transversa (F Smith) x

Carabidae Chaenius spp OphioneaW- aJasciataSchmidt-Goebel

x

Ophionea ishii ishii Habu x x X

Gryllidav Anaxipta longipennis (Saussur-) x xMetioche ittaticollis (Sial) x x x

Tettigoniidae Conocephalus longipennis (de Haan) x x x

aNo rearing made on rainfed lowland bNo rearing made on rainfed lowland or Irrigated lowland

MrasmiaAmong the Laval parasites the braconid Cardio- lowlands and parasitizes oriental maize borer OsIriniachilesphilijpphensis is most adapted to an upland environ- furnacalis (Guenee) larvae The leaffolder pupal parasitesment It also parasitizes Hydelepta indicata a common not only occur equally in all environments but also haveleaffolder of legumes The ichneumonids Trichomma cna- wvide host ranges- parasitizing species of OsIrinia Marucaphalocrosis and Temehcha stangli occur more in the Hydelepta and Honiona - in maize and legumeslowlands T stangli is most adapted to irrigated lowlands The principal egg parasite of yeltow stem borer (YSB)and also parasitizes Chilo and Scirpophaga stem borer Telenomus rowani is widely adapted to all environmentslarvae T cnaphalocrosis is most prevalent in rainfed and has an unusual alternate host -Tabanus eggs (25)

32 IRPS No 123 January 1987

Tetrastictu schoenobii specializes only in YSB and occurs mainly in irrigated lowlands -- further proof that YSB is most adapted to the wetter environments

Larval parasites of YSB are highly environment-specific The braconid Stenohracon nicevillei occurs only in upland rice but also attacks the pink stem borer (PSB) Tieinchwa phiiiplPwnsis an ichneumonid occurs in upland rice and also parasitizes the larvae of (7hiu stqpresaisinatro-morpha accepta melaihoracica like 7 stangi is mnost adapted to the irrigated lowlands

The dark-headed stem borer(l)HSB) (ih pochit3sus was not collected in all rice environments and the gold fringed borer Chilo auricilius nd PSB were collected only in upland rice Xaantiholiplastenrnaloran ichneumonid larval parasite appears to be widely adapted whereas TrihomaslopisapanitI)cteiiaa pteromalid pupal parasite occurs mostly in wetlands and also parasitizes larvae of Chilo suppressais and Pelhuidas mathias

Armyworms Myit hinin and Spodujipera have two main larval parasites - a tachinid Zrgohothriaatropivora in rainfed environments and a braconid Cotesia spp in irrigated lowlands

The butterfly pests Pelopidas and 4elanitishave tricho-grammatid egg parasites found only in irrigated lowland environments

Two larval parasites show environmental preferences --Argrroph av nigrotihialis a tachinid in rainfed habitats and Trichonialopsisapantelkcnela in irrigated ricefields A limgrotibialisalsoattacks the sweet potato hornwormAgrius convolvtdi and T apanteloctena parasitizes DH S1B

Predators as a group show more distinct environmental preferences The aquatic damselflies Coenagrionidae are most abundant in the lowlands Cvriorhinus is also most adapted to the lowlands The nabid Stenonabis tagalicus occurs only in the uplands and isalso prevalent on legumes The coccinellid lady beetles wasps and ants are more prevalent in upland areas AlicrasIpis cit-ea larvae prey ona variety of aphids attacking legumes and maize Etunienes campanonia vespid mud wasp makes nests in trees and is therefore most adapted to the more botanically diverse uplands Soil-dwelling ants cannot tolerate flooding

Upland environments are habitats to arboreal carabid beetles Three species of Chlaeniusbeetles prey on leaffolder larvae and are perhaps more important than parasites (27)

The gryllids and tettigoniids are widely adapted egg predators not only on rice but also on maize and legumes They feed on eggs of most insects that are laid on leaves

The spidercommunity of upland rice environments isrich in species At one site (Batangas Philippines) 31 species have been recorded (23) Of the 176 spider species recorded in Philippine ricefields about one-half (82) occur in upland rice The spiderspecies of upland rice environments overlap more with those of rainfed lowland than with those of irrigated lowland rice (24) The same study showed that of the three environments irrigated rice has (ie greatest spider species diversity foiiowed by rainfed lowland and upland

However of the 10 most prevalent ricefield spiders 9 were abundant in upland rice showing wide adaptation All three spider guilds -- orb-web space-web and huting spiders shywere prevalent in upland ricefields There were however differences in environmental prelerences for some spider species

Of the orb-wcavers Tetragniathamanlihulatawas partishycila rlv abundant 7 japonica was low and Ltucatge dehcorata was absent in upland rice

As in other environments Aiypeiwnaifirtosana was the most dominant space-web spider however its relative numbers were lowerthan for lowland sites Among hunting spiders Ox|opes javanus and Lcosa leatcost fgla were more abundant and Lvcosa lpseudoanmnlata was less abundant in upland compared with lowland rice envishyroniments

Extensive rearing of rice insect pests in upland rainfed lowland and irrigated sites over an 8-yr period in the Philippines provided insight into the effectiveness of parasites as natural enemies (Table 7 8) Most of toe data are from large sample sizes taken over at least one crop which overcomes the pitfalls expressed by Van Driesche (238)

VSB egg parasitization was surprisingly similar across all environments Larval parasitization was highly dynamic among sites and even years within the same site but showed somewhat lowerlevels in uplandsthan in irrigated wetlands This would have been predicted for egg parasites as well since YSB is adapted to the lowlands and so must be its parasites This isevidence that Telenomus rowani is as good at dispersing as its main hosts Trowani clings onto the anal tufts of female YSB and parasitizes eggs as they are laid This phoresy may explain its wide adaptation The more specialized lowland YSB larval parasites therefore appear more effective than their upland counterparts with more alternate hosts

Low collection levels of PSB larvae especially in the wetlands make comparisons between environments difficult Low levels of larval parasifization were recorded in both rainfed environments Except in the Cagayan rainfed wetland site in 1980 low levels ef larval parasitization were evident at irrigated and rainfed sitesalike No differences by environment were evident Higher levels of parasitization occurred in Sarawak during a 1967 outbreak also from tachinids Rothschild (197) indicated that higher parasishytization occurred in upland rice areas

Skipper and green horned caterpillar larval parasitization rates were generally low and showed no environmental effect htighest activity occurred in Cagayan in 1980 (48 parasitization) Although larval parasitization levels of RLF were variable year to year activity appeared greater in the irrigated lowlands than in the uplands Again the highest levels were recorded in Cagayan in 1980 Cagayan has extensive grasslands and fallow around the rice areas and a relatively long rainfall period Rainfall in 1930 was good for crop growth and apparently for increase of larval parasites

IRPS No 123 January 1987 33

Table 7 Parasitization rates of rice planthoppers and green leafhopper in upland rainfed lowland and irrigated lowland environments Philipshypines 1976-840

Environment and site Year Egg parasitization () Nymphaladult parasitization ()

BPH W13P GLH BPII WBPH GLIH

UplandBatangas

Rainfed lowland

1976 1979 1980 1982

15 17 12

2(47)8(39)

4(51)

9(1800)14 (212)

12(304)

8 (101)6 (127)

9(116)

Iloilo

Cagayan

1976 1977 1978 1981 1980 1981 1982

0 (69)

11

44 48 40

-

-

46 55 44

0 (471)

11

52 56 49

0 (200)9 (214)

10 (1259)8(235)6 (900) 3(800)

-7 (462)

11 (270)3 (313)5 (900) 1 (800)

4 (600)10 (725)10 (362)12(870)5 (900) 3 (800)

Pangasinan

Irrigatedlowland

1984 1976 1978 1979 1982

13 (436) 13(379) 14 (457)

6(14)7 (197)

17(198) 11 (486) 18 (275)

3 (81)10 (274)3 (116)

16(871) 19(88)

28(211)9 (764)5 (647)7 (1204) 8(305)

Laguna 1977 11 7 21 1978 31 1979 14 4 4 28 1980 7 8 20 1981 1983

56 7(278)

26 16 (1650)

16 27(1921)

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R I Apostol unpublished 1lo1lo and Pangasinan 136 Cagayan BCanapi unpublished Liuna 175 I C Pantua unpublished 118 234 43 Carino and Shepard unpublished BPH = brown planthopper Nila parvara htgens W131Pi =whitebacked planthopper Sogatelia furcifera GLH = green leafhopper Nephotettrix virescens Figiires in parentheses are sample sizes

Tauher et al (230) reported that internal external and genetic factors influence the seasonal activity of parasitoids

Hopper egg parasites but not nymphaladult parasites were also parlicularly abundant at the Cagayan site possibly the egg parasites have more alternative hopper prey in the weedy areas than nymphaladult paisites

The seasonal dynamics of hopper egg parasites can be seen for upland rice in Batangas(Fig I 2) and for rainfed lowland lice in Pangasinan( 135)and Cagayan( 105) Levels of BPHWBPI-L and GLH egg parasitization were corn-parable in upland and irrigated lowland environments Except for some high rates of paraMsitization early in the season the parasitization rate tended to be steady in all envilonments However hopper nymphaladult parasi-tizution levels were more similar in the rainfed environments and lower than in irrigated rice

Overall parasites do not appear highly effective by themselves against upland or even lowland pests Among natural enemies parasites have attracted the most attention because they can be more readily assesscd

Predators and pathogens lend themselves better to management practices Predators the most important group of natural enemies in riccfields are difficult to quantify On the other hand pathogens appear less im-portant but can be readily cultured and disseminated

The natural enemy commulnity of upland rice is rich in species and differs significantly from that of lowland rice

These beneficial organisms must be conserved by applying insecticides judiciously particularly sprays Seed or soil placement of chemicals minimizes exposure to natural enemies The strategy to derive the greatest benefit from natural enemies isto allow the greatest number of beneficial species to thrive Then perhaps at least one species will be effective against each pest at any tine overcoming the variable abundance of each species within and between years

Programs to introduce exotic species or mass produce indigenous ones are ambitious and expensive because they require trained people on a sustained not ad hoc basis

Chemical control Insecticides offer rapid and efficient control of upland insect pests (190) but-should be used only after other control measures have been considered Insecticides are rarely used on upland rice because of cost (32) Upland rice yields are normally too low to justify the expenditure Also most upland rice is grown as a cash crop Spraying upland rice is more difficult than lowland rice because water is less accessible Government subsidy programs to provide pestishycides to tribal peoples for upland rice production have created problems of toxicity to humans because isolated people have had no experience to be able to handle pesticides safely

However ins-eticide use has beenjustified when a crop is

0Z

Table 8 Parasitization rates of lepidopterous pests in upland rainfed lowland and irrigated lowland environments Philippines 1976-84a

Parasitization (5)

Environment and site Year YSB GFB PSB Earcutting Swarming Skipper Greenborned RLF Brown larva larva caterpillar caterpillar caterpillar caterpillar larva semilooperEgg Larva larva larva larva larva larva

UplandBatangas 1977 - 0 (57) - 0 (63) - 0 (71) - 0 (106) 11(142) 2 (101)1978 43 (4697) 8(451) 5 (38) 3 (40) 6 (61) 2(58) 15 (57) 13 (129) 13 (1286) 4 (30)1979 59(4135) 12(320) 13 (11) 5(28) 9(45) 6(46) 7(23) 8(160) 9(1152) 8(26)

Rainfed lowland Iloilo 1976 - 0 (10) 0(10) 0 (7) 0 (19) 2 13) 15 (180) 5 (76)1978 0 (54) 0 (3) 0 (17) 9 (64) 4 (78) 3 (73) 0 (121) 0 (191)1979 64 (2350) 16 (84) 6 (17) 0(6) 18 (68) 25 (104)

1981 56 (1322) 18(304) 0(5) 12(51) 18(138)Pangasinan 1976 60 (7 7 )b 62(82) 0(111) 5 (39)

1978 58 (53)b 5(51) Cagayan 1979 59(713) 8 (101) 6 (34) 0(7)

1980 0(11) 4 (28) 15 (286)

62 ( 48(177) 61(140)1981 0(5) 12 (2) 13(45) 14 (92)

Irrigatedlowland Laguna 1974 62 8

1974-75 50 41 1975 43 72 1978 96 (623) 21(24) 33 (60)1979 83(2017) 18(110) 8(62) 11 (89) 22(400)1980 78 (1625) 20(75) 0 (16) 4 (72)1983 60

9 (80) 7 (270) 30

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R F Apostol unpublished Iloilo and Pangasnan 136 Cagayan B Canapi unpublished Laguna 175 P CPantua unpublished 118 234 143 Carino and Shepard unpublished YSB = yellow stem borer ScirpophagaincertulasGFB = gold fringed borer Chilo auricilius PSB = pink stem borerSesamia inferens earcutting caterpillar Alythimna separata swarming caterpillar Spcdoptera mauritia acronyctoides skipper Pelopidas mathias reen horned caterpillar Melanitls ledaismene RLF = rice leaffoldersCnaphalocrocismedinalis and Marasmiaspp brown semilooper Mocisfrugalis Figures in parentheses are sample sizes Egg masses

IRPS No 123 January 1987 35

Table 9 Effect of seeding rate and insecticide on plant density and Yield losses from insect pests in upland rice often areyield of Dagge upland rice Batangas Philippines 1976a comparable to those of lowland rainfed or irrigated rice

Seeding rate Insecticide Dosage Plant density Yield (0-Y30o) but usually from different pests The principal (kg aiha) seed treatment (kg aiha) (noIra-row) (tha) groups are soil pests feeding on sown seed and rootsat 14 DE followed by seedling pests These groups have largely gone

50 None - 35 a 20 a unnoticed leading researchers to conclude that stem borers50 Carbofuran F 10 38 ab 23 a and seed bugs are the most important pests (17) Yield loss 10 beCabofuranF100 None - 4551 c 23 a trials that pinpoint major pests need to includc seed

100 Dieldrin WP 10 52 cd 22 a treatments Granules applied in the furrow may not give150 None - 59 d 22 a adequate protection from many seed pests

aAv of 8 fields DE = days after crop emergence In a column means The floristic and pedological diversity of upland rice followed by a common letter are not significantly different at the environments is matched by the diversity of arthropods5level by D)MRT F = flowable Wi = wettable powder both pests and their natural enemies Irrigation has homoshy

genized lowland rice environments Flooded soils tend tothreatened by epidemics of locusts or armyworms New have the same properties and to eliminate soil pests The technology stressing efficiency has minimized insecticide spatial and temporal dominance of cultivated rice has cost benefited specialist pests

Seed treatment is normally inexpensive and can be Upland rice has no specialist arthropod species The economicallyjustified in -nanycases(66) Treating seed with monophagous pests on upland rice prefer the lowlands systemic insecticide (216) but not chlorinated hydrocarbons where rice evolved Rice in the uplands exists only because (247) is most effective in controlling seedling maggots of man and even though lowland rice predates upland riceFoliar sprays have to be repeated to obtain control and specialist species evolved Wheat maize and sorghum do often are washed off by frequent early season rains not have monophagous pests leading us to conclude that Granular insecticides are effective but required dosages are there is no evolutionary advantage for inssects to have too expensive to justify except under heavy infestation narrow host ranges in upland graminaceae(247) When directed against first-instar larvae banding Pests have adapted to the uplands not only by having granules in seed furrows (177) can be effective against white wider plant host ranges but also by having long life cyclesgrubs at low dosages (025 kg aii ha) (134) Seed treatments undergoing dormancy or being dispersive Each pest has a are not promising for white grub control (248) unique life history strategy suited to highly unpredictable

Seed treatment is also effective ainst ants and may be environments In agivenyear someinsect specics an better cheaper than increasing the seeding rate In Batangas exploit upland rice as a food source but no pest is a senious treating 50 kg seed ha was as effective as increasing the one every year on upland rice seeding rate to 100 kgha and using untreated seeds (Table The low yielding potential of upland rice means less 9) In Brazil seed treatments control termites and other soil quantity and lower quality of food for pests to exploit but insects including the lessercornstalk borer(66) Domiciano also less incentive for farmers to undertake control (58) however found seed treatments inconsistent in their The most effective control would be early and synchroshyeffect against high populations of lesser corn stalk borer nous planting of early-maturing upland rice varieties to

Baiting can be an inexpensive way of controlling ants prevent insect pest buildup Breeding for insect resistance seedling maggot flies mole crickets and field crickets Baits should initially focus on stem borers because they are verymade of locally available material and impregnated with difficult to control by other means Biocontrol efforts insecticide can be sparingly distributed taking adantage of should first stress conservation of natural enemies preshythe pests ability to disperse and encounter bait sites dators and pathogens rather than parasites Efficient

The new low-volume hand-held controlled-droplet chemical control includes seed treatment or baiting for seed sprayers may offer an advantage over high-volume knap- seedling and some root pests and using controlled dropletsack sprayes for controlling foliar pests Upland rice sprayers for spot treatments against foliar pests and seed canopies are more open than those in lowland rice and bugs when economic thresholds are reached better droplet penetration should result Controlled droplet applicators would be ideal for seedbugs and defoliators

CONCLUSION

Upland rice is attacked by a wider array of insect pests than is lowland rice mainly beca use of the addition of soil pests Generally population levels of lowland rice pests are lower in upland than lowland rice leading many people to conclude that insect pests are not important on upland rice

36 IRPS No 123 January 1987

REFERENCES CITED

1Abraham C C and K S R Mony 1978 Occurrence ofLeptocorisa acuta Fabr (Coreidae Hemiptera) as a pest of nutmeg trees J Bombay Nat ttistSoc 74553

2 Agyen-Sampong M 1975 HlierogrlphusdiaganensisKrauss (Orthop-tera Acrididae) a new pest of rice in Northern Ghana Ghana J Agric Sci 8249-253

3 Agyen-Samnpong M 1976 Varietal resistance and other inle-ratcd pest control components as factors of rice varietal improvement in West Africa Prcsented at the Second varietal improvement serinar 14-18 Sep 1976 Bouak Ivory Coast West Africa Rice Development Association Monrovia Liberia 18 p

4 Agyen-Sampong M 1977 Outbreak of Ihccrorrdnisorae in rice seedlings near Rokupr Sierra Leone lit Rice Res Newsl 2(6)16-17

5 Agyen-Sampong M 1980 Dirty panicles and rice yield reduction caused by hugs IntRice Res Newsl 5(l)11-12

6 Agyen-Sampong M 1982 The major pest problems of irrigated upland and mangrove swanip rice ecosystems ir the humid tropical Guinea savanna and Salel climate zones Pages 318-328 in Pro-ceedings training course cn concepts techniques and application of irtegrated pest management in rice inWest Africa 10-20 Jan 1982 Fendall Liberia West Africa Rice l)evelopioent Association Monrovia Liberia 505 p

7 Akibo-Betts V I and S A Raymundo 1978 Aphids as rice pests in Sierra Leone lit Rice Res Newsl 3(6)15-16

8 Akinsola F A 1975 Present status of different rice stemborers in parts of Nigeria iicL Entomo Newsl 328

9 Akinsola FA 1979 Ihe biology and ecology of rice stemborers in Nigeria Ph 1) thesis University ol lbadan Nigeria 217 p

10 Akinsola E A 1979 Guidelines of integrated management of insect pests of upland rice in West Africa Presented at the Seminar on integrated management of rice diseases and insect pests at Bobo-Dioulasso Upper Volta 17-22 Sep 1979 West Africa Rice Develop-ment Association Monrovia Liberia

II Akinsola IA 1984 Insect pests of upland rice in Africa Pages 301-305 in An overview of upland rice research Proceedings of the 1982 lIouak Iwsrv Coast upland rice workshop International Rice Research Institute 13O Box 933 Manila Philippines

12 Alam M S V TJohn and Kaung Zan 1985 Insect pests and diseases of rice in Africa Pages 67-82 in Rice improvement in Eastern Central and Southern Africa proceedings of the international rice workshop at Iusaka Zambia 9-19 April 1984

13 Alticri M A A van Schoonhovcn and J D Doll 1977 The ecological role of weeds in insect pest management systems a review illustiated with bean (Phaseohsvulgaris L)cropping systems PANS (Pest Artie News Summ) 23195-205

14 Anmante F 1967 The leaf-cutting ant Atia capiguaraa pest of pastures [in Portuguese] Biol6gico (Sao Paulo) 33113-120 (Rev Appl Entomol A56207)

15Andrade A C and D Puzzi 1951 Preliminary results of experi-merits on the control of Sculptcoris iastantus on sugarcane [in Portuguese] Bioulgico (Sao Paulo) 1744-49 (Rev Appl Entomol A41421422)

16 Appert J 1970 Maliarplia separaelha (rice stem borer) new observations of entomological problems of rice in Madagascar [in French] Agron Trop (Paris) 25329-334

17 Ariadeau Mand Z Harahap 1986 Relevant upland rice breeding objectives Pages 189-197 in Progress in upland rice research Proceedings of the 1985 Jakarta conference International Rice Research Institute 1O Box 933 Manila Philippines

18Asar S and Kisimoto 1975 Seasonal occurrence of the smaller brown planthopper in upland rice [in Japanese] Proc Kanto-Tosan Plant Prot Soc 2228

19 Bakker W 1974 Characterization and ecological aspects of rice yellow mottle virus in Kenya Meded Landbouwhogesh Wage-ningen 152 p

20 Balut F F 1970 White grubs attacking rice Oryza ativa L [in Portuguese] Biol6gico (So Paulo) 36321-322

21 Banerjee S N and L M Pramanik 1967 The lepidopterous stalk borers of rice and their life cycles in the tropics pages 103-124 in The major insect pests ofthe rice plant Proceedings ofa symposium at the International Rice Research Institute September 1964 The Johns Hopkins Press Baltimore Maryland USA

22 Barrion A T and J A Litsinger 1980 Ants a natural enemy of cliaiarac in dryland rice IntRice Res Newsl

5(4)22-23 23 Barrion A T and J A litsinger 1981 The spider fauna of

Philippine rice agroecosystem 1Dryland Philipp Entomnol 5139-166

24 Barrion A T and 1 A Iitsinger 1984 The spider fauna of Philippine rice agroccosystcms II Welland Philipp Eiitomol 611-37

25 Barrion A T1 and J A Litsinger 1984 lihanus (Diptera Tabanidae) eggs an alternative host of rice stei borer (SI3) egg parasite Telenoonis digns (Iliymenoptcra Scelionidae) lit Rice Res Newvsl 9(6)19

26 Barrion A T and J A Litsinger 1985 Identilication of rice leaffolders by wing markings Int Rice Res Newsl 10( 1)24

27 Barrion A T and J A Litsinger 1985 Chctiosspp (Coleoptera Carabidae) a leaffolder ILF)predator lit Rice Res Newsl 10(1)21

28 Bertels A 1970 Insect pests of rice and theircontrol [in Portuguese] Pelotas Brazil Instituto de Expcrimentacao Agropccuararias do Sul Circular 43 24 p

29 Bonzi S M 1982 Ctilo (flisilineus J de Joannis (Lepidoptera Pyralidae) stem borer of irrigated cereals in Upper Volta LioFrench] Agron Trop Riu Rizic Cult Vivrieres Trop 37207-209

30 Bourke T V T L Fenner J N LStihick G Baker EHassan D F OSullivan and C S li 1977 Insect pest survey for the year ending 30th June 1969 Papta New Guinea )epartment of Agrishyculture Stock and Fisheries Port Moresby 57 p

31 Bowden 1 1973 Migration of pests in the tropics Meded Rijksfac Landbouwwet Gent 38785-796

32 Breniere J 1973 The main rice parasites in West Africa and their control Pages 27-42 in Proceedings seminar on plant protection for the rice crop West Africa Rice Development Association Monrovia Liberia

33 Breniere J 1976 The principal insect pests of rice in West Africa and their control West Africa Rice Development Association Monrovia Liberia 52 p

34 Brown ES 1962 The African arinyworn Spoiolptera eXI)Ina (Walker) (Lepidoptera Noctuidae) a review of the literature Cornshymonwealth Institute of Entomology London UK57 p

35 Brown F B 1968 Upland rice in Latin America FAO-International Rice Commission 12th Session of the Working Party on Rice Production and Protection 2-14 Sept Peradeniya Sri lanka Food and Agriculture Organization Rome Italy 6 p

36 Buyckx F J 1 1962 Disease and insect pests of crops inCongo Rwanda and Burundi [iil French] Publications dc LInstilut National pour lestude Agrononiquedu Congo M Weissenbruck S A Brussels Belgium 708 p

37 Cariiio F 0 1981 Role of natural enentics inpopulation suppression and pest management of green rice lealhoppers Ph D thesis University ofthe Philippines at Los Bafitos PhilippinesLaguna 136 -

38 Carpente A J 1983 The development of rice cultivation and extension of vegetable productin in Zanzibar Final Report Vol II1 October 1983 Food and Agricultme Organization Ronte Italy

39 Carroll C P and S1 Risch 1983 Tropical annual cropping systems ant ecology Environ Manage 751-57

4(0 Cates R G 198 1 Host plant predictability and the feeding patterns of monophagous oligophagous and polyphagous insect herbivores Oecologia (Berl) 48319-326

IRPS No 123 January 1987 37

41 Cendcafia S M and F B Caora 1967 Insect pests of rice in the 62 Ene 1 C 1981 I)iiorldhol)ierusihilis Slater (licteroplera Philippines Pages 591-616 in The najor insect pests ofthe rice plant lygacidue Illissinac) a new rice pest front Nigeria lit Rice Res Proceedings of a symposium at the International Rice Research Newsl N3)15 Institute September 1964 The Johns Hopkins Press Baltimore 63 Fugenio T S 1965 rhe effect oi e-vCgetation on rice production Maryland USA 729 p and the possibility of overcoming it by leaf pruning MS thesis

42 Centro Ihternacional de Agricklittira Tropical 1984 Upland rice in University ol the Philippines at Los Baijos ILaguna Ihilippins 69 p Iatin Anerica Pages 93-119 hi An os ervies of uplaind rice research 04 Farrow R A 1975 The Alrican nmigratory locust ii is main Proceedings of the 1982 Boinakamp Ivory Coast upland rice workshop outbreak area of the middle Niger qlinlitatise studies of solitary International Rice Research Institute P0 Blox 933 Manila populations in relation to environmental factors Locusla No II Philippines 198 p

43 Chandra G 1979 Taxonomy ad hn riononlics ofthe imect parasites of 65 Fennal R G 1943 Food crop investigation Windward and rice Icalloppers and planthoppers in the Pihilippines and their Ieeward Islands First report Report onl work dunring the periodiiiportairce iii natural biol)gical cointrol Philipp Fntoltol 4119-139 April 1942-April 1943 Imperial College of Tropical Agriculture

44 Chang T T 1976 The originii colut ion enlIivation dissemination Trinidad 16 p (Rev of Appl Fintornol A3222-25) and diersificition of Asian and African rices -uphytica 25425-44 1 66 Ferreira F 1983 Integrated pest ritaageiret of upland rice [ill

45 Chatierjee P B 1979 Rice leaffolder attacks in India lit Rice Res Portuguese English summary] Pages 323-341 in Upland riceculttre Newsl 4(3)21 factors affecting production E Ferreira T Yamada and F-

46 Cheng Cbien-pan 19(4 Upland rice in Taiwan International Rice Malmolta eds Institutoda Potassae Foslato Institnto Internacional Cnmmission Wiorking Party oit Rice Production ard Protection I()th da Potassa Piracicaha Sao Paulo lraid Meeting 3-10 March 1964 Manila Philippines Food and Agricul- 67 Ferreira E arid F dl Silva Martins 1979 Occurrence of ture Organiaion Rome Italy 5 p Sogatohs orvziola (Muir) inupland rice in Goias Ilrail lit Rice

47 Chcrrett 1 M and 1) 1Peregrine 1976 A review of the status of Rcs Ncwssl 4(4)18 leaf-cutting ants and their control Ann Appl Biol 84124-128 68 Ferreira E and 1F da Silva Martiris 1984 Insect pestof rice ill

48 Chipeta F M 1974 Pests of rice in Malawi Pages 90-95 in Brazil and their control Jill Portugucsej FMIlRAtPA-CNPAF Proceedings Sith eastern Africa cereals research conference 10-15 Goiania Brmiil 67 p March 1974 DRB Mialda ed Ministry of Agriculture lilongwe 69 Ferreira FI F ca Silha Miartins S S Nero and 1 It P de Malawi Carvallio 1982 M tltiplc regression analysis todeterimine the effect of

49 Chu Yau-i 1979 The d atiaging period of the rice seedling fly insect populatiois ott upland rice yield [inIPortuguese Pesqui (Atlterigotiavxigu Stcin) (NItiscide Diptera) oil cori) at Fast Java Agropecu Bras 17671-675 Indonesia Plant Prot Bill 21403413 70 Ferreira F 1F da Silva Martins and F 1PZimmermann 1979

50 Conklin H C 1957 IHaninoo agriculture a report on aln integral Rice varietal resistance to tle lesser corn stalk borer [in Port uguese] systent of shi fti rig cult ivationi in the Pltilippincs Food and Agriculture Pesqui Agropccu Biras 14317-321 Organization Rome Italy 209 p 71 Fleming W F 1968 Biological control of lie hapanese beetle U S

51 Cramer H II 1967 Plant proection and world crop production Dep Agric RLs Sers [ech Bull 1383 78 p Pilaniensclhutz-Nachr 201-524 72 Food and Agriculture Organiation of the United Nations 1982

52 l)ani R C and Majunidar 1978 Prelininary observations on tie Country report surmimary Pages 27-37 in Report on the first incidence of root aphids on differeit rice cultnrcs Sci Cult 4488-89 FAOIUN)IPhail and regional training coiuirse in iniproved ctltural

53 Davidson R I 1979 lffects of root feeding on foliage yield Pages practices for upland rice 5-23 Oct 1981 Northern Region Agrculturd117-120 in T K Ci osb and R B Pottinger eds Proceedings of the Developririt Center Chiang Mai Food and Agricuilture Organiiashy2nd Australian conlerence on grassland invertebrate ecology Govt tion Rome Italy 37 n Printer Wellington New Zealand 73 Franco do Ariaral S 195 1 Residual action of insecticides in turfy

54 l)ean G 1W 1976 Rice insect pests inlaos Int Rice Res Newsl soil--the results of a field experiment [in Portuguese] Biol6gico (Sao 1(2)15 Paulo) 17209-214 (Rev Appl Entomol A41423)

55 I)ean G J W 1978 Insect pests of rice in Laos PANS 24(3)280-289 74 Freeman ) 1970 Report onithe Iban london School of -coroiics 56 )ingle H 1984 Behavior genes and lifc histories conplex adapta- Monographs on Social Anthropology No 41 Athlane Press

tions to uncertain environments Pages 169-194 in A new ecology London UK 317 p novel approaches to interactive systems P W Price C N Slobod- 75 Fuchs T W1 A Harding and 1W Smithlr 1979 Induction and chikoff and W S Gaud cdsl ohn Wiley New York USA termination oldiapause in the sugarcane borer Ann Flitonlol Soc

57 Dobelmann-JP and M Falais 1968 Upland rice cultivation ill tie Am 72271-274 North-western part of Madagascar in 1966-1967 [in French English 76 (hbriel B1) 1975 A review tf the major insect pests of some tpland summary] Trop Agron 2313-21 crops in tlc Philippines Pages 86-111 in Ad Htoc panel of experts on

58 Doniiciano N L 1979 Chemical control of tie lesser corn stalk pest disease arnd weed problems in some rainfed crops Bangkok (lolsonno~paloIgnowslhts) (Zeller 1848) (I-epid(ptera Ihycitid ae) Thailand Food aid Agriculture Organization Roire Italy

on rice in Ponta Gressa Parara 1977-1978 [in Portuguese] Pages 77 Garg D K and N K Shal 1983 White grub outbreak on rainfed 291-303 in Anais 1 Reuniao de tecnicos cmiirizicultura do estado de dryland rice in Uttar Pradeslh Int Rice Res Ncwsl 81(4)16-17 Sao Paulo-Campinas Cati Brazil 78 Garg I) K and IP Tandon 1982 Major insect pests of rice on hilly

59 Dunsmore 1 R 1970 Investigations on tlie varieties pests and tracts of Uttar Pradesh India lilt Rice Res Newsl 7(11)11-12diseases of upland rice (hill padi) in Sarawak Malaysia Int Rice 79 Garrity 1) P 1983 Upland rice research an international bibliograpy Comm Newsl 1929-35 1975-1982 Iternational Rice Research hnstitnte PO Box 933

60 Elias 1R1965 The new decline of rice seedlings is not caused by Manila Philippines 121 p parasites [in Portuguese] Biol6gico (Sao Paulo) 313-6 80 Ghauri M S K 1982 A new subspecies of Iinorh)rone bull

61 Empresa Brasiliera de Pesquisa Agropecuaria 1984 Upland rice itt cornlus Slater (Hemiptera Iygaeidae) front Papua New Guinea on Brazil Pages 121-134 it Av overview of upland rice research rice and carpet grass Ihll lEntoniol Res 72133-137 Proceedings ofthe 1982 Blouak Ivory Coast Upland rice workshop 81 Gonzales F 0 PArregoces R I ihernandef and OT Parada International Rice Research Institute PO Box 933 Manila 1983 Insect and imite pests of rice arid their control in Latin America Philippines [in Spanish] la Federacion Nacional de Arrocerts (Fedearroz)

Bogota Colombia 60 n

38 IRPS No 123 January 1987

82 Grist D H and R J A W Lever 1969 Pests of rice Longmans London UK520 p

83 Hale P R 1979 Two subterranean pests of upland rice in Papua New Guirnca IntRice Res Newsl 4(2)16-17

84 Hale P R and J T Hale 1975 Insect pests of rice in the New Guinea Islands Rice Entomol Newsl 34

85 Harris W V 1969 Termites as pests of crops and trees Common-wealth Institute of Entomology London UK 41 p

86 Hattori 1 1968 A method to distinguish between grass leaf roller (Ctaplha rocis medinalisGuenee) and the rice leaf roller (Susurtia exigua Butler) [in Japanese] Plant Prot (Japan) 23167-170

87 Henry I E and 1A Onsager 1984 Experimental control of the mormon cricket Iahrosshnph by NJsena locustae(Aficrospora Microsporidae) a protozoan parasite of grasshoppers (Ort Acri-didae) Entomophaga 27197-201

88 Hidaka 1 P Vungsilabulr and SKadkao 1974 Studies on ecology and control of the rice gall midge in Thailand Tech Bull 6 TropicalAgricultural Research Center Tokyo Japan 113 p

89 Hincklcy A 1) 1963 Ecology and control of rice planthoppers in Fiji Bull Entomrol Res 53467 481

90 Hirao J 1982 Ecology and chtInical control of the rice leaffolder Jpn Pestic Inf 4114-17

91 Ho DT 1984 Sten borer incifience in rice ecosystems in KenyaEast Africa Pages 307-320 in A overview of upland rice research Proceedings of the 1982 Bouac Ivory Coast upland rice workshop International Rice Research Institute PO Box 933 Manila Philippines

92 Ho )T J 1Njokah and J GKibuka 1983 Studies on rice sten borers in Kenya with emphasis on ifaliarpiaipiraellRag Insect Sci Appl 465-73

93 Ichihara I T Ito A NIatsuo and M Sawada 1971 Chemical control of the rice striped stem borer in upland rice grown in plastic mulch [iii Japanese] Proc Kanto-Tosan Plant Prot Soc 1882

94 IclhiharaIand A Matsuo 1969 Damage caused by rice striped stett borer to upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1686-87

95 Ichhara I atid A MIatsun11970 Conside ration oft lie extraordinary prevalence of rice striped stem borer in upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1778-79

96 Institut de Recherches Agrolnliqes Iropicale et des Cultures Vivrieres 1978 Insect resistance Pages 12-13 in Upland rice 1978 atnual report Bouak Ivory Coast 27 p

97 International Institute for Tropical Agriculture 1983 Aphids itt rice roots I ITA Res Briefs 4(1-3)4-5

98 International Institite fi Tropical Agriculture 1983 Hopperburn in rice caused by brown plathopper IITA Res Briefs 4(l-3)6-7

99 International Rice Research Institute 1975 Annual report for 1974 PO Box 933 Manila Philippines p 216

100 International Rice Restarch Institute 1976 Annual repiort for 1975 PO Box 933 Manila Philippines p 228

101 International Rice Research Institute 1979 Cropping systems program Component technology development and evaluation Page 404 i Annual report for 1978 11O Box 933 Manila Philippines

102 International Rice Research Institute Department of Entomology 1981) Insecticid evaluation 1979 Tables 73 74 PO Box 933Manila Philipes

103 International Rice Research Institute Department of Entonmology 1981 Insecticide evalomtion 1980 Table 81 POBox 933 Manila Philippines

104 International Rice Research Institute 1981 Report of a meeting of Asian rice-based cropping systems entomologists Eleventh cropping systems working group meeting 18-22 May 1981 PO Box 933 Manila Philippines p 132 133 144

105 International Rice Research Institute 1984 Cropping systems program Pages 368-373 in Annual report for 1983 PO Box 933 Manila Philippines

106 International Rice Research Institutc 1984 Upland rice in Asia Pages 45-68 in An overview of upland rice research Proceedings of the 1982 Bouakamp Ivory Coast upland rice workiiop loternational Rice Research Institute 11O Box 933 Manila Philippines

107 Irshad M 1977 Grasshoppers in rice in Pakistan lit Rice Res Newsl 2(3)8-9

108 Ito R and TOno 1964 Upland lice its culture and varieties in Japan lIternational Rice Cormmission loth Meeting of the Working Party on Rice Production and Protection 3-10 March 1964 Manila Philippines Food and Agriculture Organization Rome Italy 12 p

109 Jeffrey M I and V A Dyck 1983 Effect of moonlight on lightshytrap catches of brown planthopper lit Rice Res Newsl 8(6) 13

110 Jennings P R and T Alicia Pineda 1983 Management ofthe rice plnthoppers Sogatodsorivi(coIl in Latin America Presented at the meeting on stable plant resistance FAO and lIndocsian I)inectorate of Food Crops Protection Bali lndonesia 29 Novcniber-2 l)ccemshyber 9 p

IllJimenez Gomez J A 1979 Rice insect pests i lla ios ()riCntales lil Spanish] Pages 99-103 in Rice ciltivation Instituto (olotibiatto Agropecuario Compndio No 29

112 Jongeleen F JJ Soesanto and A Mockidjo 1979 Studies oti pathogens of two species of Scarabacidac froinIndonesia Z Angew Enlomnol 87412-417

113 Jouvenaz 1) PC S lofgren and W A Baiks 1981 Biological control of imported fire antsa review of cirrent knowledge Bull Entomol Soc Am 27203-208

114 Kainoh Y K Shiniiu S MarUani Y Taiaki 1980 lost finding behavior of tie rice bu lplotori+JchinvittjI)allas (liciniptera Corcidac) with special reference to diet pattern ol aggregation and feeding on the rice plant Appl Entomol Zool 15225-233

115 Kakkar K 0 P Sharma and P I Sharma 1977 Note on Ihteronyrchus liodresRedt (Coleopicra I)ynastidae) as asporadic pest of paddy and maie inHimachal Pradesh Corr Agric 156-57

116 Kalode M131 79 Major insect pest problems and their control in upland rice (Unpublishcd mineogriph CWI oJNaaiat lni Chandaratat (198 1)original not seen)

117 Kalshoven L i 98 I The pests of crops in I ldoniesia (translated by P A vatn der Laan) 1 T Ichtiar B3aru-vati Iloeve lakarta Indonesia 701 p

118 Kamal N Q 1981 Suppression of whitebacked planto ner Sogatella fircaifra (Horvath) aLId rice lealfolder (nhal -cis licdb lis (Guenee) populations by natural cnclmie hlI) tiesis Grcgorio Araneta University Foundation Victoncta Park Metto Manila Philippines 179 Ip

119 Karim A N M Reaul 1975 A species ofliealy bug Trioni i70s ceres new oii rice in the Philippines Rice Entoniol Newsl 247

120 Katanyukul Wand C Chandaratat 1981 Insect pestsof uplaid rice and their control Report firstFAOUNlFi Tliailand training course on improved cultural practices for tplad rice Chiang Mai Thailand 5-23 Oct 1981 Food and Agriculture Organiation Rome Italy 22 p

121 Kern F 1955 lhletwioa(I)uninemts) bhhit Biturm(Coleoptera) a new pest of sugarcane inVenezuela [inSpanish with Etnglish sunmary] Agron Trop (Maracay) 527-29 (Rev Appl Eitoniol A4572)

122 Khan K M and S Ghai 1974 White grubs ald their control in India Pesticides (1Bombay) 819-25

123 Kiritani K 1975 Integrated pest management of rice pests Pages 66-70 itt Proceedings platning workshop ott cooperative hIeld research in pest managementP SMotooka ed East-West Center Honololu Hawaii USA

124 Kisimoto R 1967 Ecology of insect vectors forecasting and chemical control Pages 243-255 in The virus diseases of tle rice plant Proceedings of a symposium atthe International Rice Research Institute April 1967 The Johns Hopkins Press altimore Maryshyland USA

125 Kisimoto R and V A Dyck 1976 Climate and rice insects Pages 367-391 in Climate and rice International Rice Research Institute PO Box 933 Manila Philippines

126 Kureha S S Kobayashi SAsari and T Ishihara 1977 Life history and control of the mole cricket Grrllotalpa africana Palisot de Beauvois [in Japanese English translation 7520 1RRI library] Proc Kanto-Tosan Plant Prot Soc 21192-197

127 Kushwaha K S 1981 The black beetle as a pest of rice in Haryana India Int Rice Res Newsl 6(4)14

128 Lawani S M 1982 A review of tile effects of various agronomic practices on cereal stem borers Trop Pest Manage 28266-276

129 Leuck D B 1966 Biology of the lesser cornstalk borer in South Georgia J Econ Entomol 59 797-801

130 lever R J A W 1955 Insect pests of the rice plant in Malaya Int Rice Comm Newsl 1612-21

131 Li Ching-Sing and Shin-Foon Chiu 1951 The study ofthe rice gall midge Iachydiplosis orvza Wood Mason [in Chinese with English summary] 1Taiwan Agric Res 21-13

132 Iitsinger J A 1984 Assesament of the need-based application for rice Proceedings TechnologyTransfer Workshop 15-16 March 1984 International Rice Research Institute 1PO Box 933 Manila Philippines 6 p

133 Litsinger J A 1984 Environmental diversity and insect pest abundance with reference to the Pacific Pages 285-307 in Proceedings sub-regional training course on methods of controlling diseases insects and other pests of plants in the South Pacific 4-20 Oct 1982 Vaini Kingdom of Tonga Nukualofa

134 Litsinger J A RF Apostol and M BObusan 1983 White grub Leucopholis irrorata (Coleoptera Scarabaeidae) pest status popula-tion dynamics and chemical conllol in arice-maize cropping pattern in the Philippines J Econ Entoinol 761133-1138

135 Litsinger J A C de la Cruz F Raymundo A T Barrion M D Lumaban J P Bandong M S Venugopal F Paragna and E Balete 1982 Insect pests and insecticide reponse of dry-seeded bunded rice Pages 179-193 in Report of workshop on cropping systems research in Asia International Rice Research Institute Los Bafos Philippine

136 Litsinger J A M 1) Luinaban J P Bandong A T Barrion W Katanyukul and N Panda 1978 Insect pest management in rainfcd lowland rice agroecosystms of tropical Asia International Rice Research Conference 17-21 April 1978 International Rice Research Institute Los Bafios Philippines

137 Litsinger J A and K Moody 1976 Integrated pest management in multiple cropping systems Pages 293-317 in R I Papendick P A Sanchez and G B Triplett eds Multiple cropping American Society of Agronomy Madison Wisconsin USA

138 Loevinsohn M E 1984 The ecology and control of rice pests in relation to the intensity and synchrony of cultivation Ph D thesis University of London 36) p

139 Loevinsohn M E J A Litsinger N Panda M A Austria R F Apostol C Andrion and J Bandong 1982 A basis for predicting rice insect pest abundance in cropping systems Paper presented at the 1982 International rice research conference on rice and cropping systems 19-23 April 1982 International Rice Research Institute Los Bafios Iaguna Philippines 25 p

140 Lopez A W 1930 Repot of the Entomology Department Pages 145-172 in Annual report of the Philippine Sugar Association 1929-30 Manila

141 Lopez AW 1931 Entomology Department Pages 227-266 in Annual report of the director or research 1930-31 Philippine Sugar Association Manila

142 Lopez A W 1932 Entomology Deparment Pages 252-279 in Annual report of the Philippine Sugar Association 1931-32 Manila

143 Lungibill P and G G Ainslee 1917 The lessercornstalk borer US Dep Agric Agric Res Serv Tech Bull 539 27 p

IRPS No 123 January 1987 39

144 McGuire J U Jr and BS Crandall 1967 Survey of insect pests and plant diseases ofselected food crops of Mexico Central America and Panama U SDepartment of Agriculture International Agriculshytural Development Service Agricultuial Researrh Service October 1967 11 p

145 McNeill Sand T R ESouthwood 1978 The role of nitrogen in the development of insectplant relations Pages 77-9 in lliochemical aspects of plant and animal coevolution J B Ilarbornu ed Academic Press London UK

146 Maebara H 1955 Spatial distribution of lastinstiarlanaeof purplish stem borer (Sesamia iqferens Walker) hibernating in an upland rice field Bull Kagoshima Univ Fac Agric 438-41

147 Mallick E H and N G Hajra 1977 Response of rice plant to defoliation treatments at different developmental stages Indian Agric 2151-56

148 Mansingh A 1971 Physiological classification of dormancies in insects Canl Entomol 103983-10)9

149 Martell C G 1974 Catalogue of plant feeding insects of Mexico [in Spanish] Fitofilo 27(69)1-176

150 Martinez 0 J 1982 Principal rice pests in Ilanos orientalis [in Spanish] P~ages 80-94 in Curso deact nalization de conocimientos en arroz Villaniencis Instituto (olonmbiano Agropecuario

151 Martins J F da Silva F Ferrcira A S Prabhu F 1 P Zimmermann 1980 Use of pesticides for the control of the main subterranean upland rice pests esqui Agropecu Bras 1553-u

152 Martins I F da Silva F Fe eira rod BdamSilveira Pinheiro I Q2 Damage simulation caused by defoliators of upland rice [in Portuguese with English summary] Pesqui Agropecu Bras 171113-1119

153 Martins 1F da Silva N(iyuen van Tan and 1Ida Silveira Pinheiro 1981 Stem borer resistance in upland rice associated with plant morphological characters [in Portuuce Iesqui Agropecu Bras 16187-192

154 Masaki S 1980 Sumnmer diapause Ann Rev Entomiol 251-25 155 Masuzawa T 11 Suwa and F Nakasoji 1983 l)ifkleences of

oviposition preferencc and survival rate of two skipper butterflies Parnara guttata and PIchlpidai iathiais (Iepidoptera HIesperiidac) on rice plant and cogon grass New Entomol 3247-56

156 Maurya D M and C P Vaish 1984 Upland rice in India Pages 69-92 in An overview oftpland rice research Proceedings ofthe 1982 Bouakamp Ivory Coast upland rice workshop International Rice Research Institute IO Box 933 Mnila Philippines

157 Menon M G R R N Katiyai S Kuar and S S Misra 1970 Termites attacking the paddy crop Sci Cult 36294-295

158 Miyahara Y and 1 Nishikubo 1967 Behavior of young nymphs of the red rice root aphid Iholahsipiun ri liabhdontinalisSasaki in relation to movement from the leaf to the roots of upland rice [in Japanese] Pioc Assoc Plant Prot Kyushu 1363-66

159 Mo ljoa Tjien 1952 White grub of upland rice (Holotrichiahelleri Brsk) [in Indonesian with English summary] Contrib Agric Res Stn Bogor No 13134 p

160 Mochida 0 and T Suryana 1979 The occurrence of the brown planthopper and ragged stunt on lowland and upland rice ill Central Java and logyakarta in tile late of January 1979 Entomnolgy Survey Report No 15 Central Research Institute of Agriculture (CRIA) Sukamandi Indonesia I February 1979 9 p

161Monet C 1979 Varietal resistance to upland rice pests [inI French] Pages 35-39 in Annual report 1977 Institut des Savanes Department des Cultures Vivrieres Bouak6 Ivory Coast

162 Monte 0 1942 A caterpillar ofrice-fields[in Portuguese] Biol6gico (Sao Paulo) 8161-163 (Rev Appl Entomol A31414)

163 Munroe I) D 1979 Pest management strategy for hill rice in four resettlement areas in Sarawak Malaysia Pages 89-102 i1 Guidelines for integrated control of rice insect pests Food and Agriculture Organization Plant Production and Protection Paper 14 Rome Italy 115 p

40 IRPS No 123 January 1987

164 Nakasuji F M Ishii and Jen-Zon Ito 1984 Rice skippers in Taiwan and their life histories ZZrC (Taipei) Tech Bull 82 10 p

165 Navas I) 1976 Fall armyworum in rice Pages 99-106 il Ecological animal control by habitat management Proc Tall Timbers Con-ference 28 Feb-I Mar 1974 Fall Timbers Res Stn Misc Publ 6 Tallahassee Horida USA

166 Nickel 1 1 1961 Biological control of rice stem borer A feasibility study Tcch Bull 2 International Rice Rcarch Institute PO Box 933 Manila Philippines IIl p

167 Nigan P M and R A Vernia 1985 Insect pests of uplaild rice in Uttar Pradesh Int Rice Res Newsl 10(6)22

168 Nilakhe S S 1985 -cological observations on spittle bugs with emphasis on their occirence in rice Pcsqtui Agropecu Bras 20407-414

169 Nixon G EJ 195 1 lie association of ats with aphids and coccids Commonwealth Iusti lcte of Entonology Iondon UK 36 p

170 Nurdin F an Ishak Manti 1985 Study oit seedling fly of upland rice Interlational Upland Rice Moitoring Tour Report 26 lebruary-2 MarLI Sukaranti West Sumatra

171 Oka 1 N 1979 Ciltural conttol of the brown planthopper Pages 357-369 in Brown planthopper threat to rice production in Asia International Rice Research Institutc 1p0 Box 933 Manila Philippines

172 Ohni M and A Vilnaii 1975 Rice insect pests of Italy biology of Sipha grcria (Kaltenbach) and its control [in Italian] Riso (Milan) 2459-71

173 Otanes F Q atnd 1 I Sison 194 1 Pests of rice Philipp J Agric 12211-259

174 Painter R I 1951 Insect icsistance in crop plants University Press of Kansits Lawrence Kansas USA 520 p

175 Pantua P C and 1A litsinger 1981 Arthropod abundance in continuous vs biannual rice cropping systeis Int Rice Res Ncwsl 6(2)20-21

176 Pathak M ) and V A l)yck 1975 Sttudics ott insect pests of upland rice Pages 186-191 in Major research in upland rice International Rice Research Institute PO Box 933 Ilanila Philippines

177 Patil B R B M IHasabe and 1) S Ajri 198 I Chemical control of white gritb lIditriihinhorraia Fabr intfesting groundntut paddy atnd sorghum Indian 1Entomol 43232-236

178 Pawar A 1) 1974 (ollection and identification of insect pests of rice rlerminal repirt lInternational Rice Research Institute PO Box 933 Manila Philippines 72 p (tnimco)

179 IerrajU A aid A S Reedy 1965 Observations ott diapausc itt tie steti borer ohrice Rice News Teller (lai-Mar)9-13

180 Pont A C 1972 A review of the Oriental species of theri ona Rondani (I)iptera Muscidac) of econtomic importance Pages 27-102 in Control of sorglium shoot fly M G lomtani and W R Young eds Oxford atd IH1 Ptthl Co New I)clhi India 324 p

181 Prasad S C 1PChand aid M 1PSingh 1983 Incidence of rice gall midge in the upland plateau region of Hihar Oryza 20235-236

182 Rai P S 198 0 Nisaga simpex damage to rice in the hill tracts of South India Int Rice kles News[ 5(I)12

183 Rathorc Y S 1969 Gonocplhahli dcprssunt Fabr as itpredator of (tilo zonwIlhis (Swinhoc) and other moths in field cage at Patnagar Uttar Pradesh Indian 1Entotnol 3127

184 Reed W 1983 Crop losses causcd by insect pists in the developing world Proceedings 10th Internatial Congress of Plant Protection lrighton UK 174-80

185 Reedy M V 1981 Ateactogasterinhens(Walker)a new root-grub pest of tipland paddy and inaize Curr Sci 501076

186 Reveno M A 1968 Catalogue of the insect pests of economic crops in Colombia [in Spaiish] Asociacion Latino Americana de Entomo-logia (AIAE) 129-33

187 Reyes G M 1957 Rice dwarf disease in the Philippines Food and Agriculture Organiiation Plant Prot Bull 617-19

188 Rice S E A A Grigarick and M 0 Way 1982 Effect of leaf and panicle feeding byarmyworm (Lepidoptera Noctuidae) larvae on rice grain yield Environ Entomol 7593-595

189 Risch S J 1) Andow and M A Altieri 1983 Agioccosystem diversity arnd pest control data tentative conclusions and nizw research dirctions Environ Entomol 12625-629

190 Rivera C T 1956 Three organic insecticides in thecontrol of insects affecting upland rice Philipp Agric 39465-472

191 Roffrey 1 1972 Locusta outbreaks in the Philippines Acrida 1177-188

192 Rojas R 1) 1941 The effect of pruning at varying ages upon the yield of Ramairio Phipp Agric 29851-860

193 Roonwal M i 1979 Ternite life and termite control in tropical South Asia Scientilic Pnbl odhpr India 139 p

194 Rosetto C1 F da Silva Martins N C Schmidt and L EAzzini 1978 Damage caused by pasturespittle bugs 4Dvoi)lavopictaandDJ schach) on rice lBragantia Nota 6 (Matcli)25-27

195 Rosztto C1- S S Neto I- Amantc I) Iink 1Garcia-Vieira D M de SoLza N V Bansa t and A M de Olivcira 1971 Rice pests of Brazil [in Portuguese] Pages 149-238 inrFA()-International Rice Commission 2nd meeting of the Rice Committee for the Americas 6-11 l)ecember 1971 Pelotas Rio Grande do Sul Brazil

196 Rothschild G H L 1969 Obseratiois on the armyworm Spodoptera matritia acronrctoiiles Gn (Lep Noctuidae) Bull Entomol Res 59143-160

197 Rothschild G 1- L 1974 Parasites of the armworm Spolopiera mauritiaacroncitoihes in Malaysia Fntornopliaga 19293-299

198 Sachan i N S K Gangwar and 1N Katujar 1980 Field crickets damaging cages in North Eastern IHill Region Indian J Entomol 42526-527

199 Sands 1) P A 1977 The biology and ecology of Lepocorisa (Hemiptera Alydidac) in Papua New Guinea Res Bull 18 Departshyment of Primary Industry Port Moresby Papua New Guinea 104 p

200 Sarawak 1968 titomology division Pages 97-106 in Report of the research branch of the Department of Altriculture 1966 Kuching (Rev AIppl Eintomol A56603)

201 Satpathy JM M S Das and K Naik 1977 Effect of multiple and mixed cropping on the incidence of scine important pestsI Entomol Res 178-85

202 Sauer IHF G 1939 Notes Ott Elasmpnlpualtsligniosellths Zeller (Lep Pyr) a serious pest ofcereals in Sao Paolo [in Portuguese] Arq Inst Biol 10199-206 (Rev Appl Entotnol A28378)

203 Satinders 1L A 1 SKing and C L Vargas S 1983 Crop pests of Central Anerica with t bibliography [in Spanish] Centro Agroshynotnico Tropical de Investigacion y Ensenanza Turrialba Costa Rica (CATIE) Tec Bol 9 90 p

204 Sauphanor B 1985 Some factors of upland rice tolerttce to stem borers in West Africa Inseu Sci Appl 6429434

205 Sen P atnd S Chakravorty 1976 Stentflics in Haritighata Nadia West Bengal India [it Rice Res Newsl 1(12) 15

206 Senapai B and 1M Satpathy 1983 The seedling fly Atherigona orrzae Malloch is a pest of rice in Orissa (India) J Entomol Res 777-79

207 Shhjahan M S Alatn M Ah nad and BN Islam 1983 A new scarabaeid beetle pest of dryland rice in Bangladesh Int Rice Res Ncwsl 8(I)16-17

208 Sharma G L S C Modgol and R C Gautam 1979 Effect of depth of submergence ott incidence of bacterial blight and white grub infestation in transplanted rice Int Rice Res Newsl 4(6)12-13

209 Sharma S 1985 Scarabaeid beetle outbreak itt upland rice Int Rice Res Newsl 1(1(l)24

210 Shiraki T 1917 The paddy borer Schoenobhits incertulas Walker [achaku Agric Exp Stn Formosa 256 p

211 Silva A de Brito and 11P Magalhaes 1981 Insect pests of rice [in Portuguese] Circular Tecnica No 22 EMBRAPA Centro de Pesquisa t ecuaria de Tropico Unido (CPAJU) Belem Brazil 14 p

212 Singh S A 1979 Emergence of rice leaf butterfly as a rice pest in Manipur India int Rice Res Newsl 4(4)16-17

213 SinghS A 1983 Changing trends of insect pestsof rice in Manipur Int Rice Res Newsl 8(f)12

214 Siwi S S and P H van Doesburg 1984 Lenovorisa Latreille in Indonesia (Heteroptera Coreidae Alydinae) Zool Meded (Leiden) 58117-129

215 Soejitno J 1977 Reaction of some upland rice varieties toseedling fly eltherikonaexigua Stein [in Indonesian] Siri Hana I Penyakit 630-35 Lembga I osat Fenclilian Pertanian Bogor Indonesia

216 Socnardi 1967 Insect pests of rice in Indonesia Pages675-683 in The major inse pests of thu rice plant Proceedings of a symposium at the Internatital Rice Rec Wi itt S pternber 1964 The Johns Hopkins Press Baltimore Maryland USA

217 Soto P E A T Petcz and I W Buddenhagen 1976 Survey of insect pests and diseases of rice in different ecological zones in Nigeria Rice Entomol Ncwsl 435-46

218 Soto 11E and Z Ziddiqi 1976 Screening for resistance to African rice insect pests Varietal improsetnent sCmi n r Bouak Ivory Coast 13-17 Sept 1976 West Africa Rice l)evelopment Association Monrovia Iiberia

219 Squire F A 1972 Entomological problems in Bolivia PANS (PestArtic News Sumr) 18249-268

220 Stapely 1 II 1978 The rice leaf roller in the Solomon Islands Int Rice Res Newsl 3(l)13

221 Suenaga H 1937 On aphids attacking the roots of rice Oyo-I)oubts Zasshi 9150-152 [in Japanese] (Rev Appl Entomol A25745)

222 Suenaga H 1952 On the seasonal abundance of three subterranean aphids on upland rice Bull Kyushu Agric Exp Sti 1249-262

223 Sutynso Yas and Dandi Soekaria 1964 Beberapa hana yang ttenperang tana man padi tanah kering [in Indonesian] Rapat kerja Padi Tanah Kering 6 sd 8 Juily 1964 Kaliurang 9 p

224 Tallamy I) W and R F Denno 1983 Alternative life history patterns in risky environments in example from lace bugs Pages 129-147 in Insect life history patterns R F l)cnio and If Dingle eds Spring-Verlag 225 p

225 Tamura 1 1939 Observations on some insect pests of field crops [in Japanese] 1 Plant Prot 26414-420 (Rev Appl EntomolA27-556)

226 Tamura and Suzuki 1964 Influences of the early-transplanting practice in rice culture upon development of the rice stem borer Chilo siql)ressalisWalker Bull Ilokuriku Natil Agric Exp Stn 761-94

227 Tan J P 1924 The rice root aphid Drowhi ihirsoia Philipp Agric 13277-288

228 Tanaka 1 1957 Taxonomy and distribution of some subterranean aphids injurious to upland rice in Japan with description of a new species [in Japanese English summary] Botyu-Kagaku 22168-176

229 Tanaka T 1961 The rice root aphids their ecology and control Spec [full College of Agriculture Utsunomiya University Japan F3 p

230 Tauber M IC A Tauber R Nechols and 1Obrycki 1983 Seasonal activity of parasitoids control by external internal and genetic factors Pages 87-108 itt I)iapause and life cycle strategies in insects V K Brown and I Ilodek eds )r W Junk The Hague Netherlands

231 Taylor W F 1972 Effects of artificial defoliation (simulating pest damage) on varieties of upland rice Exp Agric 879-83

232 feotia T P S and S Natud 1966 Biotoomics of the rice skipper Parnaraniathiis Fabricius (Lepidoptera Hesperiidac) Indian J Entomol 28181-186

233 Teran F 0 1971 Rice pests of Santa Cruz [in Spanish Ministerio de Agricultura Sita Cruz Bolivia Bull 45 8 p

234 Than Itun 1976 Population dynamics of the yellow rice borer Tryporza itcertulas (Walker) and its damage to the rice plant MS thesis University of the Philippines at Ios Bafios Laguna Philippines 92 p

235 Uichanco L B 1911 Pests Pages 55-74 it Rice Philipp Agric 11-200

IRPS No 123 January 1987 41

236 van der Laan P A 195 1 Life history of the rice seedling fly Contrib Gen Agric Res Stn Boor Indonesia 118 15 p

237 van der Laan 11A 1959 Correlation between rainfall in the dry season and the occurrence of the white rice borer (Scirpolhaga innotala WIk) in lava Entomol Fxp Appl 212-20

238 Van Driesche R G 1983 Meaning of percent parasitismin studies of insect parasitism Fnviron Entomol 121 I1- 122

239 Venugopal M S and A litsingcr 1984 Fffect ofcarbofuran on rice growth Prot Ecol 7313-317

240 Vietnam Directorate of Rural Affairs 1962 Annual work progress report on crop improvement program of rice sugarcane vegetable and field crops (lor the period from July 1961 to June 1962) (11) 299 p (Rev Appl Entomol A51526-527)

241 Wan M T K 1972 Observations on rice leaf- and planthoppers in Sarawak (Malaysian Borneo) Malay Agric 148308-335

242 West Africa Rice I)esclopnient Association 1978 Annual research report Vol 2 Statistics Monrovia Liberia

243 West Africa Rice Development Association 1979 Annual research report p47 Monrovia Liberia

244 Williams ) F and CS lofgren 1981 Fli I illy l-468 anew bait toxicant for control of the red imported lire at Fla Entomol 64472477

245 Williams I) 1 1970 The mealybugs OFlomoptera Coccoidea Iscudococcidac) of sugar-cane rice and sorghum Bull Entomol Res 0109-188

246 Williams D) 1 1982 Pulinaria icervi (Signoret) and its allies ott sugar cane and other grasses Btll Entoinol Res 72111-117

247 Wirjosuiardjo S 1976 Study ofchenmical cont rot of insect pests ott upland rice Imiu Pertanian (Jogjakarta) 1323-329

248 Wirjosuhardjo S and N Wirarta 1976 Chemical control of the white grub tIolorithiatlltriin upland rice in M uneng Irobolinggo Indonesia) [in Indonesian] Imut Pertanian (Jogjakarta) 1330-332

249 Yano K T Miyake and V F Eastop 1983 The biology and economic importance of rice aphids (Henuiptera Aphid idae) a review Bull Entomol Res 73539-566

250 Yelp D and P Rhodes 19(4 lhe success of drilled dryland rice iii the Fiji Islands International Rice Commission Working Party on Rice Production and Protection Manila Philippines 3-10 March1964 Food and Agriculture Organization Rome Italy 13 p

251 Young W R 1981 Fifty-five years of research on the sorghum shootfly Insect Sci Appi 23-9

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Page 31: UPLANDRICE. INSECTl--1i M T -E!R.EOLOU t RTANCE ,AND …delphacid.s3.amazonaws.com › 6958.pdf · The significance of abiotic factors in upland rice insect ecology will be apparent

IRPS No 123 January 1987 31

Table 6 cntinued

Natural enemy Environment

Host stage Species Upland Rainfed lowland Irrigated lowland

Larva-pupa Gold fringed borer Chilo attriciiuslchneumonidae

Xanthopimpla stemmator (Thunberg) x aPupa x

Pteromalidae Trichoinalopsis apanteloctena (Crawford) ax

Pink stem borerb Sesamnia inferensParasite

Larva Braconidae

Stenobracon nicevillei (Bingham) x b b Armyworms Mvthinna SpodopteraLarva

Tachinidac Zygobothriaarroptvora (Robineau-Desvoidy) x x

Braconidac Cotesia spp

X Skipper Pelopidas mathias and green horned caterpillar Melanitis leda ismeneF-gg

TrichogramrnatidacTrichogrannaspp x

Scelionidae Undetermined Scelionid

x Larva

Tachinidae Argyroph lax nitrotibialis (Baranov) X x

Pteromalidae Trichomnalopsis apanteloctena (Crawford)

X Predator

CoenagrionidaeAgriocnenis femina femina Brauer x xIschnura senegalensis tambur

Miridae CrtorhinsliidipennisReuter x x

Nabidac Stcnonabis tagalicus (Stal) x

VeliidaeAlicrovelia douglasi atrolineata (Bergroth) x

Coccinellidae Micraspis crocea (Mulsant) X x x

Vespidae Eumenes campaniformis (Fabricius) x

Formicidac Diaecamna spp xOdontoponera transversa (F Smith) x

Carabidae Chaenius spp OphioneaW- aJasciataSchmidt-Goebel

x

Ophionea ishii ishii Habu x x X

Gryllidav Anaxipta longipennis (Saussur-) x xMetioche ittaticollis (Sial) x x x

Tettigoniidae Conocephalus longipennis (de Haan) x x x

aNo rearing made on rainfed lowland bNo rearing made on rainfed lowland or Irrigated lowland

MrasmiaAmong the Laval parasites the braconid Cardio- lowlands and parasitizes oriental maize borer OsIriniachilesphilijpphensis is most adapted to an upland environ- furnacalis (Guenee) larvae The leaffolder pupal parasitesment It also parasitizes Hydelepta indicata a common not only occur equally in all environments but also haveleaffolder of legumes The ichneumonids Trichomma cna- wvide host ranges- parasitizing species of OsIrinia Marucaphalocrosis and Temehcha stangli occur more in the Hydelepta and Honiona - in maize and legumeslowlands T stangli is most adapted to irrigated lowlands The principal egg parasite of yeltow stem borer (YSB)and also parasitizes Chilo and Scirpophaga stem borer Telenomus rowani is widely adapted to all environmentslarvae T cnaphalocrosis is most prevalent in rainfed and has an unusual alternate host -Tabanus eggs (25)

32 IRPS No 123 January 1987

Tetrastictu schoenobii specializes only in YSB and occurs mainly in irrigated lowlands -- further proof that YSB is most adapted to the wetter environments

Larval parasites of YSB are highly environment-specific The braconid Stenohracon nicevillei occurs only in upland rice but also attacks the pink stem borer (PSB) Tieinchwa phiiiplPwnsis an ichneumonid occurs in upland rice and also parasitizes the larvae of (7hiu stqpresaisinatro-morpha accepta melaihoracica like 7 stangi is mnost adapted to the irrigated lowlands

The dark-headed stem borer(l)HSB) (ih pochit3sus was not collected in all rice environments and the gold fringed borer Chilo auricilius nd PSB were collected only in upland rice Xaantiholiplastenrnaloran ichneumonid larval parasite appears to be widely adapted whereas TrihomaslopisapanitI)cteiiaa pteromalid pupal parasite occurs mostly in wetlands and also parasitizes larvae of Chilo suppressais and Pelhuidas mathias

Armyworms Myit hinin and Spodujipera have two main larval parasites - a tachinid Zrgohothriaatropivora in rainfed environments and a braconid Cotesia spp in irrigated lowlands

The butterfly pests Pelopidas and 4elanitishave tricho-grammatid egg parasites found only in irrigated lowland environments

Two larval parasites show environmental preferences --Argrroph av nigrotihialis a tachinid in rainfed habitats and Trichonialopsisapantelkcnela in irrigated ricefields A limgrotibialisalsoattacks the sweet potato hornwormAgrius convolvtdi and T apanteloctena parasitizes DH S1B

Predators as a group show more distinct environmental preferences The aquatic damselflies Coenagrionidae are most abundant in the lowlands Cvriorhinus is also most adapted to the lowlands The nabid Stenonabis tagalicus occurs only in the uplands and isalso prevalent on legumes The coccinellid lady beetles wasps and ants are more prevalent in upland areas AlicrasIpis cit-ea larvae prey ona variety of aphids attacking legumes and maize Etunienes campanonia vespid mud wasp makes nests in trees and is therefore most adapted to the more botanically diverse uplands Soil-dwelling ants cannot tolerate flooding

Upland environments are habitats to arboreal carabid beetles Three species of Chlaeniusbeetles prey on leaffolder larvae and are perhaps more important than parasites (27)

The gryllids and tettigoniids are widely adapted egg predators not only on rice but also on maize and legumes They feed on eggs of most insects that are laid on leaves

The spidercommunity of upland rice environments isrich in species At one site (Batangas Philippines) 31 species have been recorded (23) Of the 176 spider species recorded in Philippine ricefields about one-half (82) occur in upland rice The spiderspecies of upland rice environments overlap more with those of rainfed lowland than with those of irrigated lowland rice (24) The same study showed that of the three environments irrigated rice has (ie greatest spider species diversity foiiowed by rainfed lowland and upland

However of the 10 most prevalent ricefield spiders 9 were abundant in upland rice showing wide adaptation All three spider guilds -- orb-web space-web and huting spiders shywere prevalent in upland ricefields There were however differences in environmental prelerences for some spider species

Of the orb-wcavers Tetragniathamanlihulatawas partishycila rlv abundant 7 japonica was low and Ltucatge dehcorata was absent in upland rice

As in other environments Aiypeiwnaifirtosana was the most dominant space-web spider however its relative numbers were lowerthan for lowland sites Among hunting spiders Ox|opes javanus and Lcosa leatcost fgla were more abundant and Lvcosa lpseudoanmnlata was less abundant in upland compared with lowland rice envishyroniments

Extensive rearing of rice insect pests in upland rainfed lowland and irrigated sites over an 8-yr period in the Philippines provided insight into the effectiveness of parasites as natural enemies (Table 7 8) Most of toe data are from large sample sizes taken over at least one crop which overcomes the pitfalls expressed by Van Driesche (238)

VSB egg parasitization was surprisingly similar across all environments Larval parasitization was highly dynamic among sites and even years within the same site but showed somewhat lowerlevels in uplandsthan in irrigated wetlands This would have been predicted for egg parasites as well since YSB is adapted to the lowlands and so must be its parasites This isevidence that Telenomus rowani is as good at dispersing as its main hosts Trowani clings onto the anal tufts of female YSB and parasitizes eggs as they are laid This phoresy may explain its wide adaptation The more specialized lowland YSB larval parasites therefore appear more effective than their upland counterparts with more alternate hosts

Low collection levels of PSB larvae especially in the wetlands make comparisons between environments difficult Low levels of larval parasifization were recorded in both rainfed environments Except in the Cagayan rainfed wetland site in 1980 low levels ef larval parasitization were evident at irrigated and rainfed sitesalike No differences by environment were evident Higher levels of parasitization occurred in Sarawak during a 1967 outbreak also from tachinids Rothschild (197) indicated that higher parasishytization occurred in upland rice areas

Skipper and green horned caterpillar larval parasitization rates were generally low and showed no environmental effect htighest activity occurred in Cagayan in 1980 (48 parasitization) Although larval parasitization levels of RLF were variable year to year activity appeared greater in the irrigated lowlands than in the uplands Again the highest levels were recorded in Cagayan in 1980 Cagayan has extensive grasslands and fallow around the rice areas and a relatively long rainfall period Rainfall in 1930 was good for crop growth and apparently for increase of larval parasites

IRPS No 123 January 1987 33

Table 7 Parasitization rates of rice planthoppers and green leafhopper in upland rainfed lowland and irrigated lowland environments Philipshypines 1976-840

Environment and site Year Egg parasitization () Nymphaladult parasitization ()

BPH W13P GLH BPII WBPH GLIH

UplandBatangas

Rainfed lowland

1976 1979 1980 1982

15 17 12

2(47)8(39)

4(51)

9(1800)14 (212)

12(304)

8 (101)6 (127)

9(116)

Iloilo

Cagayan

1976 1977 1978 1981 1980 1981 1982

0 (69)

11

44 48 40

-

-

46 55 44

0 (471)

11

52 56 49

0 (200)9 (214)

10 (1259)8(235)6 (900) 3(800)

-7 (462)

11 (270)3 (313)5 (900) 1 (800)

4 (600)10 (725)10 (362)12(870)5 (900) 3 (800)

Pangasinan

Irrigatedlowland

1984 1976 1978 1979 1982

13 (436) 13(379) 14 (457)

6(14)7 (197)

17(198) 11 (486) 18 (275)

3 (81)10 (274)3 (116)

16(871) 19(88)

28(211)9 (764)5 (647)7 (1204) 8(305)

Laguna 1977 11 7 21 1978 31 1979 14 4 4 28 1980 7 8 20 1981 1983

56 7(278)

26 16 (1650)

16 27(1921)

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R I Apostol unpublished 1lo1lo and Pangasinan 136 Cagayan BCanapi unpublished Liuna 175 I C Pantua unpublished 118 234 43 Carino and Shepard unpublished BPH = brown planthopper Nila parvara htgens W131Pi =whitebacked planthopper Sogatelia furcifera GLH = green leafhopper Nephotettrix virescens Figiires in parentheses are sample sizes

Tauher et al (230) reported that internal external and genetic factors influence the seasonal activity of parasitoids

Hopper egg parasites but not nymphaladult parasites were also parlicularly abundant at the Cagayan site possibly the egg parasites have more alternative hopper prey in the weedy areas than nymphaladult paisites

The seasonal dynamics of hopper egg parasites can be seen for upland rice in Batangas(Fig I 2) and for rainfed lowland lice in Pangasinan( 135)and Cagayan( 105) Levels of BPHWBPI-L and GLH egg parasitization were corn-parable in upland and irrigated lowland environments Except for some high rates of paraMsitization early in the season the parasitization rate tended to be steady in all envilonments However hopper nymphaladult parasi-tizution levels were more similar in the rainfed environments and lower than in irrigated rice

Overall parasites do not appear highly effective by themselves against upland or even lowland pests Among natural enemies parasites have attracted the most attention because they can be more readily assesscd

Predators and pathogens lend themselves better to management practices Predators the most important group of natural enemies in riccfields are difficult to quantify On the other hand pathogens appear less im-portant but can be readily cultured and disseminated

The natural enemy commulnity of upland rice is rich in species and differs significantly from that of lowland rice

These beneficial organisms must be conserved by applying insecticides judiciously particularly sprays Seed or soil placement of chemicals minimizes exposure to natural enemies The strategy to derive the greatest benefit from natural enemies isto allow the greatest number of beneficial species to thrive Then perhaps at least one species will be effective against each pest at any tine overcoming the variable abundance of each species within and between years

Programs to introduce exotic species or mass produce indigenous ones are ambitious and expensive because they require trained people on a sustained not ad hoc basis

Chemical control Insecticides offer rapid and efficient control of upland insect pests (190) but-should be used only after other control measures have been considered Insecticides are rarely used on upland rice because of cost (32) Upland rice yields are normally too low to justify the expenditure Also most upland rice is grown as a cash crop Spraying upland rice is more difficult than lowland rice because water is less accessible Government subsidy programs to provide pestishycides to tribal peoples for upland rice production have created problems of toxicity to humans because isolated people have had no experience to be able to handle pesticides safely

However ins-eticide use has beenjustified when a crop is

0Z

Table 8 Parasitization rates of lepidopterous pests in upland rainfed lowland and irrigated lowland environments Philippines 1976-84a

Parasitization (5)

Environment and site Year YSB GFB PSB Earcutting Swarming Skipper Greenborned RLF Brown larva larva caterpillar caterpillar caterpillar caterpillar larva semilooperEgg Larva larva larva larva larva larva

UplandBatangas 1977 - 0 (57) - 0 (63) - 0 (71) - 0 (106) 11(142) 2 (101)1978 43 (4697) 8(451) 5 (38) 3 (40) 6 (61) 2(58) 15 (57) 13 (129) 13 (1286) 4 (30)1979 59(4135) 12(320) 13 (11) 5(28) 9(45) 6(46) 7(23) 8(160) 9(1152) 8(26)

Rainfed lowland Iloilo 1976 - 0 (10) 0(10) 0 (7) 0 (19) 2 13) 15 (180) 5 (76)1978 0 (54) 0 (3) 0 (17) 9 (64) 4 (78) 3 (73) 0 (121) 0 (191)1979 64 (2350) 16 (84) 6 (17) 0(6) 18 (68) 25 (104)

1981 56 (1322) 18(304) 0(5) 12(51) 18(138)Pangasinan 1976 60 (7 7 )b 62(82) 0(111) 5 (39)

1978 58 (53)b 5(51) Cagayan 1979 59(713) 8 (101) 6 (34) 0(7)

1980 0(11) 4 (28) 15 (286)

62 ( 48(177) 61(140)1981 0(5) 12 (2) 13(45) 14 (92)

Irrigatedlowland Laguna 1974 62 8

1974-75 50 41 1975 43 72 1978 96 (623) 21(24) 33 (60)1979 83(2017) 18(110) 8(62) 11 (89) 22(400)1980 78 (1625) 20(75) 0 (16) 4 (72)1983 60

9 (80) 7 (270) 30

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R F Apostol unpublished Iloilo and Pangasnan 136 Cagayan B Canapi unpublished Laguna 175 P CPantua unpublished 118 234 143 Carino and Shepard unpublished YSB = yellow stem borer ScirpophagaincertulasGFB = gold fringed borer Chilo auricilius PSB = pink stem borerSesamia inferens earcutting caterpillar Alythimna separata swarming caterpillar Spcdoptera mauritia acronyctoides skipper Pelopidas mathias reen horned caterpillar Melanitls ledaismene RLF = rice leaffoldersCnaphalocrocismedinalis and Marasmiaspp brown semilooper Mocisfrugalis Figures in parentheses are sample sizes Egg masses

IRPS No 123 January 1987 35

Table 9 Effect of seeding rate and insecticide on plant density and Yield losses from insect pests in upland rice often areyield of Dagge upland rice Batangas Philippines 1976a comparable to those of lowland rainfed or irrigated rice

Seeding rate Insecticide Dosage Plant density Yield (0-Y30o) but usually from different pests The principal (kg aiha) seed treatment (kg aiha) (noIra-row) (tha) groups are soil pests feeding on sown seed and rootsat 14 DE followed by seedling pests These groups have largely gone

50 None - 35 a 20 a unnoticed leading researchers to conclude that stem borers50 Carbofuran F 10 38 ab 23 a and seed bugs are the most important pests (17) Yield loss 10 beCabofuranF100 None - 4551 c 23 a trials that pinpoint major pests need to includc seed

100 Dieldrin WP 10 52 cd 22 a treatments Granules applied in the furrow may not give150 None - 59 d 22 a adequate protection from many seed pests

aAv of 8 fields DE = days after crop emergence In a column means The floristic and pedological diversity of upland rice followed by a common letter are not significantly different at the environments is matched by the diversity of arthropods5level by D)MRT F = flowable Wi = wettable powder both pests and their natural enemies Irrigation has homoshy

genized lowland rice environments Flooded soils tend tothreatened by epidemics of locusts or armyworms New have the same properties and to eliminate soil pests The technology stressing efficiency has minimized insecticide spatial and temporal dominance of cultivated rice has cost benefited specialist pests

Seed treatment is normally inexpensive and can be Upland rice has no specialist arthropod species The economicallyjustified in -nanycases(66) Treating seed with monophagous pests on upland rice prefer the lowlands systemic insecticide (216) but not chlorinated hydrocarbons where rice evolved Rice in the uplands exists only because (247) is most effective in controlling seedling maggots of man and even though lowland rice predates upland riceFoliar sprays have to be repeated to obtain control and specialist species evolved Wheat maize and sorghum do often are washed off by frequent early season rains not have monophagous pests leading us to conclude that Granular insecticides are effective but required dosages are there is no evolutionary advantage for inssects to have too expensive to justify except under heavy infestation narrow host ranges in upland graminaceae(247) When directed against first-instar larvae banding Pests have adapted to the uplands not only by having granules in seed furrows (177) can be effective against white wider plant host ranges but also by having long life cyclesgrubs at low dosages (025 kg aii ha) (134) Seed treatments undergoing dormancy or being dispersive Each pest has a are not promising for white grub control (248) unique life history strategy suited to highly unpredictable

Seed treatment is also effective ainst ants and may be environments In agivenyear someinsect specics an better cheaper than increasing the seeding rate In Batangas exploit upland rice as a food source but no pest is a senious treating 50 kg seed ha was as effective as increasing the one every year on upland rice seeding rate to 100 kgha and using untreated seeds (Table The low yielding potential of upland rice means less 9) In Brazil seed treatments control termites and other soil quantity and lower quality of food for pests to exploit but insects including the lessercornstalk borer(66) Domiciano also less incentive for farmers to undertake control (58) however found seed treatments inconsistent in their The most effective control would be early and synchroshyeffect against high populations of lesser corn stalk borer nous planting of early-maturing upland rice varieties to

Baiting can be an inexpensive way of controlling ants prevent insect pest buildup Breeding for insect resistance seedling maggot flies mole crickets and field crickets Baits should initially focus on stem borers because they are verymade of locally available material and impregnated with difficult to control by other means Biocontrol efforts insecticide can be sparingly distributed taking adantage of should first stress conservation of natural enemies preshythe pests ability to disperse and encounter bait sites dators and pathogens rather than parasites Efficient

The new low-volume hand-held controlled-droplet chemical control includes seed treatment or baiting for seed sprayers may offer an advantage over high-volume knap- seedling and some root pests and using controlled dropletsack sprayes for controlling foliar pests Upland rice sprayers for spot treatments against foliar pests and seed canopies are more open than those in lowland rice and bugs when economic thresholds are reached better droplet penetration should result Controlled droplet applicators would be ideal for seedbugs and defoliators

CONCLUSION

Upland rice is attacked by a wider array of insect pests than is lowland rice mainly beca use of the addition of soil pests Generally population levels of lowland rice pests are lower in upland than lowland rice leading many people to conclude that insect pests are not important on upland rice

36 IRPS No 123 January 1987

REFERENCES CITED

1Abraham C C and K S R Mony 1978 Occurrence ofLeptocorisa acuta Fabr (Coreidae Hemiptera) as a pest of nutmeg trees J Bombay Nat ttistSoc 74553

2 Agyen-Sampong M 1975 HlierogrlphusdiaganensisKrauss (Orthop-tera Acrididae) a new pest of rice in Northern Ghana Ghana J Agric Sci 8249-253

3 Agyen-Samnpong M 1976 Varietal resistance and other inle-ratcd pest control components as factors of rice varietal improvement in West Africa Prcsented at the Second varietal improvement serinar 14-18 Sep 1976 Bouak Ivory Coast West Africa Rice Development Association Monrovia Liberia 18 p

4 Agyen-Sampong M 1977 Outbreak of Ihccrorrdnisorae in rice seedlings near Rokupr Sierra Leone lit Rice Res Newsl 2(6)16-17

5 Agyen-Sampong M 1980 Dirty panicles and rice yield reduction caused by hugs IntRice Res Newsl 5(l)11-12

6 Agyen-Sampong M 1982 The major pest problems of irrigated upland and mangrove swanip rice ecosystems ir the humid tropical Guinea savanna and Salel climate zones Pages 318-328 in Pro-ceedings training course cn concepts techniques and application of irtegrated pest management in rice inWest Africa 10-20 Jan 1982 Fendall Liberia West Africa Rice l)evelopioent Association Monrovia Liberia 505 p

7 Akibo-Betts V I and S A Raymundo 1978 Aphids as rice pests in Sierra Leone lit Rice Res Newsl 3(6)15-16

8 Akinsola F A 1975 Present status of different rice stemborers in parts of Nigeria iicL Entomo Newsl 328

9 Akinsola FA 1979 Ihe biology and ecology of rice stemborers in Nigeria Ph 1) thesis University ol lbadan Nigeria 217 p

10 Akinsola E A 1979 Guidelines of integrated management of insect pests of upland rice in West Africa Presented at the Seminar on integrated management of rice diseases and insect pests at Bobo-Dioulasso Upper Volta 17-22 Sep 1979 West Africa Rice Develop-ment Association Monrovia Liberia

II Akinsola IA 1984 Insect pests of upland rice in Africa Pages 301-305 in An overview of upland rice research Proceedings of the 1982 lIouak Iwsrv Coast upland rice workshop International Rice Research Institute 13O Box 933 Manila Philippines

12 Alam M S V TJohn and Kaung Zan 1985 Insect pests and diseases of rice in Africa Pages 67-82 in Rice improvement in Eastern Central and Southern Africa proceedings of the international rice workshop at Iusaka Zambia 9-19 April 1984

13 Alticri M A A van Schoonhovcn and J D Doll 1977 The ecological role of weeds in insect pest management systems a review illustiated with bean (Phaseohsvulgaris L)cropping systems PANS (Pest Artie News Summ) 23195-205

14 Anmante F 1967 The leaf-cutting ant Atia capiguaraa pest of pastures [in Portuguese] Biol6gico (Sao Paulo) 33113-120 (Rev Appl Entomol A56207)

15Andrade A C and D Puzzi 1951 Preliminary results of experi-merits on the control of Sculptcoris iastantus on sugarcane [in Portuguese] Bioulgico (Sao Paulo) 1744-49 (Rev Appl Entomol A41421422)

16 Appert J 1970 Maliarplia separaelha (rice stem borer) new observations of entomological problems of rice in Madagascar [in French] Agron Trop (Paris) 25329-334

17 Ariadeau Mand Z Harahap 1986 Relevant upland rice breeding objectives Pages 189-197 in Progress in upland rice research Proceedings of the 1985 Jakarta conference International Rice Research Institute 1O Box 933 Manila Philippines

18Asar S and Kisimoto 1975 Seasonal occurrence of the smaller brown planthopper in upland rice [in Japanese] Proc Kanto-Tosan Plant Prot Soc 2228

19 Bakker W 1974 Characterization and ecological aspects of rice yellow mottle virus in Kenya Meded Landbouwhogesh Wage-ningen 152 p

20 Balut F F 1970 White grubs attacking rice Oryza ativa L [in Portuguese] Biol6gico (So Paulo) 36321-322

21 Banerjee S N and L M Pramanik 1967 The lepidopterous stalk borers of rice and their life cycles in the tropics pages 103-124 in The major insect pests ofthe rice plant Proceedings ofa symposium at the International Rice Research Institute September 1964 The Johns Hopkins Press Baltimore Maryland USA

22 Barrion A T and J A Litsinger 1980 Ants a natural enemy of cliaiarac in dryland rice IntRice Res Newsl

5(4)22-23 23 Barrion A T and J A litsinger 1981 The spider fauna of

Philippine rice agroecosystem 1Dryland Philipp Entomnol 5139-166

24 Barrion A T and 1 A Iitsinger 1984 The spider fauna of Philippine rice agroccosystcms II Welland Philipp Eiitomol 611-37

25 Barrion A T1 and J A Litsinger 1984 lihanus (Diptera Tabanidae) eggs an alternative host of rice stei borer (SI3) egg parasite Telenoonis digns (Iliymenoptcra Scelionidae) lit Rice Res Newvsl 9(6)19

26 Barrion A T and J A Litsinger 1985 Identilication of rice leaffolders by wing markings Int Rice Res Newsl 10( 1)24

27 Barrion A T and J A Litsinger 1985 Chctiosspp (Coleoptera Carabidae) a leaffolder ILF)predator lit Rice Res Newsl 10(1)21

28 Bertels A 1970 Insect pests of rice and theircontrol [in Portuguese] Pelotas Brazil Instituto de Expcrimentacao Agropccuararias do Sul Circular 43 24 p

29 Bonzi S M 1982 Ctilo (flisilineus J de Joannis (Lepidoptera Pyralidae) stem borer of irrigated cereals in Upper Volta LioFrench] Agron Trop Riu Rizic Cult Vivrieres Trop 37207-209

30 Bourke T V T L Fenner J N LStihick G Baker EHassan D F OSullivan and C S li 1977 Insect pest survey for the year ending 30th June 1969 Papta New Guinea )epartment of Agrishyculture Stock and Fisheries Port Moresby 57 p

31 Bowden 1 1973 Migration of pests in the tropics Meded Rijksfac Landbouwwet Gent 38785-796

32 Breniere J 1973 The main rice parasites in West Africa and their control Pages 27-42 in Proceedings seminar on plant protection for the rice crop West Africa Rice Development Association Monrovia Liberia

33 Breniere J 1976 The principal insect pests of rice in West Africa and their control West Africa Rice Development Association Monrovia Liberia 52 p

34 Brown ES 1962 The African arinyworn Spoiolptera eXI)Ina (Walker) (Lepidoptera Noctuidae) a review of the literature Cornshymonwealth Institute of Entomology London UK57 p

35 Brown F B 1968 Upland rice in Latin America FAO-International Rice Commission 12th Session of the Working Party on Rice Production and Protection 2-14 Sept Peradeniya Sri lanka Food and Agriculture Organization Rome Italy 6 p

36 Buyckx F J 1 1962 Disease and insect pests of crops inCongo Rwanda and Burundi [iil French] Publications dc LInstilut National pour lestude Agrononiquedu Congo M Weissenbruck S A Brussels Belgium 708 p

37 Cariiio F 0 1981 Role of natural enentics inpopulation suppression and pest management of green rice lealhoppers Ph D thesis University ofthe Philippines at Los Bafitos PhilippinesLaguna 136 -

38 Carpente A J 1983 The development of rice cultivation and extension of vegetable productin in Zanzibar Final Report Vol II1 October 1983 Food and Agricultme Organization Ronte Italy

39 Carroll C P and S1 Risch 1983 Tropical annual cropping systems ant ecology Environ Manage 751-57

4(0 Cates R G 198 1 Host plant predictability and the feeding patterns of monophagous oligophagous and polyphagous insect herbivores Oecologia (Berl) 48319-326

IRPS No 123 January 1987 37

41 Cendcafia S M and F B Caora 1967 Insect pests of rice in the 62 Ene 1 C 1981 I)iiorldhol)ierusihilis Slater (licteroplera Philippines Pages 591-616 in The najor insect pests ofthe rice plant lygacidue Illissinac) a new rice pest front Nigeria lit Rice Res Proceedings of a symposium at the International Rice Research Newsl N3)15 Institute September 1964 The Johns Hopkins Press Baltimore 63 Fugenio T S 1965 rhe effect oi e-vCgetation on rice production Maryland USA 729 p and the possibility of overcoming it by leaf pruning MS thesis

42 Centro Ihternacional de Agricklittira Tropical 1984 Upland rice in University ol the Philippines at Los Baijos ILaguna Ihilippins 69 p Iatin Anerica Pages 93-119 hi An os ervies of uplaind rice research 04 Farrow R A 1975 The Alrican nmigratory locust ii is main Proceedings of the 1982 Boinakamp Ivory Coast upland rice workshop outbreak area of the middle Niger qlinlitatise studies of solitary International Rice Research Institute P0 Blox 933 Manila populations in relation to environmental factors Locusla No II Philippines 198 p

43 Chandra G 1979 Taxonomy ad hn riononlics ofthe imect parasites of 65 Fennal R G 1943 Food crop investigation Windward and rice Icalloppers and planthoppers in the Pihilippines and their Ieeward Islands First report Report onl work dunring the periodiiiportairce iii natural biol)gical cointrol Philipp Fntoltol 4119-139 April 1942-April 1943 Imperial College of Tropical Agriculture

44 Chang T T 1976 The originii colut ion enlIivation dissemination Trinidad 16 p (Rev of Appl Fintornol A3222-25) and diersificition of Asian and African rices -uphytica 25425-44 1 66 Ferreira F 1983 Integrated pest ritaageiret of upland rice [ill

45 Chatierjee P B 1979 Rice leaffolder attacks in India lit Rice Res Portuguese English summary] Pages 323-341 in Upland riceculttre Newsl 4(3)21 factors affecting production E Ferreira T Yamada and F-

46 Cheng Cbien-pan 19(4 Upland rice in Taiwan International Rice Malmolta eds Institutoda Potassae Foslato Institnto Internacional Cnmmission Wiorking Party oit Rice Production ard Protection I()th da Potassa Piracicaha Sao Paulo lraid Meeting 3-10 March 1964 Manila Philippines Food and Agricul- 67 Ferreira E arid F dl Silva Martins 1979 Occurrence of ture Organiaion Rome Italy 5 p Sogatohs orvziola (Muir) inupland rice in Goias Ilrail lit Rice

47 Chcrrett 1 M and 1) 1Peregrine 1976 A review of the status of Rcs Ncwssl 4(4)18 leaf-cutting ants and their control Ann Appl Biol 84124-128 68 Ferreira E and 1F da Silva Martiris 1984 Insect pestof rice ill

48 Chipeta F M 1974 Pests of rice in Malawi Pages 90-95 in Brazil and their control Jill Portugucsej FMIlRAtPA-CNPAF Proceedings Sith eastern Africa cereals research conference 10-15 Goiania Brmiil 67 p March 1974 DRB Mialda ed Ministry of Agriculture lilongwe 69 Ferreira FI F ca Silha Miartins S S Nero and 1 It P de Malawi Carvallio 1982 M tltiplc regression analysis todeterimine the effect of

49 Chu Yau-i 1979 The d atiaging period of the rice seedling fly insect populatiois ott upland rice yield [inIPortuguese Pesqui (Atlterigotiavxigu Stcin) (NItiscide Diptera) oil cori) at Fast Java Agropecu Bras 17671-675 Indonesia Plant Prot Bill 21403413 70 Ferreira F 1F da Silva Martins and F 1PZimmermann 1979

50 Conklin H C 1957 IHaninoo agriculture a report on aln integral Rice varietal resistance to tle lesser corn stalk borer [in Port uguese] systent of shi fti rig cult ivationi in the Pltilippincs Food and Agriculture Pesqui Agropccu Biras 14317-321 Organization Rome Italy 209 p 71 Fleming W F 1968 Biological control of lie hapanese beetle U S

51 Cramer H II 1967 Plant proection and world crop production Dep Agric RLs Sers [ech Bull 1383 78 p Pilaniensclhutz-Nachr 201-524 72 Food and Agriculture Organiation of the United Nations 1982

52 l)ani R C and Majunidar 1978 Prelininary observations on tie Country report surmimary Pages 27-37 in Report on the first incidence of root aphids on differeit rice cultnrcs Sci Cult 4488-89 FAOIUN)IPhail and regional training coiuirse in iniproved ctltural

53 Davidson R I 1979 lffects of root feeding on foliage yield Pages practices for upland rice 5-23 Oct 1981 Northern Region Agrculturd117-120 in T K Ci osb and R B Pottinger eds Proceedings of the Developririt Center Chiang Mai Food and Agricuilture Organiiashy2nd Australian conlerence on grassland invertebrate ecology Govt tion Rome Italy 37 n Printer Wellington New Zealand 73 Franco do Ariaral S 195 1 Residual action of insecticides in turfy

54 l)ean G 1W 1976 Rice insect pests inlaos Int Rice Res Newsl soil--the results of a field experiment [in Portuguese] Biol6gico (Sao 1(2)15 Paulo) 17209-214 (Rev Appl Entomol A41423)

55 I)ean G J W 1978 Insect pests of rice in Laos PANS 24(3)280-289 74 Freeman ) 1970 Report onithe Iban london School of -coroiics 56 )ingle H 1984 Behavior genes and lifc histories conplex adapta- Monographs on Social Anthropology No 41 Athlane Press

tions to uncertain environments Pages 169-194 in A new ecology London UK 317 p novel approaches to interactive systems P W Price C N Slobod- 75 Fuchs T W1 A Harding and 1W Smithlr 1979 Induction and chikoff and W S Gaud cdsl ohn Wiley New York USA termination oldiapause in the sugarcane borer Ann Flitonlol Soc

57 Dobelmann-JP and M Falais 1968 Upland rice cultivation ill tie Am 72271-274 North-western part of Madagascar in 1966-1967 [in French English 76 (hbriel B1) 1975 A review tf the major insect pests of some tpland summary] Trop Agron 2313-21 crops in tlc Philippines Pages 86-111 in Ad Htoc panel of experts on

58 Doniiciano N L 1979 Chemical control of tie lesser corn stalk pest disease arnd weed problems in some rainfed crops Bangkok (lolsonno~paloIgnowslhts) (Zeller 1848) (I-epid(ptera Ihycitid ae) Thailand Food aid Agriculture Organization Roire Italy

on rice in Ponta Gressa Parara 1977-1978 [in Portuguese] Pages 77 Garg D K and N K Shal 1983 White grub outbreak on rainfed 291-303 in Anais 1 Reuniao de tecnicos cmiirizicultura do estado de dryland rice in Uttar Pradeslh Int Rice Res Ncwsl 81(4)16-17 Sao Paulo-Campinas Cati Brazil 78 Garg I) K and IP Tandon 1982 Major insect pests of rice on hilly

59 Dunsmore 1 R 1970 Investigations on tlie varieties pests and tracts of Uttar Pradesh India lilt Rice Res Newsl 7(11)11-12diseases of upland rice (hill padi) in Sarawak Malaysia Int Rice 79 Garrity 1) P 1983 Upland rice research an international bibliograpy Comm Newsl 1929-35 1975-1982 Iternational Rice Research hnstitnte PO Box 933

60 Elias 1R1965 The new decline of rice seedlings is not caused by Manila Philippines 121 p parasites [in Portuguese] Biol6gico (Sao Paulo) 313-6 80 Ghauri M S K 1982 A new subspecies of Iinorh)rone bull

61 Empresa Brasiliera de Pesquisa Agropecuaria 1984 Upland rice itt cornlus Slater (Hemiptera Iygaeidae) front Papua New Guinea on Brazil Pages 121-134 it Av overview of upland rice research rice and carpet grass Ihll lEntoniol Res 72133-137 Proceedings ofthe 1982 Blouak Ivory Coast Upland rice workshop 81 Gonzales F 0 PArregoces R I ihernandef and OT Parada International Rice Research Institute PO Box 933 Manila 1983 Insect and imite pests of rice arid their control in Latin America Philippines [in Spanish] la Federacion Nacional de Arrocerts (Fedearroz)

Bogota Colombia 60 n

38 IRPS No 123 January 1987

82 Grist D H and R J A W Lever 1969 Pests of rice Longmans London UK520 p

83 Hale P R 1979 Two subterranean pests of upland rice in Papua New Guirnca IntRice Res Newsl 4(2)16-17

84 Hale P R and J T Hale 1975 Insect pests of rice in the New Guinea Islands Rice Entomol Newsl 34

85 Harris W V 1969 Termites as pests of crops and trees Common-wealth Institute of Entomology London UK 41 p

86 Hattori 1 1968 A method to distinguish between grass leaf roller (Ctaplha rocis medinalisGuenee) and the rice leaf roller (Susurtia exigua Butler) [in Japanese] Plant Prot (Japan) 23167-170

87 Henry I E and 1A Onsager 1984 Experimental control of the mormon cricket Iahrosshnph by NJsena locustae(Aficrospora Microsporidae) a protozoan parasite of grasshoppers (Ort Acri-didae) Entomophaga 27197-201

88 Hidaka 1 P Vungsilabulr and SKadkao 1974 Studies on ecology and control of the rice gall midge in Thailand Tech Bull 6 TropicalAgricultural Research Center Tokyo Japan 113 p

89 Hincklcy A 1) 1963 Ecology and control of rice planthoppers in Fiji Bull Entomrol Res 53467 481

90 Hirao J 1982 Ecology and chtInical control of the rice leaffolder Jpn Pestic Inf 4114-17

91 Ho DT 1984 Sten borer incifience in rice ecosystems in KenyaEast Africa Pages 307-320 in A overview of upland rice research Proceedings of the 1982 Bouac Ivory Coast upland rice workshop International Rice Research Institute PO Box 933 Manila Philippines

92 Ho )T J 1Njokah and J GKibuka 1983 Studies on rice sten borers in Kenya with emphasis on ifaliarpiaipiraellRag Insect Sci Appl 465-73

93 Ichihara I T Ito A NIatsuo and M Sawada 1971 Chemical control of the rice striped stem borer in upland rice grown in plastic mulch [iii Japanese] Proc Kanto-Tosan Plant Prot Soc 1882

94 IclhiharaIand A Matsuo 1969 Damage caused by rice striped stett borer to upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1686-87

95 Ichhara I atid A MIatsun11970 Conside ration oft lie extraordinary prevalence of rice striped stem borer in upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1778-79

96 Institut de Recherches Agrolnliqes Iropicale et des Cultures Vivrieres 1978 Insect resistance Pages 12-13 in Upland rice 1978 atnual report Bouak Ivory Coast 27 p

97 International Institute for Tropical Agriculture 1983 Aphids itt rice roots I ITA Res Briefs 4(1-3)4-5

98 International Institite fi Tropical Agriculture 1983 Hopperburn in rice caused by brown plathopper IITA Res Briefs 4(l-3)6-7

99 International Rice Research Institute 1975 Annual report for 1974 PO Box 933 Manila Philippines p 216

100 International Rice Restarch Institute 1976 Annual repiort for 1975 PO Box 933 Manila Philippines p 228

101 International Rice Research Institute 1979 Cropping systems program Component technology development and evaluation Page 404 i Annual report for 1978 11O Box 933 Manila Philippines

102 International Rice Research Institute Department of Entomology 1981) Insecticid evaluation 1979 Tables 73 74 PO Box 933Manila Philipes

103 International Rice Research Institute Department of Entonmology 1981 Insecticide evalomtion 1980 Table 81 POBox 933 Manila Philippines

104 International Rice Research Institute 1981 Report of a meeting of Asian rice-based cropping systems entomologists Eleventh cropping systems working group meeting 18-22 May 1981 PO Box 933 Manila Philippines p 132 133 144

105 International Rice Research Institute 1984 Cropping systems program Pages 368-373 in Annual report for 1983 PO Box 933 Manila Philippines

106 International Rice Research Institutc 1984 Upland rice in Asia Pages 45-68 in An overview of upland rice research Proceedings of the 1982 Bouakamp Ivory Coast upland rice workiiop loternational Rice Research Institute 11O Box 933 Manila Philippines

107 Irshad M 1977 Grasshoppers in rice in Pakistan lit Rice Res Newsl 2(3)8-9

108 Ito R and TOno 1964 Upland lice its culture and varieties in Japan lIternational Rice Cormmission loth Meeting of the Working Party on Rice Production and Protection 3-10 March 1964 Manila Philippines Food and Agriculture Organization Rome Italy 12 p

109 Jeffrey M I and V A Dyck 1983 Effect of moonlight on lightshytrap catches of brown planthopper lit Rice Res Newsl 8(6) 13

110 Jennings P R and T Alicia Pineda 1983 Management ofthe rice plnthoppers Sogatodsorivi(coIl in Latin America Presented at the meeting on stable plant resistance FAO and lIndocsian I)inectorate of Food Crops Protection Bali lndonesia 29 Novcniber-2 l)ccemshyber 9 p

IllJimenez Gomez J A 1979 Rice insect pests i lla ios ()riCntales lil Spanish] Pages 99-103 in Rice ciltivation Instituto (olotibiatto Agropecuario Compndio No 29

112 Jongeleen F JJ Soesanto and A Mockidjo 1979 Studies oti pathogens of two species of Scarabacidac froinIndonesia Z Angew Enlomnol 87412-417

113 Jouvenaz 1) PC S lofgren and W A Baiks 1981 Biological control of imported fire antsa review of cirrent knowledge Bull Entomol Soc Am 27203-208

114 Kainoh Y K Shiniiu S MarUani Y Taiaki 1980 lost finding behavior of tie rice bu lplotori+JchinvittjI)allas (liciniptera Corcidac) with special reference to diet pattern ol aggregation and feeding on the rice plant Appl Entomol Zool 15225-233

115 Kakkar K 0 P Sharma and P I Sharma 1977 Note on Ihteronyrchus liodresRedt (Coleopicra I)ynastidae) as asporadic pest of paddy and maie inHimachal Pradesh Corr Agric 156-57

116 Kalode M131 79 Major insect pest problems and their control in upland rice (Unpublishcd mineogriph CWI oJNaaiat lni Chandaratat (198 1)original not seen)

117 Kalshoven L i 98 I The pests of crops in I ldoniesia (translated by P A vatn der Laan) 1 T Ichtiar B3aru-vati Iloeve lakarta Indonesia 701 p

118 Kamal N Q 1981 Suppression of whitebacked planto ner Sogatella fircaifra (Horvath) aLId rice lealfolder (nhal -cis licdb lis (Guenee) populations by natural cnclmie hlI) tiesis Grcgorio Araneta University Foundation Victoncta Park Metto Manila Philippines 179 Ip

119 Karim A N M Reaul 1975 A species ofliealy bug Trioni i70s ceres new oii rice in the Philippines Rice Entoniol Newsl 247

120 Katanyukul Wand C Chandaratat 1981 Insect pestsof uplaid rice and their control Report firstFAOUNlFi Tliailand training course on improved cultural practices for tplad rice Chiang Mai Thailand 5-23 Oct 1981 Food and Agriculture Organiation Rome Italy 22 p

121 Kern F 1955 lhletwioa(I)uninemts) bhhit Biturm(Coleoptera) a new pest of sugarcane inVenezuela [inSpanish with Etnglish sunmary] Agron Trop (Maracay) 527-29 (Rev Appl Eitoniol A4572)

122 Khan K M and S Ghai 1974 White grubs ald their control in India Pesticides (1Bombay) 819-25

123 Kiritani K 1975 Integrated pest management of rice pests Pages 66-70 itt Proceedings platning workshop ott cooperative hIeld research in pest managementP SMotooka ed East-West Center Honololu Hawaii USA

124 Kisimoto R 1967 Ecology of insect vectors forecasting and chemical control Pages 243-255 in The virus diseases of tle rice plant Proceedings of a symposium atthe International Rice Research Institute April 1967 The Johns Hopkins Press altimore Maryshyland USA

125 Kisimoto R and V A Dyck 1976 Climate and rice insects Pages 367-391 in Climate and rice International Rice Research Institute PO Box 933 Manila Philippines

126 Kureha S S Kobayashi SAsari and T Ishihara 1977 Life history and control of the mole cricket Grrllotalpa africana Palisot de Beauvois [in Japanese English translation 7520 1RRI library] Proc Kanto-Tosan Plant Prot Soc 21192-197

127 Kushwaha K S 1981 The black beetle as a pest of rice in Haryana India Int Rice Res Newsl 6(4)14

128 Lawani S M 1982 A review of tile effects of various agronomic practices on cereal stem borers Trop Pest Manage 28266-276

129 Leuck D B 1966 Biology of the lesser cornstalk borer in South Georgia J Econ Entomol 59 797-801

130 lever R J A W 1955 Insect pests of the rice plant in Malaya Int Rice Comm Newsl 1612-21

131 Li Ching-Sing and Shin-Foon Chiu 1951 The study ofthe rice gall midge Iachydiplosis orvza Wood Mason [in Chinese with English summary] 1Taiwan Agric Res 21-13

132 Iitsinger J A 1984 Assesament of the need-based application for rice Proceedings TechnologyTransfer Workshop 15-16 March 1984 International Rice Research Institute 1PO Box 933 Manila Philippines 6 p

133 Litsinger J A 1984 Environmental diversity and insect pest abundance with reference to the Pacific Pages 285-307 in Proceedings sub-regional training course on methods of controlling diseases insects and other pests of plants in the South Pacific 4-20 Oct 1982 Vaini Kingdom of Tonga Nukualofa

134 Litsinger J A RF Apostol and M BObusan 1983 White grub Leucopholis irrorata (Coleoptera Scarabaeidae) pest status popula-tion dynamics and chemical conllol in arice-maize cropping pattern in the Philippines J Econ Entoinol 761133-1138

135 Litsinger J A C de la Cruz F Raymundo A T Barrion M D Lumaban J P Bandong M S Venugopal F Paragna and E Balete 1982 Insect pests and insecticide reponse of dry-seeded bunded rice Pages 179-193 in Report of workshop on cropping systems research in Asia International Rice Research Institute Los Bafos Philippine

136 Litsinger J A M 1) Luinaban J P Bandong A T Barrion W Katanyukul and N Panda 1978 Insect pest management in rainfcd lowland rice agroecosystms of tropical Asia International Rice Research Conference 17-21 April 1978 International Rice Research Institute Los Bafios Philippines

137 Litsinger J A and K Moody 1976 Integrated pest management in multiple cropping systems Pages 293-317 in R I Papendick P A Sanchez and G B Triplett eds Multiple cropping American Society of Agronomy Madison Wisconsin USA

138 Loevinsohn M E 1984 The ecology and control of rice pests in relation to the intensity and synchrony of cultivation Ph D thesis University of London 36) p

139 Loevinsohn M E J A Litsinger N Panda M A Austria R F Apostol C Andrion and J Bandong 1982 A basis for predicting rice insect pest abundance in cropping systems Paper presented at the 1982 International rice research conference on rice and cropping systems 19-23 April 1982 International Rice Research Institute Los Bafios Iaguna Philippines 25 p

140 Lopez A W 1930 Repot of the Entomology Department Pages 145-172 in Annual report of the Philippine Sugar Association 1929-30 Manila

141 Lopez AW 1931 Entomology Department Pages 227-266 in Annual report of the director or research 1930-31 Philippine Sugar Association Manila

142 Lopez A W 1932 Entomology Deparment Pages 252-279 in Annual report of the Philippine Sugar Association 1931-32 Manila

143 Lungibill P and G G Ainslee 1917 The lessercornstalk borer US Dep Agric Agric Res Serv Tech Bull 539 27 p

IRPS No 123 January 1987 39

144 McGuire J U Jr and BS Crandall 1967 Survey of insect pests and plant diseases ofselected food crops of Mexico Central America and Panama U SDepartment of Agriculture International Agriculshytural Development Service Agricultuial Researrh Service October 1967 11 p

145 McNeill Sand T R ESouthwood 1978 The role of nitrogen in the development of insectplant relations Pages 77-9 in lliochemical aspects of plant and animal coevolution J B Ilarbornu ed Academic Press London UK

146 Maebara H 1955 Spatial distribution of lastinstiarlanaeof purplish stem borer (Sesamia iqferens Walker) hibernating in an upland rice field Bull Kagoshima Univ Fac Agric 438-41

147 Mallick E H and N G Hajra 1977 Response of rice plant to defoliation treatments at different developmental stages Indian Agric 2151-56

148 Mansingh A 1971 Physiological classification of dormancies in insects Canl Entomol 103983-10)9

149 Martell C G 1974 Catalogue of plant feeding insects of Mexico [in Spanish] Fitofilo 27(69)1-176

150 Martinez 0 J 1982 Principal rice pests in Ilanos orientalis [in Spanish] P~ages 80-94 in Curso deact nalization de conocimientos en arroz Villaniencis Instituto (olonmbiano Agropecuario

151 Martins J F da Silva F Ferrcira A S Prabhu F 1 P Zimmermann 1980 Use of pesticides for the control of the main subterranean upland rice pests esqui Agropecu Bras 1553-u

152 Martins I F da Silva F Fe eira rod BdamSilveira Pinheiro I Q2 Damage simulation caused by defoliators of upland rice [in Portuguese with English summary] Pesqui Agropecu Bras 171113-1119

153 Martins 1F da Silva N(iyuen van Tan and 1Ida Silveira Pinheiro 1981 Stem borer resistance in upland rice associated with plant morphological characters [in Portuuce Iesqui Agropecu Bras 16187-192

154 Masaki S 1980 Sumnmer diapause Ann Rev Entomiol 251-25 155 Masuzawa T 11 Suwa and F Nakasoji 1983 l)ifkleences of

oviposition preferencc and survival rate of two skipper butterflies Parnara guttata and PIchlpidai iathiais (Iepidoptera HIesperiidac) on rice plant and cogon grass New Entomol 3247-56

156 Maurya D M and C P Vaish 1984 Upland rice in India Pages 69-92 in An overview oftpland rice research Proceedings ofthe 1982 Bouakamp Ivory Coast upland rice workshop International Rice Research Institute IO Box 933 Mnila Philippines

157 Menon M G R R N Katiyai S Kuar and S S Misra 1970 Termites attacking the paddy crop Sci Cult 36294-295

158 Miyahara Y and 1 Nishikubo 1967 Behavior of young nymphs of the red rice root aphid Iholahsipiun ri liabhdontinalisSasaki in relation to movement from the leaf to the roots of upland rice [in Japanese] Pioc Assoc Plant Prot Kyushu 1363-66

159 Mo ljoa Tjien 1952 White grub of upland rice (Holotrichiahelleri Brsk) [in Indonesian with English summary] Contrib Agric Res Stn Bogor No 13134 p

160 Mochida 0 and T Suryana 1979 The occurrence of the brown planthopper and ragged stunt on lowland and upland rice ill Central Java and logyakarta in tile late of January 1979 Entomnolgy Survey Report No 15 Central Research Institute of Agriculture (CRIA) Sukamandi Indonesia I February 1979 9 p

161Monet C 1979 Varietal resistance to upland rice pests [inI French] Pages 35-39 in Annual report 1977 Institut des Savanes Department des Cultures Vivrieres Bouak6 Ivory Coast

162 Monte 0 1942 A caterpillar ofrice-fields[in Portuguese] Biol6gico (Sao Paulo) 8161-163 (Rev Appl Entomol A31414)

163 Munroe I) D 1979 Pest management strategy for hill rice in four resettlement areas in Sarawak Malaysia Pages 89-102 i1 Guidelines for integrated control of rice insect pests Food and Agriculture Organization Plant Production and Protection Paper 14 Rome Italy 115 p

40 IRPS No 123 January 1987

164 Nakasuji F M Ishii and Jen-Zon Ito 1984 Rice skippers in Taiwan and their life histories ZZrC (Taipei) Tech Bull 82 10 p

165 Navas I) 1976 Fall armyworum in rice Pages 99-106 il Ecological animal control by habitat management Proc Tall Timbers Con-ference 28 Feb-I Mar 1974 Fall Timbers Res Stn Misc Publ 6 Tallahassee Horida USA

166 Nickel 1 1 1961 Biological control of rice stem borer A feasibility study Tcch Bull 2 International Rice Rcarch Institute PO Box 933 Manila Philippines IIl p

167 Nigan P M and R A Vernia 1985 Insect pests of uplaild rice in Uttar Pradesh Int Rice Res Newsl 10(6)22

168 Nilakhe S S 1985 -cological observations on spittle bugs with emphasis on their occirence in rice Pcsqtui Agropecu Bras 20407-414

169 Nixon G EJ 195 1 lie association of ats with aphids and coccids Commonwealth Iusti lcte of Entonology Iondon UK 36 p

170 Nurdin F an Ishak Manti 1985 Study oit seedling fly of upland rice Interlational Upland Rice Moitoring Tour Report 26 lebruary-2 MarLI Sukaranti West Sumatra

171 Oka 1 N 1979 Ciltural conttol of the brown planthopper Pages 357-369 in Brown planthopper threat to rice production in Asia International Rice Research Institutc 1p0 Box 933 Manila Philippines

172 Ohni M and A Vilnaii 1975 Rice insect pests of Italy biology of Sipha grcria (Kaltenbach) and its control [in Italian] Riso (Milan) 2459-71

173 Otanes F Q atnd 1 I Sison 194 1 Pests of rice Philipp J Agric 12211-259

174 Painter R I 1951 Insect icsistance in crop plants University Press of Kansits Lawrence Kansas USA 520 p

175 Pantua P C and 1A litsinger 1981 Arthropod abundance in continuous vs biannual rice cropping systeis Int Rice Res Ncwsl 6(2)20-21

176 Pathak M ) and V A l)yck 1975 Sttudics ott insect pests of upland rice Pages 186-191 in Major research in upland rice International Rice Research Institute PO Box 933 Ilanila Philippines

177 Patil B R B M IHasabe and 1) S Ajri 198 I Chemical control of white gritb lIditriihinhorraia Fabr intfesting groundntut paddy atnd sorghum Indian 1Entomol 43232-236

178 Pawar A 1) 1974 (ollection and identification of insect pests of rice rlerminal repirt lInternational Rice Research Institute PO Box 933 Manila Philippines 72 p (tnimco)

179 IerrajU A aid A S Reedy 1965 Observations ott diapausc itt tie steti borer ohrice Rice News Teller (lai-Mar)9-13

180 Pont A C 1972 A review of the Oriental species of theri ona Rondani (I)iptera Muscidac) of econtomic importance Pages 27-102 in Control of sorglium shoot fly M G lomtani and W R Young eds Oxford atd IH1 Ptthl Co New I)clhi India 324 p

181 Prasad S C 1PChand aid M 1PSingh 1983 Incidence of rice gall midge in the upland plateau region of Hihar Oryza 20235-236

182 Rai P S 198 0 Nisaga simpex damage to rice in the hill tracts of South India Int Rice kles News[ 5(I)12

183 Rathorc Y S 1969 Gonocplhahli dcprssunt Fabr as itpredator of (tilo zonwIlhis (Swinhoc) and other moths in field cage at Patnagar Uttar Pradesh Indian 1Entotnol 3127

184 Reed W 1983 Crop losses causcd by insect pists in the developing world Proceedings 10th Internatial Congress of Plant Protection lrighton UK 174-80

185 Reedy M V 1981 Ateactogasterinhens(Walker)a new root-grub pest of tipland paddy and inaize Curr Sci 501076

186 Reveno M A 1968 Catalogue of the insect pests of economic crops in Colombia [in Spaiish] Asociacion Latino Americana de Entomo-logia (AIAE) 129-33

187 Reyes G M 1957 Rice dwarf disease in the Philippines Food and Agriculture Organiiation Plant Prot Bull 617-19

188 Rice S E A A Grigarick and M 0 Way 1982 Effect of leaf and panicle feeding byarmyworm (Lepidoptera Noctuidae) larvae on rice grain yield Environ Entomol 7593-595

189 Risch S J 1) Andow and M A Altieri 1983 Agioccosystem diversity arnd pest control data tentative conclusions and nizw research dirctions Environ Entomol 12625-629

190 Rivera C T 1956 Three organic insecticides in thecontrol of insects affecting upland rice Philipp Agric 39465-472

191 Roffrey 1 1972 Locusta outbreaks in the Philippines Acrida 1177-188

192 Rojas R 1) 1941 The effect of pruning at varying ages upon the yield of Ramairio Phipp Agric 29851-860

193 Roonwal M i 1979 Ternite life and termite control in tropical South Asia Scientilic Pnbl odhpr India 139 p

194 Rosetto C1 F da Silva Martins N C Schmidt and L EAzzini 1978 Damage caused by pasturespittle bugs 4Dvoi)lavopictaandDJ schach) on rice lBragantia Nota 6 (Matcli)25-27

195 Rosztto C1- S S Neto I- Amantc I) Iink 1Garcia-Vieira D M de SoLza N V Bansa t and A M de Olivcira 1971 Rice pests of Brazil [in Portuguese] Pages 149-238 inrFA()-International Rice Commission 2nd meeting of the Rice Committee for the Americas 6-11 l)ecember 1971 Pelotas Rio Grande do Sul Brazil

196 Rothschild G H L 1969 Obseratiois on the armyworm Spodoptera matritia acronrctoiiles Gn (Lep Noctuidae) Bull Entomol Res 59143-160

197 Rothschild G 1- L 1974 Parasites of the armworm Spolopiera mauritiaacroncitoihes in Malaysia Fntornopliaga 19293-299

198 Sachan i N S K Gangwar and 1N Katujar 1980 Field crickets damaging cages in North Eastern IHill Region Indian J Entomol 42526-527

199 Sands 1) P A 1977 The biology and ecology of Lepocorisa (Hemiptera Alydidac) in Papua New Guinea Res Bull 18 Departshyment of Primary Industry Port Moresby Papua New Guinea 104 p

200 Sarawak 1968 titomology division Pages 97-106 in Report of the research branch of the Department of Altriculture 1966 Kuching (Rev AIppl Eintomol A56603)

201 Satpathy JM M S Das and K Naik 1977 Effect of multiple and mixed cropping on the incidence of scine important pestsI Entomol Res 178-85

202 Sauer IHF G 1939 Notes Ott Elasmpnlpualtsligniosellths Zeller (Lep Pyr) a serious pest ofcereals in Sao Paolo [in Portuguese] Arq Inst Biol 10199-206 (Rev Appl Entotnol A28378)

203 Satinders 1L A 1 SKing and C L Vargas S 1983 Crop pests of Central Anerica with t bibliography [in Spanish] Centro Agroshynotnico Tropical de Investigacion y Ensenanza Turrialba Costa Rica (CATIE) Tec Bol 9 90 p

204 Sauphanor B 1985 Some factors of upland rice tolerttce to stem borers in West Africa Inseu Sci Appl 6429434

205 Sen P atnd S Chakravorty 1976 Stentflics in Haritighata Nadia West Bengal India [it Rice Res Newsl 1(12) 15

206 Senapai B and 1M Satpathy 1983 The seedling fly Atherigona orrzae Malloch is a pest of rice in Orissa (India) J Entomol Res 777-79

207 Shhjahan M S Alatn M Ah nad and BN Islam 1983 A new scarabaeid beetle pest of dryland rice in Bangladesh Int Rice Res Ncwsl 8(I)16-17

208 Sharma G L S C Modgol and R C Gautam 1979 Effect of depth of submergence ott incidence of bacterial blight and white grub infestation in transplanted rice Int Rice Res Newsl 4(6)12-13

209 Sharma S 1985 Scarabaeid beetle outbreak itt upland rice Int Rice Res Newsl 1(1(l)24

210 Shiraki T 1917 The paddy borer Schoenobhits incertulas Walker [achaku Agric Exp Stn Formosa 256 p

211 Silva A de Brito and 11P Magalhaes 1981 Insect pests of rice [in Portuguese] Circular Tecnica No 22 EMBRAPA Centro de Pesquisa t ecuaria de Tropico Unido (CPAJU) Belem Brazil 14 p

212 Singh S A 1979 Emergence of rice leaf butterfly as a rice pest in Manipur India int Rice Res Newsl 4(4)16-17

213 SinghS A 1983 Changing trends of insect pestsof rice in Manipur Int Rice Res Newsl 8(f)12

214 Siwi S S and P H van Doesburg 1984 Lenovorisa Latreille in Indonesia (Heteroptera Coreidae Alydinae) Zool Meded (Leiden) 58117-129

215 Soejitno J 1977 Reaction of some upland rice varieties toseedling fly eltherikonaexigua Stein [in Indonesian] Siri Hana I Penyakit 630-35 Lembga I osat Fenclilian Pertanian Bogor Indonesia

216 Socnardi 1967 Insect pests of rice in Indonesia Pages675-683 in The major inse pests of thu rice plant Proceedings of a symposium at the Internatital Rice Rec Wi itt S pternber 1964 The Johns Hopkins Press Baltimore Maryland USA

217 Soto P E A T Petcz and I W Buddenhagen 1976 Survey of insect pests and diseases of rice in different ecological zones in Nigeria Rice Entomol Ncwsl 435-46

218 Soto 11E and Z Ziddiqi 1976 Screening for resistance to African rice insect pests Varietal improsetnent sCmi n r Bouak Ivory Coast 13-17 Sept 1976 West Africa Rice l)evelopment Association Monrovia Iiberia

219 Squire F A 1972 Entomological problems in Bolivia PANS (PestArtic News Sumr) 18249-268

220 Stapely 1 II 1978 The rice leaf roller in the Solomon Islands Int Rice Res Newsl 3(l)13

221 Suenaga H 1937 On aphids attacking the roots of rice Oyo-I)oubts Zasshi 9150-152 [in Japanese] (Rev Appl Entomol A25745)

222 Suenaga H 1952 On the seasonal abundance of three subterranean aphids on upland rice Bull Kyushu Agric Exp Sti 1249-262

223 Sutynso Yas and Dandi Soekaria 1964 Beberapa hana yang ttenperang tana man padi tanah kering [in Indonesian] Rapat kerja Padi Tanah Kering 6 sd 8 Juily 1964 Kaliurang 9 p

224 Tallamy I) W and R F Denno 1983 Alternative life history patterns in risky environments in example from lace bugs Pages 129-147 in Insect life history patterns R F l)cnio and If Dingle eds Spring-Verlag 225 p

225 Tamura 1 1939 Observations on some insect pests of field crops [in Japanese] 1 Plant Prot 26414-420 (Rev Appl EntomolA27-556)

226 Tamura and Suzuki 1964 Influences of the early-transplanting practice in rice culture upon development of the rice stem borer Chilo siql)ressalisWalker Bull Ilokuriku Natil Agric Exp Stn 761-94

227 Tan J P 1924 The rice root aphid Drowhi ihirsoia Philipp Agric 13277-288

228 Tanaka 1 1957 Taxonomy and distribution of some subterranean aphids injurious to upland rice in Japan with description of a new species [in Japanese English summary] Botyu-Kagaku 22168-176

229 Tanaka T 1961 The rice root aphids their ecology and control Spec [full College of Agriculture Utsunomiya University Japan F3 p

230 Tauber M IC A Tauber R Nechols and 1Obrycki 1983 Seasonal activity of parasitoids control by external internal and genetic factors Pages 87-108 itt I)iapause and life cycle strategies in insects V K Brown and I Ilodek eds )r W Junk The Hague Netherlands

231 Taylor W F 1972 Effects of artificial defoliation (simulating pest damage) on varieties of upland rice Exp Agric 879-83

232 feotia T P S and S Natud 1966 Biotoomics of the rice skipper Parnaraniathiis Fabricius (Lepidoptera Hesperiidac) Indian J Entomol 28181-186

233 Teran F 0 1971 Rice pests of Santa Cruz [in Spanish Ministerio de Agricultura Sita Cruz Bolivia Bull 45 8 p

234 Than Itun 1976 Population dynamics of the yellow rice borer Tryporza itcertulas (Walker) and its damage to the rice plant MS thesis University of the Philippines at Ios Bafios Laguna Philippines 92 p

235 Uichanco L B 1911 Pests Pages 55-74 it Rice Philipp Agric 11-200

IRPS No 123 January 1987 41

236 van der Laan P A 195 1 Life history of the rice seedling fly Contrib Gen Agric Res Stn Boor Indonesia 118 15 p

237 van der Laan 11A 1959 Correlation between rainfall in the dry season and the occurrence of the white rice borer (Scirpolhaga innotala WIk) in lava Entomol Fxp Appl 212-20

238 Van Driesche R G 1983 Meaning of percent parasitismin studies of insect parasitism Fnviron Entomol 121 I1- 122

239 Venugopal M S and A litsingcr 1984 Fffect ofcarbofuran on rice growth Prot Ecol 7313-317

240 Vietnam Directorate of Rural Affairs 1962 Annual work progress report on crop improvement program of rice sugarcane vegetable and field crops (lor the period from July 1961 to June 1962) (11) 299 p (Rev Appl Entomol A51526-527)

241 Wan M T K 1972 Observations on rice leaf- and planthoppers in Sarawak (Malaysian Borneo) Malay Agric 148308-335

242 West Africa Rice I)esclopnient Association 1978 Annual research report Vol 2 Statistics Monrovia Liberia

243 West Africa Rice Development Association 1979 Annual research report p47 Monrovia Liberia

244 Williams ) F and CS lofgren 1981 Fli I illy l-468 anew bait toxicant for control of the red imported lire at Fla Entomol 64472477

245 Williams I) 1 1970 The mealybugs OFlomoptera Coccoidea Iscudococcidac) of sugar-cane rice and sorghum Bull Entomol Res 0109-188

246 Williams D) 1 1982 Pulinaria icervi (Signoret) and its allies ott sugar cane and other grasses Btll Entoinol Res 72111-117

247 Wirjosuiardjo S 1976 Study ofchenmical cont rot of insect pests ott upland rice Imiu Pertanian (Jogjakarta) 1323-329

248 Wirjosuhardjo S and N Wirarta 1976 Chemical control of the white grub tIolorithiatlltriin upland rice in M uneng Irobolinggo Indonesia) [in Indonesian] Imut Pertanian (Jogjakarta) 1330-332

249 Yano K T Miyake and V F Eastop 1983 The biology and economic importance of rice aphids (Henuiptera Aphid idae) a review Bull Entomol Res 73539-566

250 Yelp D and P Rhodes 19(4 lhe success of drilled dryland rice iii the Fiji Islands International Rice Commission Working Party on Rice Production and Protection Manila Philippines 3-10 March1964 Food and Agriculture Organization Rome Italy 13 p

251 Young W R 1981 Fifty-five years of research on the sorghum shootfly Insect Sci Appi 23-9

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No 106 Rice grassy stunt virus 2 a new strain of rice grassy stunt in tire literature

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axial-flow pump

Page 32: UPLANDRICE. INSECTl--1i M T -E!R.EOLOU t RTANCE ,AND …delphacid.s3.amazonaws.com › 6958.pdf · The significance of abiotic factors in upland rice insect ecology will be apparent

32 IRPS No 123 January 1987

Tetrastictu schoenobii specializes only in YSB and occurs mainly in irrigated lowlands -- further proof that YSB is most adapted to the wetter environments

Larval parasites of YSB are highly environment-specific The braconid Stenohracon nicevillei occurs only in upland rice but also attacks the pink stem borer (PSB) Tieinchwa phiiiplPwnsis an ichneumonid occurs in upland rice and also parasitizes the larvae of (7hiu stqpresaisinatro-morpha accepta melaihoracica like 7 stangi is mnost adapted to the irrigated lowlands

The dark-headed stem borer(l)HSB) (ih pochit3sus was not collected in all rice environments and the gold fringed borer Chilo auricilius nd PSB were collected only in upland rice Xaantiholiplastenrnaloran ichneumonid larval parasite appears to be widely adapted whereas TrihomaslopisapanitI)cteiiaa pteromalid pupal parasite occurs mostly in wetlands and also parasitizes larvae of Chilo suppressais and Pelhuidas mathias

Armyworms Myit hinin and Spodujipera have two main larval parasites - a tachinid Zrgohothriaatropivora in rainfed environments and a braconid Cotesia spp in irrigated lowlands

The butterfly pests Pelopidas and 4elanitishave tricho-grammatid egg parasites found only in irrigated lowland environments

Two larval parasites show environmental preferences --Argrroph av nigrotihialis a tachinid in rainfed habitats and Trichonialopsisapantelkcnela in irrigated ricefields A limgrotibialisalsoattacks the sweet potato hornwormAgrius convolvtdi and T apanteloctena parasitizes DH S1B

Predators as a group show more distinct environmental preferences The aquatic damselflies Coenagrionidae are most abundant in the lowlands Cvriorhinus is also most adapted to the lowlands The nabid Stenonabis tagalicus occurs only in the uplands and isalso prevalent on legumes The coccinellid lady beetles wasps and ants are more prevalent in upland areas AlicrasIpis cit-ea larvae prey ona variety of aphids attacking legumes and maize Etunienes campanonia vespid mud wasp makes nests in trees and is therefore most adapted to the more botanically diverse uplands Soil-dwelling ants cannot tolerate flooding

Upland environments are habitats to arboreal carabid beetles Three species of Chlaeniusbeetles prey on leaffolder larvae and are perhaps more important than parasites (27)

The gryllids and tettigoniids are widely adapted egg predators not only on rice but also on maize and legumes They feed on eggs of most insects that are laid on leaves

The spidercommunity of upland rice environments isrich in species At one site (Batangas Philippines) 31 species have been recorded (23) Of the 176 spider species recorded in Philippine ricefields about one-half (82) occur in upland rice The spiderspecies of upland rice environments overlap more with those of rainfed lowland than with those of irrigated lowland rice (24) The same study showed that of the three environments irrigated rice has (ie greatest spider species diversity foiiowed by rainfed lowland and upland

However of the 10 most prevalent ricefield spiders 9 were abundant in upland rice showing wide adaptation All three spider guilds -- orb-web space-web and huting spiders shywere prevalent in upland ricefields There were however differences in environmental prelerences for some spider species

Of the orb-wcavers Tetragniathamanlihulatawas partishycila rlv abundant 7 japonica was low and Ltucatge dehcorata was absent in upland rice

As in other environments Aiypeiwnaifirtosana was the most dominant space-web spider however its relative numbers were lowerthan for lowland sites Among hunting spiders Ox|opes javanus and Lcosa leatcost fgla were more abundant and Lvcosa lpseudoanmnlata was less abundant in upland compared with lowland rice envishyroniments

Extensive rearing of rice insect pests in upland rainfed lowland and irrigated sites over an 8-yr period in the Philippines provided insight into the effectiveness of parasites as natural enemies (Table 7 8) Most of toe data are from large sample sizes taken over at least one crop which overcomes the pitfalls expressed by Van Driesche (238)

VSB egg parasitization was surprisingly similar across all environments Larval parasitization was highly dynamic among sites and even years within the same site but showed somewhat lowerlevels in uplandsthan in irrigated wetlands This would have been predicted for egg parasites as well since YSB is adapted to the lowlands and so must be its parasites This isevidence that Telenomus rowani is as good at dispersing as its main hosts Trowani clings onto the anal tufts of female YSB and parasitizes eggs as they are laid This phoresy may explain its wide adaptation The more specialized lowland YSB larval parasites therefore appear more effective than their upland counterparts with more alternate hosts

Low collection levels of PSB larvae especially in the wetlands make comparisons between environments difficult Low levels of larval parasifization were recorded in both rainfed environments Except in the Cagayan rainfed wetland site in 1980 low levels ef larval parasitization were evident at irrigated and rainfed sitesalike No differences by environment were evident Higher levels of parasitization occurred in Sarawak during a 1967 outbreak also from tachinids Rothschild (197) indicated that higher parasishytization occurred in upland rice areas

Skipper and green horned caterpillar larval parasitization rates were generally low and showed no environmental effect htighest activity occurred in Cagayan in 1980 (48 parasitization) Although larval parasitization levels of RLF were variable year to year activity appeared greater in the irrigated lowlands than in the uplands Again the highest levels were recorded in Cagayan in 1980 Cagayan has extensive grasslands and fallow around the rice areas and a relatively long rainfall period Rainfall in 1930 was good for crop growth and apparently for increase of larval parasites

IRPS No 123 January 1987 33

Table 7 Parasitization rates of rice planthoppers and green leafhopper in upland rainfed lowland and irrigated lowland environments Philipshypines 1976-840

Environment and site Year Egg parasitization () Nymphaladult parasitization ()

BPH W13P GLH BPII WBPH GLIH

UplandBatangas

Rainfed lowland

1976 1979 1980 1982

15 17 12

2(47)8(39)

4(51)

9(1800)14 (212)

12(304)

8 (101)6 (127)

9(116)

Iloilo

Cagayan

1976 1977 1978 1981 1980 1981 1982

0 (69)

11

44 48 40

-

-

46 55 44

0 (471)

11

52 56 49

0 (200)9 (214)

10 (1259)8(235)6 (900) 3(800)

-7 (462)

11 (270)3 (313)5 (900) 1 (800)

4 (600)10 (725)10 (362)12(870)5 (900) 3 (800)

Pangasinan

Irrigatedlowland

1984 1976 1978 1979 1982

13 (436) 13(379) 14 (457)

6(14)7 (197)

17(198) 11 (486) 18 (275)

3 (81)10 (274)3 (116)

16(871) 19(88)

28(211)9 (764)5 (647)7 (1204) 8(305)

Laguna 1977 11 7 21 1978 31 1979 14 4 4 28 1980 7 8 20 1981 1983

56 7(278)

26 16 (1650)

16 27(1921)

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R I Apostol unpublished 1lo1lo and Pangasinan 136 Cagayan BCanapi unpublished Liuna 175 I C Pantua unpublished 118 234 43 Carino and Shepard unpublished BPH = brown planthopper Nila parvara htgens W131Pi =whitebacked planthopper Sogatelia furcifera GLH = green leafhopper Nephotettrix virescens Figiires in parentheses are sample sizes

Tauher et al (230) reported that internal external and genetic factors influence the seasonal activity of parasitoids

Hopper egg parasites but not nymphaladult parasites were also parlicularly abundant at the Cagayan site possibly the egg parasites have more alternative hopper prey in the weedy areas than nymphaladult paisites

The seasonal dynamics of hopper egg parasites can be seen for upland rice in Batangas(Fig I 2) and for rainfed lowland lice in Pangasinan( 135)and Cagayan( 105) Levels of BPHWBPI-L and GLH egg parasitization were corn-parable in upland and irrigated lowland environments Except for some high rates of paraMsitization early in the season the parasitization rate tended to be steady in all envilonments However hopper nymphaladult parasi-tizution levels were more similar in the rainfed environments and lower than in irrigated rice

Overall parasites do not appear highly effective by themselves against upland or even lowland pests Among natural enemies parasites have attracted the most attention because they can be more readily assesscd

Predators and pathogens lend themselves better to management practices Predators the most important group of natural enemies in riccfields are difficult to quantify On the other hand pathogens appear less im-portant but can be readily cultured and disseminated

The natural enemy commulnity of upland rice is rich in species and differs significantly from that of lowland rice

These beneficial organisms must be conserved by applying insecticides judiciously particularly sprays Seed or soil placement of chemicals minimizes exposure to natural enemies The strategy to derive the greatest benefit from natural enemies isto allow the greatest number of beneficial species to thrive Then perhaps at least one species will be effective against each pest at any tine overcoming the variable abundance of each species within and between years

Programs to introduce exotic species or mass produce indigenous ones are ambitious and expensive because they require trained people on a sustained not ad hoc basis

Chemical control Insecticides offer rapid and efficient control of upland insect pests (190) but-should be used only after other control measures have been considered Insecticides are rarely used on upland rice because of cost (32) Upland rice yields are normally too low to justify the expenditure Also most upland rice is grown as a cash crop Spraying upland rice is more difficult than lowland rice because water is less accessible Government subsidy programs to provide pestishycides to tribal peoples for upland rice production have created problems of toxicity to humans because isolated people have had no experience to be able to handle pesticides safely

However ins-eticide use has beenjustified when a crop is

0Z

Table 8 Parasitization rates of lepidopterous pests in upland rainfed lowland and irrigated lowland environments Philippines 1976-84a

Parasitization (5)

Environment and site Year YSB GFB PSB Earcutting Swarming Skipper Greenborned RLF Brown larva larva caterpillar caterpillar caterpillar caterpillar larva semilooperEgg Larva larva larva larva larva larva

UplandBatangas 1977 - 0 (57) - 0 (63) - 0 (71) - 0 (106) 11(142) 2 (101)1978 43 (4697) 8(451) 5 (38) 3 (40) 6 (61) 2(58) 15 (57) 13 (129) 13 (1286) 4 (30)1979 59(4135) 12(320) 13 (11) 5(28) 9(45) 6(46) 7(23) 8(160) 9(1152) 8(26)

Rainfed lowland Iloilo 1976 - 0 (10) 0(10) 0 (7) 0 (19) 2 13) 15 (180) 5 (76)1978 0 (54) 0 (3) 0 (17) 9 (64) 4 (78) 3 (73) 0 (121) 0 (191)1979 64 (2350) 16 (84) 6 (17) 0(6) 18 (68) 25 (104)

1981 56 (1322) 18(304) 0(5) 12(51) 18(138)Pangasinan 1976 60 (7 7 )b 62(82) 0(111) 5 (39)

1978 58 (53)b 5(51) Cagayan 1979 59(713) 8 (101) 6 (34) 0(7)

1980 0(11) 4 (28) 15 (286)

62 ( 48(177) 61(140)1981 0(5) 12 (2) 13(45) 14 (92)

Irrigatedlowland Laguna 1974 62 8

1974-75 50 41 1975 43 72 1978 96 (623) 21(24) 33 (60)1979 83(2017) 18(110) 8(62) 11 (89) 22(400)1980 78 (1625) 20(75) 0 (16) 4 (72)1983 60

9 (80) 7 (270) 30

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R F Apostol unpublished Iloilo and Pangasnan 136 Cagayan B Canapi unpublished Laguna 175 P CPantua unpublished 118 234 143 Carino and Shepard unpublished YSB = yellow stem borer ScirpophagaincertulasGFB = gold fringed borer Chilo auricilius PSB = pink stem borerSesamia inferens earcutting caterpillar Alythimna separata swarming caterpillar Spcdoptera mauritia acronyctoides skipper Pelopidas mathias reen horned caterpillar Melanitls ledaismene RLF = rice leaffoldersCnaphalocrocismedinalis and Marasmiaspp brown semilooper Mocisfrugalis Figures in parentheses are sample sizes Egg masses

IRPS No 123 January 1987 35

Table 9 Effect of seeding rate and insecticide on plant density and Yield losses from insect pests in upland rice often areyield of Dagge upland rice Batangas Philippines 1976a comparable to those of lowland rainfed or irrigated rice

Seeding rate Insecticide Dosage Plant density Yield (0-Y30o) but usually from different pests The principal (kg aiha) seed treatment (kg aiha) (noIra-row) (tha) groups are soil pests feeding on sown seed and rootsat 14 DE followed by seedling pests These groups have largely gone

50 None - 35 a 20 a unnoticed leading researchers to conclude that stem borers50 Carbofuran F 10 38 ab 23 a and seed bugs are the most important pests (17) Yield loss 10 beCabofuranF100 None - 4551 c 23 a trials that pinpoint major pests need to includc seed

100 Dieldrin WP 10 52 cd 22 a treatments Granules applied in the furrow may not give150 None - 59 d 22 a adequate protection from many seed pests

aAv of 8 fields DE = days after crop emergence In a column means The floristic and pedological diversity of upland rice followed by a common letter are not significantly different at the environments is matched by the diversity of arthropods5level by D)MRT F = flowable Wi = wettable powder both pests and their natural enemies Irrigation has homoshy

genized lowland rice environments Flooded soils tend tothreatened by epidemics of locusts or armyworms New have the same properties and to eliminate soil pests The technology stressing efficiency has minimized insecticide spatial and temporal dominance of cultivated rice has cost benefited specialist pests

Seed treatment is normally inexpensive and can be Upland rice has no specialist arthropod species The economicallyjustified in -nanycases(66) Treating seed with monophagous pests on upland rice prefer the lowlands systemic insecticide (216) but not chlorinated hydrocarbons where rice evolved Rice in the uplands exists only because (247) is most effective in controlling seedling maggots of man and even though lowland rice predates upland riceFoliar sprays have to be repeated to obtain control and specialist species evolved Wheat maize and sorghum do often are washed off by frequent early season rains not have monophagous pests leading us to conclude that Granular insecticides are effective but required dosages are there is no evolutionary advantage for inssects to have too expensive to justify except under heavy infestation narrow host ranges in upland graminaceae(247) When directed against first-instar larvae banding Pests have adapted to the uplands not only by having granules in seed furrows (177) can be effective against white wider plant host ranges but also by having long life cyclesgrubs at low dosages (025 kg aii ha) (134) Seed treatments undergoing dormancy or being dispersive Each pest has a are not promising for white grub control (248) unique life history strategy suited to highly unpredictable

Seed treatment is also effective ainst ants and may be environments In agivenyear someinsect specics an better cheaper than increasing the seeding rate In Batangas exploit upland rice as a food source but no pest is a senious treating 50 kg seed ha was as effective as increasing the one every year on upland rice seeding rate to 100 kgha and using untreated seeds (Table The low yielding potential of upland rice means less 9) In Brazil seed treatments control termites and other soil quantity and lower quality of food for pests to exploit but insects including the lessercornstalk borer(66) Domiciano also less incentive for farmers to undertake control (58) however found seed treatments inconsistent in their The most effective control would be early and synchroshyeffect against high populations of lesser corn stalk borer nous planting of early-maturing upland rice varieties to

Baiting can be an inexpensive way of controlling ants prevent insect pest buildup Breeding for insect resistance seedling maggot flies mole crickets and field crickets Baits should initially focus on stem borers because they are verymade of locally available material and impregnated with difficult to control by other means Biocontrol efforts insecticide can be sparingly distributed taking adantage of should first stress conservation of natural enemies preshythe pests ability to disperse and encounter bait sites dators and pathogens rather than parasites Efficient

The new low-volume hand-held controlled-droplet chemical control includes seed treatment or baiting for seed sprayers may offer an advantage over high-volume knap- seedling and some root pests and using controlled dropletsack sprayes for controlling foliar pests Upland rice sprayers for spot treatments against foliar pests and seed canopies are more open than those in lowland rice and bugs when economic thresholds are reached better droplet penetration should result Controlled droplet applicators would be ideal for seedbugs and defoliators

CONCLUSION

Upland rice is attacked by a wider array of insect pests than is lowland rice mainly beca use of the addition of soil pests Generally population levels of lowland rice pests are lower in upland than lowland rice leading many people to conclude that insect pests are not important on upland rice

36 IRPS No 123 January 1987

REFERENCES CITED

1Abraham C C and K S R Mony 1978 Occurrence ofLeptocorisa acuta Fabr (Coreidae Hemiptera) as a pest of nutmeg trees J Bombay Nat ttistSoc 74553

2 Agyen-Sampong M 1975 HlierogrlphusdiaganensisKrauss (Orthop-tera Acrididae) a new pest of rice in Northern Ghana Ghana J Agric Sci 8249-253

3 Agyen-Samnpong M 1976 Varietal resistance and other inle-ratcd pest control components as factors of rice varietal improvement in West Africa Prcsented at the Second varietal improvement serinar 14-18 Sep 1976 Bouak Ivory Coast West Africa Rice Development Association Monrovia Liberia 18 p

4 Agyen-Sampong M 1977 Outbreak of Ihccrorrdnisorae in rice seedlings near Rokupr Sierra Leone lit Rice Res Newsl 2(6)16-17

5 Agyen-Sampong M 1980 Dirty panicles and rice yield reduction caused by hugs IntRice Res Newsl 5(l)11-12

6 Agyen-Sampong M 1982 The major pest problems of irrigated upland and mangrove swanip rice ecosystems ir the humid tropical Guinea savanna and Salel climate zones Pages 318-328 in Pro-ceedings training course cn concepts techniques and application of irtegrated pest management in rice inWest Africa 10-20 Jan 1982 Fendall Liberia West Africa Rice l)evelopioent Association Monrovia Liberia 505 p

7 Akibo-Betts V I and S A Raymundo 1978 Aphids as rice pests in Sierra Leone lit Rice Res Newsl 3(6)15-16

8 Akinsola F A 1975 Present status of different rice stemborers in parts of Nigeria iicL Entomo Newsl 328

9 Akinsola FA 1979 Ihe biology and ecology of rice stemborers in Nigeria Ph 1) thesis University ol lbadan Nigeria 217 p

10 Akinsola E A 1979 Guidelines of integrated management of insect pests of upland rice in West Africa Presented at the Seminar on integrated management of rice diseases and insect pests at Bobo-Dioulasso Upper Volta 17-22 Sep 1979 West Africa Rice Develop-ment Association Monrovia Liberia

II Akinsola IA 1984 Insect pests of upland rice in Africa Pages 301-305 in An overview of upland rice research Proceedings of the 1982 lIouak Iwsrv Coast upland rice workshop International Rice Research Institute 13O Box 933 Manila Philippines

12 Alam M S V TJohn and Kaung Zan 1985 Insect pests and diseases of rice in Africa Pages 67-82 in Rice improvement in Eastern Central and Southern Africa proceedings of the international rice workshop at Iusaka Zambia 9-19 April 1984

13 Alticri M A A van Schoonhovcn and J D Doll 1977 The ecological role of weeds in insect pest management systems a review illustiated with bean (Phaseohsvulgaris L)cropping systems PANS (Pest Artie News Summ) 23195-205

14 Anmante F 1967 The leaf-cutting ant Atia capiguaraa pest of pastures [in Portuguese] Biol6gico (Sao Paulo) 33113-120 (Rev Appl Entomol A56207)

15Andrade A C and D Puzzi 1951 Preliminary results of experi-merits on the control of Sculptcoris iastantus on sugarcane [in Portuguese] Bioulgico (Sao Paulo) 1744-49 (Rev Appl Entomol A41421422)

16 Appert J 1970 Maliarplia separaelha (rice stem borer) new observations of entomological problems of rice in Madagascar [in French] Agron Trop (Paris) 25329-334

17 Ariadeau Mand Z Harahap 1986 Relevant upland rice breeding objectives Pages 189-197 in Progress in upland rice research Proceedings of the 1985 Jakarta conference International Rice Research Institute 1O Box 933 Manila Philippines

18Asar S and Kisimoto 1975 Seasonal occurrence of the smaller brown planthopper in upland rice [in Japanese] Proc Kanto-Tosan Plant Prot Soc 2228

19 Bakker W 1974 Characterization and ecological aspects of rice yellow mottle virus in Kenya Meded Landbouwhogesh Wage-ningen 152 p

20 Balut F F 1970 White grubs attacking rice Oryza ativa L [in Portuguese] Biol6gico (So Paulo) 36321-322

21 Banerjee S N and L M Pramanik 1967 The lepidopterous stalk borers of rice and their life cycles in the tropics pages 103-124 in The major insect pests ofthe rice plant Proceedings ofa symposium at the International Rice Research Institute September 1964 The Johns Hopkins Press Baltimore Maryland USA

22 Barrion A T and J A Litsinger 1980 Ants a natural enemy of cliaiarac in dryland rice IntRice Res Newsl

5(4)22-23 23 Barrion A T and J A litsinger 1981 The spider fauna of

Philippine rice agroecosystem 1Dryland Philipp Entomnol 5139-166

24 Barrion A T and 1 A Iitsinger 1984 The spider fauna of Philippine rice agroccosystcms II Welland Philipp Eiitomol 611-37

25 Barrion A T1 and J A Litsinger 1984 lihanus (Diptera Tabanidae) eggs an alternative host of rice stei borer (SI3) egg parasite Telenoonis digns (Iliymenoptcra Scelionidae) lit Rice Res Newvsl 9(6)19

26 Barrion A T and J A Litsinger 1985 Identilication of rice leaffolders by wing markings Int Rice Res Newsl 10( 1)24

27 Barrion A T and J A Litsinger 1985 Chctiosspp (Coleoptera Carabidae) a leaffolder ILF)predator lit Rice Res Newsl 10(1)21

28 Bertels A 1970 Insect pests of rice and theircontrol [in Portuguese] Pelotas Brazil Instituto de Expcrimentacao Agropccuararias do Sul Circular 43 24 p

29 Bonzi S M 1982 Ctilo (flisilineus J de Joannis (Lepidoptera Pyralidae) stem borer of irrigated cereals in Upper Volta LioFrench] Agron Trop Riu Rizic Cult Vivrieres Trop 37207-209

30 Bourke T V T L Fenner J N LStihick G Baker EHassan D F OSullivan and C S li 1977 Insect pest survey for the year ending 30th June 1969 Papta New Guinea )epartment of Agrishyculture Stock and Fisheries Port Moresby 57 p

31 Bowden 1 1973 Migration of pests in the tropics Meded Rijksfac Landbouwwet Gent 38785-796

32 Breniere J 1973 The main rice parasites in West Africa and their control Pages 27-42 in Proceedings seminar on plant protection for the rice crop West Africa Rice Development Association Monrovia Liberia

33 Breniere J 1976 The principal insect pests of rice in West Africa and their control West Africa Rice Development Association Monrovia Liberia 52 p

34 Brown ES 1962 The African arinyworn Spoiolptera eXI)Ina (Walker) (Lepidoptera Noctuidae) a review of the literature Cornshymonwealth Institute of Entomology London UK57 p

35 Brown F B 1968 Upland rice in Latin America FAO-International Rice Commission 12th Session of the Working Party on Rice Production and Protection 2-14 Sept Peradeniya Sri lanka Food and Agriculture Organization Rome Italy 6 p

36 Buyckx F J 1 1962 Disease and insect pests of crops inCongo Rwanda and Burundi [iil French] Publications dc LInstilut National pour lestude Agrononiquedu Congo M Weissenbruck S A Brussels Belgium 708 p

37 Cariiio F 0 1981 Role of natural enentics inpopulation suppression and pest management of green rice lealhoppers Ph D thesis University ofthe Philippines at Los Bafitos PhilippinesLaguna 136 -

38 Carpente A J 1983 The development of rice cultivation and extension of vegetable productin in Zanzibar Final Report Vol II1 October 1983 Food and Agricultme Organization Ronte Italy

39 Carroll C P and S1 Risch 1983 Tropical annual cropping systems ant ecology Environ Manage 751-57

4(0 Cates R G 198 1 Host plant predictability and the feeding patterns of monophagous oligophagous and polyphagous insect herbivores Oecologia (Berl) 48319-326

IRPS No 123 January 1987 37

41 Cendcafia S M and F B Caora 1967 Insect pests of rice in the 62 Ene 1 C 1981 I)iiorldhol)ierusihilis Slater (licteroplera Philippines Pages 591-616 in The najor insect pests ofthe rice plant lygacidue Illissinac) a new rice pest front Nigeria lit Rice Res Proceedings of a symposium at the International Rice Research Newsl N3)15 Institute September 1964 The Johns Hopkins Press Baltimore 63 Fugenio T S 1965 rhe effect oi e-vCgetation on rice production Maryland USA 729 p and the possibility of overcoming it by leaf pruning MS thesis

42 Centro Ihternacional de Agricklittira Tropical 1984 Upland rice in University ol the Philippines at Los Baijos ILaguna Ihilippins 69 p Iatin Anerica Pages 93-119 hi An os ervies of uplaind rice research 04 Farrow R A 1975 The Alrican nmigratory locust ii is main Proceedings of the 1982 Boinakamp Ivory Coast upland rice workshop outbreak area of the middle Niger qlinlitatise studies of solitary International Rice Research Institute P0 Blox 933 Manila populations in relation to environmental factors Locusla No II Philippines 198 p

43 Chandra G 1979 Taxonomy ad hn riononlics ofthe imect parasites of 65 Fennal R G 1943 Food crop investigation Windward and rice Icalloppers and planthoppers in the Pihilippines and their Ieeward Islands First report Report onl work dunring the periodiiiportairce iii natural biol)gical cointrol Philipp Fntoltol 4119-139 April 1942-April 1943 Imperial College of Tropical Agriculture

44 Chang T T 1976 The originii colut ion enlIivation dissemination Trinidad 16 p (Rev of Appl Fintornol A3222-25) and diersificition of Asian and African rices -uphytica 25425-44 1 66 Ferreira F 1983 Integrated pest ritaageiret of upland rice [ill

45 Chatierjee P B 1979 Rice leaffolder attacks in India lit Rice Res Portuguese English summary] Pages 323-341 in Upland riceculttre Newsl 4(3)21 factors affecting production E Ferreira T Yamada and F-

46 Cheng Cbien-pan 19(4 Upland rice in Taiwan International Rice Malmolta eds Institutoda Potassae Foslato Institnto Internacional Cnmmission Wiorking Party oit Rice Production ard Protection I()th da Potassa Piracicaha Sao Paulo lraid Meeting 3-10 March 1964 Manila Philippines Food and Agricul- 67 Ferreira E arid F dl Silva Martins 1979 Occurrence of ture Organiaion Rome Italy 5 p Sogatohs orvziola (Muir) inupland rice in Goias Ilrail lit Rice

47 Chcrrett 1 M and 1) 1Peregrine 1976 A review of the status of Rcs Ncwssl 4(4)18 leaf-cutting ants and their control Ann Appl Biol 84124-128 68 Ferreira E and 1F da Silva Martiris 1984 Insect pestof rice ill

48 Chipeta F M 1974 Pests of rice in Malawi Pages 90-95 in Brazil and their control Jill Portugucsej FMIlRAtPA-CNPAF Proceedings Sith eastern Africa cereals research conference 10-15 Goiania Brmiil 67 p March 1974 DRB Mialda ed Ministry of Agriculture lilongwe 69 Ferreira FI F ca Silha Miartins S S Nero and 1 It P de Malawi Carvallio 1982 M tltiplc regression analysis todeterimine the effect of

49 Chu Yau-i 1979 The d atiaging period of the rice seedling fly insect populatiois ott upland rice yield [inIPortuguese Pesqui (Atlterigotiavxigu Stcin) (NItiscide Diptera) oil cori) at Fast Java Agropecu Bras 17671-675 Indonesia Plant Prot Bill 21403413 70 Ferreira F 1F da Silva Martins and F 1PZimmermann 1979

50 Conklin H C 1957 IHaninoo agriculture a report on aln integral Rice varietal resistance to tle lesser corn stalk borer [in Port uguese] systent of shi fti rig cult ivationi in the Pltilippincs Food and Agriculture Pesqui Agropccu Biras 14317-321 Organization Rome Italy 209 p 71 Fleming W F 1968 Biological control of lie hapanese beetle U S

51 Cramer H II 1967 Plant proection and world crop production Dep Agric RLs Sers [ech Bull 1383 78 p Pilaniensclhutz-Nachr 201-524 72 Food and Agriculture Organiation of the United Nations 1982

52 l)ani R C and Majunidar 1978 Prelininary observations on tie Country report surmimary Pages 27-37 in Report on the first incidence of root aphids on differeit rice cultnrcs Sci Cult 4488-89 FAOIUN)IPhail and regional training coiuirse in iniproved ctltural

53 Davidson R I 1979 lffects of root feeding on foliage yield Pages practices for upland rice 5-23 Oct 1981 Northern Region Agrculturd117-120 in T K Ci osb and R B Pottinger eds Proceedings of the Developririt Center Chiang Mai Food and Agricuilture Organiiashy2nd Australian conlerence on grassland invertebrate ecology Govt tion Rome Italy 37 n Printer Wellington New Zealand 73 Franco do Ariaral S 195 1 Residual action of insecticides in turfy

54 l)ean G 1W 1976 Rice insect pests inlaos Int Rice Res Newsl soil--the results of a field experiment [in Portuguese] Biol6gico (Sao 1(2)15 Paulo) 17209-214 (Rev Appl Entomol A41423)

55 I)ean G J W 1978 Insect pests of rice in Laos PANS 24(3)280-289 74 Freeman ) 1970 Report onithe Iban london School of -coroiics 56 )ingle H 1984 Behavior genes and lifc histories conplex adapta- Monographs on Social Anthropology No 41 Athlane Press

tions to uncertain environments Pages 169-194 in A new ecology London UK 317 p novel approaches to interactive systems P W Price C N Slobod- 75 Fuchs T W1 A Harding and 1W Smithlr 1979 Induction and chikoff and W S Gaud cdsl ohn Wiley New York USA termination oldiapause in the sugarcane borer Ann Flitonlol Soc

57 Dobelmann-JP and M Falais 1968 Upland rice cultivation ill tie Am 72271-274 North-western part of Madagascar in 1966-1967 [in French English 76 (hbriel B1) 1975 A review tf the major insect pests of some tpland summary] Trop Agron 2313-21 crops in tlc Philippines Pages 86-111 in Ad Htoc panel of experts on

58 Doniiciano N L 1979 Chemical control of tie lesser corn stalk pest disease arnd weed problems in some rainfed crops Bangkok (lolsonno~paloIgnowslhts) (Zeller 1848) (I-epid(ptera Ihycitid ae) Thailand Food aid Agriculture Organization Roire Italy

on rice in Ponta Gressa Parara 1977-1978 [in Portuguese] Pages 77 Garg D K and N K Shal 1983 White grub outbreak on rainfed 291-303 in Anais 1 Reuniao de tecnicos cmiirizicultura do estado de dryland rice in Uttar Pradeslh Int Rice Res Ncwsl 81(4)16-17 Sao Paulo-Campinas Cati Brazil 78 Garg I) K and IP Tandon 1982 Major insect pests of rice on hilly

59 Dunsmore 1 R 1970 Investigations on tlie varieties pests and tracts of Uttar Pradesh India lilt Rice Res Newsl 7(11)11-12diseases of upland rice (hill padi) in Sarawak Malaysia Int Rice 79 Garrity 1) P 1983 Upland rice research an international bibliograpy Comm Newsl 1929-35 1975-1982 Iternational Rice Research hnstitnte PO Box 933

60 Elias 1R1965 The new decline of rice seedlings is not caused by Manila Philippines 121 p parasites [in Portuguese] Biol6gico (Sao Paulo) 313-6 80 Ghauri M S K 1982 A new subspecies of Iinorh)rone bull

61 Empresa Brasiliera de Pesquisa Agropecuaria 1984 Upland rice itt cornlus Slater (Hemiptera Iygaeidae) front Papua New Guinea on Brazil Pages 121-134 it Av overview of upland rice research rice and carpet grass Ihll lEntoniol Res 72133-137 Proceedings ofthe 1982 Blouak Ivory Coast Upland rice workshop 81 Gonzales F 0 PArregoces R I ihernandef and OT Parada International Rice Research Institute PO Box 933 Manila 1983 Insect and imite pests of rice arid their control in Latin America Philippines [in Spanish] la Federacion Nacional de Arrocerts (Fedearroz)

Bogota Colombia 60 n

38 IRPS No 123 January 1987

82 Grist D H and R J A W Lever 1969 Pests of rice Longmans London UK520 p

83 Hale P R 1979 Two subterranean pests of upland rice in Papua New Guirnca IntRice Res Newsl 4(2)16-17

84 Hale P R and J T Hale 1975 Insect pests of rice in the New Guinea Islands Rice Entomol Newsl 34

85 Harris W V 1969 Termites as pests of crops and trees Common-wealth Institute of Entomology London UK 41 p

86 Hattori 1 1968 A method to distinguish between grass leaf roller (Ctaplha rocis medinalisGuenee) and the rice leaf roller (Susurtia exigua Butler) [in Japanese] Plant Prot (Japan) 23167-170

87 Henry I E and 1A Onsager 1984 Experimental control of the mormon cricket Iahrosshnph by NJsena locustae(Aficrospora Microsporidae) a protozoan parasite of grasshoppers (Ort Acri-didae) Entomophaga 27197-201

88 Hidaka 1 P Vungsilabulr and SKadkao 1974 Studies on ecology and control of the rice gall midge in Thailand Tech Bull 6 TropicalAgricultural Research Center Tokyo Japan 113 p

89 Hincklcy A 1) 1963 Ecology and control of rice planthoppers in Fiji Bull Entomrol Res 53467 481

90 Hirao J 1982 Ecology and chtInical control of the rice leaffolder Jpn Pestic Inf 4114-17

91 Ho DT 1984 Sten borer incifience in rice ecosystems in KenyaEast Africa Pages 307-320 in A overview of upland rice research Proceedings of the 1982 Bouac Ivory Coast upland rice workshop International Rice Research Institute PO Box 933 Manila Philippines

92 Ho )T J 1Njokah and J GKibuka 1983 Studies on rice sten borers in Kenya with emphasis on ifaliarpiaipiraellRag Insect Sci Appl 465-73

93 Ichihara I T Ito A NIatsuo and M Sawada 1971 Chemical control of the rice striped stem borer in upland rice grown in plastic mulch [iii Japanese] Proc Kanto-Tosan Plant Prot Soc 1882

94 IclhiharaIand A Matsuo 1969 Damage caused by rice striped stett borer to upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1686-87

95 Ichhara I atid A MIatsun11970 Conside ration oft lie extraordinary prevalence of rice striped stem borer in upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1778-79

96 Institut de Recherches Agrolnliqes Iropicale et des Cultures Vivrieres 1978 Insect resistance Pages 12-13 in Upland rice 1978 atnual report Bouak Ivory Coast 27 p

97 International Institute for Tropical Agriculture 1983 Aphids itt rice roots I ITA Res Briefs 4(1-3)4-5

98 International Institite fi Tropical Agriculture 1983 Hopperburn in rice caused by brown plathopper IITA Res Briefs 4(l-3)6-7

99 International Rice Research Institute 1975 Annual report for 1974 PO Box 933 Manila Philippines p 216

100 International Rice Restarch Institute 1976 Annual repiort for 1975 PO Box 933 Manila Philippines p 228

101 International Rice Research Institute 1979 Cropping systems program Component technology development and evaluation Page 404 i Annual report for 1978 11O Box 933 Manila Philippines

102 International Rice Research Institute Department of Entomology 1981) Insecticid evaluation 1979 Tables 73 74 PO Box 933Manila Philipes

103 International Rice Research Institute Department of Entonmology 1981 Insecticide evalomtion 1980 Table 81 POBox 933 Manila Philippines

104 International Rice Research Institute 1981 Report of a meeting of Asian rice-based cropping systems entomologists Eleventh cropping systems working group meeting 18-22 May 1981 PO Box 933 Manila Philippines p 132 133 144

105 International Rice Research Institute 1984 Cropping systems program Pages 368-373 in Annual report for 1983 PO Box 933 Manila Philippines

106 International Rice Research Institutc 1984 Upland rice in Asia Pages 45-68 in An overview of upland rice research Proceedings of the 1982 Bouakamp Ivory Coast upland rice workiiop loternational Rice Research Institute 11O Box 933 Manila Philippines

107 Irshad M 1977 Grasshoppers in rice in Pakistan lit Rice Res Newsl 2(3)8-9

108 Ito R and TOno 1964 Upland lice its culture and varieties in Japan lIternational Rice Cormmission loth Meeting of the Working Party on Rice Production and Protection 3-10 March 1964 Manila Philippines Food and Agriculture Organization Rome Italy 12 p

109 Jeffrey M I and V A Dyck 1983 Effect of moonlight on lightshytrap catches of brown planthopper lit Rice Res Newsl 8(6) 13

110 Jennings P R and T Alicia Pineda 1983 Management ofthe rice plnthoppers Sogatodsorivi(coIl in Latin America Presented at the meeting on stable plant resistance FAO and lIndocsian I)inectorate of Food Crops Protection Bali lndonesia 29 Novcniber-2 l)ccemshyber 9 p

IllJimenez Gomez J A 1979 Rice insect pests i lla ios ()riCntales lil Spanish] Pages 99-103 in Rice ciltivation Instituto (olotibiatto Agropecuario Compndio No 29

112 Jongeleen F JJ Soesanto and A Mockidjo 1979 Studies oti pathogens of two species of Scarabacidac froinIndonesia Z Angew Enlomnol 87412-417

113 Jouvenaz 1) PC S lofgren and W A Baiks 1981 Biological control of imported fire antsa review of cirrent knowledge Bull Entomol Soc Am 27203-208

114 Kainoh Y K Shiniiu S MarUani Y Taiaki 1980 lost finding behavior of tie rice bu lplotori+JchinvittjI)allas (liciniptera Corcidac) with special reference to diet pattern ol aggregation and feeding on the rice plant Appl Entomol Zool 15225-233

115 Kakkar K 0 P Sharma and P I Sharma 1977 Note on Ihteronyrchus liodresRedt (Coleopicra I)ynastidae) as asporadic pest of paddy and maie inHimachal Pradesh Corr Agric 156-57

116 Kalode M131 79 Major insect pest problems and their control in upland rice (Unpublishcd mineogriph CWI oJNaaiat lni Chandaratat (198 1)original not seen)

117 Kalshoven L i 98 I The pests of crops in I ldoniesia (translated by P A vatn der Laan) 1 T Ichtiar B3aru-vati Iloeve lakarta Indonesia 701 p

118 Kamal N Q 1981 Suppression of whitebacked planto ner Sogatella fircaifra (Horvath) aLId rice lealfolder (nhal -cis licdb lis (Guenee) populations by natural cnclmie hlI) tiesis Grcgorio Araneta University Foundation Victoncta Park Metto Manila Philippines 179 Ip

119 Karim A N M Reaul 1975 A species ofliealy bug Trioni i70s ceres new oii rice in the Philippines Rice Entoniol Newsl 247

120 Katanyukul Wand C Chandaratat 1981 Insect pestsof uplaid rice and their control Report firstFAOUNlFi Tliailand training course on improved cultural practices for tplad rice Chiang Mai Thailand 5-23 Oct 1981 Food and Agriculture Organiation Rome Italy 22 p

121 Kern F 1955 lhletwioa(I)uninemts) bhhit Biturm(Coleoptera) a new pest of sugarcane inVenezuela [inSpanish with Etnglish sunmary] Agron Trop (Maracay) 527-29 (Rev Appl Eitoniol A4572)

122 Khan K M and S Ghai 1974 White grubs ald their control in India Pesticides (1Bombay) 819-25

123 Kiritani K 1975 Integrated pest management of rice pests Pages 66-70 itt Proceedings platning workshop ott cooperative hIeld research in pest managementP SMotooka ed East-West Center Honololu Hawaii USA

124 Kisimoto R 1967 Ecology of insect vectors forecasting and chemical control Pages 243-255 in The virus diseases of tle rice plant Proceedings of a symposium atthe International Rice Research Institute April 1967 The Johns Hopkins Press altimore Maryshyland USA

125 Kisimoto R and V A Dyck 1976 Climate and rice insects Pages 367-391 in Climate and rice International Rice Research Institute PO Box 933 Manila Philippines

126 Kureha S S Kobayashi SAsari and T Ishihara 1977 Life history and control of the mole cricket Grrllotalpa africana Palisot de Beauvois [in Japanese English translation 7520 1RRI library] Proc Kanto-Tosan Plant Prot Soc 21192-197

127 Kushwaha K S 1981 The black beetle as a pest of rice in Haryana India Int Rice Res Newsl 6(4)14

128 Lawani S M 1982 A review of tile effects of various agronomic practices on cereal stem borers Trop Pest Manage 28266-276

129 Leuck D B 1966 Biology of the lesser cornstalk borer in South Georgia J Econ Entomol 59 797-801

130 lever R J A W 1955 Insect pests of the rice plant in Malaya Int Rice Comm Newsl 1612-21

131 Li Ching-Sing and Shin-Foon Chiu 1951 The study ofthe rice gall midge Iachydiplosis orvza Wood Mason [in Chinese with English summary] 1Taiwan Agric Res 21-13

132 Iitsinger J A 1984 Assesament of the need-based application for rice Proceedings TechnologyTransfer Workshop 15-16 March 1984 International Rice Research Institute 1PO Box 933 Manila Philippines 6 p

133 Litsinger J A 1984 Environmental diversity and insect pest abundance with reference to the Pacific Pages 285-307 in Proceedings sub-regional training course on methods of controlling diseases insects and other pests of plants in the South Pacific 4-20 Oct 1982 Vaini Kingdom of Tonga Nukualofa

134 Litsinger J A RF Apostol and M BObusan 1983 White grub Leucopholis irrorata (Coleoptera Scarabaeidae) pest status popula-tion dynamics and chemical conllol in arice-maize cropping pattern in the Philippines J Econ Entoinol 761133-1138

135 Litsinger J A C de la Cruz F Raymundo A T Barrion M D Lumaban J P Bandong M S Venugopal F Paragna and E Balete 1982 Insect pests and insecticide reponse of dry-seeded bunded rice Pages 179-193 in Report of workshop on cropping systems research in Asia International Rice Research Institute Los Bafos Philippine

136 Litsinger J A M 1) Luinaban J P Bandong A T Barrion W Katanyukul and N Panda 1978 Insect pest management in rainfcd lowland rice agroecosystms of tropical Asia International Rice Research Conference 17-21 April 1978 International Rice Research Institute Los Bafios Philippines

137 Litsinger J A and K Moody 1976 Integrated pest management in multiple cropping systems Pages 293-317 in R I Papendick P A Sanchez and G B Triplett eds Multiple cropping American Society of Agronomy Madison Wisconsin USA

138 Loevinsohn M E 1984 The ecology and control of rice pests in relation to the intensity and synchrony of cultivation Ph D thesis University of London 36) p

139 Loevinsohn M E J A Litsinger N Panda M A Austria R F Apostol C Andrion and J Bandong 1982 A basis for predicting rice insect pest abundance in cropping systems Paper presented at the 1982 International rice research conference on rice and cropping systems 19-23 April 1982 International Rice Research Institute Los Bafios Iaguna Philippines 25 p

140 Lopez A W 1930 Repot of the Entomology Department Pages 145-172 in Annual report of the Philippine Sugar Association 1929-30 Manila

141 Lopez AW 1931 Entomology Department Pages 227-266 in Annual report of the director or research 1930-31 Philippine Sugar Association Manila

142 Lopez A W 1932 Entomology Deparment Pages 252-279 in Annual report of the Philippine Sugar Association 1931-32 Manila

143 Lungibill P and G G Ainslee 1917 The lessercornstalk borer US Dep Agric Agric Res Serv Tech Bull 539 27 p

IRPS No 123 January 1987 39

144 McGuire J U Jr and BS Crandall 1967 Survey of insect pests and plant diseases ofselected food crops of Mexico Central America and Panama U SDepartment of Agriculture International Agriculshytural Development Service Agricultuial Researrh Service October 1967 11 p

145 McNeill Sand T R ESouthwood 1978 The role of nitrogen in the development of insectplant relations Pages 77-9 in lliochemical aspects of plant and animal coevolution J B Ilarbornu ed Academic Press London UK

146 Maebara H 1955 Spatial distribution of lastinstiarlanaeof purplish stem borer (Sesamia iqferens Walker) hibernating in an upland rice field Bull Kagoshima Univ Fac Agric 438-41

147 Mallick E H and N G Hajra 1977 Response of rice plant to defoliation treatments at different developmental stages Indian Agric 2151-56

148 Mansingh A 1971 Physiological classification of dormancies in insects Canl Entomol 103983-10)9

149 Martell C G 1974 Catalogue of plant feeding insects of Mexico [in Spanish] Fitofilo 27(69)1-176

150 Martinez 0 J 1982 Principal rice pests in Ilanos orientalis [in Spanish] P~ages 80-94 in Curso deact nalization de conocimientos en arroz Villaniencis Instituto (olonmbiano Agropecuario

151 Martins J F da Silva F Ferrcira A S Prabhu F 1 P Zimmermann 1980 Use of pesticides for the control of the main subterranean upland rice pests esqui Agropecu Bras 1553-u

152 Martins I F da Silva F Fe eira rod BdamSilveira Pinheiro I Q2 Damage simulation caused by defoliators of upland rice [in Portuguese with English summary] Pesqui Agropecu Bras 171113-1119

153 Martins 1F da Silva N(iyuen van Tan and 1Ida Silveira Pinheiro 1981 Stem borer resistance in upland rice associated with plant morphological characters [in Portuuce Iesqui Agropecu Bras 16187-192

154 Masaki S 1980 Sumnmer diapause Ann Rev Entomiol 251-25 155 Masuzawa T 11 Suwa and F Nakasoji 1983 l)ifkleences of

oviposition preferencc and survival rate of two skipper butterflies Parnara guttata and PIchlpidai iathiais (Iepidoptera HIesperiidac) on rice plant and cogon grass New Entomol 3247-56

156 Maurya D M and C P Vaish 1984 Upland rice in India Pages 69-92 in An overview oftpland rice research Proceedings ofthe 1982 Bouakamp Ivory Coast upland rice workshop International Rice Research Institute IO Box 933 Mnila Philippines

157 Menon M G R R N Katiyai S Kuar and S S Misra 1970 Termites attacking the paddy crop Sci Cult 36294-295

158 Miyahara Y and 1 Nishikubo 1967 Behavior of young nymphs of the red rice root aphid Iholahsipiun ri liabhdontinalisSasaki in relation to movement from the leaf to the roots of upland rice [in Japanese] Pioc Assoc Plant Prot Kyushu 1363-66

159 Mo ljoa Tjien 1952 White grub of upland rice (Holotrichiahelleri Brsk) [in Indonesian with English summary] Contrib Agric Res Stn Bogor No 13134 p

160 Mochida 0 and T Suryana 1979 The occurrence of the brown planthopper and ragged stunt on lowland and upland rice ill Central Java and logyakarta in tile late of January 1979 Entomnolgy Survey Report No 15 Central Research Institute of Agriculture (CRIA) Sukamandi Indonesia I February 1979 9 p

161Monet C 1979 Varietal resistance to upland rice pests [inI French] Pages 35-39 in Annual report 1977 Institut des Savanes Department des Cultures Vivrieres Bouak6 Ivory Coast

162 Monte 0 1942 A caterpillar ofrice-fields[in Portuguese] Biol6gico (Sao Paulo) 8161-163 (Rev Appl Entomol A31414)

163 Munroe I) D 1979 Pest management strategy for hill rice in four resettlement areas in Sarawak Malaysia Pages 89-102 i1 Guidelines for integrated control of rice insect pests Food and Agriculture Organization Plant Production and Protection Paper 14 Rome Italy 115 p

40 IRPS No 123 January 1987

164 Nakasuji F M Ishii and Jen-Zon Ito 1984 Rice skippers in Taiwan and their life histories ZZrC (Taipei) Tech Bull 82 10 p

165 Navas I) 1976 Fall armyworum in rice Pages 99-106 il Ecological animal control by habitat management Proc Tall Timbers Con-ference 28 Feb-I Mar 1974 Fall Timbers Res Stn Misc Publ 6 Tallahassee Horida USA

166 Nickel 1 1 1961 Biological control of rice stem borer A feasibility study Tcch Bull 2 International Rice Rcarch Institute PO Box 933 Manila Philippines IIl p

167 Nigan P M and R A Vernia 1985 Insect pests of uplaild rice in Uttar Pradesh Int Rice Res Newsl 10(6)22

168 Nilakhe S S 1985 -cological observations on spittle bugs with emphasis on their occirence in rice Pcsqtui Agropecu Bras 20407-414

169 Nixon G EJ 195 1 lie association of ats with aphids and coccids Commonwealth Iusti lcte of Entonology Iondon UK 36 p

170 Nurdin F an Ishak Manti 1985 Study oit seedling fly of upland rice Interlational Upland Rice Moitoring Tour Report 26 lebruary-2 MarLI Sukaranti West Sumatra

171 Oka 1 N 1979 Ciltural conttol of the brown planthopper Pages 357-369 in Brown planthopper threat to rice production in Asia International Rice Research Institutc 1p0 Box 933 Manila Philippines

172 Ohni M and A Vilnaii 1975 Rice insect pests of Italy biology of Sipha grcria (Kaltenbach) and its control [in Italian] Riso (Milan) 2459-71

173 Otanes F Q atnd 1 I Sison 194 1 Pests of rice Philipp J Agric 12211-259

174 Painter R I 1951 Insect icsistance in crop plants University Press of Kansits Lawrence Kansas USA 520 p

175 Pantua P C and 1A litsinger 1981 Arthropod abundance in continuous vs biannual rice cropping systeis Int Rice Res Ncwsl 6(2)20-21

176 Pathak M ) and V A l)yck 1975 Sttudics ott insect pests of upland rice Pages 186-191 in Major research in upland rice International Rice Research Institute PO Box 933 Ilanila Philippines

177 Patil B R B M IHasabe and 1) S Ajri 198 I Chemical control of white gritb lIditriihinhorraia Fabr intfesting groundntut paddy atnd sorghum Indian 1Entomol 43232-236

178 Pawar A 1) 1974 (ollection and identification of insect pests of rice rlerminal repirt lInternational Rice Research Institute PO Box 933 Manila Philippines 72 p (tnimco)

179 IerrajU A aid A S Reedy 1965 Observations ott diapausc itt tie steti borer ohrice Rice News Teller (lai-Mar)9-13

180 Pont A C 1972 A review of the Oriental species of theri ona Rondani (I)iptera Muscidac) of econtomic importance Pages 27-102 in Control of sorglium shoot fly M G lomtani and W R Young eds Oxford atd IH1 Ptthl Co New I)clhi India 324 p

181 Prasad S C 1PChand aid M 1PSingh 1983 Incidence of rice gall midge in the upland plateau region of Hihar Oryza 20235-236

182 Rai P S 198 0 Nisaga simpex damage to rice in the hill tracts of South India Int Rice kles News[ 5(I)12

183 Rathorc Y S 1969 Gonocplhahli dcprssunt Fabr as itpredator of (tilo zonwIlhis (Swinhoc) and other moths in field cage at Patnagar Uttar Pradesh Indian 1Entotnol 3127

184 Reed W 1983 Crop losses causcd by insect pists in the developing world Proceedings 10th Internatial Congress of Plant Protection lrighton UK 174-80

185 Reedy M V 1981 Ateactogasterinhens(Walker)a new root-grub pest of tipland paddy and inaize Curr Sci 501076

186 Reveno M A 1968 Catalogue of the insect pests of economic crops in Colombia [in Spaiish] Asociacion Latino Americana de Entomo-logia (AIAE) 129-33

187 Reyes G M 1957 Rice dwarf disease in the Philippines Food and Agriculture Organiiation Plant Prot Bull 617-19

188 Rice S E A A Grigarick and M 0 Way 1982 Effect of leaf and panicle feeding byarmyworm (Lepidoptera Noctuidae) larvae on rice grain yield Environ Entomol 7593-595

189 Risch S J 1) Andow and M A Altieri 1983 Agioccosystem diversity arnd pest control data tentative conclusions and nizw research dirctions Environ Entomol 12625-629

190 Rivera C T 1956 Three organic insecticides in thecontrol of insects affecting upland rice Philipp Agric 39465-472

191 Roffrey 1 1972 Locusta outbreaks in the Philippines Acrida 1177-188

192 Rojas R 1) 1941 The effect of pruning at varying ages upon the yield of Ramairio Phipp Agric 29851-860

193 Roonwal M i 1979 Ternite life and termite control in tropical South Asia Scientilic Pnbl odhpr India 139 p

194 Rosetto C1 F da Silva Martins N C Schmidt and L EAzzini 1978 Damage caused by pasturespittle bugs 4Dvoi)lavopictaandDJ schach) on rice lBragantia Nota 6 (Matcli)25-27

195 Rosztto C1- S S Neto I- Amantc I) Iink 1Garcia-Vieira D M de SoLza N V Bansa t and A M de Olivcira 1971 Rice pests of Brazil [in Portuguese] Pages 149-238 inrFA()-International Rice Commission 2nd meeting of the Rice Committee for the Americas 6-11 l)ecember 1971 Pelotas Rio Grande do Sul Brazil

196 Rothschild G H L 1969 Obseratiois on the armyworm Spodoptera matritia acronrctoiiles Gn (Lep Noctuidae) Bull Entomol Res 59143-160

197 Rothschild G 1- L 1974 Parasites of the armworm Spolopiera mauritiaacroncitoihes in Malaysia Fntornopliaga 19293-299

198 Sachan i N S K Gangwar and 1N Katujar 1980 Field crickets damaging cages in North Eastern IHill Region Indian J Entomol 42526-527

199 Sands 1) P A 1977 The biology and ecology of Lepocorisa (Hemiptera Alydidac) in Papua New Guinea Res Bull 18 Departshyment of Primary Industry Port Moresby Papua New Guinea 104 p

200 Sarawak 1968 titomology division Pages 97-106 in Report of the research branch of the Department of Altriculture 1966 Kuching (Rev AIppl Eintomol A56603)

201 Satpathy JM M S Das and K Naik 1977 Effect of multiple and mixed cropping on the incidence of scine important pestsI Entomol Res 178-85

202 Sauer IHF G 1939 Notes Ott Elasmpnlpualtsligniosellths Zeller (Lep Pyr) a serious pest ofcereals in Sao Paolo [in Portuguese] Arq Inst Biol 10199-206 (Rev Appl Entotnol A28378)

203 Satinders 1L A 1 SKing and C L Vargas S 1983 Crop pests of Central Anerica with t bibliography [in Spanish] Centro Agroshynotnico Tropical de Investigacion y Ensenanza Turrialba Costa Rica (CATIE) Tec Bol 9 90 p

204 Sauphanor B 1985 Some factors of upland rice tolerttce to stem borers in West Africa Inseu Sci Appl 6429434

205 Sen P atnd S Chakravorty 1976 Stentflics in Haritighata Nadia West Bengal India [it Rice Res Newsl 1(12) 15

206 Senapai B and 1M Satpathy 1983 The seedling fly Atherigona orrzae Malloch is a pest of rice in Orissa (India) J Entomol Res 777-79

207 Shhjahan M S Alatn M Ah nad and BN Islam 1983 A new scarabaeid beetle pest of dryland rice in Bangladesh Int Rice Res Ncwsl 8(I)16-17

208 Sharma G L S C Modgol and R C Gautam 1979 Effect of depth of submergence ott incidence of bacterial blight and white grub infestation in transplanted rice Int Rice Res Newsl 4(6)12-13

209 Sharma S 1985 Scarabaeid beetle outbreak itt upland rice Int Rice Res Newsl 1(1(l)24

210 Shiraki T 1917 The paddy borer Schoenobhits incertulas Walker [achaku Agric Exp Stn Formosa 256 p

211 Silva A de Brito and 11P Magalhaes 1981 Insect pests of rice [in Portuguese] Circular Tecnica No 22 EMBRAPA Centro de Pesquisa t ecuaria de Tropico Unido (CPAJU) Belem Brazil 14 p

212 Singh S A 1979 Emergence of rice leaf butterfly as a rice pest in Manipur India int Rice Res Newsl 4(4)16-17

213 SinghS A 1983 Changing trends of insect pestsof rice in Manipur Int Rice Res Newsl 8(f)12

214 Siwi S S and P H van Doesburg 1984 Lenovorisa Latreille in Indonesia (Heteroptera Coreidae Alydinae) Zool Meded (Leiden) 58117-129

215 Soejitno J 1977 Reaction of some upland rice varieties toseedling fly eltherikonaexigua Stein [in Indonesian] Siri Hana I Penyakit 630-35 Lembga I osat Fenclilian Pertanian Bogor Indonesia

216 Socnardi 1967 Insect pests of rice in Indonesia Pages675-683 in The major inse pests of thu rice plant Proceedings of a symposium at the Internatital Rice Rec Wi itt S pternber 1964 The Johns Hopkins Press Baltimore Maryland USA

217 Soto P E A T Petcz and I W Buddenhagen 1976 Survey of insect pests and diseases of rice in different ecological zones in Nigeria Rice Entomol Ncwsl 435-46

218 Soto 11E and Z Ziddiqi 1976 Screening for resistance to African rice insect pests Varietal improsetnent sCmi n r Bouak Ivory Coast 13-17 Sept 1976 West Africa Rice l)evelopment Association Monrovia Iiberia

219 Squire F A 1972 Entomological problems in Bolivia PANS (PestArtic News Sumr) 18249-268

220 Stapely 1 II 1978 The rice leaf roller in the Solomon Islands Int Rice Res Newsl 3(l)13

221 Suenaga H 1937 On aphids attacking the roots of rice Oyo-I)oubts Zasshi 9150-152 [in Japanese] (Rev Appl Entomol A25745)

222 Suenaga H 1952 On the seasonal abundance of three subterranean aphids on upland rice Bull Kyushu Agric Exp Sti 1249-262

223 Sutynso Yas and Dandi Soekaria 1964 Beberapa hana yang ttenperang tana man padi tanah kering [in Indonesian] Rapat kerja Padi Tanah Kering 6 sd 8 Juily 1964 Kaliurang 9 p

224 Tallamy I) W and R F Denno 1983 Alternative life history patterns in risky environments in example from lace bugs Pages 129-147 in Insect life history patterns R F l)cnio and If Dingle eds Spring-Verlag 225 p

225 Tamura 1 1939 Observations on some insect pests of field crops [in Japanese] 1 Plant Prot 26414-420 (Rev Appl EntomolA27-556)

226 Tamura and Suzuki 1964 Influences of the early-transplanting practice in rice culture upon development of the rice stem borer Chilo siql)ressalisWalker Bull Ilokuriku Natil Agric Exp Stn 761-94

227 Tan J P 1924 The rice root aphid Drowhi ihirsoia Philipp Agric 13277-288

228 Tanaka 1 1957 Taxonomy and distribution of some subterranean aphids injurious to upland rice in Japan with description of a new species [in Japanese English summary] Botyu-Kagaku 22168-176

229 Tanaka T 1961 The rice root aphids their ecology and control Spec [full College of Agriculture Utsunomiya University Japan F3 p

230 Tauber M IC A Tauber R Nechols and 1Obrycki 1983 Seasonal activity of parasitoids control by external internal and genetic factors Pages 87-108 itt I)iapause and life cycle strategies in insects V K Brown and I Ilodek eds )r W Junk The Hague Netherlands

231 Taylor W F 1972 Effects of artificial defoliation (simulating pest damage) on varieties of upland rice Exp Agric 879-83

232 feotia T P S and S Natud 1966 Biotoomics of the rice skipper Parnaraniathiis Fabricius (Lepidoptera Hesperiidac) Indian J Entomol 28181-186

233 Teran F 0 1971 Rice pests of Santa Cruz [in Spanish Ministerio de Agricultura Sita Cruz Bolivia Bull 45 8 p

234 Than Itun 1976 Population dynamics of the yellow rice borer Tryporza itcertulas (Walker) and its damage to the rice plant MS thesis University of the Philippines at Ios Bafios Laguna Philippines 92 p

235 Uichanco L B 1911 Pests Pages 55-74 it Rice Philipp Agric 11-200

IRPS No 123 January 1987 41

236 van der Laan P A 195 1 Life history of the rice seedling fly Contrib Gen Agric Res Stn Boor Indonesia 118 15 p

237 van der Laan 11A 1959 Correlation between rainfall in the dry season and the occurrence of the white rice borer (Scirpolhaga innotala WIk) in lava Entomol Fxp Appl 212-20

238 Van Driesche R G 1983 Meaning of percent parasitismin studies of insect parasitism Fnviron Entomol 121 I1- 122

239 Venugopal M S and A litsingcr 1984 Fffect ofcarbofuran on rice growth Prot Ecol 7313-317

240 Vietnam Directorate of Rural Affairs 1962 Annual work progress report on crop improvement program of rice sugarcane vegetable and field crops (lor the period from July 1961 to June 1962) (11) 299 p (Rev Appl Entomol A51526-527)

241 Wan M T K 1972 Observations on rice leaf- and planthoppers in Sarawak (Malaysian Borneo) Malay Agric 148308-335

242 West Africa Rice I)esclopnient Association 1978 Annual research report Vol 2 Statistics Monrovia Liberia

243 West Africa Rice Development Association 1979 Annual research report p47 Monrovia Liberia

244 Williams ) F and CS lofgren 1981 Fli I illy l-468 anew bait toxicant for control of the red imported lire at Fla Entomol 64472477

245 Williams I) 1 1970 The mealybugs OFlomoptera Coccoidea Iscudococcidac) of sugar-cane rice and sorghum Bull Entomol Res 0109-188

246 Williams D) 1 1982 Pulinaria icervi (Signoret) and its allies ott sugar cane and other grasses Btll Entoinol Res 72111-117

247 Wirjosuiardjo S 1976 Study ofchenmical cont rot of insect pests ott upland rice Imiu Pertanian (Jogjakarta) 1323-329

248 Wirjosuhardjo S and N Wirarta 1976 Chemical control of the white grub tIolorithiatlltriin upland rice in M uneng Irobolinggo Indonesia) [in Indonesian] Imut Pertanian (Jogjakarta) 1330-332

249 Yano K T Miyake and V F Eastop 1983 The biology and economic importance of rice aphids (Henuiptera Aphid idae) a review Bull Entomol Res 73539-566

250 Yelp D and P Rhodes 19(4 lhe success of drilled dryland rice iii the Fiji Islands International Rice Commission Working Party on Rice Production and Protection Manila Philippines 3-10 March1964 Food and Agriculture Organization Rome Italy 13 p

251 Young W R 1981 Fifty-five years of research on the sorghum shootfly Insect Sci Appi 23-9

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Page 33: UPLANDRICE. INSECTl--1i M T -E!R.EOLOU t RTANCE ,AND …delphacid.s3.amazonaws.com › 6958.pdf · The significance of abiotic factors in upland rice insect ecology will be apparent

IRPS No 123 January 1987 33

Table 7 Parasitization rates of rice planthoppers and green leafhopper in upland rainfed lowland and irrigated lowland environments Philipshypines 1976-840

Environment and site Year Egg parasitization () Nymphaladult parasitization ()

BPH W13P GLH BPII WBPH GLIH

UplandBatangas

Rainfed lowland

1976 1979 1980 1982

15 17 12

2(47)8(39)

4(51)

9(1800)14 (212)

12(304)

8 (101)6 (127)

9(116)

Iloilo

Cagayan

1976 1977 1978 1981 1980 1981 1982

0 (69)

11

44 48 40

-

-

46 55 44

0 (471)

11

52 56 49

0 (200)9 (214)

10 (1259)8(235)6 (900) 3(800)

-7 (462)

11 (270)3 (313)5 (900) 1 (800)

4 (600)10 (725)10 (362)12(870)5 (900) 3 (800)

Pangasinan

Irrigatedlowland

1984 1976 1978 1979 1982

13 (436) 13(379) 14 (457)

6(14)7 (197)

17(198) 11 (486) 18 (275)

3 (81)10 (274)3 (116)

16(871) 19(88)

28(211)9 (764)5 (647)7 (1204) 8(305)

Laguna 1977 11 7 21 1978 31 1979 14 4 4 28 1980 7 8 20 1981 1983

56 7(278)

26 16 (1650)

16 27(1921)

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R I Apostol unpublished 1lo1lo and Pangasinan 136 Cagayan BCanapi unpublished Liuna 175 I C Pantua unpublished 118 234 43 Carino and Shepard unpublished BPH = brown planthopper Nila parvara htgens W131Pi =whitebacked planthopper Sogatelia furcifera GLH = green leafhopper Nephotettrix virescens Figiires in parentheses are sample sizes

Tauher et al (230) reported that internal external and genetic factors influence the seasonal activity of parasitoids

Hopper egg parasites but not nymphaladult parasites were also parlicularly abundant at the Cagayan site possibly the egg parasites have more alternative hopper prey in the weedy areas than nymphaladult paisites

The seasonal dynamics of hopper egg parasites can be seen for upland rice in Batangas(Fig I 2) and for rainfed lowland lice in Pangasinan( 135)and Cagayan( 105) Levels of BPHWBPI-L and GLH egg parasitization were corn-parable in upland and irrigated lowland environments Except for some high rates of paraMsitization early in the season the parasitization rate tended to be steady in all envilonments However hopper nymphaladult parasi-tizution levels were more similar in the rainfed environments and lower than in irrigated rice

Overall parasites do not appear highly effective by themselves against upland or even lowland pests Among natural enemies parasites have attracted the most attention because they can be more readily assesscd

Predators and pathogens lend themselves better to management practices Predators the most important group of natural enemies in riccfields are difficult to quantify On the other hand pathogens appear less im-portant but can be readily cultured and disseminated

The natural enemy commulnity of upland rice is rich in species and differs significantly from that of lowland rice

These beneficial organisms must be conserved by applying insecticides judiciously particularly sprays Seed or soil placement of chemicals minimizes exposure to natural enemies The strategy to derive the greatest benefit from natural enemies isto allow the greatest number of beneficial species to thrive Then perhaps at least one species will be effective against each pest at any tine overcoming the variable abundance of each species within and between years

Programs to introduce exotic species or mass produce indigenous ones are ambitious and expensive because they require trained people on a sustained not ad hoc basis

Chemical control Insecticides offer rapid and efficient control of upland insect pests (190) but-should be used only after other control measures have been considered Insecticides are rarely used on upland rice because of cost (32) Upland rice yields are normally too low to justify the expenditure Also most upland rice is grown as a cash crop Spraying upland rice is more difficult than lowland rice because water is less accessible Government subsidy programs to provide pestishycides to tribal peoples for upland rice production have created problems of toxicity to humans because isolated people have had no experience to be able to handle pesticides safely

However ins-eticide use has beenjustified when a crop is

0Z

Table 8 Parasitization rates of lepidopterous pests in upland rainfed lowland and irrigated lowland environments Philippines 1976-84a

Parasitization (5)

Environment and site Year YSB GFB PSB Earcutting Swarming Skipper Greenborned RLF Brown larva larva caterpillar caterpillar caterpillar caterpillar larva semilooperEgg Larva larva larva larva larva larva

UplandBatangas 1977 - 0 (57) - 0 (63) - 0 (71) - 0 (106) 11(142) 2 (101)1978 43 (4697) 8(451) 5 (38) 3 (40) 6 (61) 2(58) 15 (57) 13 (129) 13 (1286) 4 (30)1979 59(4135) 12(320) 13 (11) 5(28) 9(45) 6(46) 7(23) 8(160) 9(1152) 8(26)

Rainfed lowland Iloilo 1976 - 0 (10) 0(10) 0 (7) 0 (19) 2 13) 15 (180) 5 (76)1978 0 (54) 0 (3) 0 (17) 9 (64) 4 (78) 3 (73) 0 (121) 0 (191)1979 64 (2350) 16 (84) 6 (17) 0(6) 18 (68) 25 (104)

1981 56 (1322) 18(304) 0(5) 12(51) 18(138)Pangasinan 1976 60 (7 7 )b 62(82) 0(111) 5 (39)

1978 58 (53)b 5(51) Cagayan 1979 59(713) 8 (101) 6 (34) 0(7)

1980 0(11) 4 (28) 15 (286)

62 ( 48(177) 61(140)1981 0(5) 12 (2) 13(45) 14 (92)

Irrigatedlowland Laguna 1974 62 8

1974-75 50 41 1975 43 72 1978 96 (623) 21(24) 33 (60)1979 83(2017) 18(110) 8(62) 11 (89) 22(400)1980 78 (1625) 20(75) 0 (16) 4 (72)1983 60

9 (80) 7 (270) 30

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R F Apostol unpublished Iloilo and Pangasnan 136 Cagayan B Canapi unpublished Laguna 175 P CPantua unpublished 118 234 143 Carino and Shepard unpublished YSB = yellow stem borer ScirpophagaincertulasGFB = gold fringed borer Chilo auricilius PSB = pink stem borerSesamia inferens earcutting caterpillar Alythimna separata swarming caterpillar Spcdoptera mauritia acronyctoides skipper Pelopidas mathias reen horned caterpillar Melanitls ledaismene RLF = rice leaffoldersCnaphalocrocismedinalis and Marasmiaspp brown semilooper Mocisfrugalis Figures in parentheses are sample sizes Egg masses

IRPS No 123 January 1987 35

Table 9 Effect of seeding rate and insecticide on plant density and Yield losses from insect pests in upland rice often areyield of Dagge upland rice Batangas Philippines 1976a comparable to those of lowland rainfed or irrigated rice

Seeding rate Insecticide Dosage Plant density Yield (0-Y30o) but usually from different pests The principal (kg aiha) seed treatment (kg aiha) (noIra-row) (tha) groups are soil pests feeding on sown seed and rootsat 14 DE followed by seedling pests These groups have largely gone

50 None - 35 a 20 a unnoticed leading researchers to conclude that stem borers50 Carbofuran F 10 38 ab 23 a and seed bugs are the most important pests (17) Yield loss 10 beCabofuranF100 None - 4551 c 23 a trials that pinpoint major pests need to includc seed

100 Dieldrin WP 10 52 cd 22 a treatments Granules applied in the furrow may not give150 None - 59 d 22 a adequate protection from many seed pests

aAv of 8 fields DE = days after crop emergence In a column means The floristic and pedological diversity of upland rice followed by a common letter are not significantly different at the environments is matched by the diversity of arthropods5level by D)MRT F = flowable Wi = wettable powder both pests and their natural enemies Irrigation has homoshy

genized lowland rice environments Flooded soils tend tothreatened by epidemics of locusts or armyworms New have the same properties and to eliminate soil pests The technology stressing efficiency has minimized insecticide spatial and temporal dominance of cultivated rice has cost benefited specialist pests

Seed treatment is normally inexpensive and can be Upland rice has no specialist arthropod species The economicallyjustified in -nanycases(66) Treating seed with monophagous pests on upland rice prefer the lowlands systemic insecticide (216) but not chlorinated hydrocarbons where rice evolved Rice in the uplands exists only because (247) is most effective in controlling seedling maggots of man and even though lowland rice predates upland riceFoliar sprays have to be repeated to obtain control and specialist species evolved Wheat maize and sorghum do often are washed off by frequent early season rains not have monophagous pests leading us to conclude that Granular insecticides are effective but required dosages are there is no evolutionary advantage for inssects to have too expensive to justify except under heavy infestation narrow host ranges in upland graminaceae(247) When directed against first-instar larvae banding Pests have adapted to the uplands not only by having granules in seed furrows (177) can be effective against white wider plant host ranges but also by having long life cyclesgrubs at low dosages (025 kg aii ha) (134) Seed treatments undergoing dormancy or being dispersive Each pest has a are not promising for white grub control (248) unique life history strategy suited to highly unpredictable

Seed treatment is also effective ainst ants and may be environments In agivenyear someinsect specics an better cheaper than increasing the seeding rate In Batangas exploit upland rice as a food source but no pest is a senious treating 50 kg seed ha was as effective as increasing the one every year on upland rice seeding rate to 100 kgha and using untreated seeds (Table The low yielding potential of upland rice means less 9) In Brazil seed treatments control termites and other soil quantity and lower quality of food for pests to exploit but insects including the lessercornstalk borer(66) Domiciano also less incentive for farmers to undertake control (58) however found seed treatments inconsistent in their The most effective control would be early and synchroshyeffect against high populations of lesser corn stalk borer nous planting of early-maturing upland rice varieties to

Baiting can be an inexpensive way of controlling ants prevent insect pest buildup Breeding for insect resistance seedling maggot flies mole crickets and field crickets Baits should initially focus on stem borers because they are verymade of locally available material and impregnated with difficult to control by other means Biocontrol efforts insecticide can be sparingly distributed taking adantage of should first stress conservation of natural enemies preshythe pests ability to disperse and encounter bait sites dators and pathogens rather than parasites Efficient

The new low-volume hand-held controlled-droplet chemical control includes seed treatment or baiting for seed sprayers may offer an advantage over high-volume knap- seedling and some root pests and using controlled dropletsack sprayes for controlling foliar pests Upland rice sprayers for spot treatments against foliar pests and seed canopies are more open than those in lowland rice and bugs when economic thresholds are reached better droplet penetration should result Controlled droplet applicators would be ideal for seedbugs and defoliators

CONCLUSION

Upland rice is attacked by a wider array of insect pests than is lowland rice mainly beca use of the addition of soil pests Generally population levels of lowland rice pests are lower in upland than lowland rice leading many people to conclude that insect pests are not important on upland rice

36 IRPS No 123 January 1987

REFERENCES CITED

1Abraham C C and K S R Mony 1978 Occurrence ofLeptocorisa acuta Fabr (Coreidae Hemiptera) as a pest of nutmeg trees J Bombay Nat ttistSoc 74553

2 Agyen-Sampong M 1975 HlierogrlphusdiaganensisKrauss (Orthop-tera Acrididae) a new pest of rice in Northern Ghana Ghana J Agric Sci 8249-253

3 Agyen-Samnpong M 1976 Varietal resistance and other inle-ratcd pest control components as factors of rice varietal improvement in West Africa Prcsented at the Second varietal improvement serinar 14-18 Sep 1976 Bouak Ivory Coast West Africa Rice Development Association Monrovia Liberia 18 p

4 Agyen-Sampong M 1977 Outbreak of Ihccrorrdnisorae in rice seedlings near Rokupr Sierra Leone lit Rice Res Newsl 2(6)16-17

5 Agyen-Sampong M 1980 Dirty panicles and rice yield reduction caused by hugs IntRice Res Newsl 5(l)11-12

6 Agyen-Sampong M 1982 The major pest problems of irrigated upland and mangrove swanip rice ecosystems ir the humid tropical Guinea savanna and Salel climate zones Pages 318-328 in Pro-ceedings training course cn concepts techniques and application of irtegrated pest management in rice inWest Africa 10-20 Jan 1982 Fendall Liberia West Africa Rice l)evelopioent Association Monrovia Liberia 505 p

7 Akibo-Betts V I and S A Raymundo 1978 Aphids as rice pests in Sierra Leone lit Rice Res Newsl 3(6)15-16

8 Akinsola F A 1975 Present status of different rice stemborers in parts of Nigeria iicL Entomo Newsl 328

9 Akinsola FA 1979 Ihe biology and ecology of rice stemborers in Nigeria Ph 1) thesis University ol lbadan Nigeria 217 p

10 Akinsola E A 1979 Guidelines of integrated management of insect pests of upland rice in West Africa Presented at the Seminar on integrated management of rice diseases and insect pests at Bobo-Dioulasso Upper Volta 17-22 Sep 1979 West Africa Rice Develop-ment Association Monrovia Liberia

II Akinsola IA 1984 Insect pests of upland rice in Africa Pages 301-305 in An overview of upland rice research Proceedings of the 1982 lIouak Iwsrv Coast upland rice workshop International Rice Research Institute 13O Box 933 Manila Philippines

12 Alam M S V TJohn and Kaung Zan 1985 Insect pests and diseases of rice in Africa Pages 67-82 in Rice improvement in Eastern Central and Southern Africa proceedings of the international rice workshop at Iusaka Zambia 9-19 April 1984

13 Alticri M A A van Schoonhovcn and J D Doll 1977 The ecological role of weeds in insect pest management systems a review illustiated with bean (Phaseohsvulgaris L)cropping systems PANS (Pest Artie News Summ) 23195-205

14 Anmante F 1967 The leaf-cutting ant Atia capiguaraa pest of pastures [in Portuguese] Biol6gico (Sao Paulo) 33113-120 (Rev Appl Entomol A56207)

15Andrade A C and D Puzzi 1951 Preliminary results of experi-merits on the control of Sculptcoris iastantus on sugarcane [in Portuguese] Bioulgico (Sao Paulo) 1744-49 (Rev Appl Entomol A41421422)

16 Appert J 1970 Maliarplia separaelha (rice stem borer) new observations of entomological problems of rice in Madagascar [in French] Agron Trop (Paris) 25329-334

17 Ariadeau Mand Z Harahap 1986 Relevant upland rice breeding objectives Pages 189-197 in Progress in upland rice research Proceedings of the 1985 Jakarta conference International Rice Research Institute 1O Box 933 Manila Philippines

18Asar S and Kisimoto 1975 Seasonal occurrence of the smaller brown planthopper in upland rice [in Japanese] Proc Kanto-Tosan Plant Prot Soc 2228

19 Bakker W 1974 Characterization and ecological aspects of rice yellow mottle virus in Kenya Meded Landbouwhogesh Wage-ningen 152 p

20 Balut F F 1970 White grubs attacking rice Oryza ativa L [in Portuguese] Biol6gico (So Paulo) 36321-322

21 Banerjee S N and L M Pramanik 1967 The lepidopterous stalk borers of rice and their life cycles in the tropics pages 103-124 in The major insect pests ofthe rice plant Proceedings ofa symposium at the International Rice Research Institute September 1964 The Johns Hopkins Press Baltimore Maryland USA

22 Barrion A T and J A Litsinger 1980 Ants a natural enemy of cliaiarac in dryland rice IntRice Res Newsl

5(4)22-23 23 Barrion A T and J A litsinger 1981 The spider fauna of

Philippine rice agroecosystem 1Dryland Philipp Entomnol 5139-166

24 Barrion A T and 1 A Iitsinger 1984 The spider fauna of Philippine rice agroccosystcms II Welland Philipp Eiitomol 611-37

25 Barrion A T1 and J A Litsinger 1984 lihanus (Diptera Tabanidae) eggs an alternative host of rice stei borer (SI3) egg parasite Telenoonis digns (Iliymenoptcra Scelionidae) lit Rice Res Newvsl 9(6)19

26 Barrion A T and J A Litsinger 1985 Identilication of rice leaffolders by wing markings Int Rice Res Newsl 10( 1)24

27 Barrion A T and J A Litsinger 1985 Chctiosspp (Coleoptera Carabidae) a leaffolder ILF)predator lit Rice Res Newsl 10(1)21

28 Bertels A 1970 Insect pests of rice and theircontrol [in Portuguese] Pelotas Brazil Instituto de Expcrimentacao Agropccuararias do Sul Circular 43 24 p

29 Bonzi S M 1982 Ctilo (flisilineus J de Joannis (Lepidoptera Pyralidae) stem borer of irrigated cereals in Upper Volta LioFrench] Agron Trop Riu Rizic Cult Vivrieres Trop 37207-209

30 Bourke T V T L Fenner J N LStihick G Baker EHassan D F OSullivan and C S li 1977 Insect pest survey for the year ending 30th June 1969 Papta New Guinea )epartment of Agrishyculture Stock and Fisheries Port Moresby 57 p

31 Bowden 1 1973 Migration of pests in the tropics Meded Rijksfac Landbouwwet Gent 38785-796

32 Breniere J 1973 The main rice parasites in West Africa and their control Pages 27-42 in Proceedings seminar on plant protection for the rice crop West Africa Rice Development Association Monrovia Liberia

33 Breniere J 1976 The principal insect pests of rice in West Africa and their control West Africa Rice Development Association Monrovia Liberia 52 p

34 Brown ES 1962 The African arinyworn Spoiolptera eXI)Ina (Walker) (Lepidoptera Noctuidae) a review of the literature Cornshymonwealth Institute of Entomology London UK57 p

35 Brown F B 1968 Upland rice in Latin America FAO-International Rice Commission 12th Session of the Working Party on Rice Production and Protection 2-14 Sept Peradeniya Sri lanka Food and Agriculture Organization Rome Italy 6 p

36 Buyckx F J 1 1962 Disease and insect pests of crops inCongo Rwanda and Burundi [iil French] Publications dc LInstilut National pour lestude Agrononiquedu Congo M Weissenbruck S A Brussels Belgium 708 p

37 Cariiio F 0 1981 Role of natural enentics inpopulation suppression and pest management of green rice lealhoppers Ph D thesis University ofthe Philippines at Los Bafitos PhilippinesLaguna 136 -

38 Carpente A J 1983 The development of rice cultivation and extension of vegetable productin in Zanzibar Final Report Vol II1 October 1983 Food and Agricultme Organization Ronte Italy

39 Carroll C P and S1 Risch 1983 Tropical annual cropping systems ant ecology Environ Manage 751-57

4(0 Cates R G 198 1 Host plant predictability and the feeding patterns of monophagous oligophagous and polyphagous insect herbivores Oecologia (Berl) 48319-326

IRPS No 123 January 1987 37

41 Cendcafia S M and F B Caora 1967 Insect pests of rice in the 62 Ene 1 C 1981 I)iiorldhol)ierusihilis Slater (licteroplera Philippines Pages 591-616 in The najor insect pests ofthe rice plant lygacidue Illissinac) a new rice pest front Nigeria lit Rice Res Proceedings of a symposium at the International Rice Research Newsl N3)15 Institute September 1964 The Johns Hopkins Press Baltimore 63 Fugenio T S 1965 rhe effect oi e-vCgetation on rice production Maryland USA 729 p and the possibility of overcoming it by leaf pruning MS thesis

42 Centro Ihternacional de Agricklittira Tropical 1984 Upland rice in University ol the Philippines at Los Baijos ILaguna Ihilippins 69 p Iatin Anerica Pages 93-119 hi An os ervies of uplaind rice research 04 Farrow R A 1975 The Alrican nmigratory locust ii is main Proceedings of the 1982 Boinakamp Ivory Coast upland rice workshop outbreak area of the middle Niger qlinlitatise studies of solitary International Rice Research Institute P0 Blox 933 Manila populations in relation to environmental factors Locusla No II Philippines 198 p

43 Chandra G 1979 Taxonomy ad hn riononlics ofthe imect parasites of 65 Fennal R G 1943 Food crop investigation Windward and rice Icalloppers and planthoppers in the Pihilippines and their Ieeward Islands First report Report onl work dunring the periodiiiportairce iii natural biol)gical cointrol Philipp Fntoltol 4119-139 April 1942-April 1943 Imperial College of Tropical Agriculture

44 Chang T T 1976 The originii colut ion enlIivation dissemination Trinidad 16 p (Rev of Appl Fintornol A3222-25) and diersificition of Asian and African rices -uphytica 25425-44 1 66 Ferreira F 1983 Integrated pest ritaageiret of upland rice [ill

45 Chatierjee P B 1979 Rice leaffolder attacks in India lit Rice Res Portuguese English summary] Pages 323-341 in Upland riceculttre Newsl 4(3)21 factors affecting production E Ferreira T Yamada and F-

46 Cheng Cbien-pan 19(4 Upland rice in Taiwan International Rice Malmolta eds Institutoda Potassae Foslato Institnto Internacional Cnmmission Wiorking Party oit Rice Production ard Protection I()th da Potassa Piracicaha Sao Paulo lraid Meeting 3-10 March 1964 Manila Philippines Food and Agricul- 67 Ferreira E arid F dl Silva Martins 1979 Occurrence of ture Organiaion Rome Italy 5 p Sogatohs orvziola (Muir) inupland rice in Goias Ilrail lit Rice

47 Chcrrett 1 M and 1) 1Peregrine 1976 A review of the status of Rcs Ncwssl 4(4)18 leaf-cutting ants and their control Ann Appl Biol 84124-128 68 Ferreira E and 1F da Silva Martiris 1984 Insect pestof rice ill

48 Chipeta F M 1974 Pests of rice in Malawi Pages 90-95 in Brazil and their control Jill Portugucsej FMIlRAtPA-CNPAF Proceedings Sith eastern Africa cereals research conference 10-15 Goiania Brmiil 67 p March 1974 DRB Mialda ed Ministry of Agriculture lilongwe 69 Ferreira FI F ca Silha Miartins S S Nero and 1 It P de Malawi Carvallio 1982 M tltiplc regression analysis todeterimine the effect of

49 Chu Yau-i 1979 The d atiaging period of the rice seedling fly insect populatiois ott upland rice yield [inIPortuguese Pesqui (Atlterigotiavxigu Stcin) (NItiscide Diptera) oil cori) at Fast Java Agropecu Bras 17671-675 Indonesia Plant Prot Bill 21403413 70 Ferreira F 1F da Silva Martins and F 1PZimmermann 1979

50 Conklin H C 1957 IHaninoo agriculture a report on aln integral Rice varietal resistance to tle lesser corn stalk borer [in Port uguese] systent of shi fti rig cult ivationi in the Pltilippincs Food and Agriculture Pesqui Agropccu Biras 14317-321 Organization Rome Italy 209 p 71 Fleming W F 1968 Biological control of lie hapanese beetle U S

51 Cramer H II 1967 Plant proection and world crop production Dep Agric RLs Sers [ech Bull 1383 78 p Pilaniensclhutz-Nachr 201-524 72 Food and Agriculture Organiation of the United Nations 1982

52 l)ani R C and Majunidar 1978 Prelininary observations on tie Country report surmimary Pages 27-37 in Report on the first incidence of root aphids on differeit rice cultnrcs Sci Cult 4488-89 FAOIUN)IPhail and regional training coiuirse in iniproved ctltural

53 Davidson R I 1979 lffects of root feeding on foliage yield Pages practices for upland rice 5-23 Oct 1981 Northern Region Agrculturd117-120 in T K Ci osb and R B Pottinger eds Proceedings of the Developririt Center Chiang Mai Food and Agricuilture Organiiashy2nd Australian conlerence on grassland invertebrate ecology Govt tion Rome Italy 37 n Printer Wellington New Zealand 73 Franco do Ariaral S 195 1 Residual action of insecticides in turfy

54 l)ean G 1W 1976 Rice insect pests inlaos Int Rice Res Newsl soil--the results of a field experiment [in Portuguese] Biol6gico (Sao 1(2)15 Paulo) 17209-214 (Rev Appl Entomol A41423)

55 I)ean G J W 1978 Insect pests of rice in Laos PANS 24(3)280-289 74 Freeman ) 1970 Report onithe Iban london School of -coroiics 56 )ingle H 1984 Behavior genes and lifc histories conplex adapta- Monographs on Social Anthropology No 41 Athlane Press

tions to uncertain environments Pages 169-194 in A new ecology London UK 317 p novel approaches to interactive systems P W Price C N Slobod- 75 Fuchs T W1 A Harding and 1W Smithlr 1979 Induction and chikoff and W S Gaud cdsl ohn Wiley New York USA termination oldiapause in the sugarcane borer Ann Flitonlol Soc

57 Dobelmann-JP and M Falais 1968 Upland rice cultivation ill tie Am 72271-274 North-western part of Madagascar in 1966-1967 [in French English 76 (hbriel B1) 1975 A review tf the major insect pests of some tpland summary] Trop Agron 2313-21 crops in tlc Philippines Pages 86-111 in Ad Htoc panel of experts on

58 Doniiciano N L 1979 Chemical control of tie lesser corn stalk pest disease arnd weed problems in some rainfed crops Bangkok (lolsonno~paloIgnowslhts) (Zeller 1848) (I-epid(ptera Ihycitid ae) Thailand Food aid Agriculture Organization Roire Italy

on rice in Ponta Gressa Parara 1977-1978 [in Portuguese] Pages 77 Garg D K and N K Shal 1983 White grub outbreak on rainfed 291-303 in Anais 1 Reuniao de tecnicos cmiirizicultura do estado de dryland rice in Uttar Pradeslh Int Rice Res Ncwsl 81(4)16-17 Sao Paulo-Campinas Cati Brazil 78 Garg I) K and IP Tandon 1982 Major insect pests of rice on hilly

59 Dunsmore 1 R 1970 Investigations on tlie varieties pests and tracts of Uttar Pradesh India lilt Rice Res Newsl 7(11)11-12diseases of upland rice (hill padi) in Sarawak Malaysia Int Rice 79 Garrity 1) P 1983 Upland rice research an international bibliograpy Comm Newsl 1929-35 1975-1982 Iternational Rice Research hnstitnte PO Box 933

60 Elias 1R1965 The new decline of rice seedlings is not caused by Manila Philippines 121 p parasites [in Portuguese] Biol6gico (Sao Paulo) 313-6 80 Ghauri M S K 1982 A new subspecies of Iinorh)rone bull

61 Empresa Brasiliera de Pesquisa Agropecuaria 1984 Upland rice itt cornlus Slater (Hemiptera Iygaeidae) front Papua New Guinea on Brazil Pages 121-134 it Av overview of upland rice research rice and carpet grass Ihll lEntoniol Res 72133-137 Proceedings ofthe 1982 Blouak Ivory Coast Upland rice workshop 81 Gonzales F 0 PArregoces R I ihernandef and OT Parada International Rice Research Institute PO Box 933 Manila 1983 Insect and imite pests of rice arid their control in Latin America Philippines [in Spanish] la Federacion Nacional de Arrocerts (Fedearroz)

Bogota Colombia 60 n

38 IRPS No 123 January 1987

82 Grist D H and R J A W Lever 1969 Pests of rice Longmans London UK520 p

83 Hale P R 1979 Two subterranean pests of upland rice in Papua New Guirnca IntRice Res Newsl 4(2)16-17

84 Hale P R and J T Hale 1975 Insect pests of rice in the New Guinea Islands Rice Entomol Newsl 34

85 Harris W V 1969 Termites as pests of crops and trees Common-wealth Institute of Entomology London UK 41 p

86 Hattori 1 1968 A method to distinguish between grass leaf roller (Ctaplha rocis medinalisGuenee) and the rice leaf roller (Susurtia exigua Butler) [in Japanese] Plant Prot (Japan) 23167-170

87 Henry I E and 1A Onsager 1984 Experimental control of the mormon cricket Iahrosshnph by NJsena locustae(Aficrospora Microsporidae) a protozoan parasite of grasshoppers (Ort Acri-didae) Entomophaga 27197-201

88 Hidaka 1 P Vungsilabulr and SKadkao 1974 Studies on ecology and control of the rice gall midge in Thailand Tech Bull 6 TropicalAgricultural Research Center Tokyo Japan 113 p

89 Hincklcy A 1) 1963 Ecology and control of rice planthoppers in Fiji Bull Entomrol Res 53467 481

90 Hirao J 1982 Ecology and chtInical control of the rice leaffolder Jpn Pestic Inf 4114-17

91 Ho DT 1984 Sten borer incifience in rice ecosystems in KenyaEast Africa Pages 307-320 in A overview of upland rice research Proceedings of the 1982 Bouac Ivory Coast upland rice workshop International Rice Research Institute PO Box 933 Manila Philippines

92 Ho )T J 1Njokah and J GKibuka 1983 Studies on rice sten borers in Kenya with emphasis on ifaliarpiaipiraellRag Insect Sci Appl 465-73

93 Ichihara I T Ito A NIatsuo and M Sawada 1971 Chemical control of the rice striped stem borer in upland rice grown in plastic mulch [iii Japanese] Proc Kanto-Tosan Plant Prot Soc 1882

94 IclhiharaIand A Matsuo 1969 Damage caused by rice striped stett borer to upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1686-87

95 Ichhara I atid A MIatsun11970 Conside ration oft lie extraordinary prevalence of rice striped stem borer in upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1778-79

96 Institut de Recherches Agrolnliqes Iropicale et des Cultures Vivrieres 1978 Insect resistance Pages 12-13 in Upland rice 1978 atnual report Bouak Ivory Coast 27 p

97 International Institute for Tropical Agriculture 1983 Aphids itt rice roots I ITA Res Briefs 4(1-3)4-5

98 International Institite fi Tropical Agriculture 1983 Hopperburn in rice caused by brown plathopper IITA Res Briefs 4(l-3)6-7

99 International Rice Research Institute 1975 Annual report for 1974 PO Box 933 Manila Philippines p 216

100 International Rice Restarch Institute 1976 Annual repiort for 1975 PO Box 933 Manila Philippines p 228

101 International Rice Research Institute 1979 Cropping systems program Component technology development and evaluation Page 404 i Annual report for 1978 11O Box 933 Manila Philippines

102 International Rice Research Institute Department of Entomology 1981) Insecticid evaluation 1979 Tables 73 74 PO Box 933Manila Philipes

103 International Rice Research Institute Department of Entonmology 1981 Insecticide evalomtion 1980 Table 81 POBox 933 Manila Philippines

104 International Rice Research Institute 1981 Report of a meeting of Asian rice-based cropping systems entomologists Eleventh cropping systems working group meeting 18-22 May 1981 PO Box 933 Manila Philippines p 132 133 144

105 International Rice Research Institute 1984 Cropping systems program Pages 368-373 in Annual report for 1983 PO Box 933 Manila Philippines

106 International Rice Research Institutc 1984 Upland rice in Asia Pages 45-68 in An overview of upland rice research Proceedings of the 1982 Bouakamp Ivory Coast upland rice workiiop loternational Rice Research Institute 11O Box 933 Manila Philippines

107 Irshad M 1977 Grasshoppers in rice in Pakistan lit Rice Res Newsl 2(3)8-9

108 Ito R and TOno 1964 Upland lice its culture and varieties in Japan lIternational Rice Cormmission loth Meeting of the Working Party on Rice Production and Protection 3-10 March 1964 Manila Philippines Food and Agriculture Organization Rome Italy 12 p

109 Jeffrey M I and V A Dyck 1983 Effect of moonlight on lightshytrap catches of brown planthopper lit Rice Res Newsl 8(6) 13

110 Jennings P R and T Alicia Pineda 1983 Management ofthe rice plnthoppers Sogatodsorivi(coIl in Latin America Presented at the meeting on stable plant resistance FAO and lIndocsian I)inectorate of Food Crops Protection Bali lndonesia 29 Novcniber-2 l)ccemshyber 9 p

IllJimenez Gomez J A 1979 Rice insect pests i lla ios ()riCntales lil Spanish] Pages 99-103 in Rice ciltivation Instituto (olotibiatto Agropecuario Compndio No 29

112 Jongeleen F JJ Soesanto and A Mockidjo 1979 Studies oti pathogens of two species of Scarabacidac froinIndonesia Z Angew Enlomnol 87412-417

113 Jouvenaz 1) PC S lofgren and W A Baiks 1981 Biological control of imported fire antsa review of cirrent knowledge Bull Entomol Soc Am 27203-208

114 Kainoh Y K Shiniiu S MarUani Y Taiaki 1980 lost finding behavior of tie rice bu lplotori+JchinvittjI)allas (liciniptera Corcidac) with special reference to diet pattern ol aggregation and feeding on the rice plant Appl Entomol Zool 15225-233

115 Kakkar K 0 P Sharma and P I Sharma 1977 Note on Ihteronyrchus liodresRedt (Coleopicra I)ynastidae) as asporadic pest of paddy and maie inHimachal Pradesh Corr Agric 156-57

116 Kalode M131 79 Major insect pest problems and their control in upland rice (Unpublishcd mineogriph CWI oJNaaiat lni Chandaratat (198 1)original not seen)

117 Kalshoven L i 98 I The pests of crops in I ldoniesia (translated by P A vatn der Laan) 1 T Ichtiar B3aru-vati Iloeve lakarta Indonesia 701 p

118 Kamal N Q 1981 Suppression of whitebacked planto ner Sogatella fircaifra (Horvath) aLId rice lealfolder (nhal -cis licdb lis (Guenee) populations by natural cnclmie hlI) tiesis Grcgorio Araneta University Foundation Victoncta Park Metto Manila Philippines 179 Ip

119 Karim A N M Reaul 1975 A species ofliealy bug Trioni i70s ceres new oii rice in the Philippines Rice Entoniol Newsl 247

120 Katanyukul Wand C Chandaratat 1981 Insect pestsof uplaid rice and their control Report firstFAOUNlFi Tliailand training course on improved cultural practices for tplad rice Chiang Mai Thailand 5-23 Oct 1981 Food and Agriculture Organiation Rome Italy 22 p

121 Kern F 1955 lhletwioa(I)uninemts) bhhit Biturm(Coleoptera) a new pest of sugarcane inVenezuela [inSpanish with Etnglish sunmary] Agron Trop (Maracay) 527-29 (Rev Appl Eitoniol A4572)

122 Khan K M and S Ghai 1974 White grubs ald their control in India Pesticides (1Bombay) 819-25

123 Kiritani K 1975 Integrated pest management of rice pests Pages 66-70 itt Proceedings platning workshop ott cooperative hIeld research in pest managementP SMotooka ed East-West Center Honololu Hawaii USA

124 Kisimoto R 1967 Ecology of insect vectors forecasting and chemical control Pages 243-255 in The virus diseases of tle rice plant Proceedings of a symposium atthe International Rice Research Institute April 1967 The Johns Hopkins Press altimore Maryshyland USA

125 Kisimoto R and V A Dyck 1976 Climate and rice insects Pages 367-391 in Climate and rice International Rice Research Institute PO Box 933 Manila Philippines

126 Kureha S S Kobayashi SAsari and T Ishihara 1977 Life history and control of the mole cricket Grrllotalpa africana Palisot de Beauvois [in Japanese English translation 7520 1RRI library] Proc Kanto-Tosan Plant Prot Soc 21192-197

127 Kushwaha K S 1981 The black beetle as a pest of rice in Haryana India Int Rice Res Newsl 6(4)14

128 Lawani S M 1982 A review of tile effects of various agronomic practices on cereal stem borers Trop Pest Manage 28266-276

129 Leuck D B 1966 Biology of the lesser cornstalk borer in South Georgia J Econ Entomol 59 797-801

130 lever R J A W 1955 Insect pests of the rice plant in Malaya Int Rice Comm Newsl 1612-21

131 Li Ching-Sing and Shin-Foon Chiu 1951 The study ofthe rice gall midge Iachydiplosis orvza Wood Mason [in Chinese with English summary] 1Taiwan Agric Res 21-13

132 Iitsinger J A 1984 Assesament of the need-based application for rice Proceedings TechnologyTransfer Workshop 15-16 March 1984 International Rice Research Institute 1PO Box 933 Manila Philippines 6 p

133 Litsinger J A 1984 Environmental diversity and insect pest abundance with reference to the Pacific Pages 285-307 in Proceedings sub-regional training course on methods of controlling diseases insects and other pests of plants in the South Pacific 4-20 Oct 1982 Vaini Kingdom of Tonga Nukualofa

134 Litsinger J A RF Apostol and M BObusan 1983 White grub Leucopholis irrorata (Coleoptera Scarabaeidae) pest status popula-tion dynamics and chemical conllol in arice-maize cropping pattern in the Philippines J Econ Entoinol 761133-1138

135 Litsinger J A C de la Cruz F Raymundo A T Barrion M D Lumaban J P Bandong M S Venugopal F Paragna and E Balete 1982 Insect pests and insecticide reponse of dry-seeded bunded rice Pages 179-193 in Report of workshop on cropping systems research in Asia International Rice Research Institute Los Bafos Philippine

136 Litsinger J A M 1) Luinaban J P Bandong A T Barrion W Katanyukul and N Panda 1978 Insect pest management in rainfcd lowland rice agroecosystms of tropical Asia International Rice Research Conference 17-21 April 1978 International Rice Research Institute Los Bafios Philippines

137 Litsinger J A and K Moody 1976 Integrated pest management in multiple cropping systems Pages 293-317 in R I Papendick P A Sanchez and G B Triplett eds Multiple cropping American Society of Agronomy Madison Wisconsin USA

138 Loevinsohn M E 1984 The ecology and control of rice pests in relation to the intensity and synchrony of cultivation Ph D thesis University of London 36) p

139 Loevinsohn M E J A Litsinger N Panda M A Austria R F Apostol C Andrion and J Bandong 1982 A basis for predicting rice insect pest abundance in cropping systems Paper presented at the 1982 International rice research conference on rice and cropping systems 19-23 April 1982 International Rice Research Institute Los Bafios Iaguna Philippines 25 p

140 Lopez A W 1930 Repot of the Entomology Department Pages 145-172 in Annual report of the Philippine Sugar Association 1929-30 Manila

141 Lopez AW 1931 Entomology Department Pages 227-266 in Annual report of the director or research 1930-31 Philippine Sugar Association Manila

142 Lopez A W 1932 Entomology Deparment Pages 252-279 in Annual report of the Philippine Sugar Association 1931-32 Manila

143 Lungibill P and G G Ainslee 1917 The lessercornstalk borer US Dep Agric Agric Res Serv Tech Bull 539 27 p

IRPS No 123 January 1987 39

144 McGuire J U Jr and BS Crandall 1967 Survey of insect pests and plant diseases ofselected food crops of Mexico Central America and Panama U SDepartment of Agriculture International Agriculshytural Development Service Agricultuial Researrh Service October 1967 11 p

145 McNeill Sand T R ESouthwood 1978 The role of nitrogen in the development of insectplant relations Pages 77-9 in lliochemical aspects of plant and animal coevolution J B Ilarbornu ed Academic Press London UK

146 Maebara H 1955 Spatial distribution of lastinstiarlanaeof purplish stem borer (Sesamia iqferens Walker) hibernating in an upland rice field Bull Kagoshima Univ Fac Agric 438-41

147 Mallick E H and N G Hajra 1977 Response of rice plant to defoliation treatments at different developmental stages Indian Agric 2151-56

148 Mansingh A 1971 Physiological classification of dormancies in insects Canl Entomol 103983-10)9

149 Martell C G 1974 Catalogue of plant feeding insects of Mexico [in Spanish] Fitofilo 27(69)1-176

150 Martinez 0 J 1982 Principal rice pests in Ilanos orientalis [in Spanish] P~ages 80-94 in Curso deact nalization de conocimientos en arroz Villaniencis Instituto (olonmbiano Agropecuario

151 Martins J F da Silva F Ferrcira A S Prabhu F 1 P Zimmermann 1980 Use of pesticides for the control of the main subterranean upland rice pests esqui Agropecu Bras 1553-u

152 Martins I F da Silva F Fe eira rod BdamSilveira Pinheiro I Q2 Damage simulation caused by defoliators of upland rice [in Portuguese with English summary] Pesqui Agropecu Bras 171113-1119

153 Martins 1F da Silva N(iyuen van Tan and 1Ida Silveira Pinheiro 1981 Stem borer resistance in upland rice associated with plant morphological characters [in Portuuce Iesqui Agropecu Bras 16187-192

154 Masaki S 1980 Sumnmer diapause Ann Rev Entomiol 251-25 155 Masuzawa T 11 Suwa and F Nakasoji 1983 l)ifkleences of

oviposition preferencc and survival rate of two skipper butterflies Parnara guttata and PIchlpidai iathiais (Iepidoptera HIesperiidac) on rice plant and cogon grass New Entomol 3247-56

156 Maurya D M and C P Vaish 1984 Upland rice in India Pages 69-92 in An overview oftpland rice research Proceedings ofthe 1982 Bouakamp Ivory Coast upland rice workshop International Rice Research Institute IO Box 933 Mnila Philippines

157 Menon M G R R N Katiyai S Kuar and S S Misra 1970 Termites attacking the paddy crop Sci Cult 36294-295

158 Miyahara Y and 1 Nishikubo 1967 Behavior of young nymphs of the red rice root aphid Iholahsipiun ri liabhdontinalisSasaki in relation to movement from the leaf to the roots of upland rice [in Japanese] Pioc Assoc Plant Prot Kyushu 1363-66

159 Mo ljoa Tjien 1952 White grub of upland rice (Holotrichiahelleri Brsk) [in Indonesian with English summary] Contrib Agric Res Stn Bogor No 13134 p

160 Mochida 0 and T Suryana 1979 The occurrence of the brown planthopper and ragged stunt on lowland and upland rice ill Central Java and logyakarta in tile late of January 1979 Entomnolgy Survey Report No 15 Central Research Institute of Agriculture (CRIA) Sukamandi Indonesia I February 1979 9 p

161Monet C 1979 Varietal resistance to upland rice pests [inI French] Pages 35-39 in Annual report 1977 Institut des Savanes Department des Cultures Vivrieres Bouak6 Ivory Coast

162 Monte 0 1942 A caterpillar ofrice-fields[in Portuguese] Biol6gico (Sao Paulo) 8161-163 (Rev Appl Entomol A31414)

163 Munroe I) D 1979 Pest management strategy for hill rice in four resettlement areas in Sarawak Malaysia Pages 89-102 i1 Guidelines for integrated control of rice insect pests Food and Agriculture Organization Plant Production and Protection Paper 14 Rome Italy 115 p

40 IRPS No 123 January 1987

164 Nakasuji F M Ishii and Jen-Zon Ito 1984 Rice skippers in Taiwan and their life histories ZZrC (Taipei) Tech Bull 82 10 p

165 Navas I) 1976 Fall armyworum in rice Pages 99-106 il Ecological animal control by habitat management Proc Tall Timbers Con-ference 28 Feb-I Mar 1974 Fall Timbers Res Stn Misc Publ 6 Tallahassee Horida USA

166 Nickel 1 1 1961 Biological control of rice stem borer A feasibility study Tcch Bull 2 International Rice Rcarch Institute PO Box 933 Manila Philippines IIl p

167 Nigan P M and R A Vernia 1985 Insect pests of uplaild rice in Uttar Pradesh Int Rice Res Newsl 10(6)22

168 Nilakhe S S 1985 -cological observations on spittle bugs with emphasis on their occirence in rice Pcsqtui Agropecu Bras 20407-414

169 Nixon G EJ 195 1 lie association of ats with aphids and coccids Commonwealth Iusti lcte of Entonology Iondon UK 36 p

170 Nurdin F an Ishak Manti 1985 Study oit seedling fly of upland rice Interlational Upland Rice Moitoring Tour Report 26 lebruary-2 MarLI Sukaranti West Sumatra

171 Oka 1 N 1979 Ciltural conttol of the brown planthopper Pages 357-369 in Brown planthopper threat to rice production in Asia International Rice Research Institutc 1p0 Box 933 Manila Philippines

172 Ohni M and A Vilnaii 1975 Rice insect pests of Italy biology of Sipha grcria (Kaltenbach) and its control [in Italian] Riso (Milan) 2459-71

173 Otanes F Q atnd 1 I Sison 194 1 Pests of rice Philipp J Agric 12211-259

174 Painter R I 1951 Insect icsistance in crop plants University Press of Kansits Lawrence Kansas USA 520 p

175 Pantua P C and 1A litsinger 1981 Arthropod abundance in continuous vs biannual rice cropping systeis Int Rice Res Ncwsl 6(2)20-21

176 Pathak M ) and V A l)yck 1975 Sttudics ott insect pests of upland rice Pages 186-191 in Major research in upland rice International Rice Research Institute PO Box 933 Ilanila Philippines

177 Patil B R B M IHasabe and 1) S Ajri 198 I Chemical control of white gritb lIditriihinhorraia Fabr intfesting groundntut paddy atnd sorghum Indian 1Entomol 43232-236

178 Pawar A 1) 1974 (ollection and identification of insect pests of rice rlerminal repirt lInternational Rice Research Institute PO Box 933 Manila Philippines 72 p (tnimco)

179 IerrajU A aid A S Reedy 1965 Observations ott diapausc itt tie steti borer ohrice Rice News Teller (lai-Mar)9-13

180 Pont A C 1972 A review of the Oriental species of theri ona Rondani (I)iptera Muscidac) of econtomic importance Pages 27-102 in Control of sorglium shoot fly M G lomtani and W R Young eds Oxford atd IH1 Ptthl Co New I)clhi India 324 p

181 Prasad S C 1PChand aid M 1PSingh 1983 Incidence of rice gall midge in the upland plateau region of Hihar Oryza 20235-236

182 Rai P S 198 0 Nisaga simpex damage to rice in the hill tracts of South India Int Rice kles News[ 5(I)12

183 Rathorc Y S 1969 Gonocplhahli dcprssunt Fabr as itpredator of (tilo zonwIlhis (Swinhoc) and other moths in field cage at Patnagar Uttar Pradesh Indian 1Entotnol 3127

184 Reed W 1983 Crop losses causcd by insect pists in the developing world Proceedings 10th Internatial Congress of Plant Protection lrighton UK 174-80

185 Reedy M V 1981 Ateactogasterinhens(Walker)a new root-grub pest of tipland paddy and inaize Curr Sci 501076

186 Reveno M A 1968 Catalogue of the insect pests of economic crops in Colombia [in Spaiish] Asociacion Latino Americana de Entomo-logia (AIAE) 129-33

187 Reyes G M 1957 Rice dwarf disease in the Philippines Food and Agriculture Organiiation Plant Prot Bull 617-19

188 Rice S E A A Grigarick and M 0 Way 1982 Effect of leaf and panicle feeding byarmyworm (Lepidoptera Noctuidae) larvae on rice grain yield Environ Entomol 7593-595

189 Risch S J 1) Andow and M A Altieri 1983 Agioccosystem diversity arnd pest control data tentative conclusions and nizw research dirctions Environ Entomol 12625-629

190 Rivera C T 1956 Three organic insecticides in thecontrol of insects affecting upland rice Philipp Agric 39465-472

191 Roffrey 1 1972 Locusta outbreaks in the Philippines Acrida 1177-188

192 Rojas R 1) 1941 The effect of pruning at varying ages upon the yield of Ramairio Phipp Agric 29851-860

193 Roonwal M i 1979 Ternite life and termite control in tropical South Asia Scientilic Pnbl odhpr India 139 p

194 Rosetto C1 F da Silva Martins N C Schmidt and L EAzzini 1978 Damage caused by pasturespittle bugs 4Dvoi)lavopictaandDJ schach) on rice lBragantia Nota 6 (Matcli)25-27

195 Rosztto C1- S S Neto I- Amantc I) Iink 1Garcia-Vieira D M de SoLza N V Bansa t and A M de Olivcira 1971 Rice pests of Brazil [in Portuguese] Pages 149-238 inrFA()-International Rice Commission 2nd meeting of the Rice Committee for the Americas 6-11 l)ecember 1971 Pelotas Rio Grande do Sul Brazil

196 Rothschild G H L 1969 Obseratiois on the armyworm Spodoptera matritia acronrctoiiles Gn (Lep Noctuidae) Bull Entomol Res 59143-160

197 Rothschild G 1- L 1974 Parasites of the armworm Spolopiera mauritiaacroncitoihes in Malaysia Fntornopliaga 19293-299

198 Sachan i N S K Gangwar and 1N Katujar 1980 Field crickets damaging cages in North Eastern IHill Region Indian J Entomol 42526-527

199 Sands 1) P A 1977 The biology and ecology of Lepocorisa (Hemiptera Alydidac) in Papua New Guinea Res Bull 18 Departshyment of Primary Industry Port Moresby Papua New Guinea 104 p

200 Sarawak 1968 titomology division Pages 97-106 in Report of the research branch of the Department of Altriculture 1966 Kuching (Rev AIppl Eintomol A56603)

201 Satpathy JM M S Das and K Naik 1977 Effect of multiple and mixed cropping on the incidence of scine important pestsI Entomol Res 178-85

202 Sauer IHF G 1939 Notes Ott Elasmpnlpualtsligniosellths Zeller (Lep Pyr) a serious pest ofcereals in Sao Paolo [in Portuguese] Arq Inst Biol 10199-206 (Rev Appl Entotnol A28378)

203 Satinders 1L A 1 SKing and C L Vargas S 1983 Crop pests of Central Anerica with t bibliography [in Spanish] Centro Agroshynotnico Tropical de Investigacion y Ensenanza Turrialba Costa Rica (CATIE) Tec Bol 9 90 p

204 Sauphanor B 1985 Some factors of upland rice tolerttce to stem borers in West Africa Inseu Sci Appl 6429434

205 Sen P atnd S Chakravorty 1976 Stentflics in Haritighata Nadia West Bengal India [it Rice Res Newsl 1(12) 15

206 Senapai B and 1M Satpathy 1983 The seedling fly Atherigona orrzae Malloch is a pest of rice in Orissa (India) J Entomol Res 777-79

207 Shhjahan M S Alatn M Ah nad and BN Islam 1983 A new scarabaeid beetle pest of dryland rice in Bangladesh Int Rice Res Ncwsl 8(I)16-17

208 Sharma G L S C Modgol and R C Gautam 1979 Effect of depth of submergence ott incidence of bacterial blight and white grub infestation in transplanted rice Int Rice Res Newsl 4(6)12-13

209 Sharma S 1985 Scarabaeid beetle outbreak itt upland rice Int Rice Res Newsl 1(1(l)24

210 Shiraki T 1917 The paddy borer Schoenobhits incertulas Walker [achaku Agric Exp Stn Formosa 256 p

211 Silva A de Brito and 11P Magalhaes 1981 Insect pests of rice [in Portuguese] Circular Tecnica No 22 EMBRAPA Centro de Pesquisa t ecuaria de Tropico Unido (CPAJU) Belem Brazil 14 p

212 Singh S A 1979 Emergence of rice leaf butterfly as a rice pest in Manipur India int Rice Res Newsl 4(4)16-17

213 SinghS A 1983 Changing trends of insect pestsof rice in Manipur Int Rice Res Newsl 8(f)12

214 Siwi S S and P H van Doesburg 1984 Lenovorisa Latreille in Indonesia (Heteroptera Coreidae Alydinae) Zool Meded (Leiden) 58117-129

215 Soejitno J 1977 Reaction of some upland rice varieties toseedling fly eltherikonaexigua Stein [in Indonesian] Siri Hana I Penyakit 630-35 Lembga I osat Fenclilian Pertanian Bogor Indonesia

216 Socnardi 1967 Insect pests of rice in Indonesia Pages675-683 in The major inse pests of thu rice plant Proceedings of a symposium at the Internatital Rice Rec Wi itt S pternber 1964 The Johns Hopkins Press Baltimore Maryland USA

217 Soto P E A T Petcz and I W Buddenhagen 1976 Survey of insect pests and diseases of rice in different ecological zones in Nigeria Rice Entomol Ncwsl 435-46

218 Soto 11E and Z Ziddiqi 1976 Screening for resistance to African rice insect pests Varietal improsetnent sCmi n r Bouak Ivory Coast 13-17 Sept 1976 West Africa Rice l)evelopment Association Monrovia Iiberia

219 Squire F A 1972 Entomological problems in Bolivia PANS (PestArtic News Sumr) 18249-268

220 Stapely 1 II 1978 The rice leaf roller in the Solomon Islands Int Rice Res Newsl 3(l)13

221 Suenaga H 1937 On aphids attacking the roots of rice Oyo-I)oubts Zasshi 9150-152 [in Japanese] (Rev Appl Entomol A25745)

222 Suenaga H 1952 On the seasonal abundance of three subterranean aphids on upland rice Bull Kyushu Agric Exp Sti 1249-262

223 Sutynso Yas and Dandi Soekaria 1964 Beberapa hana yang ttenperang tana man padi tanah kering [in Indonesian] Rapat kerja Padi Tanah Kering 6 sd 8 Juily 1964 Kaliurang 9 p

224 Tallamy I) W and R F Denno 1983 Alternative life history patterns in risky environments in example from lace bugs Pages 129-147 in Insect life history patterns R F l)cnio and If Dingle eds Spring-Verlag 225 p

225 Tamura 1 1939 Observations on some insect pests of field crops [in Japanese] 1 Plant Prot 26414-420 (Rev Appl EntomolA27-556)

226 Tamura and Suzuki 1964 Influences of the early-transplanting practice in rice culture upon development of the rice stem borer Chilo siql)ressalisWalker Bull Ilokuriku Natil Agric Exp Stn 761-94

227 Tan J P 1924 The rice root aphid Drowhi ihirsoia Philipp Agric 13277-288

228 Tanaka 1 1957 Taxonomy and distribution of some subterranean aphids injurious to upland rice in Japan with description of a new species [in Japanese English summary] Botyu-Kagaku 22168-176

229 Tanaka T 1961 The rice root aphids their ecology and control Spec [full College of Agriculture Utsunomiya University Japan F3 p

230 Tauber M IC A Tauber R Nechols and 1Obrycki 1983 Seasonal activity of parasitoids control by external internal and genetic factors Pages 87-108 itt I)iapause and life cycle strategies in insects V K Brown and I Ilodek eds )r W Junk The Hague Netherlands

231 Taylor W F 1972 Effects of artificial defoliation (simulating pest damage) on varieties of upland rice Exp Agric 879-83

232 feotia T P S and S Natud 1966 Biotoomics of the rice skipper Parnaraniathiis Fabricius (Lepidoptera Hesperiidac) Indian J Entomol 28181-186

233 Teran F 0 1971 Rice pests of Santa Cruz [in Spanish Ministerio de Agricultura Sita Cruz Bolivia Bull 45 8 p

234 Than Itun 1976 Population dynamics of the yellow rice borer Tryporza itcertulas (Walker) and its damage to the rice plant MS thesis University of the Philippines at Ios Bafios Laguna Philippines 92 p

235 Uichanco L B 1911 Pests Pages 55-74 it Rice Philipp Agric 11-200

IRPS No 123 January 1987 41

236 van der Laan P A 195 1 Life history of the rice seedling fly Contrib Gen Agric Res Stn Boor Indonesia 118 15 p

237 van der Laan 11A 1959 Correlation between rainfall in the dry season and the occurrence of the white rice borer (Scirpolhaga innotala WIk) in lava Entomol Fxp Appl 212-20

238 Van Driesche R G 1983 Meaning of percent parasitismin studies of insect parasitism Fnviron Entomol 121 I1- 122

239 Venugopal M S and A litsingcr 1984 Fffect ofcarbofuran on rice growth Prot Ecol 7313-317

240 Vietnam Directorate of Rural Affairs 1962 Annual work progress report on crop improvement program of rice sugarcane vegetable and field crops (lor the period from July 1961 to June 1962) (11) 299 p (Rev Appl Entomol A51526-527)

241 Wan M T K 1972 Observations on rice leaf- and planthoppers in Sarawak (Malaysian Borneo) Malay Agric 148308-335

242 West Africa Rice I)esclopnient Association 1978 Annual research report Vol 2 Statistics Monrovia Liberia

243 West Africa Rice Development Association 1979 Annual research report p47 Monrovia Liberia

244 Williams ) F and CS lofgren 1981 Fli I illy l-468 anew bait toxicant for control of the red imported lire at Fla Entomol 64472477

245 Williams I) 1 1970 The mealybugs OFlomoptera Coccoidea Iscudococcidac) of sugar-cane rice and sorghum Bull Entomol Res 0109-188

246 Williams D) 1 1982 Pulinaria icervi (Signoret) and its allies ott sugar cane and other grasses Btll Entoinol Res 72111-117

247 Wirjosuiardjo S 1976 Study ofchenmical cont rot of insect pests ott upland rice Imiu Pertanian (Jogjakarta) 1323-329

248 Wirjosuhardjo S and N Wirarta 1976 Chemical control of the white grub tIolorithiatlltriin upland rice in M uneng Irobolinggo Indonesia) [in Indonesian] Imut Pertanian (Jogjakarta) 1330-332

249 Yano K T Miyake and V F Eastop 1983 The biology and economic importance of rice aphids (Henuiptera Aphid idae) a review Bull Entomol Res 73539-566

250 Yelp D and P Rhodes 19(4 lhe success of drilled dryland rice iii the Fiji Islands International Rice Commission Working Party on Rice Production and Protection Manila Philippines 3-10 March1964 Food and Agriculture Organization Rome Italy 13 p

251 Young W R 1981 Fifty-five years of research on the sorghum shootfly Insect Sci Appi 23-9

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axial-flow pump

Page 34: UPLANDRICE. INSECTl--1i M T -E!R.EOLOU t RTANCE ,AND …delphacid.s3.amazonaws.com › 6958.pdf · The significance of abiotic factors in upland rice insect ecology will be apparent

0Z

Table 8 Parasitization rates of lepidopterous pests in upland rainfed lowland and irrigated lowland environments Philippines 1976-84a

Parasitization (5)

Environment and site Year YSB GFB PSB Earcutting Swarming Skipper Greenborned RLF Brown larva larva caterpillar caterpillar caterpillar caterpillar larva semilooperEgg Larva larva larva larva larva larva

UplandBatangas 1977 - 0 (57) - 0 (63) - 0 (71) - 0 (106) 11(142) 2 (101)1978 43 (4697) 8(451) 5 (38) 3 (40) 6 (61) 2(58) 15 (57) 13 (129) 13 (1286) 4 (30)1979 59(4135) 12(320) 13 (11) 5(28) 9(45) 6(46) 7(23) 8(160) 9(1152) 8(26)

Rainfed lowland Iloilo 1976 - 0 (10) 0(10) 0 (7) 0 (19) 2 13) 15 (180) 5 (76)1978 0 (54) 0 (3) 0 (17) 9 (64) 4 (78) 3 (73) 0 (121) 0 (191)1979 64 (2350) 16 (84) 6 (17) 0(6) 18 (68) 25 (104)

1981 56 (1322) 18(304) 0(5) 12(51) 18(138)Pangasinan 1976 60 (7 7 )b 62(82) 0(111) 5 (39)

1978 58 (53)b 5(51) Cagayan 1979 59(713) 8 (101) 6 (34) 0(7)

1980 0(11) 4 (28) 15 (286)

62 ( 48(177) 61(140)1981 0(5) 12 (2) 13(45) 14 (92)

Irrigatedlowland Laguna 1974 62 8

1974-75 50 41 1975 43 72 1978 96 (623) 21(24) 33 (60)1979 83(2017) 18(110) 8(62) 11 (89) 22(400)1980 78 (1625) 20(75) 0 (16) 4 (72)1983 60

9 (80) 7 (270) 30

aSources of data All sites A T Barrion unpublished 1977-82 Batangas R F Apostol unpublished Iloilo and Pangasnan 136 Cagayan B Canapi unpublished Laguna 175 P CPantua unpublished 118 234 143 Carino and Shepard unpublished YSB = yellow stem borer ScirpophagaincertulasGFB = gold fringed borer Chilo auricilius PSB = pink stem borerSesamia inferens earcutting caterpillar Alythimna separata swarming caterpillar Spcdoptera mauritia acronyctoides skipper Pelopidas mathias reen horned caterpillar Melanitls ledaismene RLF = rice leaffoldersCnaphalocrocismedinalis and Marasmiaspp brown semilooper Mocisfrugalis Figures in parentheses are sample sizes Egg masses

IRPS No 123 January 1987 35

Table 9 Effect of seeding rate and insecticide on plant density and Yield losses from insect pests in upland rice often areyield of Dagge upland rice Batangas Philippines 1976a comparable to those of lowland rainfed or irrigated rice

Seeding rate Insecticide Dosage Plant density Yield (0-Y30o) but usually from different pests The principal (kg aiha) seed treatment (kg aiha) (noIra-row) (tha) groups are soil pests feeding on sown seed and rootsat 14 DE followed by seedling pests These groups have largely gone

50 None - 35 a 20 a unnoticed leading researchers to conclude that stem borers50 Carbofuran F 10 38 ab 23 a and seed bugs are the most important pests (17) Yield loss 10 beCabofuranF100 None - 4551 c 23 a trials that pinpoint major pests need to includc seed

100 Dieldrin WP 10 52 cd 22 a treatments Granules applied in the furrow may not give150 None - 59 d 22 a adequate protection from many seed pests

aAv of 8 fields DE = days after crop emergence In a column means The floristic and pedological diversity of upland rice followed by a common letter are not significantly different at the environments is matched by the diversity of arthropods5level by D)MRT F = flowable Wi = wettable powder both pests and their natural enemies Irrigation has homoshy

genized lowland rice environments Flooded soils tend tothreatened by epidemics of locusts or armyworms New have the same properties and to eliminate soil pests The technology stressing efficiency has minimized insecticide spatial and temporal dominance of cultivated rice has cost benefited specialist pests

Seed treatment is normally inexpensive and can be Upland rice has no specialist arthropod species The economicallyjustified in -nanycases(66) Treating seed with monophagous pests on upland rice prefer the lowlands systemic insecticide (216) but not chlorinated hydrocarbons where rice evolved Rice in the uplands exists only because (247) is most effective in controlling seedling maggots of man and even though lowland rice predates upland riceFoliar sprays have to be repeated to obtain control and specialist species evolved Wheat maize and sorghum do often are washed off by frequent early season rains not have monophagous pests leading us to conclude that Granular insecticides are effective but required dosages are there is no evolutionary advantage for inssects to have too expensive to justify except under heavy infestation narrow host ranges in upland graminaceae(247) When directed against first-instar larvae banding Pests have adapted to the uplands not only by having granules in seed furrows (177) can be effective against white wider plant host ranges but also by having long life cyclesgrubs at low dosages (025 kg aii ha) (134) Seed treatments undergoing dormancy or being dispersive Each pest has a are not promising for white grub control (248) unique life history strategy suited to highly unpredictable

Seed treatment is also effective ainst ants and may be environments In agivenyear someinsect specics an better cheaper than increasing the seeding rate In Batangas exploit upland rice as a food source but no pest is a senious treating 50 kg seed ha was as effective as increasing the one every year on upland rice seeding rate to 100 kgha and using untreated seeds (Table The low yielding potential of upland rice means less 9) In Brazil seed treatments control termites and other soil quantity and lower quality of food for pests to exploit but insects including the lessercornstalk borer(66) Domiciano also less incentive for farmers to undertake control (58) however found seed treatments inconsistent in their The most effective control would be early and synchroshyeffect against high populations of lesser corn stalk borer nous planting of early-maturing upland rice varieties to

Baiting can be an inexpensive way of controlling ants prevent insect pest buildup Breeding for insect resistance seedling maggot flies mole crickets and field crickets Baits should initially focus on stem borers because they are verymade of locally available material and impregnated with difficult to control by other means Biocontrol efforts insecticide can be sparingly distributed taking adantage of should first stress conservation of natural enemies preshythe pests ability to disperse and encounter bait sites dators and pathogens rather than parasites Efficient

The new low-volume hand-held controlled-droplet chemical control includes seed treatment or baiting for seed sprayers may offer an advantage over high-volume knap- seedling and some root pests and using controlled dropletsack sprayes for controlling foliar pests Upland rice sprayers for spot treatments against foliar pests and seed canopies are more open than those in lowland rice and bugs when economic thresholds are reached better droplet penetration should result Controlled droplet applicators would be ideal for seedbugs and defoliators

CONCLUSION

Upland rice is attacked by a wider array of insect pests than is lowland rice mainly beca use of the addition of soil pests Generally population levels of lowland rice pests are lower in upland than lowland rice leading many people to conclude that insect pests are not important on upland rice

36 IRPS No 123 January 1987

REFERENCES CITED

1Abraham C C and K S R Mony 1978 Occurrence ofLeptocorisa acuta Fabr (Coreidae Hemiptera) as a pest of nutmeg trees J Bombay Nat ttistSoc 74553

2 Agyen-Sampong M 1975 HlierogrlphusdiaganensisKrauss (Orthop-tera Acrididae) a new pest of rice in Northern Ghana Ghana J Agric Sci 8249-253

3 Agyen-Samnpong M 1976 Varietal resistance and other inle-ratcd pest control components as factors of rice varietal improvement in West Africa Prcsented at the Second varietal improvement serinar 14-18 Sep 1976 Bouak Ivory Coast West Africa Rice Development Association Monrovia Liberia 18 p

4 Agyen-Sampong M 1977 Outbreak of Ihccrorrdnisorae in rice seedlings near Rokupr Sierra Leone lit Rice Res Newsl 2(6)16-17

5 Agyen-Sampong M 1980 Dirty panicles and rice yield reduction caused by hugs IntRice Res Newsl 5(l)11-12

6 Agyen-Sampong M 1982 The major pest problems of irrigated upland and mangrove swanip rice ecosystems ir the humid tropical Guinea savanna and Salel climate zones Pages 318-328 in Pro-ceedings training course cn concepts techniques and application of irtegrated pest management in rice inWest Africa 10-20 Jan 1982 Fendall Liberia West Africa Rice l)evelopioent Association Monrovia Liberia 505 p

7 Akibo-Betts V I and S A Raymundo 1978 Aphids as rice pests in Sierra Leone lit Rice Res Newsl 3(6)15-16

8 Akinsola F A 1975 Present status of different rice stemborers in parts of Nigeria iicL Entomo Newsl 328

9 Akinsola FA 1979 Ihe biology and ecology of rice stemborers in Nigeria Ph 1) thesis University ol lbadan Nigeria 217 p

10 Akinsola E A 1979 Guidelines of integrated management of insect pests of upland rice in West Africa Presented at the Seminar on integrated management of rice diseases and insect pests at Bobo-Dioulasso Upper Volta 17-22 Sep 1979 West Africa Rice Develop-ment Association Monrovia Liberia

II Akinsola IA 1984 Insect pests of upland rice in Africa Pages 301-305 in An overview of upland rice research Proceedings of the 1982 lIouak Iwsrv Coast upland rice workshop International Rice Research Institute 13O Box 933 Manila Philippines

12 Alam M S V TJohn and Kaung Zan 1985 Insect pests and diseases of rice in Africa Pages 67-82 in Rice improvement in Eastern Central and Southern Africa proceedings of the international rice workshop at Iusaka Zambia 9-19 April 1984

13 Alticri M A A van Schoonhovcn and J D Doll 1977 The ecological role of weeds in insect pest management systems a review illustiated with bean (Phaseohsvulgaris L)cropping systems PANS (Pest Artie News Summ) 23195-205

14 Anmante F 1967 The leaf-cutting ant Atia capiguaraa pest of pastures [in Portuguese] Biol6gico (Sao Paulo) 33113-120 (Rev Appl Entomol A56207)

15Andrade A C and D Puzzi 1951 Preliminary results of experi-merits on the control of Sculptcoris iastantus on sugarcane [in Portuguese] Bioulgico (Sao Paulo) 1744-49 (Rev Appl Entomol A41421422)

16 Appert J 1970 Maliarplia separaelha (rice stem borer) new observations of entomological problems of rice in Madagascar [in French] Agron Trop (Paris) 25329-334

17 Ariadeau Mand Z Harahap 1986 Relevant upland rice breeding objectives Pages 189-197 in Progress in upland rice research Proceedings of the 1985 Jakarta conference International Rice Research Institute 1O Box 933 Manila Philippines

18Asar S and Kisimoto 1975 Seasonal occurrence of the smaller brown planthopper in upland rice [in Japanese] Proc Kanto-Tosan Plant Prot Soc 2228

19 Bakker W 1974 Characterization and ecological aspects of rice yellow mottle virus in Kenya Meded Landbouwhogesh Wage-ningen 152 p

20 Balut F F 1970 White grubs attacking rice Oryza ativa L [in Portuguese] Biol6gico (So Paulo) 36321-322

21 Banerjee S N and L M Pramanik 1967 The lepidopterous stalk borers of rice and their life cycles in the tropics pages 103-124 in The major insect pests ofthe rice plant Proceedings ofa symposium at the International Rice Research Institute September 1964 The Johns Hopkins Press Baltimore Maryland USA

22 Barrion A T and J A Litsinger 1980 Ants a natural enemy of cliaiarac in dryland rice IntRice Res Newsl

5(4)22-23 23 Barrion A T and J A litsinger 1981 The spider fauna of

Philippine rice agroecosystem 1Dryland Philipp Entomnol 5139-166

24 Barrion A T and 1 A Iitsinger 1984 The spider fauna of Philippine rice agroccosystcms II Welland Philipp Eiitomol 611-37

25 Barrion A T1 and J A Litsinger 1984 lihanus (Diptera Tabanidae) eggs an alternative host of rice stei borer (SI3) egg parasite Telenoonis digns (Iliymenoptcra Scelionidae) lit Rice Res Newvsl 9(6)19

26 Barrion A T and J A Litsinger 1985 Identilication of rice leaffolders by wing markings Int Rice Res Newsl 10( 1)24

27 Barrion A T and J A Litsinger 1985 Chctiosspp (Coleoptera Carabidae) a leaffolder ILF)predator lit Rice Res Newsl 10(1)21

28 Bertels A 1970 Insect pests of rice and theircontrol [in Portuguese] Pelotas Brazil Instituto de Expcrimentacao Agropccuararias do Sul Circular 43 24 p

29 Bonzi S M 1982 Ctilo (flisilineus J de Joannis (Lepidoptera Pyralidae) stem borer of irrigated cereals in Upper Volta LioFrench] Agron Trop Riu Rizic Cult Vivrieres Trop 37207-209

30 Bourke T V T L Fenner J N LStihick G Baker EHassan D F OSullivan and C S li 1977 Insect pest survey for the year ending 30th June 1969 Papta New Guinea )epartment of Agrishyculture Stock and Fisheries Port Moresby 57 p

31 Bowden 1 1973 Migration of pests in the tropics Meded Rijksfac Landbouwwet Gent 38785-796

32 Breniere J 1973 The main rice parasites in West Africa and their control Pages 27-42 in Proceedings seminar on plant protection for the rice crop West Africa Rice Development Association Monrovia Liberia

33 Breniere J 1976 The principal insect pests of rice in West Africa and their control West Africa Rice Development Association Monrovia Liberia 52 p

34 Brown ES 1962 The African arinyworn Spoiolptera eXI)Ina (Walker) (Lepidoptera Noctuidae) a review of the literature Cornshymonwealth Institute of Entomology London UK57 p

35 Brown F B 1968 Upland rice in Latin America FAO-International Rice Commission 12th Session of the Working Party on Rice Production and Protection 2-14 Sept Peradeniya Sri lanka Food and Agriculture Organization Rome Italy 6 p

36 Buyckx F J 1 1962 Disease and insect pests of crops inCongo Rwanda and Burundi [iil French] Publications dc LInstilut National pour lestude Agrononiquedu Congo M Weissenbruck S A Brussels Belgium 708 p

37 Cariiio F 0 1981 Role of natural enentics inpopulation suppression and pest management of green rice lealhoppers Ph D thesis University ofthe Philippines at Los Bafitos PhilippinesLaguna 136 -

38 Carpente A J 1983 The development of rice cultivation and extension of vegetable productin in Zanzibar Final Report Vol II1 October 1983 Food and Agricultme Organization Ronte Italy

39 Carroll C P and S1 Risch 1983 Tropical annual cropping systems ant ecology Environ Manage 751-57

4(0 Cates R G 198 1 Host plant predictability and the feeding patterns of monophagous oligophagous and polyphagous insect herbivores Oecologia (Berl) 48319-326

IRPS No 123 January 1987 37

41 Cendcafia S M and F B Caora 1967 Insect pests of rice in the 62 Ene 1 C 1981 I)iiorldhol)ierusihilis Slater (licteroplera Philippines Pages 591-616 in The najor insect pests ofthe rice plant lygacidue Illissinac) a new rice pest front Nigeria lit Rice Res Proceedings of a symposium at the International Rice Research Newsl N3)15 Institute September 1964 The Johns Hopkins Press Baltimore 63 Fugenio T S 1965 rhe effect oi e-vCgetation on rice production Maryland USA 729 p and the possibility of overcoming it by leaf pruning MS thesis

42 Centro Ihternacional de Agricklittira Tropical 1984 Upland rice in University ol the Philippines at Los Baijos ILaguna Ihilippins 69 p Iatin Anerica Pages 93-119 hi An os ervies of uplaind rice research 04 Farrow R A 1975 The Alrican nmigratory locust ii is main Proceedings of the 1982 Boinakamp Ivory Coast upland rice workshop outbreak area of the middle Niger qlinlitatise studies of solitary International Rice Research Institute P0 Blox 933 Manila populations in relation to environmental factors Locusla No II Philippines 198 p

43 Chandra G 1979 Taxonomy ad hn riononlics ofthe imect parasites of 65 Fennal R G 1943 Food crop investigation Windward and rice Icalloppers and planthoppers in the Pihilippines and their Ieeward Islands First report Report onl work dunring the periodiiiportairce iii natural biol)gical cointrol Philipp Fntoltol 4119-139 April 1942-April 1943 Imperial College of Tropical Agriculture

44 Chang T T 1976 The originii colut ion enlIivation dissemination Trinidad 16 p (Rev of Appl Fintornol A3222-25) and diersificition of Asian and African rices -uphytica 25425-44 1 66 Ferreira F 1983 Integrated pest ritaageiret of upland rice [ill

45 Chatierjee P B 1979 Rice leaffolder attacks in India lit Rice Res Portuguese English summary] Pages 323-341 in Upland riceculttre Newsl 4(3)21 factors affecting production E Ferreira T Yamada and F-

46 Cheng Cbien-pan 19(4 Upland rice in Taiwan International Rice Malmolta eds Institutoda Potassae Foslato Institnto Internacional Cnmmission Wiorking Party oit Rice Production ard Protection I()th da Potassa Piracicaha Sao Paulo lraid Meeting 3-10 March 1964 Manila Philippines Food and Agricul- 67 Ferreira E arid F dl Silva Martins 1979 Occurrence of ture Organiaion Rome Italy 5 p Sogatohs orvziola (Muir) inupland rice in Goias Ilrail lit Rice

47 Chcrrett 1 M and 1) 1Peregrine 1976 A review of the status of Rcs Ncwssl 4(4)18 leaf-cutting ants and their control Ann Appl Biol 84124-128 68 Ferreira E and 1F da Silva Martiris 1984 Insect pestof rice ill

48 Chipeta F M 1974 Pests of rice in Malawi Pages 90-95 in Brazil and their control Jill Portugucsej FMIlRAtPA-CNPAF Proceedings Sith eastern Africa cereals research conference 10-15 Goiania Brmiil 67 p March 1974 DRB Mialda ed Ministry of Agriculture lilongwe 69 Ferreira FI F ca Silha Miartins S S Nero and 1 It P de Malawi Carvallio 1982 M tltiplc regression analysis todeterimine the effect of

49 Chu Yau-i 1979 The d atiaging period of the rice seedling fly insect populatiois ott upland rice yield [inIPortuguese Pesqui (Atlterigotiavxigu Stcin) (NItiscide Diptera) oil cori) at Fast Java Agropecu Bras 17671-675 Indonesia Plant Prot Bill 21403413 70 Ferreira F 1F da Silva Martins and F 1PZimmermann 1979

50 Conklin H C 1957 IHaninoo agriculture a report on aln integral Rice varietal resistance to tle lesser corn stalk borer [in Port uguese] systent of shi fti rig cult ivationi in the Pltilippincs Food and Agriculture Pesqui Agropccu Biras 14317-321 Organization Rome Italy 209 p 71 Fleming W F 1968 Biological control of lie hapanese beetle U S

51 Cramer H II 1967 Plant proection and world crop production Dep Agric RLs Sers [ech Bull 1383 78 p Pilaniensclhutz-Nachr 201-524 72 Food and Agriculture Organiation of the United Nations 1982

52 l)ani R C and Majunidar 1978 Prelininary observations on tie Country report surmimary Pages 27-37 in Report on the first incidence of root aphids on differeit rice cultnrcs Sci Cult 4488-89 FAOIUN)IPhail and regional training coiuirse in iniproved ctltural

53 Davidson R I 1979 lffects of root feeding on foliage yield Pages practices for upland rice 5-23 Oct 1981 Northern Region Agrculturd117-120 in T K Ci osb and R B Pottinger eds Proceedings of the Developririt Center Chiang Mai Food and Agricuilture Organiiashy2nd Australian conlerence on grassland invertebrate ecology Govt tion Rome Italy 37 n Printer Wellington New Zealand 73 Franco do Ariaral S 195 1 Residual action of insecticides in turfy

54 l)ean G 1W 1976 Rice insect pests inlaos Int Rice Res Newsl soil--the results of a field experiment [in Portuguese] Biol6gico (Sao 1(2)15 Paulo) 17209-214 (Rev Appl Entomol A41423)

55 I)ean G J W 1978 Insect pests of rice in Laos PANS 24(3)280-289 74 Freeman ) 1970 Report onithe Iban london School of -coroiics 56 )ingle H 1984 Behavior genes and lifc histories conplex adapta- Monographs on Social Anthropology No 41 Athlane Press

tions to uncertain environments Pages 169-194 in A new ecology London UK 317 p novel approaches to interactive systems P W Price C N Slobod- 75 Fuchs T W1 A Harding and 1W Smithlr 1979 Induction and chikoff and W S Gaud cdsl ohn Wiley New York USA termination oldiapause in the sugarcane borer Ann Flitonlol Soc

57 Dobelmann-JP and M Falais 1968 Upland rice cultivation ill tie Am 72271-274 North-western part of Madagascar in 1966-1967 [in French English 76 (hbriel B1) 1975 A review tf the major insect pests of some tpland summary] Trop Agron 2313-21 crops in tlc Philippines Pages 86-111 in Ad Htoc panel of experts on

58 Doniiciano N L 1979 Chemical control of tie lesser corn stalk pest disease arnd weed problems in some rainfed crops Bangkok (lolsonno~paloIgnowslhts) (Zeller 1848) (I-epid(ptera Ihycitid ae) Thailand Food aid Agriculture Organization Roire Italy

on rice in Ponta Gressa Parara 1977-1978 [in Portuguese] Pages 77 Garg D K and N K Shal 1983 White grub outbreak on rainfed 291-303 in Anais 1 Reuniao de tecnicos cmiirizicultura do estado de dryland rice in Uttar Pradeslh Int Rice Res Ncwsl 81(4)16-17 Sao Paulo-Campinas Cati Brazil 78 Garg I) K and IP Tandon 1982 Major insect pests of rice on hilly

59 Dunsmore 1 R 1970 Investigations on tlie varieties pests and tracts of Uttar Pradesh India lilt Rice Res Newsl 7(11)11-12diseases of upland rice (hill padi) in Sarawak Malaysia Int Rice 79 Garrity 1) P 1983 Upland rice research an international bibliograpy Comm Newsl 1929-35 1975-1982 Iternational Rice Research hnstitnte PO Box 933

60 Elias 1R1965 The new decline of rice seedlings is not caused by Manila Philippines 121 p parasites [in Portuguese] Biol6gico (Sao Paulo) 313-6 80 Ghauri M S K 1982 A new subspecies of Iinorh)rone bull

61 Empresa Brasiliera de Pesquisa Agropecuaria 1984 Upland rice itt cornlus Slater (Hemiptera Iygaeidae) front Papua New Guinea on Brazil Pages 121-134 it Av overview of upland rice research rice and carpet grass Ihll lEntoniol Res 72133-137 Proceedings ofthe 1982 Blouak Ivory Coast Upland rice workshop 81 Gonzales F 0 PArregoces R I ihernandef and OT Parada International Rice Research Institute PO Box 933 Manila 1983 Insect and imite pests of rice arid their control in Latin America Philippines [in Spanish] la Federacion Nacional de Arrocerts (Fedearroz)

Bogota Colombia 60 n

38 IRPS No 123 January 1987

82 Grist D H and R J A W Lever 1969 Pests of rice Longmans London UK520 p

83 Hale P R 1979 Two subterranean pests of upland rice in Papua New Guirnca IntRice Res Newsl 4(2)16-17

84 Hale P R and J T Hale 1975 Insect pests of rice in the New Guinea Islands Rice Entomol Newsl 34

85 Harris W V 1969 Termites as pests of crops and trees Common-wealth Institute of Entomology London UK 41 p

86 Hattori 1 1968 A method to distinguish between grass leaf roller (Ctaplha rocis medinalisGuenee) and the rice leaf roller (Susurtia exigua Butler) [in Japanese] Plant Prot (Japan) 23167-170

87 Henry I E and 1A Onsager 1984 Experimental control of the mormon cricket Iahrosshnph by NJsena locustae(Aficrospora Microsporidae) a protozoan parasite of grasshoppers (Ort Acri-didae) Entomophaga 27197-201

88 Hidaka 1 P Vungsilabulr and SKadkao 1974 Studies on ecology and control of the rice gall midge in Thailand Tech Bull 6 TropicalAgricultural Research Center Tokyo Japan 113 p

89 Hincklcy A 1) 1963 Ecology and control of rice planthoppers in Fiji Bull Entomrol Res 53467 481

90 Hirao J 1982 Ecology and chtInical control of the rice leaffolder Jpn Pestic Inf 4114-17

91 Ho DT 1984 Sten borer incifience in rice ecosystems in KenyaEast Africa Pages 307-320 in A overview of upland rice research Proceedings of the 1982 Bouac Ivory Coast upland rice workshop International Rice Research Institute PO Box 933 Manila Philippines

92 Ho )T J 1Njokah and J GKibuka 1983 Studies on rice sten borers in Kenya with emphasis on ifaliarpiaipiraellRag Insect Sci Appl 465-73

93 Ichihara I T Ito A NIatsuo and M Sawada 1971 Chemical control of the rice striped stem borer in upland rice grown in plastic mulch [iii Japanese] Proc Kanto-Tosan Plant Prot Soc 1882

94 IclhiharaIand A Matsuo 1969 Damage caused by rice striped stett borer to upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1686-87

95 Ichhara I atid A MIatsun11970 Conside ration oft lie extraordinary prevalence of rice striped stem borer in upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1778-79

96 Institut de Recherches Agrolnliqes Iropicale et des Cultures Vivrieres 1978 Insect resistance Pages 12-13 in Upland rice 1978 atnual report Bouak Ivory Coast 27 p

97 International Institute for Tropical Agriculture 1983 Aphids itt rice roots I ITA Res Briefs 4(1-3)4-5

98 International Institite fi Tropical Agriculture 1983 Hopperburn in rice caused by brown plathopper IITA Res Briefs 4(l-3)6-7

99 International Rice Research Institute 1975 Annual report for 1974 PO Box 933 Manila Philippines p 216

100 International Rice Restarch Institute 1976 Annual repiort for 1975 PO Box 933 Manila Philippines p 228

101 International Rice Research Institute 1979 Cropping systems program Component technology development and evaluation Page 404 i Annual report for 1978 11O Box 933 Manila Philippines

102 International Rice Research Institute Department of Entomology 1981) Insecticid evaluation 1979 Tables 73 74 PO Box 933Manila Philipes

103 International Rice Research Institute Department of Entonmology 1981 Insecticide evalomtion 1980 Table 81 POBox 933 Manila Philippines

104 International Rice Research Institute 1981 Report of a meeting of Asian rice-based cropping systems entomologists Eleventh cropping systems working group meeting 18-22 May 1981 PO Box 933 Manila Philippines p 132 133 144

105 International Rice Research Institute 1984 Cropping systems program Pages 368-373 in Annual report for 1983 PO Box 933 Manila Philippines

106 International Rice Research Institutc 1984 Upland rice in Asia Pages 45-68 in An overview of upland rice research Proceedings of the 1982 Bouakamp Ivory Coast upland rice workiiop loternational Rice Research Institute 11O Box 933 Manila Philippines

107 Irshad M 1977 Grasshoppers in rice in Pakistan lit Rice Res Newsl 2(3)8-9

108 Ito R and TOno 1964 Upland lice its culture and varieties in Japan lIternational Rice Cormmission loth Meeting of the Working Party on Rice Production and Protection 3-10 March 1964 Manila Philippines Food and Agriculture Organization Rome Italy 12 p

109 Jeffrey M I and V A Dyck 1983 Effect of moonlight on lightshytrap catches of brown planthopper lit Rice Res Newsl 8(6) 13

110 Jennings P R and T Alicia Pineda 1983 Management ofthe rice plnthoppers Sogatodsorivi(coIl in Latin America Presented at the meeting on stable plant resistance FAO and lIndocsian I)inectorate of Food Crops Protection Bali lndonesia 29 Novcniber-2 l)ccemshyber 9 p

IllJimenez Gomez J A 1979 Rice insect pests i lla ios ()riCntales lil Spanish] Pages 99-103 in Rice ciltivation Instituto (olotibiatto Agropecuario Compndio No 29

112 Jongeleen F JJ Soesanto and A Mockidjo 1979 Studies oti pathogens of two species of Scarabacidac froinIndonesia Z Angew Enlomnol 87412-417

113 Jouvenaz 1) PC S lofgren and W A Baiks 1981 Biological control of imported fire antsa review of cirrent knowledge Bull Entomol Soc Am 27203-208

114 Kainoh Y K Shiniiu S MarUani Y Taiaki 1980 lost finding behavior of tie rice bu lplotori+JchinvittjI)allas (liciniptera Corcidac) with special reference to diet pattern ol aggregation and feeding on the rice plant Appl Entomol Zool 15225-233

115 Kakkar K 0 P Sharma and P I Sharma 1977 Note on Ihteronyrchus liodresRedt (Coleopicra I)ynastidae) as asporadic pest of paddy and maie inHimachal Pradesh Corr Agric 156-57

116 Kalode M131 79 Major insect pest problems and their control in upland rice (Unpublishcd mineogriph CWI oJNaaiat lni Chandaratat (198 1)original not seen)

117 Kalshoven L i 98 I The pests of crops in I ldoniesia (translated by P A vatn der Laan) 1 T Ichtiar B3aru-vati Iloeve lakarta Indonesia 701 p

118 Kamal N Q 1981 Suppression of whitebacked planto ner Sogatella fircaifra (Horvath) aLId rice lealfolder (nhal -cis licdb lis (Guenee) populations by natural cnclmie hlI) tiesis Grcgorio Araneta University Foundation Victoncta Park Metto Manila Philippines 179 Ip

119 Karim A N M Reaul 1975 A species ofliealy bug Trioni i70s ceres new oii rice in the Philippines Rice Entoniol Newsl 247

120 Katanyukul Wand C Chandaratat 1981 Insect pestsof uplaid rice and their control Report firstFAOUNlFi Tliailand training course on improved cultural practices for tplad rice Chiang Mai Thailand 5-23 Oct 1981 Food and Agriculture Organiation Rome Italy 22 p

121 Kern F 1955 lhletwioa(I)uninemts) bhhit Biturm(Coleoptera) a new pest of sugarcane inVenezuela [inSpanish with Etnglish sunmary] Agron Trop (Maracay) 527-29 (Rev Appl Eitoniol A4572)

122 Khan K M and S Ghai 1974 White grubs ald their control in India Pesticides (1Bombay) 819-25

123 Kiritani K 1975 Integrated pest management of rice pests Pages 66-70 itt Proceedings platning workshop ott cooperative hIeld research in pest managementP SMotooka ed East-West Center Honololu Hawaii USA

124 Kisimoto R 1967 Ecology of insect vectors forecasting and chemical control Pages 243-255 in The virus diseases of tle rice plant Proceedings of a symposium atthe International Rice Research Institute April 1967 The Johns Hopkins Press altimore Maryshyland USA

125 Kisimoto R and V A Dyck 1976 Climate and rice insects Pages 367-391 in Climate and rice International Rice Research Institute PO Box 933 Manila Philippines

126 Kureha S S Kobayashi SAsari and T Ishihara 1977 Life history and control of the mole cricket Grrllotalpa africana Palisot de Beauvois [in Japanese English translation 7520 1RRI library] Proc Kanto-Tosan Plant Prot Soc 21192-197

127 Kushwaha K S 1981 The black beetle as a pest of rice in Haryana India Int Rice Res Newsl 6(4)14

128 Lawani S M 1982 A review of tile effects of various agronomic practices on cereal stem borers Trop Pest Manage 28266-276

129 Leuck D B 1966 Biology of the lesser cornstalk borer in South Georgia J Econ Entomol 59 797-801

130 lever R J A W 1955 Insect pests of the rice plant in Malaya Int Rice Comm Newsl 1612-21

131 Li Ching-Sing and Shin-Foon Chiu 1951 The study ofthe rice gall midge Iachydiplosis orvza Wood Mason [in Chinese with English summary] 1Taiwan Agric Res 21-13

132 Iitsinger J A 1984 Assesament of the need-based application for rice Proceedings TechnologyTransfer Workshop 15-16 March 1984 International Rice Research Institute 1PO Box 933 Manila Philippines 6 p

133 Litsinger J A 1984 Environmental diversity and insect pest abundance with reference to the Pacific Pages 285-307 in Proceedings sub-regional training course on methods of controlling diseases insects and other pests of plants in the South Pacific 4-20 Oct 1982 Vaini Kingdom of Tonga Nukualofa

134 Litsinger J A RF Apostol and M BObusan 1983 White grub Leucopholis irrorata (Coleoptera Scarabaeidae) pest status popula-tion dynamics and chemical conllol in arice-maize cropping pattern in the Philippines J Econ Entoinol 761133-1138

135 Litsinger J A C de la Cruz F Raymundo A T Barrion M D Lumaban J P Bandong M S Venugopal F Paragna and E Balete 1982 Insect pests and insecticide reponse of dry-seeded bunded rice Pages 179-193 in Report of workshop on cropping systems research in Asia International Rice Research Institute Los Bafos Philippine

136 Litsinger J A M 1) Luinaban J P Bandong A T Barrion W Katanyukul and N Panda 1978 Insect pest management in rainfcd lowland rice agroecosystms of tropical Asia International Rice Research Conference 17-21 April 1978 International Rice Research Institute Los Bafios Philippines

137 Litsinger J A and K Moody 1976 Integrated pest management in multiple cropping systems Pages 293-317 in R I Papendick P A Sanchez and G B Triplett eds Multiple cropping American Society of Agronomy Madison Wisconsin USA

138 Loevinsohn M E 1984 The ecology and control of rice pests in relation to the intensity and synchrony of cultivation Ph D thesis University of London 36) p

139 Loevinsohn M E J A Litsinger N Panda M A Austria R F Apostol C Andrion and J Bandong 1982 A basis for predicting rice insect pest abundance in cropping systems Paper presented at the 1982 International rice research conference on rice and cropping systems 19-23 April 1982 International Rice Research Institute Los Bafios Iaguna Philippines 25 p

140 Lopez A W 1930 Repot of the Entomology Department Pages 145-172 in Annual report of the Philippine Sugar Association 1929-30 Manila

141 Lopez AW 1931 Entomology Department Pages 227-266 in Annual report of the director or research 1930-31 Philippine Sugar Association Manila

142 Lopez A W 1932 Entomology Deparment Pages 252-279 in Annual report of the Philippine Sugar Association 1931-32 Manila

143 Lungibill P and G G Ainslee 1917 The lessercornstalk borer US Dep Agric Agric Res Serv Tech Bull 539 27 p

IRPS No 123 January 1987 39

144 McGuire J U Jr and BS Crandall 1967 Survey of insect pests and plant diseases ofselected food crops of Mexico Central America and Panama U SDepartment of Agriculture International Agriculshytural Development Service Agricultuial Researrh Service October 1967 11 p

145 McNeill Sand T R ESouthwood 1978 The role of nitrogen in the development of insectplant relations Pages 77-9 in lliochemical aspects of plant and animal coevolution J B Ilarbornu ed Academic Press London UK

146 Maebara H 1955 Spatial distribution of lastinstiarlanaeof purplish stem borer (Sesamia iqferens Walker) hibernating in an upland rice field Bull Kagoshima Univ Fac Agric 438-41

147 Mallick E H and N G Hajra 1977 Response of rice plant to defoliation treatments at different developmental stages Indian Agric 2151-56

148 Mansingh A 1971 Physiological classification of dormancies in insects Canl Entomol 103983-10)9

149 Martell C G 1974 Catalogue of plant feeding insects of Mexico [in Spanish] Fitofilo 27(69)1-176

150 Martinez 0 J 1982 Principal rice pests in Ilanos orientalis [in Spanish] P~ages 80-94 in Curso deact nalization de conocimientos en arroz Villaniencis Instituto (olonmbiano Agropecuario

151 Martins J F da Silva F Ferrcira A S Prabhu F 1 P Zimmermann 1980 Use of pesticides for the control of the main subterranean upland rice pests esqui Agropecu Bras 1553-u

152 Martins I F da Silva F Fe eira rod BdamSilveira Pinheiro I Q2 Damage simulation caused by defoliators of upland rice [in Portuguese with English summary] Pesqui Agropecu Bras 171113-1119

153 Martins 1F da Silva N(iyuen van Tan and 1Ida Silveira Pinheiro 1981 Stem borer resistance in upland rice associated with plant morphological characters [in Portuuce Iesqui Agropecu Bras 16187-192

154 Masaki S 1980 Sumnmer diapause Ann Rev Entomiol 251-25 155 Masuzawa T 11 Suwa and F Nakasoji 1983 l)ifkleences of

oviposition preferencc and survival rate of two skipper butterflies Parnara guttata and PIchlpidai iathiais (Iepidoptera HIesperiidac) on rice plant and cogon grass New Entomol 3247-56

156 Maurya D M and C P Vaish 1984 Upland rice in India Pages 69-92 in An overview oftpland rice research Proceedings ofthe 1982 Bouakamp Ivory Coast upland rice workshop International Rice Research Institute IO Box 933 Mnila Philippines

157 Menon M G R R N Katiyai S Kuar and S S Misra 1970 Termites attacking the paddy crop Sci Cult 36294-295

158 Miyahara Y and 1 Nishikubo 1967 Behavior of young nymphs of the red rice root aphid Iholahsipiun ri liabhdontinalisSasaki in relation to movement from the leaf to the roots of upland rice [in Japanese] Pioc Assoc Plant Prot Kyushu 1363-66

159 Mo ljoa Tjien 1952 White grub of upland rice (Holotrichiahelleri Brsk) [in Indonesian with English summary] Contrib Agric Res Stn Bogor No 13134 p

160 Mochida 0 and T Suryana 1979 The occurrence of the brown planthopper and ragged stunt on lowland and upland rice ill Central Java and logyakarta in tile late of January 1979 Entomnolgy Survey Report No 15 Central Research Institute of Agriculture (CRIA) Sukamandi Indonesia I February 1979 9 p

161Monet C 1979 Varietal resistance to upland rice pests [inI French] Pages 35-39 in Annual report 1977 Institut des Savanes Department des Cultures Vivrieres Bouak6 Ivory Coast

162 Monte 0 1942 A caterpillar ofrice-fields[in Portuguese] Biol6gico (Sao Paulo) 8161-163 (Rev Appl Entomol A31414)

163 Munroe I) D 1979 Pest management strategy for hill rice in four resettlement areas in Sarawak Malaysia Pages 89-102 i1 Guidelines for integrated control of rice insect pests Food and Agriculture Organization Plant Production and Protection Paper 14 Rome Italy 115 p

40 IRPS No 123 January 1987

164 Nakasuji F M Ishii and Jen-Zon Ito 1984 Rice skippers in Taiwan and their life histories ZZrC (Taipei) Tech Bull 82 10 p

165 Navas I) 1976 Fall armyworum in rice Pages 99-106 il Ecological animal control by habitat management Proc Tall Timbers Con-ference 28 Feb-I Mar 1974 Fall Timbers Res Stn Misc Publ 6 Tallahassee Horida USA

166 Nickel 1 1 1961 Biological control of rice stem borer A feasibility study Tcch Bull 2 International Rice Rcarch Institute PO Box 933 Manila Philippines IIl p

167 Nigan P M and R A Vernia 1985 Insect pests of uplaild rice in Uttar Pradesh Int Rice Res Newsl 10(6)22

168 Nilakhe S S 1985 -cological observations on spittle bugs with emphasis on their occirence in rice Pcsqtui Agropecu Bras 20407-414

169 Nixon G EJ 195 1 lie association of ats with aphids and coccids Commonwealth Iusti lcte of Entonology Iondon UK 36 p

170 Nurdin F an Ishak Manti 1985 Study oit seedling fly of upland rice Interlational Upland Rice Moitoring Tour Report 26 lebruary-2 MarLI Sukaranti West Sumatra

171 Oka 1 N 1979 Ciltural conttol of the brown planthopper Pages 357-369 in Brown planthopper threat to rice production in Asia International Rice Research Institutc 1p0 Box 933 Manila Philippines

172 Ohni M and A Vilnaii 1975 Rice insect pests of Italy biology of Sipha grcria (Kaltenbach) and its control [in Italian] Riso (Milan) 2459-71

173 Otanes F Q atnd 1 I Sison 194 1 Pests of rice Philipp J Agric 12211-259

174 Painter R I 1951 Insect icsistance in crop plants University Press of Kansits Lawrence Kansas USA 520 p

175 Pantua P C and 1A litsinger 1981 Arthropod abundance in continuous vs biannual rice cropping systeis Int Rice Res Ncwsl 6(2)20-21

176 Pathak M ) and V A l)yck 1975 Sttudics ott insect pests of upland rice Pages 186-191 in Major research in upland rice International Rice Research Institute PO Box 933 Ilanila Philippines

177 Patil B R B M IHasabe and 1) S Ajri 198 I Chemical control of white gritb lIditriihinhorraia Fabr intfesting groundntut paddy atnd sorghum Indian 1Entomol 43232-236

178 Pawar A 1) 1974 (ollection and identification of insect pests of rice rlerminal repirt lInternational Rice Research Institute PO Box 933 Manila Philippines 72 p (tnimco)

179 IerrajU A aid A S Reedy 1965 Observations ott diapausc itt tie steti borer ohrice Rice News Teller (lai-Mar)9-13

180 Pont A C 1972 A review of the Oriental species of theri ona Rondani (I)iptera Muscidac) of econtomic importance Pages 27-102 in Control of sorglium shoot fly M G lomtani and W R Young eds Oxford atd IH1 Ptthl Co New I)clhi India 324 p

181 Prasad S C 1PChand aid M 1PSingh 1983 Incidence of rice gall midge in the upland plateau region of Hihar Oryza 20235-236

182 Rai P S 198 0 Nisaga simpex damage to rice in the hill tracts of South India Int Rice kles News[ 5(I)12

183 Rathorc Y S 1969 Gonocplhahli dcprssunt Fabr as itpredator of (tilo zonwIlhis (Swinhoc) and other moths in field cage at Patnagar Uttar Pradesh Indian 1Entotnol 3127

184 Reed W 1983 Crop losses causcd by insect pists in the developing world Proceedings 10th Internatial Congress of Plant Protection lrighton UK 174-80

185 Reedy M V 1981 Ateactogasterinhens(Walker)a new root-grub pest of tipland paddy and inaize Curr Sci 501076

186 Reveno M A 1968 Catalogue of the insect pests of economic crops in Colombia [in Spaiish] Asociacion Latino Americana de Entomo-logia (AIAE) 129-33

187 Reyes G M 1957 Rice dwarf disease in the Philippines Food and Agriculture Organiiation Plant Prot Bull 617-19

188 Rice S E A A Grigarick and M 0 Way 1982 Effect of leaf and panicle feeding byarmyworm (Lepidoptera Noctuidae) larvae on rice grain yield Environ Entomol 7593-595

189 Risch S J 1) Andow and M A Altieri 1983 Agioccosystem diversity arnd pest control data tentative conclusions and nizw research dirctions Environ Entomol 12625-629

190 Rivera C T 1956 Three organic insecticides in thecontrol of insects affecting upland rice Philipp Agric 39465-472

191 Roffrey 1 1972 Locusta outbreaks in the Philippines Acrida 1177-188

192 Rojas R 1) 1941 The effect of pruning at varying ages upon the yield of Ramairio Phipp Agric 29851-860

193 Roonwal M i 1979 Ternite life and termite control in tropical South Asia Scientilic Pnbl odhpr India 139 p

194 Rosetto C1 F da Silva Martins N C Schmidt and L EAzzini 1978 Damage caused by pasturespittle bugs 4Dvoi)lavopictaandDJ schach) on rice lBragantia Nota 6 (Matcli)25-27

195 Rosztto C1- S S Neto I- Amantc I) Iink 1Garcia-Vieira D M de SoLza N V Bansa t and A M de Olivcira 1971 Rice pests of Brazil [in Portuguese] Pages 149-238 inrFA()-International Rice Commission 2nd meeting of the Rice Committee for the Americas 6-11 l)ecember 1971 Pelotas Rio Grande do Sul Brazil

196 Rothschild G H L 1969 Obseratiois on the armyworm Spodoptera matritia acronrctoiiles Gn (Lep Noctuidae) Bull Entomol Res 59143-160

197 Rothschild G 1- L 1974 Parasites of the armworm Spolopiera mauritiaacroncitoihes in Malaysia Fntornopliaga 19293-299

198 Sachan i N S K Gangwar and 1N Katujar 1980 Field crickets damaging cages in North Eastern IHill Region Indian J Entomol 42526-527

199 Sands 1) P A 1977 The biology and ecology of Lepocorisa (Hemiptera Alydidac) in Papua New Guinea Res Bull 18 Departshyment of Primary Industry Port Moresby Papua New Guinea 104 p

200 Sarawak 1968 titomology division Pages 97-106 in Report of the research branch of the Department of Altriculture 1966 Kuching (Rev AIppl Eintomol A56603)

201 Satpathy JM M S Das and K Naik 1977 Effect of multiple and mixed cropping on the incidence of scine important pestsI Entomol Res 178-85

202 Sauer IHF G 1939 Notes Ott Elasmpnlpualtsligniosellths Zeller (Lep Pyr) a serious pest ofcereals in Sao Paolo [in Portuguese] Arq Inst Biol 10199-206 (Rev Appl Entotnol A28378)

203 Satinders 1L A 1 SKing and C L Vargas S 1983 Crop pests of Central Anerica with t bibliography [in Spanish] Centro Agroshynotnico Tropical de Investigacion y Ensenanza Turrialba Costa Rica (CATIE) Tec Bol 9 90 p

204 Sauphanor B 1985 Some factors of upland rice tolerttce to stem borers in West Africa Inseu Sci Appl 6429434

205 Sen P atnd S Chakravorty 1976 Stentflics in Haritighata Nadia West Bengal India [it Rice Res Newsl 1(12) 15

206 Senapai B and 1M Satpathy 1983 The seedling fly Atherigona orrzae Malloch is a pest of rice in Orissa (India) J Entomol Res 777-79

207 Shhjahan M S Alatn M Ah nad and BN Islam 1983 A new scarabaeid beetle pest of dryland rice in Bangladesh Int Rice Res Ncwsl 8(I)16-17

208 Sharma G L S C Modgol and R C Gautam 1979 Effect of depth of submergence ott incidence of bacterial blight and white grub infestation in transplanted rice Int Rice Res Newsl 4(6)12-13

209 Sharma S 1985 Scarabaeid beetle outbreak itt upland rice Int Rice Res Newsl 1(1(l)24

210 Shiraki T 1917 The paddy borer Schoenobhits incertulas Walker [achaku Agric Exp Stn Formosa 256 p

211 Silva A de Brito and 11P Magalhaes 1981 Insect pests of rice [in Portuguese] Circular Tecnica No 22 EMBRAPA Centro de Pesquisa t ecuaria de Tropico Unido (CPAJU) Belem Brazil 14 p

212 Singh S A 1979 Emergence of rice leaf butterfly as a rice pest in Manipur India int Rice Res Newsl 4(4)16-17

213 SinghS A 1983 Changing trends of insect pestsof rice in Manipur Int Rice Res Newsl 8(f)12

214 Siwi S S and P H van Doesburg 1984 Lenovorisa Latreille in Indonesia (Heteroptera Coreidae Alydinae) Zool Meded (Leiden) 58117-129

215 Soejitno J 1977 Reaction of some upland rice varieties toseedling fly eltherikonaexigua Stein [in Indonesian] Siri Hana I Penyakit 630-35 Lembga I osat Fenclilian Pertanian Bogor Indonesia

216 Socnardi 1967 Insect pests of rice in Indonesia Pages675-683 in The major inse pests of thu rice plant Proceedings of a symposium at the Internatital Rice Rec Wi itt S pternber 1964 The Johns Hopkins Press Baltimore Maryland USA

217 Soto P E A T Petcz and I W Buddenhagen 1976 Survey of insect pests and diseases of rice in different ecological zones in Nigeria Rice Entomol Ncwsl 435-46

218 Soto 11E and Z Ziddiqi 1976 Screening for resistance to African rice insect pests Varietal improsetnent sCmi n r Bouak Ivory Coast 13-17 Sept 1976 West Africa Rice l)evelopment Association Monrovia Iiberia

219 Squire F A 1972 Entomological problems in Bolivia PANS (PestArtic News Sumr) 18249-268

220 Stapely 1 II 1978 The rice leaf roller in the Solomon Islands Int Rice Res Newsl 3(l)13

221 Suenaga H 1937 On aphids attacking the roots of rice Oyo-I)oubts Zasshi 9150-152 [in Japanese] (Rev Appl Entomol A25745)

222 Suenaga H 1952 On the seasonal abundance of three subterranean aphids on upland rice Bull Kyushu Agric Exp Sti 1249-262

223 Sutynso Yas and Dandi Soekaria 1964 Beberapa hana yang ttenperang tana man padi tanah kering [in Indonesian] Rapat kerja Padi Tanah Kering 6 sd 8 Juily 1964 Kaliurang 9 p

224 Tallamy I) W and R F Denno 1983 Alternative life history patterns in risky environments in example from lace bugs Pages 129-147 in Insect life history patterns R F l)cnio and If Dingle eds Spring-Verlag 225 p

225 Tamura 1 1939 Observations on some insect pests of field crops [in Japanese] 1 Plant Prot 26414-420 (Rev Appl EntomolA27-556)

226 Tamura and Suzuki 1964 Influences of the early-transplanting practice in rice culture upon development of the rice stem borer Chilo siql)ressalisWalker Bull Ilokuriku Natil Agric Exp Stn 761-94

227 Tan J P 1924 The rice root aphid Drowhi ihirsoia Philipp Agric 13277-288

228 Tanaka 1 1957 Taxonomy and distribution of some subterranean aphids injurious to upland rice in Japan with description of a new species [in Japanese English summary] Botyu-Kagaku 22168-176

229 Tanaka T 1961 The rice root aphids their ecology and control Spec [full College of Agriculture Utsunomiya University Japan F3 p

230 Tauber M IC A Tauber R Nechols and 1Obrycki 1983 Seasonal activity of parasitoids control by external internal and genetic factors Pages 87-108 itt I)iapause and life cycle strategies in insects V K Brown and I Ilodek eds )r W Junk The Hague Netherlands

231 Taylor W F 1972 Effects of artificial defoliation (simulating pest damage) on varieties of upland rice Exp Agric 879-83

232 feotia T P S and S Natud 1966 Biotoomics of the rice skipper Parnaraniathiis Fabricius (Lepidoptera Hesperiidac) Indian J Entomol 28181-186

233 Teran F 0 1971 Rice pests of Santa Cruz [in Spanish Ministerio de Agricultura Sita Cruz Bolivia Bull 45 8 p

234 Than Itun 1976 Population dynamics of the yellow rice borer Tryporza itcertulas (Walker) and its damage to the rice plant MS thesis University of the Philippines at Ios Bafios Laguna Philippines 92 p

235 Uichanco L B 1911 Pests Pages 55-74 it Rice Philipp Agric 11-200

IRPS No 123 January 1987 41

236 van der Laan P A 195 1 Life history of the rice seedling fly Contrib Gen Agric Res Stn Boor Indonesia 118 15 p

237 van der Laan 11A 1959 Correlation between rainfall in the dry season and the occurrence of the white rice borer (Scirpolhaga innotala WIk) in lava Entomol Fxp Appl 212-20

238 Van Driesche R G 1983 Meaning of percent parasitismin studies of insect parasitism Fnviron Entomol 121 I1- 122

239 Venugopal M S and A litsingcr 1984 Fffect ofcarbofuran on rice growth Prot Ecol 7313-317

240 Vietnam Directorate of Rural Affairs 1962 Annual work progress report on crop improvement program of rice sugarcane vegetable and field crops (lor the period from July 1961 to June 1962) (11) 299 p (Rev Appl Entomol A51526-527)

241 Wan M T K 1972 Observations on rice leaf- and planthoppers in Sarawak (Malaysian Borneo) Malay Agric 148308-335

242 West Africa Rice I)esclopnient Association 1978 Annual research report Vol 2 Statistics Monrovia Liberia

243 West Africa Rice Development Association 1979 Annual research report p47 Monrovia Liberia

244 Williams ) F and CS lofgren 1981 Fli I illy l-468 anew bait toxicant for control of the red imported lire at Fla Entomol 64472477

245 Williams I) 1 1970 The mealybugs OFlomoptera Coccoidea Iscudococcidac) of sugar-cane rice and sorghum Bull Entomol Res 0109-188

246 Williams D) 1 1982 Pulinaria icervi (Signoret) and its allies ott sugar cane and other grasses Btll Entoinol Res 72111-117

247 Wirjosuiardjo S 1976 Study ofchenmical cont rot of insect pests ott upland rice Imiu Pertanian (Jogjakarta) 1323-329

248 Wirjosuhardjo S and N Wirarta 1976 Chemical control of the white grub tIolorithiatlltriin upland rice in M uneng Irobolinggo Indonesia) [in Indonesian] Imut Pertanian (Jogjakarta) 1330-332

249 Yano K T Miyake and V F Eastop 1983 The biology and economic importance of rice aphids (Henuiptera Aphid idae) a review Bull Entomol Res 73539-566

250 Yelp D and P Rhodes 19(4 lhe success of drilled dryland rice iii the Fiji Islands International Rice Commission Working Party on Rice Production and Protection Manila Philippines 3-10 March1964 Food and Agriculture Organization Rome Italy 13 p

251 Young W R 1981 Fifty-five years of research on the sorghum shootfly Insect Sci Appi 23-9

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No 101The economics of hybrid rice production in (hina No 113 Boron toxicity in rice No 102 Rice ratooning No 114 Energy analysis rice production systems and rice research No 103 Growth and development of the (Jeep water rice plant No 115 Production risk and optimal fertilizer rates an application of the

No 104 Faridpur acomputer-assisted instruction nmodel forrainfed lowland random coefficient model No 116 Consumer demand for rice grain quality in Thailand Indonesia andrice

No 105 A readingand listening comprehcnsion test inEnglish for non native the Philippines speakers applying for tiraining at IRIRI No 117 Morphological changes in rice panicle development a review of

No 106 Rice grassy stunt virus 2 a new strain of rice grassy stunt in tire literature

Philippines No 118 IRRI-Korca collaborative project for the development of cold-No 107 Physical losses and quality deterioration in rice plstprodnCtion tolerant lines through anthei culture

systemis No 119 Problen soils as potenial areas for adverse soils-tolerant rice No 108 (olptblication ofIRRI materials a survey of translators and varieties in South and Ssuthcasi Asia

publishers No 120 Changes isinall-farm iLtireliing techii)hgy in Thailand and [he No 109 Classification of Philippine rainfall patterns Philippines No 110 Contributions of modern rice varieties to rutrition in Asia No 121 Landforms and modern rice varieties No Ill Changes in rice breeding in 10 Asian countries 1965-84 No 122 Yield stability and modern rice technology No 112 Design parameters affecting the performance of the IRRI-designed

axial-flow pump

Page 35: UPLANDRICE. INSECTl--1i M T -E!R.EOLOU t RTANCE ,AND …delphacid.s3.amazonaws.com › 6958.pdf · The significance of abiotic factors in upland rice insect ecology will be apparent

IRPS No 123 January 1987 35

Table 9 Effect of seeding rate and insecticide on plant density and Yield losses from insect pests in upland rice often areyield of Dagge upland rice Batangas Philippines 1976a comparable to those of lowland rainfed or irrigated rice

Seeding rate Insecticide Dosage Plant density Yield (0-Y30o) but usually from different pests The principal (kg aiha) seed treatment (kg aiha) (noIra-row) (tha) groups are soil pests feeding on sown seed and rootsat 14 DE followed by seedling pests These groups have largely gone

50 None - 35 a 20 a unnoticed leading researchers to conclude that stem borers50 Carbofuran F 10 38 ab 23 a and seed bugs are the most important pests (17) Yield loss 10 beCabofuranF100 None - 4551 c 23 a trials that pinpoint major pests need to includc seed

100 Dieldrin WP 10 52 cd 22 a treatments Granules applied in the furrow may not give150 None - 59 d 22 a adequate protection from many seed pests

aAv of 8 fields DE = days after crop emergence In a column means The floristic and pedological diversity of upland rice followed by a common letter are not significantly different at the environments is matched by the diversity of arthropods5level by D)MRT F = flowable Wi = wettable powder both pests and their natural enemies Irrigation has homoshy

genized lowland rice environments Flooded soils tend tothreatened by epidemics of locusts or armyworms New have the same properties and to eliminate soil pests The technology stressing efficiency has minimized insecticide spatial and temporal dominance of cultivated rice has cost benefited specialist pests

Seed treatment is normally inexpensive and can be Upland rice has no specialist arthropod species The economicallyjustified in -nanycases(66) Treating seed with monophagous pests on upland rice prefer the lowlands systemic insecticide (216) but not chlorinated hydrocarbons where rice evolved Rice in the uplands exists only because (247) is most effective in controlling seedling maggots of man and even though lowland rice predates upland riceFoliar sprays have to be repeated to obtain control and specialist species evolved Wheat maize and sorghum do often are washed off by frequent early season rains not have monophagous pests leading us to conclude that Granular insecticides are effective but required dosages are there is no evolutionary advantage for inssects to have too expensive to justify except under heavy infestation narrow host ranges in upland graminaceae(247) When directed against first-instar larvae banding Pests have adapted to the uplands not only by having granules in seed furrows (177) can be effective against white wider plant host ranges but also by having long life cyclesgrubs at low dosages (025 kg aii ha) (134) Seed treatments undergoing dormancy or being dispersive Each pest has a are not promising for white grub control (248) unique life history strategy suited to highly unpredictable

Seed treatment is also effective ainst ants and may be environments In agivenyear someinsect specics an better cheaper than increasing the seeding rate In Batangas exploit upland rice as a food source but no pest is a senious treating 50 kg seed ha was as effective as increasing the one every year on upland rice seeding rate to 100 kgha and using untreated seeds (Table The low yielding potential of upland rice means less 9) In Brazil seed treatments control termites and other soil quantity and lower quality of food for pests to exploit but insects including the lessercornstalk borer(66) Domiciano also less incentive for farmers to undertake control (58) however found seed treatments inconsistent in their The most effective control would be early and synchroshyeffect against high populations of lesser corn stalk borer nous planting of early-maturing upland rice varieties to

Baiting can be an inexpensive way of controlling ants prevent insect pest buildup Breeding for insect resistance seedling maggot flies mole crickets and field crickets Baits should initially focus on stem borers because they are verymade of locally available material and impregnated with difficult to control by other means Biocontrol efforts insecticide can be sparingly distributed taking adantage of should first stress conservation of natural enemies preshythe pests ability to disperse and encounter bait sites dators and pathogens rather than parasites Efficient

The new low-volume hand-held controlled-droplet chemical control includes seed treatment or baiting for seed sprayers may offer an advantage over high-volume knap- seedling and some root pests and using controlled dropletsack sprayes for controlling foliar pests Upland rice sprayers for spot treatments against foliar pests and seed canopies are more open than those in lowland rice and bugs when economic thresholds are reached better droplet penetration should result Controlled droplet applicators would be ideal for seedbugs and defoliators

CONCLUSION

Upland rice is attacked by a wider array of insect pests than is lowland rice mainly beca use of the addition of soil pests Generally population levels of lowland rice pests are lower in upland than lowland rice leading many people to conclude that insect pests are not important on upland rice

36 IRPS No 123 January 1987

REFERENCES CITED

1Abraham C C and K S R Mony 1978 Occurrence ofLeptocorisa acuta Fabr (Coreidae Hemiptera) as a pest of nutmeg trees J Bombay Nat ttistSoc 74553

2 Agyen-Sampong M 1975 HlierogrlphusdiaganensisKrauss (Orthop-tera Acrididae) a new pest of rice in Northern Ghana Ghana J Agric Sci 8249-253

3 Agyen-Samnpong M 1976 Varietal resistance and other inle-ratcd pest control components as factors of rice varietal improvement in West Africa Prcsented at the Second varietal improvement serinar 14-18 Sep 1976 Bouak Ivory Coast West Africa Rice Development Association Monrovia Liberia 18 p

4 Agyen-Sampong M 1977 Outbreak of Ihccrorrdnisorae in rice seedlings near Rokupr Sierra Leone lit Rice Res Newsl 2(6)16-17

5 Agyen-Sampong M 1980 Dirty panicles and rice yield reduction caused by hugs IntRice Res Newsl 5(l)11-12

6 Agyen-Sampong M 1982 The major pest problems of irrigated upland and mangrove swanip rice ecosystems ir the humid tropical Guinea savanna and Salel climate zones Pages 318-328 in Pro-ceedings training course cn concepts techniques and application of irtegrated pest management in rice inWest Africa 10-20 Jan 1982 Fendall Liberia West Africa Rice l)evelopioent Association Monrovia Liberia 505 p

7 Akibo-Betts V I and S A Raymundo 1978 Aphids as rice pests in Sierra Leone lit Rice Res Newsl 3(6)15-16

8 Akinsola F A 1975 Present status of different rice stemborers in parts of Nigeria iicL Entomo Newsl 328

9 Akinsola FA 1979 Ihe biology and ecology of rice stemborers in Nigeria Ph 1) thesis University ol lbadan Nigeria 217 p

10 Akinsola E A 1979 Guidelines of integrated management of insect pests of upland rice in West Africa Presented at the Seminar on integrated management of rice diseases and insect pests at Bobo-Dioulasso Upper Volta 17-22 Sep 1979 West Africa Rice Develop-ment Association Monrovia Liberia

II Akinsola IA 1984 Insect pests of upland rice in Africa Pages 301-305 in An overview of upland rice research Proceedings of the 1982 lIouak Iwsrv Coast upland rice workshop International Rice Research Institute 13O Box 933 Manila Philippines

12 Alam M S V TJohn and Kaung Zan 1985 Insect pests and diseases of rice in Africa Pages 67-82 in Rice improvement in Eastern Central and Southern Africa proceedings of the international rice workshop at Iusaka Zambia 9-19 April 1984

13 Alticri M A A van Schoonhovcn and J D Doll 1977 The ecological role of weeds in insect pest management systems a review illustiated with bean (Phaseohsvulgaris L)cropping systems PANS (Pest Artie News Summ) 23195-205

14 Anmante F 1967 The leaf-cutting ant Atia capiguaraa pest of pastures [in Portuguese] Biol6gico (Sao Paulo) 33113-120 (Rev Appl Entomol A56207)

15Andrade A C and D Puzzi 1951 Preliminary results of experi-merits on the control of Sculptcoris iastantus on sugarcane [in Portuguese] Bioulgico (Sao Paulo) 1744-49 (Rev Appl Entomol A41421422)

16 Appert J 1970 Maliarplia separaelha (rice stem borer) new observations of entomological problems of rice in Madagascar [in French] Agron Trop (Paris) 25329-334

17 Ariadeau Mand Z Harahap 1986 Relevant upland rice breeding objectives Pages 189-197 in Progress in upland rice research Proceedings of the 1985 Jakarta conference International Rice Research Institute 1O Box 933 Manila Philippines

18Asar S and Kisimoto 1975 Seasonal occurrence of the smaller brown planthopper in upland rice [in Japanese] Proc Kanto-Tosan Plant Prot Soc 2228

19 Bakker W 1974 Characterization and ecological aspects of rice yellow mottle virus in Kenya Meded Landbouwhogesh Wage-ningen 152 p

20 Balut F F 1970 White grubs attacking rice Oryza ativa L [in Portuguese] Biol6gico (So Paulo) 36321-322

21 Banerjee S N and L M Pramanik 1967 The lepidopterous stalk borers of rice and their life cycles in the tropics pages 103-124 in The major insect pests ofthe rice plant Proceedings ofa symposium at the International Rice Research Institute September 1964 The Johns Hopkins Press Baltimore Maryland USA

22 Barrion A T and J A Litsinger 1980 Ants a natural enemy of cliaiarac in dryland rice IntRice Res Newsl

5(4)22-23 23 Barrion A T and J A litsinger 1981 The spider fauna of

Philippine rice agroecosystem 1Dryland Philipp Entomnol 5139-166

24 Barrion A T and 1 A Iitsinger 1984 The spider fauna of Philippine rice agroccosystcms II Welland Philipp Eiitomol 611-37

25 Barrion A T1 and J A Litsinger 1984 lihanus (Diptera Tabanidae) eggs an alternative host of rice stei borer (SI3) egg parasite Telenoonis digns (Iliymenoptcra Scelionidae) lit Rice Res Newvsl 9(6)19

26 Barrion A T and J A Litsinger 1985 Identilication of rice leaffolders by wing markings Int Rice Res Newsl 10( 1)24

27 Barrion A T and J A Litsinger 1985 Chctiosspp (Coleoptera Carabidae) a leaffolder ILF)predator lit Rice Res Newsl 10(1)21

28 Bertels A 1970 Insect pests of rice and theircontrol [in Portuguese] Pelotas Brazil Instituto de Expcrimentacao Agropccuararias do Sul Circular 43 24 p

29 Bonzi S M 1982 Ctilo (flisilineus J de Joannis (Lepidoptera Pyralidae) stem borer of irrigated cereals in Upper Volta LioFrench] Agron Trop Riu Rizic Cult Vivrieres Trop 37207-209

30 Bourke T V T L Fenner J N LStihick G Baker EHassan D F OSullivan and C S li 1977 Insect pest survey for the year ending 30th June 1969 Papta New Guinea )epartment of Agrishyculture Stock and Fisheries Port Moresby 57 p

31 Bowden 1 1973 Migration of pests in the tropics Meded Rijksfac Landbouwwet Gent 38785-796

32 Breniere J 1973 The main rice parasites in West Africa and their control Pages 27-42 in Proceedings seminar on plant protection for the rice crop West Africa Rice Development Association Monrovia Liberia

33 Breniere J 1976 The principal insect pests of rice in West Africa and their control West Africa Rice Development Association Monrovia Liberia 52 p

34 Brown ES 1962 The African arinyworn Spoiolptera eXI)Ina (Walker) (Lepidoptera Noctuidae) a review of the literature Cornshymonwealth Institute of Entomology London UK57 p

35 Brown F B 1968 Upland rice in Latin America FAO-International Rice Commission 12th Session of the Working Party on Rice Production and Protection 2-14 Sept Peradeniya Sri lanka Food and Agriculture Organization Rome Italy 6 p

36 Buyckx F J 1 1962 Disease and insect pests of crops inCongo Rwanda and Burundi [iil French] Publications dc LInstilut National pour lestude Agrononiquedu Congo M Weissenbruck S A Brussels Belgium 708 p

37 Cariiio F 0 1981 Role of natural enentics inpopulation suppression and pest management of green rice lealhoppers Ph D thesis University ofthe Philippines at Los Bafitos PhilippinesLaguna 136 -

38 Carpente A J 1983 The development of rice cultivation and extension of vegetable productin in Zanzibar Final Report Vol II1 October 1983 Food and Agricultme Organization Ronte Italy

39 Carroll C P and S1 Risch 1983 Tropical annual cropping systems ant ecology Environ Manage 751-57

4(0 Cates R G 198 1 Host plant predictability and the feeding patterns of monophagous oligophagous and polyphagous insect herbivores Oecologia (Berl) 48319-326

IRPS No 123 January 1987 37

41 Cendcafia S M and F B Caora 1967 Insect pests of rice in the 62 Ene 1 C 1981 I)iiorldhol)ierusihilis Slater (licteroplera Philippines Pages 591-616 in The najor insect pests ofthe rice plant lygacidue Illissinac) a new rice pest front Nigeria lit Rice Res Proceedings of a symposium at the International Rice Research Newsl N3)15 Institute September 1964 The Johns Hopkins Press Baltimore 63 Fugenio T S 1965 rhe effect oi e-vCgetation on rice production Maryland USA 729 p and the possibility of overcoming it by leaf pruning MS thesis

42 Centro Ihternacional de Agricklittira Tropical 1984 Upland rice in University ol the Philippines at Los Baijos ILaguna Ihilippins 69 p Iatin Anerica Pages 93-119 hi An os ervies of uplaind rice research 04 Farrow R A 1975 The Alrican nmigratory locust ii is main Proceedings of the 1982 Boinakamp Ivory Coast upland rice workshop outbreak area of the middle Niger qlinlitatise studies of solitary International Rice Research Institute P0 Blox 933 Manila populations in relation to environmental factors Locusla No II Philippines 198 p

43 Chandra G 1979 Taxonomy ad hn riononlics ofthe imect parasites of 65 Fennal R G 1943 Food crop investigation Windward and rice Icalloppers and planthoppers in the Pihilippines and their Ieeward Islands First report Report onl work dunring the periodiiiportairce iii natural biol)gical cointrol Philipp Fntoltol 4119-139 April 1942-April 1943 Imperial College of Tropical Agriculture

44 Chang T T 1976 The originii colut ion enlIivation dissemination Trinidad 16 p (Rev of Appl Fintornol A3222-25) and diersificition of Asian and African rices -uphytica 25425-44 1 66 Ferreira F 1983 Integrated pest ritaageiret of upland rice [ill

45 Chatierjee P B 1979 Rice leaffolder attacks in India lit Rice Res Portuguese English summary] Pages 323-341 in Upland riceculttre Newsl 4(3)21 factors affecting production E Ferreira T Yamada and F-

46 Cheng Cbien-pan 19(4 Upland rice in Taiwan International Rice Malmolta eds Institutoda Potassae Foslato Institnto Internacional Cnmmission Wiorking Party oit Rice Production ard Protection I()th da Potassa Piracicaha Sao Paulo lraid Meeting 3-10 March 1964 Manila Philippines Food and Agricul- 67 Ferreira E arid F dl Silva Martins 1979 Occurrence of ture Organiaion Rome Italy 5 p Sogatohs orvziola (Muir) inupland rice in Goias Ilrail lit Rice

47 Chcrrett 1 M and 1) 1Peregrine 1976 A review of the status of Rcs Ncwssl 4(4)18 leaf-cutting ants and their control Ann Appl Biol 84124-128 68 Ferreira E and 1F da Silva Martiris 1984 Insect pestof rice ill

48 Chipeta F M 1974 Pests of rice in Malawi Pages 90-95 in Brazil and their control Jill Portugucsej FMIlRAtPA-CNPAF Proceedings Sith eastern Africa cereals research conference 10-15 Goiania Brmiil 67 p March 1974 DRB Mialda ed Ministry of Agriculture lilongwe 69 Ferreira FI F ca Silha Miartins S S Nero and 1 It P de Malawi Carvallio 1982 M tltiplc regression analysis todeterimine the effect of

49 Chu Yau-i 1979 The d atiaging period of the rice seedling fly insect populatiois ott upland rice yield [inIPortuguese Pesqui (Atlterigotiavxigu Stcin) (NItiscide Diptera) oil cori) at Fast Java Agropecu Bras 17671-675 Indonesia Plant Prot Bill 21403413 70 Ferreira F 1F da Silva Martins and F 1PZimmermann 1979

50 Conklin H C 1957 IHaninoo agriculture a report on aln integral Rice varietal resistance to tle lesser corn stalk borer [in Port uguese] systent of shi fti rig cult ivationi in the Pltilippincs Food and Agriculture Pesqui Agropccu Biras 14317-321 Organization Rome Italy 209 p 71 Fleming W F 1968 Biological control of lie hapanese beetle U S

51 Cramer H II 1967 Plant proection and world crop production Dep Agric RLs Sers [ech Bull 1383 78 p Pilaniensclhutz-Nachr 201-524 72 Food and Agriculture Organiation of the United Nations 1982

52 l)ani R C and Majunidar 1978 Prelininary observations on tie Country report surmimary Pages 27-37 in Report on the first incidence of root aphids on differeit rice cultnrcs Sci Cult 4488-89 FAOIUN)IPhail and regional training coiuirse in iniproved ctltural

53 Davidson R I 1979 lffects of root feeding on foliage yield Pages practices for upland rice 5-23 Oct 1981 Northern Region Agrculturd117-120 in T K Ci osb and R B Pottinger eds Proceedings of the Developririt Center Chiang Mai Food and Agricuilture Organiiashy2nd Australian conlerence on grassland invertebrate ecology Govt tion Rome Italy 37 n Printer Wellington New Zealand 73 Franco do Ariaral S 195 1 Residual action of insecticides in turfy

54 l)ean G 1W 1976 Rice insect pests inlaos Int Rice Res Newsl soil--the results of a field experiment [in Portuguese] Biol6gico (Sao 1(2)15 Paulo) 17209-214 (Rev Appl Entomol A41423)

55 I)ean G J W 1978 Insect pests of rice in Laos PANS 24(3)280-289 74 Freeman ) 1970 Report onithe Iban london School of -coroiics 56 )ingle H 1984 Behavior genes and lifc histories conplex adapta- Monographs on Social Anthropology No 41 Athlane Press

tions to uncertain environments Pages 169-194 in A new ecology London UK 317 p novel approaches to interactive systems P W Price C N Slobod- 75 Fuchs T W1 A Harding and 1W Smithlr 1979 Induction and chikoff and W S Gaud cdsl ohn Wiley New York USA termination oldiapause in the sugarcane borer Ann Flitonlol Soc

57 Dobelmann-JP and M Falais 1968 Upland rice cultivation ill tie Am 72271-274 North-western part of Madagascar in 1966-1967 [in French English 76 (hbriel B1) 1975 A review tf the major insect pests of some tpland summary] Trop Agron 2313-21 crops in tlc Philippines Pages 86-111 in Ad Htoc panel of experts on

58 Doniiciano N L 1979 Chemical control of tie lesser corn stalk pest disease arnd weed problems in some rainfed crops Bangkok (lolsonno~paloIgnowslhts) (Zeller 1848) (I-epid(ptera Ihycitid ae) Thailand Food aid Agriculture Organization Roire Italy

on rice in Ponta Gressa Parara 1977-1978 [in Portuguese] Pages 77 Garg D K and N K Shal 1983 White grub outbreak on rainfed 291-303 in Anais 1 Reuniao de tecnicos cmiirizicultura do estado de dryland rice in Uttar Pradeslh Int Rice Res Ncwsl 81(4)16-17 Sao Paulo-Campinas Cati Brazil 78 Garg I) K and IP Tandon 1982 Major insect pests of rice on hilly

59 Dunsmore 1 R 1970 Investigations on tlie varieties pests and tracts of Uttar Pradesh India lilt Rice Res Newsl 7(11)11-12diseases of upland rice (hill padi) in Sarawak Malaysia Int Rice 79 Garrity 1) P 1983 Upland rice research an international bibliograpy Comm Newsl 1929-35 1975-1982 Iternational Rice Research hnstitnte PO Box 933

60 Elias 1R1965 The new decline of rice seedlings is not caused by Manila Philippines 121 p parasites [in Portuguese] Biol6gico (Sao Paulo) 313-6 80 Ghauri M S K 1982 A new subspecies of Iinorh)rone bull

61 Empresa Brasiliera de Pesquisa Agropecuaria 1984 Upland rice itt cornlus Slater (Hemiptera Iygaeidae) front Papua New Guinea on Brazil Pages 121-134 it Av overview of upland rice research rice and carpet grass Ihll lEntoniol Res 72133-137 Proceedings ofthe 1982 Blouak Ivory Coast Upland rice workshop 81 Gonzales F 0 PArregoces R I ihernandef and OT Parada International Rice Research Institute PO Box 933 Manila 1983 Insect and imite pests of rice arid their control in Latin America Philippines [in Spanish] la Federacion Nacional de Arrocerts (Fedearroz)

Bogota Colombia 60 n

38 IRPS No 123 January 1987

82 Grist D H and R J A W Lever 1969 Pests of rice Longmans London UK520 p

83 Hale P R 1979 Two subterranean pests of upland rice in Papua New Guirnca IntRice Res Newsl 4(2)16-17

84 Hale P R and J T Hale 1975 Insect pests of rice in the New Guinea Islands Rice Entomol Newsl 34

85 Harris W V 1969 Termites as pests of crops and trees Common-wealth Institute of Entomology London UK 41 p

86 Hattori 1 1968 A method to distinguish between grass leaf roller (Ctaplha rocis medinalisGuenee) and the rice leaf roller (Susurtia exigua Butler) [in Japanese] Plant Prot (Japan) 23167-170

87 Henry I E and 1A Onsager 1984 Experimental control of the mormon cricket Iahrosshnph by NJsena locustae(Aficrospora Microsporidae) a protozoan parasite of grasshoppers (Ort Acri-didae) Entomophaga 27197-201

88 Hidaka 1 P Vungsilabulr and SKadkao 1974 Studies on ecology and control of the rice gall midge in Thailand Tech Bull 6 TropicalAgricultural Research Center Tokyo Japan 113 p

89 Hincklcy A 1) 1963 Ecology and control of rice planthoppers in Fiji Bull Entomrol Res 53467 481

90 Hirao J 1982 Ecology and chtInical control of the rice leaffolder Jpn Pestic Inf 4114-17

91 Ho DT 1984 Sten borer incifience in rice ecosystems in KenyaEast Africa Pages 307-320 in A overview of upland rice research Proceedings of the 1982 Bouac Ivory Coast upland rice workshop International Rice Research Institute PO Box 933 Manila Philippines

92 Ho )T J 1Njokah and J GKibuka 1983 Studies on rice sten borers in Kenya with emphasis on ifaliarpiaipiraellRag Insect Sci Appl 465-73

93 Ichihara I T Ito A NIatsuo and M Sawada 1971 Chemical control of the rice striped stem borer in upland rice grown in plastic mulch [iii Japanese] Proc Kanto-Tosan Plant Prot Soc 1882

94 IclhiharaIand A Matsuo 1969 Damage caused by rice striped stett borer to upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1686-87

95 Ichhara I atid A MIatsun11970 Conside ration oft lie extraordinary prevalence of rice striped stem borer in upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1778-79

96 Institut de Recherches Agrolnliqes Iropicale et des Cultures Vivrieres 1978 Insect resistance Pages 12-13 in Upland rice 1978 atnual report Bouak Ivory Coast 27 p

97 International Institute for Tropical Agriculture 1983 Aphids itt rice roots I ITA Res Briefs 4(1-3)4-5

98 International Institite fi Tropical Agriculture 1983 Hopperburn in rice caused by brown plathopper IITA Res Briefs 4(l-3)6-7

99 International Rice Research Institute 1975 Annual report for 1974 PO Box 933 Manila Philippines p 216

100 International Rice Restarch Institute 1976 Annual repiort for 1975 PO Box 933 Manila Philippines p 228

101 International Rice Research Institute 1979 Cropping systems program Component technology development and evaluation Page 404 i Annual report for 1978 11O Box 933 Manila Philippines

102 International Rice Research Institute Department of Entomology 1981) Insecticid evaluation 1979 Tables 73 74 PO Box 933Manila Philipes

103 International Rice Research Institute Department of Entonmology 1981 Insecticide evalomtion 1980 Table 81 POBox 933 Manila Philippines

104 International Rice Research Institute 1981 Report of a meeting of Asian rice-based cropping systems entomologists Eleventh cropping systems working group meeting 18-22 May 1981 PO Box 933 Manila Philippines p 132 133 144

105 International Rice Research Institute 1984 Cropping systems program Pages 368-373 in Annual report for 1983 PO Box 933 Manila Philippines

106 International Rice Research Institutc 1984 Upland rice in Asia Pages 45-68 in An overview of upland rice research Proceedings of the 1982 Bouakamp Ivory Coast upland rice workiiop loternational Rice Research Institute 11O Box 933 Manila Philippines

107 Irshad M 1977 Grasshoppers in rice in Pakistan lit Rice Res Newsl 2(3)8-9

108 Ito R and TOno 1964 Upland lice its culture and varieties in Japan lIternational Rice Cormmission loth Meeting of the Working Party on Rice Production and Protection 3-10 March 1964 Manila Philippines Food and Agriculture Organization Rome Italy 12 p

109 Jeffrey M I and V A Dyck 1983 Effect of moonlight on lightshytrap catches of brown planthopper lit Rice Res Newsl 8(6) 13

110 Jennings P R and T Alicia Pineda 1983 Management ofthe rice plnthoppers Sogatodsorivi(coIl in Latin America Presented at the meeting on stable plant resistance FAO and lIndocsian I)inectorate of Food Crops Protection Bali lndonesia 29 Novcniber-2 l)ccemshyber 9 p

IllJimenez Gomez J A 1979 Rice insect pests i lla ios ()riCntales lil Spanish] Pages 99-103 in Rice ciltivation Instituto (olotibiatto Agropecuario Compndio No 29

112 Jongeleen F JJ Soesanto and A Mockidjo 1979 Studies oti pathogens of two species of Scarabacidac froinIndonesia Z Angew Enlomnol 87412-417

113 Jouvenaz 1) PC S lofgren and W A Baiks 1981 Biological control of imported fire antsa review of cirrent knowledge Bull Entomol Soc Am 27203-208

114 Kainoh Y K Shiniiu S MarUani Y Taiaki 1980 lost finding behavior of tie rice bu lplotori+JchinvittjI)allas (liciniptera Corcidac) with special reference to diet pattern ol aggregation and feeding on the rice plant Appl Entomol Zool 15225-233

115 Kakkar K 0 P Sharma and P I Sharma 1977 Note on Ihteronyrchus liodresRedt (Coleopicra I)ynastidae) as asporadic pest of paddy and maie inHimachal Pradesh Corr Agric 156-57

116 Kalode M131 79 Major insect pest problems and their control in upland rice (Unpublishcd mineogriph CWI oJNaaiat lni Chandaratat (198 1)original not seen)

117 Kalshoven L i 98 I The pests of crops in I ldoniesia (translated by P A vatn der Laan) 1 T Ichtiar B3aru-vati Iloeve lakarta Indonesia 701 p

118 Kamal N Q 1981 Suppression of whitebacked planto ner Sogatella fircaifra (Horvath) aLId rice lealfolder (nhal -cis licdb lis (Guenee) populations by natural cnclmie hlI) tiesis Grcgorio Araneta University Foundation Victoncta Park Metto Manila Philippines 179 Ip

119 Karim A N M Reaul 1975 A species ofliealy bug Trioni i70s ceres new oii rice in the Philippines Rice Entoniol Newsl 247

120 Katanyukul Wand C Chandaratat 1981 Insect pestsof uplaid rice and their control Report firstFAOUNlFi Tliailand training course on improved cultural practices for tplad rice Chiang Mai Thailand 5-23 Oct 1981 Food and Agriculture Organiation Rome Italy 22 p

121 Kern F 1955 lhletwioa(I)uninemts) bhhit Biturm(Coleoptera) a new pest of sugarcane inVenezuela [inSpanish with Etnglish sunmary] Agron Trop (Maracay) 527-29 (Rev Appl Eitoniol A4572)

122 Khan K M and S Ghai 1974 White grubs ald their control in India Pesticides (1Bombay) 819-25

123 Kiritani K 1975 Integrated pest management of rice pests Pages 66-70 itt Proceedings platning workshop ott cooperative hIeld research in pest managementP SMotooka ed East-West Center Honololu Hawaii USA

124 Kisimoto R 1967 Ecology of insect vectors forecasting and chemical control Pages 243-255 in The virus diseases of tle rice plant Proceedings of a symposium atthe International Rice Research Institute April 1967 The Johns Hopkins Press altimore Maryshyland USA

125 Kisimoto R and V A Dyck 1976 Climate and rice insects Pages 367-391 in Climate and rice International Rice Research Institute PO Box 933 Manila Philippines

126 Kureha S S Kobayashi SAsari and T Ishihara 1977 Life history and control of the mole cricket Grrllotalpa africana Palisot de Beauvois [in Japanese English translation 7520 1RRI library] Proc Kanto-Tosan Plant Prot Soc 21192-197

127 Kushwaha K S 1981 The black beetle as a pest of rice in Haryana India Int Rice Res Newsl 6(4)14

128 Lawani S M 1982 A review of tile effects of various agronomic practices on cereal stem borers Trop Pest Manage 28266-276

129 Leuck D B 1966 Biology of the lesser cornstalk borer in South Georgia J Econ Entomol 59 797-801

130 lever R J A W 1955 Insect pests of the rice plant in Malaya Int Rice Comm Newsl 1612-21

131 Li Ching-Sing and Shin-Foon Chiu 1951 The study ofthe rice gall midge Iachydiplosis orvza Wood Mason [in Chinese with English summary] 1Taiwan Agric Res 21-13

132 Iitsinger J A 1984 Assesament of the need-based application for rice Proceedings TechnologyTransfer Workshop 15-16 March 1984 International Rice Research Institute 1PO Box 933 Manila Philippines 6 p

133 Litsinger J A 1984 Environmental diversity and insect pest abundance with reference to the Pacific Pages 285-307 in Proceedings sub-regional training course on methods of controlling diseases insects and other pests of plants in the South Pacific 4-20 Oct 1982 Vaini Kingdom of Tonga Nukualofa

134 Litsinger J A RF Apostol and M BObusan 1983 White grub Leucopholis irrorata (Coleoptera Scarabaeidae) pest status popula-tion dynamics and chemical conllol in arice-maize cropping pattern in the Philippines J Econ Entoinol 761133-1138

135 Litsinger J A C de la Cruz F Raymundo A T Barrion M D Lumaban J P Bandong M S Venugopal F Paragna and E Balete 1982 Insect pests and insecticide reponse of dry-seeded bunded rice Pages 179-193 in Report of workshop on cropping systems research in Asia International Rice Research Institute Los Bafos Philippine

136 Litsinger J A M 1) Luinaban J P Bandong A T Barrion W Katanyukul and N Panda 1978 Insect pest management in rainfcd lowland rice agroecosystms of tropical Asia International Rice Research Conference 17-21 April 1978 International Rice Research Institute Los Bafios Philippines

137 Litsinger J A and K Moody 1976 Integrated pest management in multiple cropping systems Pages 293-317 in R I Papendick P A Sanchez and G B Triplett eds Multiple cropping American Society of Agronomy Madison Wisconsin USA

138 Loevinsohn M E 1984 The ecology and control of rice pests in relation to the intensity and synchrony of cultivation Ph D thesis University of London 36) p

139 Loevinsohn M E J A Litsinger N Panda M A Austria R F Apostol C Andrion and J Bandong 1982 A basis for predicting rice insect pest abundance in cropping systems Paper presented at the 1982 International rice research conference on rice and cropping systems 19-23 April 1982 International Rice Research Institute Los Bafios Iaguna Philippines 25 p

140 Lopez A W 1930 Repot of the Entomology Department Pages 145-172 in Annual report of the Philippine Sugar Association 1929-30 Manila

141 Lopez AW 1931 Entomology Department Pages 227-266 in Annual report of the director or research 1930-31 Philippine Sugar Association Manila

142 Lopez A W 1932 Entomology Deparment Pages 252-279 in Annual report of the Philippine Sugar Association 1931-32 Manila

143 Lungibill P and G G Ainslee 1917 The lessercornstalk borer US Dep Agric Agric Res Serv Tech Bull 539 27 p

IRPS No 123 January 1987 39

144 McGuire J U Jr and BS Crandall 1967 Survey of insect pests and plant diseases ofselected food crops of Mexico Central America and Panama U SDepartment of Agriculture International Agriculshytural Development Service Agricultuial Researrh Service October 1967 11 p

145 McNeill Sand T R ESouthwood 1978 The role of nitrogen in the development of insectplant relations Pages 77-9 in lliochemical aspects of plant and animal coevolution J B Ilarbornu ed Academic Press London UK

146 Maebara H 1955 Spatial distribution of lastinstiarlanaeof purplish stem borer (Sesamia iqferens Walker) hibernating in an upland rice field Bull Kagoshima Univ Fac Agric 438-41

147 Mallick E H and N G Hajra 1977 Response of rice plant to defoliation treatments at different developmental stages Indian Agric 2151-56

148 Mansingh A 1971 Physiological classification of dormancies in insects Canl Entomol 103983-10)9

149 Martell C G 1974 Catalogue of plant feeding insects of Mexico [in Spanish] Fitofilo 27(69)1-176

150 Martinez 0 J 1982 Principal rice pests in Ilanos orientalis [in Spanish] P~ages 80-94 in Curso deact nalization de conocimientos en arroz Villaniencis Instituto (olonmbiano Agropecuario

151 Martins J F da Silva F Ferrcira A S Prabhu F 1 P Zimmermann 1980 Use of pesticides for the control of the main subterranean upland rice pests esqui Agropecu Bras 1553-u

152 Martins I F da Silva F Fe eira rod BdamSilveira Pinheiro I Q2 Damage simulation caused by defoliators of upland rice [in Portuguese with English summary] Pesqui Agropecu Bras 171113-1119

153 Martins 1F da Silva N(iyuen van Tan and 1Ida Silveira Pinheiro 1981 Stem borer resistance in upland rice associated with plant morphological characters [in Portuuce Iesqui Agropecu Bras 16187-192

154 Masaki S 1980 Sumnmer diapause Ann Rev Entomiol 251-25 155 Masuzawa T 11 Suwa and F Nakasoji 1983 l)ifkleences of

oviposition preferencc and survival rate of two skipper butterflies Parnara guttata and PIchlpidai iathiais (Iepidoptera HIesperiidac) on rice plant and cogon grass New Entomol 3247-56

156 Maurya D M and C P Vaish 1984 Upland rice in India Pages 69-92 in An overview oftpland rice research Proceedings ofthe 1982 Bouakamp Ivory Coast upland rice workshop International Rice Research Institute IO Box 933 Mnila Philippines

157 Menon M G R R N Katiyai S Kuar and S S Misra 1970 Termites attacking the paddy crop Sci Cult 36294-295

158 Miyahara Y and 1 Nishikubo 1967 Behavior of young nymphs of the red rice root aphid Iholahsipiun ri liabhdontinalisSasaki in relation to movement from the leaf to the roots of upland rice [in Japanese] Pioc Assoc Plant Prot Kyushu 1363-66

159 Mo ljoa Tjien 1952 White grub of upland rice (Holotrichiahelleri Brsk) [in Indonesian with English summary] Contrib Agric Res Stn Bogor No 13134 p

160 Mochida 0 and T Suryana 1979 The occurrence of the brown planthopper and ragged stunt on lowland and upland rice ill Central Java and logyakarta in tile late of January 1979 Entomnolgy Survey Report No 15 Central Research Institute of Agriculture (CRIA) Sukamandi Indonesia I February 1979 9 p

161Monet C 1979 Varietal resistance to upland rice pests [inI French] Pages 35-39 in Annual report 1977 Institut des Savanes Department des Cultures Vivrieres Bouak6 Ivory Coast

162 Monte 0 1942 A caterpillar ofrice-fields[in Portuguese] Biol6gico (Sao Paulo) 8161-163 (Rev Appl Entomol A31414)

163 Munroe I) D 1979 Pest management strategy for hill rice in four resettlement areas in Sarawak Malaysia Pages 89-102 i1 Guidelines for integrated control of rice insect pests Food and Agriculture Organization Plant Production and Protection Paper 14 Rome Italy 115 p

40 IRPS No 123 January 1987

164 Nakasuji F M Ishii and Jen-Zon Ito 1984 Rice skippers in Taiwan and their life histories ZZrC (Taipei) Tech Bull 82 10 p

165 Navas I) 1976 Fall armyworum in rice Pages 99-106 il Ecological animal control by habitat management Proc Tall Timbers Con-ference 28 Feb-I Mar 1974 Fall Timbers Res Stn Misc Publ 6 Tallahassee Horida USA

166 Nickel 1 1 1961 Biological control of rice stem borer A feasibility study Tcch Bull 2 International Rice Rcarch Institute PO Box 933 Manila Philippines IIl p

167 Nigan P M and R A Vernia 1985 Insect pests of uplaild rice in Uttar Pradesh Int Rice Res Newsl 10(6)22

168 Nilakhe S S 1985 -cological observations on spittle bugs with emphasis on their occirence in rice Pcsqtui Agropecu Bras 20407-414

169 Nixon G EJ 195 1 lie association of ats with aphids and coccids Commonwealth Iusti lcte of Entonology Iondon UK 36 p

170 Nurdin F an Ishak Manti 1985 Study oit seedling fly of upland rice Interlational Upland Rice Moitoring Tour Report 26 lebruary-2 MarLI Sukaranti West Sumatra

171 Oka 1 N 1979 Ciltural conttol of the brown planthopper Pages 357-369 in Brown planthopper threat to rice production in Asia International Rice Research Institutc 1p0 Box 933 Manila Philippines

172 Ohni M and A Vilnaii 1975 Rice insect pests of Italy biology of Sipha grcria (Kaltenbach) and its control [in Italian] Riso (Milan) 2459-71

173 Otanes F Q atnd 1 I Sison 194 1 Pests of rice Philipp J Agric 12211-259

174 Painter R I 1951 Insect icsistance in crop plants University Press of Kansits Lawrence Kansas USA 520 p

175 Pantua P C and 1A litsinger 1981 Arthropod abundance in continuous vs biannual rice cropping systeis Int Rice Res Ncwsl 6(2)20-21

176 Pathak M ) and V A l)yck 1975 Sttudics ott insect pests of upland rice Pages 186-191 in Major research in upland rice International Rice Research Institute PO Box 933 Ilanila Philippines

177 Patil B R B M IHasabe and 1) S Ajri 198 I Chemical control of white gritb lIditriihinhorraia Fabr intfesting groundntut paddy atnd sorghum Indian 1Entomol 43232-236

178 Pawar A 1) 1974 (ollection and identification of insect pests of rice rlerminal repirt lInternational Rice Research Institute PO Box 933 Manila Philippines 72 p (tnimco)

179 IerrajU A aid A S Reedy 1965 Observations ott diapausc itt tie steti borer ohrice Rice News Teller (lai-Mar)9-13

180 Pont A C 1972 A review of the Oriental species of theri ona Rondani (I)iptera Muscidac) of econtomic importance Pages 27-102 in Control of sorglium shoot fly M G lomtani and W R Young eds Oxford atd IH1 Ptthl Co New I)clhi India 324 p

181 Prasad S C 1PChand aid M 1PSingh 1983 Incidence of rice gall midge in the upland plateau region of Hihar Oryza 20235-236

182 Rai P S 198 0 Nisaga simpex damage to rice in the hill tracts of South India Int Rice kles News[ 5(I)12

183 Rathorc Y S 1969 Gonocplhahli dcprssunt Fabr as itpredator of (tilo zonwIlhis (Swinhoc) and other moths in field cage at Patnagar Uttar Pradesh Indian 1Entotnol 3127

184 Reed W 1983 Crop losses causcd by insect pists in the developing world Proceedings 10th Internatial Congress of Plant Protection lrighton UK 174-80

185 Reedy M V 1981 Ateactogasterinhens(Walker)a new root-grub pest of tipland paddy and inaize Curr Sci 501076

186 Reveno M A 1968 Catalogue of the insect pests of economic crops in Colombia [in Spaiish] Asociacion Latino Americana de Entomo-logia (AIAE) 129-33

187 Reyes G M 1957 Rice dwarf disease in the Philippines Food and Agriculture Organiiation Plant Prot Bull 617-19

188 Rice S E A A Grigarick and M 0 Way 1982 Effect of leaf and panicle feeding byarmyworm (Lepidoptera Noctuidae) larvae on rice grain yield Environ Entomol 7593-595

189 Risch S J 1) Andow and M A Altieri 1983 Agioccosystem diversity arnd pest control data tentative conclusions and nizw research dirctions Environ Entomol 12625-629

190 Rivera C T 1956 Three organic insecticides in thecontrol of insects affecting upland rice Philipp Agric 39465-472

191 Roffrey 1 1972 Locusta outbreaks in the Philippines Acrida 1177-188

192 Rojas R 1) 1941 The effect of pruning at varying ages upon the yield of Ramairio Phipp Agric 29851-860

193 Roonwal M i 1979 Ternite life and termite control in tropical South Asia Scientilic Pnbl odhpr India 139 p

194 Rosetto C1 F da Silva Martins N C Schmidt and L EAzzini 1978 Damage caused by pasturespittle bugs 4Dvoi)lavopictaandDJ schach) on rice lBragantia Nota 6 (Matcli)25-27

195 Rosztto C1- S S Neto I- Amantc I) Iink 1Garcia-Vieira D M de SoLza N V Bansa t and A M de Olivcira 1971 Rice pests of Brazil [in Portuguese] Pages 149-238 inrFA()-International Rice Commission 2nd meeting of the Rice Committee for the Americas 6-11 l)ecember 1971 Pelotas Rio Grande do Sul Brazil

196 Rothschild G H L 1969 Obseratiois on the armyworm Spodoptera matritia acronrctoiiles Gn (Lep Noctuidae) Bull Entomol Res 59143-160

197 Rothschild G 1- L 1974 Parasites of the armworm Spolopiera mauritiaacroncitoihes in Malaysia Fntornopliaga 19293-299

198 Sachan i N S K Gangwar and 1N Katujar 1980 Field crickets damaging cages in North Eastern IHill Region Indian J Entomol 42526-527

199 Sands 1) P A 1977 The biology and ecology of Lepocorisa (Hemiptera Alydidac) in Papua New Guinea Res Bull 18 Departshyment of Primary Industry Port Moresby Papua New Guinea 104 p

200 Sarawak 1968 titomology division Pages 97-106 in Report of the research branch of the Department of Altriculture 1966 Kuching (Rev AIppl Eintomol A56603)

201 Satpathy JM M S Das and K Naik 1977 Effect of multiple and mixed cropping on the incidence of scine important pestsI Entomol Res 178-85

202 Sauer IHF G 1939 Notes Ott Elasmpnlpualtsligniosellths Zeller (Lep Pyr) a serious pest ofcereals in Sao Paolo [in Portuguese] Arq Inst Biol 10199-206 (Rev Appl Entotnol A28378)

203 Satinders 1L A 1 SKing and C L Vargas S 1983 Crop pests of Central Anerica with t bibliography [in Spanish] Centro Agroshynotnico Tropical de Investigacion y Ensenanza Turrialba Costa Rica (CATIE) Tec Bol 9 90 p

204 Sauphanor B 1985 Some factors of upland rice tolerttce to stem borers in West Africa Inseu Sci Appl 6429434

205 Sen P atnd S Chakravorty 1976 Stentflics in Haritighata Nadia West Bengal India [it Rice Res Newsl 1(12) 15

206 Senapai B and 1M Satpathy 1983 The seedling fly Atherigona orrzae Malloch is a pest of rice in Orissa (India) J Entomol Res 777-79

207 Shhjahan M S Alatn M Ah nad and BN Islam 1983 A new scarabaeid beetle pest of dryland rice in Bangladesh Int Rice Res Ncwsl 8(I)16-17

208 Sharma G L S C Modgol and R C Gautam 1979 Effect of depth of submergence ott incidence of bacterial blight and white grub infestation in transplanted rice Int Rice Res Newsl 4(6)12-13

209 Sharma S 1985 Scarabaeid beetle outbreak itt upland rice Int Rice Res Newsl 1(1(l)24

210 Shiraki T 1917 The paddy borer Schoenobhits incertulas Walker [achaku Agric Exp Stn Formosa 256 p

211 Silva A de Brito and 11P Magalhaes 1981 Insect pests of rice [in Portuguese] Circular Tecnica No 22 EMBRAPA Centro de Pesquisa t ecuaria de Tropico Unido (CPAJU) Belem Brazil 14 p

212 Singh S A 1979 Emergence of rice leaf butterfly as a rice pest in Manipur India int Rice Res Newsl 4(4)16-17

213 SinghS A 1983 Changing trends of insect pestsof rice in Manipur Int Rice Res Newsl 8(f)12

214 Siwi S S and P H van Doesburg 1984 Lenovorisa Latreille in Indonesia (Heteroptera Coreidae Alydinae) Zool Meded (Leiden) 58117-129

215 Soejitno J 1977 Reaction of some upland rice varieties toseedling fly eltherikonaexigua Stein [in Indonesian] Siri Hana I Penyakit 630-35 Lembga I osat Fenclilian Pertanian Bogor Indonesia

216 Socnardi 1967 Insect pests of rice in Indonesia Pages675-683 in The major inse pests of thu rice plant Proceedings of a symposium at the Internatital Rice Rec Wi itt S pternber 1964 The Johns Hopkins Press Baltimore Maryland USA

217 Soto P E A T Petcz and I W Buddenhagen 1976 Survey of insect pests and diseases of rice in different ecological zones in Nigeria Rice Entomol Ncwsl 435-46

218 Soto 11E and Z Ziddiqi 1976 Screening for resistance to African rice insect pests Varietal improsetnent sCmi n r Bouak Ivory Coast 13-17 Sept 1976 West Africa Rice l)evelopment Association Monrovia Iiberia

219 Squire F A 1972 Entomological problems in Bolivia PANS (PestArtic News Sumr) 18249-268

220 Stapely 1 II 1978 The rice leaf roller in the Solomon Islands Int Rice Res Newsl 3(l)13

221 Suenaga H 1937 On aphids attacking the roots of rice Oyo-I)oubts Zasshi 9150-152 [in Japanese] (Rev Appl Entomol A25745)

222 Suenaga H 1952 On the seasonal abundance of three subterranean aphids on upland rice Bull Kyushu Agric Exp Sti 1249-262

223 Sutynso Yas and Dandi Soekaria 1964 Beberapa hana yang ttenperang tana man padi tanah kering [in Indonesian] Rapat kerja Padi Tanah Kering 6 sd 8 Juily 1964 Kaliurang 9 p

224 Tallamy I) W and R F Denno 1983 Alternative life history patterns in risky environments in example from lace bugs Pages 129-147 in Insect life history patterns R F l)cnio and If Dingle eds Spring-Verlag 225 p

225 Tamura 1 1939 Observations on some insect pests of field crops [in Japanese] 1 Plant Prot 26414-420 (Rev Appl EntomolA27-556)

226 Tamura and Suzuki 1964 Influences of the early-transplanting practice in rice culture upon development of the rice stem borer Chilo siql)ressalisWalker Bull Ilokuriku Natil Agric Exp Stn 761-94

227 Tan J P 1924 The rice root aphid Drowhi ihirsoia Philipp Agric 13277-288

228 Tanaka 1 1957 Taxonomy and distribution of some subterranean aphids injurious to upland rice in Japan with description of a new species [in Japanese English summary] Botyu-Kagaku 22168-176

229 Tanaka T 1961 The rice root aphids their ecology and control Spec [full College of Agriculture Utsunomiya University Japan F3 p

230 Tauber M IC A Tauber R Nechols and 1Obrycki 1983 Seasonal activity of parasitoids control by external internal and genetic factors Pages 87-108 itt I)iapause and life cycle strategies in insects V K Brown and I Ilodek eds )r W Junk The Hague Netherlands

231 Taylor W F 1972 Effects of artificial defoliation (simulating pest damage) on varieties of upland rice Exp Agric 879-83

232 feotia T P S and S Natud 1966 Biotoomics of the rice skipper Parnaraniathiis Fabricius (Lepidoptera Hesperiidac) Indian J Entomol 28181-186

233 Teran F 0 1971 Rice pests of Santa Cruz [in Spanish Ministerio de Agricultura Sita Cruz Bolivia Bull 45 8 p

234 Than Itun 1976 Population dynamics of the yellow rice borer Tryporza itcertulas (Walker) and its damage to the rice plant MS thesis University of the Philippines at Ios Bafios Laguna Philippines 92 p

235 Uichanco L B 1911 Pests Pages 55-74 it Rice Philipp Agric 11-200

IRPS No 123 January 1987 41

236 van der Laan P A 195 1 Life history of the rice seedling fly Contrib Gen Agric Res Stn Boor Indonesia 118 15 p

237 van der Laan 11A 1959 Correlation between rainfall in the dry season and the occurrence of the white rice borer (Scirpolhaga innotala WIk) in lava Entomol Fxp Appl 212-20

238 Van Driesche R G 1983 Meaning of percent parasitismin studies of insect parasitism Fnviron Entomol 121 I1- 122

239 Venugopal M S and A litsingcr 1984 Fffect ofcarbofuran on rice growth Prot Ecol 7313-317

240 Vietnam Directorate of Rural Affairs 1962 Annual work progress report on crop improvement program of rice sugarcane vegetable and field crops (lor the period from July 1961 to June 1962) (11) 299 p (Rev Appl Entomol A51526-527)

241 Wan M T K 1972 Observations on rice leaf- and planthoppers in Sarawak (Malaysian Borneo) Malay Agric 148308-335

242 West Africa Rice I)esclopnient Association 1978 Annual research report Vol 2 Statistics Monrovia Liberia

243 West Africa Rice Development Association 1979 Annual research report p47 Monrovia Liberia

244 Williams ) F and CS lofgren 1981 Fli I illy l-468 anew bait toxicant for control of the red imported lire at Fla Entomol 64472477

245 Williams I) 1 1970 The mealybugs OFlomoptera Coccoidea Iscudococcidac) of sugar-cane rice and sorghum Bull Entomol Res 0109-188

246 Williams D) 1 1982 Pulinaria icervi (Signoret) and its allies ott sugar cane and other grasses Btll Entoinol Res 72111-117

247 Wirjosuiardjo S 1976 Study ofchenmical cont rot of insect pests ott upland rice Imiu Pertanian (Jogjakarta) 1323-329

248 Wirjosuhardjo S and N Wirarta 1976 Chemical control of the white grub tIolorithiatlltriin upland rice in M uneng Irobolinggo Indonesia) [in Indonesian] Imut Pertanian (Jogjakarta) 1330-332

249 Yano K T Miyake and V F Eastop 1983 The biology and economic importance of rice aphids (Henuiptera Aphid idae) a review Bull Entomol Res 73539-566

250 Yelp D and P Rhodes 19(4 lhe success of drilled dryland rice iii the Fiji Islands International Rice Commission Working Party on Rice Production and Protection Manila Philippines 3-10 March1964 Food and Agriculture Organization Rome Italy 13 p

251 Young W R 1981 Fifty-five years of research on the sorghum shootfly Insect Sci Appi 23-9

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ISSN 05-3862Other papers in this series TITLES OF NUMBERS 1-45 ARE LISTED ON THE LAST PAGE OF NO46 THOSE OF NUMBERS 46-70 ARE ON THE LAST PAGE OF NO 71-80AND THOSE OF NUMBERS 71-1O0ARE ON

THE LAST PAGE OF NO 101-122

No 101The economics of hybrid rice production in (hina No 113 Boron toxicity in rice No 102 Rice ratooning No 114 Energy analysis rice production systems and rice research No 103 Growth and development of the (Jeep water rice plant No 115 Production risk and optimal fertilizer rates an application of the

No 104 Faridpur acomputer-assisted instruction nmodel forrainfed lowland random coefficient model No 116 Consumer demand for rice grain quality in Thailand Indonesia andrice

No 105 A readingand listening comprehcnsion test inEnglish for non native the Philippines speakers applying for tiraining at IRIRI No 117 Morphological changes in rice panicle development a review of

No 106 Rice grassy stunt virus 2 a new strain of rice grassy stunt in tire literature

Philippines No 118 IRRI-Korca collaborative project for the development of cold-No 107 Physical losses and quality deterioration in rice plstprodnCtion tolerant lines through anthei culture

systemis No 119 Problen soils as potenial areas for adverse soils-tolerant rice No 108 (olptblication ofIRRI materials a survey of translators and varieties in South and Ssuthcasi Asia

publishers No 120 Changes isinall-farm iLtireliing techii)hgy in Thailand and [he No 109 Classification of Philippine rainfall patterns Philippines No 110 Contributions of modern rice varieties to rutrition in Asia No 121 Landforms and modern rice varieties No Ill Changes in rice breeding in 10 Asian countries 1965-84 No 122 Yield stability and modern rice technology No 112 Design parameters affecting the performance of the IRRI-designed

axial-flow pump

Page 36: UPLANDRICE. INSECTl--1i M T -E!R.EOLOU t RTANCE ,AND …delphacid.s3.amazonaws.com › 6958.pdf · The significance of abiotic factors in upland rice insect ecology will be apparent

36 IRPS No 123 January 1987

REFERENCES CITED

1Abraham C C and K S R Mony 1978 Occurrence ofLeptocorisa acuta Fabr (Coreidae Hemiptera) as a pest of nutmeg trees J Bombay Nat ttistSoc 74553

2 Agyen-Sampong M 1975 HlierogrlphusdiaganensisKrauss (Orthop-tera Acrididae) a new pest of rice in Northern Ghana Ghana J Agric Sci 8249-253

3 Agyen-Samnpong M 1976 Varietal resistance and other inle-ratcd pest control components as factors of rice varietal improvement in West Africa Prcsented at the Second varietal improvement serinar 14-18 Sep 1976 Bouak Ivory Coast West Africa Rice Development Association Monrovia Liberia 18 p

4 Agyen-Sampong M 1977 Outbreak of Ihccrorrdnisorae in rice seedlings near Rokupr Sierra Leone lit Rice Res Newsl 2(6)16-17

5 Agyen-Sampong M 1980 Dirty panicles and rice yield reduction caused by hugs IntRice Res Newsl 5(l)11-12

6 Agyen-Sampong M 1982 The major pest problems of irrigated upland and mangrove swanip rice ecosystems ir the humid tropical Guinea savanna and Salel climate zones Pages 318-328 in Pro-ceedings training course cn concepts techniques and application of irtegrated pest management in rice inWest Africa 10-20 Jan 1982 Fendall Liberia West Africa Rice l)evelopioent Association Monrovia Liberia 505 p

7 Akibo-Betts V I and S A Raymundo 1978 Aphids as rice pests in Sierra Leone lit Rice Res Newsl 3(6)15-16

8 Akinsola F A 1975 Present status of different rice stemborers in parts of Nigeria iicL Entomo Newsl 328

9 Akinsola FA 1979 Ihe biology and ecology of rice stemborers in Nigeria Ph 1) thesis University ol lbadan Nigeria 217 p

10 Akinsola E A 1979 Guidelines of integrated management of insect pests of upland rice in West Africa Presented at the Seminar on integrated management of rice diseases and insect pests at Bobo-Dioulasso Upper Volta 17-22 Sep 1979 West Africa Rice Develop-ment Association Monrovia Liberia

II Akinsola IA 1984 Insect pests of upland rice in Africa Pages 301-305 in An overview of upland rice research Proceedings of the 1982 lIouak Iwsrv Coast upland rice workshop International Rice Research Institute 13O Box 933 Manila Philippines

12 Alam M S V TJohn and Kaung Zan 1985 Insect pests and diseases of rice in Africa Pages 67-82 in Rice improvement in Eastern Central and Southern Africa proceedings of the international rice workshop at Iusaka Zambia 9-19 April 1984

13 Alticri M A A van Schoonhovcn and J D Doll 1977 The ecological role of weeds in insect pest management systems a review illustiated with bean (Phaseohsvulgaris L)cropping systems PANS (Pest Artie News Summ) 23195-205

14 Anmante F 1967 The leaf-cutting ant Atia capiguaraa pest of pastures [in Portuguese] Biol6gico (Sao Paulo) 33113-120 (Rev Appl Entomol A56207)

15Andrade A C and D Puzzi 1951 Preliminary results of experi-merits on the control of Sculptcoris iastantus on sugarcane [in Portuguese] Bioulgico (Sao Paulo) 1744-49 (Rev Appl Entomol A41421422)

16 Appert J 1970 Maliarplia separaelha (rice stem borer) new observations of entomological problems of rice in Madagascar [in French] Agron Trop (Paris) 25329-334

17 Ariadeau Mand Z Harahap 1986 Relevant upland rice breeding objectives Pages 189-197 in Progress in upland rice research Proceedings of the 1985 Jakarta conference International Rice Research Institute 1O Box 933 Manila Philippines

18Asar S and Kisimoto 1975 Seasonal occurrence of the smaller brown planthopper in upland rice [in Japanese] Proc Kanto-Tosan Plant Prot Soc 2228

19 Bakker W 1974 Characterization and ecological aspects of rice yellow mottle virus in Kenya Meded Landbouwhogesh Wage-ningen 152 p

20 Balut F F 1970 White grubs attacking rice Oryza ativa L [in Portuguese] Biol6gico (So Paulo) 36321-322

21 Banerjee S N and L M Pramanik 1967 The lepidopterous stalk borers of rice and their life cycles in the tropics pages 103-124 in The major insect pests ofthe rice plant Proceedings ofa symposium at the International Rice Research Institute September 1964 The Johns Hopkins Press Baltimore Maryland USA

22 Barrion A T and J A Litsinger 1980 Ants a natural enemy of cliaiarac in dryland rice IntRice Res Newsl

5(4)22-23 23 Barrion A T and J A litsinger 1981 The spider fauna of

Philippine rice agroecosystem 1Dryland Philipp Entomnol 5139-166

24 Barrion A T and 1 A Iitsinger 1984 The spider fauna of Philippine rice agroccosystcms II Welland Philipp Eiitomol 611-37

25 Barrion A T1 and J A Litsinger 1984 lihanus (Diptera Tabanidae) eggs an alternative host of rice stei borer (SI3) egg parasite Telenoonis digns (Iliymenoptcra Scelionidae) lit Rice Res Newvsl 9(6)19

26 Barrion A T and J A Litsinger 1985 Identilication of rice leaffolders by wing markings Int Rice Res Newsl 10( 1)24

27 Barrion A T and J A Litsinger 1985 Chctiosspp (Coleoptera Carabidae) a leaffolder ILF)predator lit Rice Res Newsl 10(1)21

28 Bertels A 1970 Insect pests of rice and theircontrol [in Portuguese] Pelotas Brazil Instituto de Expcrimentacao Agropccuararias do Sul Circular 43 24 p

29 Bonzi S M 1982 Ctilo (flisilineus J de Joannis (Lepidoptera Pyralidae) stem borer of irrigated cereals in Upper Volta LioFrench] Agron Trop Riu Rizic Cult Vivrieres Trop 37207-209

30 Bourke T V T L Fenner J N LStihick G Baker EHassan D F OSullivan and C S li 1977 Insect pest survey for the year ending 30th June 1969 Papta New Guinea )epartment of Agrishyculture Stock and Fisheries Port Moresby 57 p

31 Bowden 1 1973 Migration of pests in the tropics Meded Rijksfac Landbouwwet Gent 38785-796

32 Breniere J 1973 The main rice parasites in West Africa and their control Pages 27-42 in Proceedings seminar on plant protection for the rice crop West Africa Rice Development Association Monrovia Liberia

33 Breniere J 1976 The principal insect pests of rice in West Africa and their control West Africa Rice Development Association Monrovia Liberia 52 p

34 Brown ES 1962 The African arinyworn Spoiolptera eXI)Ina (Walker) (Lepidoptera Noctuidae) a review of the literature Cornshymonwealth Institute of Entomology London UK57 p

35 Brown F B 1968 Upland rice in Latin America FAO-International Rice Commission 12th Session of the Working Party on Rice Production and Protection 2-14 Sept Peradeniya Sri lanka Food and Agriculture Organization Rome Italy 6 p

36 Buyckx F J 1 1962 Disease and insect pests of crops inCongo Rwanda and Burundi [iil French] Publications dc LInstilut National pour lestude Agrononiquedu Congo M Weissenbruck S A Brussels Belgium 708 p

37 Cariiio F 0 1981 Role of natural enentics inpopulation suppression and pest management of green rice lealhoppers Ph D thesis University ofthe Philippines at Los Bafitos PhilippinesLaguna 136 -

38 Carpente A J 1983 The development of rice cultivation and extension of vegetable productin in Zanzibar Final Report Vol II1 October 1983 Food and Agricultme Organization Ronte Italy

39 Carroll C P and S1 Risch 1983 Tropical annual cropping systems ant ecology Environ Manage 751-57

4(0 Cates R G 198 1 Host plant predictability and the feeding patterns of monophagous oligophagous and polyphagous insect herbivores Oecologia (Berl) 48319-326

IRPS No 123 January 1987 37

41 Cendcafia S M and F B Caora 1967 Insect pests of rice in the 62 Ene 1 C 1981 I)iiorldhol)ierusihilis Slater (licteroplera Philippines Pages 591-616 in The najor insect pests ofthe rice plant lygacidue Illissinac) a new rice pest front Nigeria lit Rice Res Proceedings of a symposium at the International Rice Research Newsl N3)15 Institute September 1964 The Johns Hopkins Press Baltimore 63 Fugenio T S 1965 rhe effect oi e-vCgetation on rice production Maryland USA 729 p and the possibility of overcoming it by leaf pruning MS thesis

42 Centro Ihternacional de Agricklittira Tropical 1984 Upland rice in University ol the Philippines at Los Baijos ILaguna Ihilippins 69 p Iatin Anerica Pages 93-119 hi An os ervies of uplaind rice research 04 Farrow R A 1975 The Alrican nmigratory locust ii is main Proceedings of the 1982 Boinakamp Ivory Coast upland rice workshop outbreak area of the middle Niger qlinlitatise studies of solitary International Rice Research Institute P0 Blox 933 Manila populations in relation to environmental factors Locusla No II Philippines 198 p

43 Chandra G 1979 Taxonomy ad hn riononlics ofthe imect parasites of 65 Fennal R G 1943 Food crop investigation Windward and rice Icalloppers and planthoppers in the Pihilippines and their Ieeward Islands First report Report onl work dunring the periodiiiportairce iii natural biol)gical cointrol Philipp Fntoltol 4119-139 April 1942-April 1943 Imperial College of Tropical Agriculture

44 Chang T T 1976 The originii colut ion enlIivation dissemination Trinidad 16 p (Rev of Appl Fintornol A3222-25) and diersificition of Asian and African rices -uphytica 25425-44 1 66 Ferreira F 1983 Integrated pest ritaageiret of upland rice [ill

45 Chatierjee P B 1979 Rice leaffolder attacks in India lit Rice Res Portuguese English summary] Pages 323-341 in Upland riceculttre Newsl 4(3)21 factors affecting production E Ferreira T Yamada and F-

46 Cheng Cbien-pan 19(4 Upland rice in Taiwan International Rice Malmolta eds Institutoda Potassae Foslato Institnto Internacional Cnmmission Wiorking Party oit Rice Production ard Protection I()th da Potassa Piracicaha Sao Paulo lraid Meeting 3-10 March 1964 Manila Philippines Food and Agricul- 67 Ferreira E arid F dl Silva Martins 1979 Occurrence of ture Organiaion Rome Italy 5 p Sogatohs orvziola (Muir) inupland rice in Goias Ilrail lit Rice

47 Chcrrett 1 M and 1) 1Peregrine 1976 A review of the status of Rcs Ncwssl 4(4)18 leaf-cutting ants and their control Ann Appl Biol 84124-128 68 Ferreira E and 1F da Silva Martiris 1984 Insect pestof rice ill

48 Chipeta F M 1974 Pests of rice in Malawi Pages 90-95 in Brazil and their control Jill Portugucsej FMIlRAtPA-CNPAF Proceedings Sith eastern Africa cereals research conference 10-15 Goiania Brmiil 67 p March 1974 DRB Mialda ed Ministry of Agriculture lilongwe 69 Ferreira FI F ca Silha Miartins S S Nero and 1 It P de Malawi Carvallio 1982 M tltiplc regression analysis todeterimine the effect of

49 Chu Yau-i 1979 The d atiaging period of the rice seedling fly insect populatiois ott upland rice yield [inIPortuguese Pesqui (Atlterigotiavxigu Stcin) (NItiscide Diptera) oil cori) at Fast Java Agropecu Bras 17671-675 Indonesia Plant Prot Bill 21403413 70 Ferreira F 1F da Silva Martins and F 1PZimmermann 1979

50 Conklin H C 1957 IHaninoo agriculture a report on aln integral Rice varietal resistance to tle lesser corn stalk borer [in Port uguese] systent of shi fti rig cult ivationi in the Pltilippincs Food and Agriculture Pesqui Agropccu Biras 14317-321 Organization Rome Italy 209 p 71 Fleming W F 1968 Biological control of lie hapanese beetle U S

51 Cramer H II 1967 Plant proection and world crop production Dep Agric RLs Sers [ech Bull 1383 78 p Pilaniensclhutz-Nachr 201-524 72 Food and Agriculture Organiation of the United Nations 1982

52 l)ani R C and Majunidar 1978 Prelininary observations on tie Country report surmimary Pages 27-37 in Report on the first incidence of root aphids on differeit rice cultnrcs Sci Cult 4488-89 FAOIUN)IPhail and regional training coiuirse in iniproved ctltural

53 Davidson R I 1979 lffects of root feeding on foliage yield Pages practices for upland rice 5-23 Oct 1981 Northern Region Agrculturd117-120 in T K Ci osb and R B Pottinger eds Proceedings of the Developririt Center Chiang Mai Food and Agricuilture Organiiashy2nd Australian conlerence on grassland invertebrate ecology Govt tion Rome Italy 37 n Printer Wellington New Zealand 73 Franco do Ariaral S 195 1 Residual action of insecticides in turfy

54 l)ean G 1W 1976 Rice insect pests inlaos Int Rice Res Newsl soil--the results of a field experiment [in Portuguese] Biol6gico (Sao 1(2)15 Paulo) 17209-214 (Rev Appl Entomol A41423)

55 I)ean G J W 1978 Insect pests of rice in Laos PANS 24(3)280-289 74 Freeman ) 1970 Report onithe Iban london School of -coroiics 56 )ingle H 1984 Behavior genes and lifc histories conplex adapta- Monographs on Social Anthropology No 41 Athlane Press

tions to uncertain environments Pages 169-194 in A new ecology London UK 317 p novel approaches to interactive systems P W Price C N Slobod- 75 Fuchs T W1 A Harding and 1W Smithlr 1979 Induction and chikoff and W S Gaud cdsl ohn Wiley New York USA termination oldiapause in the sugarcane borer Ann Flitonlol Soc

57 Dobelmann-JP and M Falais 1968 Upland rice cultivation ill tie Am 72271-274 North-western part of Madagascar in 1966-1967 [in French English 76 (hbriel B1) 1975 A review tf the major insect pests of some tpland summary] Trop Agron 2313-21 crops in tlc Philippines Pages 86-111 in Ad Htoc panel of experts on

58 Doniiciano N L 1979 Chemical control of tie lesser corn stalk pest disease arnd weed problems in some rainfed crops Bangkok (lolsonno~paloIgnowslhts) (Zeller 1848) (I-epid(ptera Ihycitid ae) Thailand Food aid Agriculture Organization Roire Italy

on rice in Ponta Gressa Parara 1977-1978 [in Portuguese] Pages 77 Garg D K and N K Shal 1983 White grub outbreak on rainfed 291-303 in Anais 1 Reuniao de tecnicos cmiirizicultura do estado de dryland rice in Uttar Pradeslh Int Rice Res Ncwsl 81(4)16-17 Sao Paulo-Campinas Cati Brazil 78 Garg I) K and IP Tandon 1982 Major insect pests of rice on hilly

59 Dunsmore 1 R 1970 Investigations on tlie varieties pests and tracts of Uttar Pradesh India lilt Rice Res Newsl 7(11)11-12diseases of upland rice (hill padi) in Sarawak Malaysia Int Rice 79 Garrity 1) P 1983 Upland rice research an international bibliograpy Comm Newsl 1929-35 1975-1982 Iternational Rice Research hnstitnte PO Box 933

60 Elias 1R1965 The new decline of rice seedlings is not caused by Manila Philippines 121 p parasites [in Portuguese] Biol6gico (Sao Paulo) 313-6 80 Ghauri M S K 1982 A new subspecies of Iinorh)rone bull

61 Empresa Brasiliera de Pesquisa Agropecuaria 1984 Upland rice itt cornlus Slater (Hemiptera Iygaeidae) front Papua New Guinea on Brazil Pages 121-134 it Av overview of upland rice research rice and carpet grass Ihll lEntoniol Res 72133-137 Proceedings ofthe 1982 Blouak Ivory Coast Upland rice workshop 81 Gonzales F 0 PArregoces R I ihernandef and OT Parada International Rice Research Institute PO Box 933 Manila 1983 Insect and imite pests of rice arid their control in Latin America Philippines [in Spanish] la Federacion Nacional de Arrocerts (Fedearroz)

Bogota Colombia 60 n

38 IRPS No 123 January 1987

82 Grist D H and R J A W Lever 1969 Pests of rice Longmans London UK520 p

83 Hale P R 1979 Two subterranean pests of upland rice in Papua New Guirnca IntRice Res Newsl 4(2)16-17

84 Hale P R and J T Hale 1975 Insect pests of rice in the New Guinea Islands Rice Entomol Newsl 34

85 Harris W V 1969 Termites as pests of crops and trees Common-wealth Institute of Entomology London UK 41 p

86 Hattori 1 1968 A method to distinguish between grass leaf roller (Ctaplha rocis medinalisGuenee) and the rice leaf roller (Susurtia exigua Butler) [in Japanese] Plant Prot (Japan) 23167-170

87 Henry I E and 1A Onsager 1984 Experimental control of the mormon cricket Iahrosshnph by NJsena locustae(Aficrospora Microsporidae) a protozoan parasite of grasshoppers (Ort Acri-didae) Entomophaga 27197-201

88 Hidaka 1 P Vungsilabulr and SKadkao 1974 Studies on ecology and control of the rice gall midge in Thailand Tech Bull 6 TropicalAgricultural Research Center Tokyo Japan 113 p

89 Hincklcy A 1) 1963 Ecology and control of rice planthoppers in Fiji Bull Entomrol Res 53467 481

90 Hirao J 1982 Ecology and chtInical control of the rice leaffolder Jpn Pestic Inf 4114-17

91 Ho DT 1984 Sten borer incifience in rice ecosystems in KenyaEast Africa Pages 307-320 in A overview of upland rice research Proceedings of the 1982 Bouac Ivory Coast upland rice workshop International Rice Research Institute PO Box 933 Manila Philippines

92 Ho )T J 1Njokah and J GKibuka 1983 Studies on rice sten borers in Kenya with emphasis on ifaliarpiaipiraellRag Insect Sci Appl 465-73

93 Ichihara I T Ito A NIatsuo and M Sawada 1971 Chemical control of the rice striped stem borer in upland rice grown in plastic mulch [iii Japanese] Proc Kanto-Tosan Plant Prot Soc 1882

94 IclhiharaIand A Matsuo 1969 Damage caused by rice striped stett borer to upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1686-87

95 Ichhara I atid A MIatsun11970 Conside ration oft lie extraordinary prevalence of rice striped stem borer in upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1778-79

96 Institut de Recherches Agrolnliqes Iropicale et des Cultures Vivrieres 1978 Insect resistance Pages 12-13 in Upland rice 1978 atnual report Bouak Ivory Coast 27 p

97 International Institute for Tropical Agriculture 1983 Aphids itt rice roots I ITA Res Briefs 4(1-3)4-5

98 International Institite fi Tropical Agriculture 1983 Hopperburn in rice caused by brown plathopper IITA Res Briefs 4(l-3)6-7

99 International Rice Research Institute 1975 Annual report for 1974 PO Box 933 Manila Philippines p 216

100 International Rice Restarch Institute 1976 Annual repiort for 1975 PO Box 933 Manila Philippines p 228

101 International Rice Research Institute 1979 Cropping systems program Component technology development and evaluation Page 404 i Annual report for 1978 11O Box 933 Manila Philippines

102 International Rice Research Institute Department of Entomology 1981) Insecticid evaluation 1979 Tables 73 74 PO Box 933Manila Philipes

103 International Rice Research Institute Department of Entonmology 1981 Insecticide evalomtion 1980 Table 81 POBox 933 Manila Philippines

104 International Rice Research Institute 1981 Report of a meeting of Asian rice-based cropping systems entomologists Eleventh cropping systems working group meeting 18-22 May 1981 PO Box 933 Manila Philippines p 132 133 144

105 International Rice Research Institute 1984 Cropping systems program Pages 368-373 in Annual report for 1983 PO Box 933 Manila Philippines

106 International Rice Research Institutc 1984 Upland rice in Asia Pages 45-68 in An overview of upland rice research Proceedings of the 1982 Bouakamp Ivory Coast upland rice workiiop loternational Rice Research Institute 11O Box 933 Manila Philippines

107 Irshad M 1977 Grasshoppers in rice in Pakistan lit Rice Res Newsl 2(3)8-9

108 Ito R and TOno 1964 Upland lice its culture and varieties in Japan lIternational Rice Cormmission loth Meeting of the Working Party on Rice Production and Protection 3-10 March 1964 Manila Philippines Food and Agriculture Organization Rome Italy 12 p

109 Jeffrey M I and V A Dyck 1983 Effect of moonlight on lightshytrap catches of brown planthopper lit Rice Res Newsl 8(6) 13

110 Jennings P R and T Alicia Pineda 1983 Management ofthe rice plnthoppers Sogatodsorivi(coIl in Latin America Presented at the meeting on stable plant resistance FAO and lIndocsian I)inectorate of Food Crops Protection Bali lndonesia 29 Novcniber-2 l)ccemshyber 9 p

IllJimenez Gomez J A 1979 Rice insect pests i lla ios ()riCntales lil Spanish] Pages 99-103 in Rice ciltivation Instituto (olotibiatto Agropecuario Compndio No 29

112 Jongeleen F JJ Soesanto and A Mockidjo 1979 Studies oti pathogens of two species of Scarabacidac froinIndonesia Z Angew Enlomnol 87412-417

113 Jouvenaz 1) PC S lofgren and W A Baiks 1981 Biological control of imported fire antsa review of cirrent knowledge Bull Entomol Soc Am 27203-208

114 Kainoh Y K Shiniiu S MarUani Y Taiaki 1980 lost finding behavior of tie rice bu lplotori+JchinvittjI)allas (liciniptera Corcidac) with special reference to diet pattern ol aggregation and feeding on the rice plant Appl Entomol Zool 15225-233

115 Kakkar K 0 P Sharma and P I Sharma 1977 Note on Ihteronyrchus liodresRedt (Coleopicra I)ynastidae) as asporadic pest of paddy and maie inHimachal Pradesh Corr Agric 156-57

116 Kalode M131 79 Major insect pest problems and their control in upland rice (Unpublishcd mineogriph CWI oJNaaiat lni Chandaratat (198 1)original not seen)

117 Kalshoven L i 98 I The pests of crops in I ldoniesia (translated by P A vatn der Laan) 1 T Ichtiar B3aru-vati Iloeve lakarta Indonesia 701 p

118 Kamal N Q 1981 Suppression of whitebacked planto ner Sogatella fircaifra (Horvath) aLId rice lealfolder (nhal -cis licdb lis (Guenee) populations by natural cnclmie hlI) tiesis Grcgorio Araneta University Foundation Victoncta Park Metto Manila Philippines 179 Ip

119 Karim A N M Reaul 1975 A species ofliealy bug Trioni i70s ceres new oii rice in the Philippines Rice Entoniol Newsl 247

120 Katanyukul Wand C Chandaratat 1981 Insect pestsof uplaid rice and their control Report firstFAOUNlFi Tliailand training course on improved cultural practices for tplad rice Chiang Mai Thailand 5-23 Oct 1981 Food and Agriculture Organiation Rome Italy 22 p

121 Kern F 1955 lhletwioa(I)uninemts) bhhit Biturm(Coleoptera) a new pest of sugarcane inVenezuela [inSpanish with Etnglish sunmary] Agron Trop (Maracay) 527-29 (Rev Appl Eitoniol A4572)

122 Khan K M and S Ghai 1974 White grubs ald their control in India Pesticides (1Bombay) 819-25

123 Kiritani K 1975 Integrated pest management of rice pests Pages 66-70 itt Proceedings platning workshop ott cooperative hIeld research in pest managementP SMotooka ed East-West Center Honololu Hawaii USA

124 Kisimoto R 1967 Ecology of insect vectors forecasting and chemical control Pages 243-255 in The virus diseases of tle rice plant Proceedings of a symposium atthe International Rice Research Institute April 1967 The Johns Hopkins Press altimore Maryshyland USA

125 Kisimoto R and V A Dyck 1976 Climate and rice insects Pages 367-391 in Climate and rice International Rice Research Institute PO Box 933 Manila Philippines

126 Kureha S S Kobayashi SAsari and T Ishihara 1977 Life history and control of the mole cricket Grrllotalpa africana Palisot de Beauvois [in Japanese English translation 7520 1RRI library] Proc Kanto-Tosan Plant Prot Soc 21192-197

127 Kushwaha K S 1981 The black beetle as a pest of rice in Haryana India Int Rice Res Newsl 6(4)14

128 Lawani S M 1982 A review of tile effects of various agronomic practices on cereal stem borers Trop Pest Manage 28266-276

129 Leuck D B 1966 Biology of the lesser cornstalk borer in South Georgia J Econ Entomol 59 797-801

130 lever R J A W 1955 Insect pests of the rice plant in Malaya Int Rice Comm Newsl 1612-21

131 Li Ching-Sing and Shin-Foon Chiu 1951 The study ofthe rice gall midge Iachydiplosis orvza Wood Mason [in Chinese with English summary] 1Taiwan Agric Res 21-13

132 Iitsinger J A 1984 Assesament of the need-based application for rice Proceedings TechnologyTransfer Workshop 15-16 March 1984 International Rice Research Institute 1PO Box 933 Manila Philippines 6 p

133 Litsinger J A 1984 Environmental diversity and insect pest abundance with reference to the Pacific Pages 285-307 in Proceedings sub-regional training course on methods of controlling diseases insects and other pests of plants in the South Pacific 4-20 Oct 1982 Vaini Kingdom of Tonga Nukualofa

134 Litsinger J A RF Apostol and M BObusan 1983 White grub Leucopholis irrorata (Coleoptera Scarabaeidae) pest status popula-tion dynamics and chemical conllol in arice-maize cropping pattern in the Philippines J Econ Entoinol 761133-1138

135 Litsinger J A C de la Cruz F Raymundo A T Barrion M D Lumaban J P Bandong M S Venugopal F Paragna and E Balete 1982 Insect pests and insecticide reponse of dry-seeded bunded rice Pages 179-193 in Report of workshop on cropping systems research in Asia International Rice Research Institute Los Bafos Philippine

136 Litsinger J A M 1) Luinaban J P Bandong A T Barrion W Katanyukul and N Panda 1978 Insect pest management in rainfcd lowland rice agroecosystms of tropical Asia International Rice Research Conference 17-21 April 1978 International Rice Research Institute Los Bafios Philippines

137 Litsinger J A and K Moody 1976 Integrated pest management in multiple cropping systems Pages 293-317 in R I Papendick P A Sanchez and G B Triplett eds Multiple cropping American Society of Agronomy Madison Wisconsin USA

138 Loevinsohn M E 1984 The ecology and control of rice pests in relation to the intensity and synchrony of cultivation Ph D thesis University of London 36) p

139 Loevinsohn M E J A Litsinger N Panda M A Austria R F Apostol C Andrion and J Bandong 1982 A basis for predicting rice insect pest abundance in cropping systems Paper presented at the 1982 International rice research conference on rice and cropping systems 19-23 April 1982 International Rice Research Institute Los Bafios Iaguna Philippines 25 p

140 Lopez A W 1930 Repot of the Entomology Department Pages 145-172 in Annual report of the Philippine Sugar Association 1929-30 Manila

141 Lopez AW 1931 Entomology Department Pages 227-266 in Annual report of the director or research 1930-31 Philippine Sugar Association Manila

142 Lopez A W 1932 Entomology Deparment Pages 252-279 in Annual report of the Philippine Sugar Association 1931-32 Manila

143 Lungibill P and G G Ainslee 1917 The lessercornstalk borer US Dep Agric Agric Res Serv Tech Bull 539 27 p

IRPS No 123 January 1987 39

144 McGuire J U Jr and BS Crandall 1967 Survey of insect pests and plant diseases ofselected food crops of Mexico Central America and Panama U SDepartment of Agriculture International Agriculshytural Development Service Agricultuial Researrh Service October 1967 11 p

145 McNeill Sand T R ESouthwood 1978 The role of nitrogen in the development of insectplant relations Pages 77-9 in lliochemical aspects of plant and animal coevolution J B Ilarbornu ed Academic Press London UK

146 Maebara H 1955 Spatial distribution of lastinstiarlanaeof purplish stem borer (Sesamia iqferens Walker) hibernating in an upland rice field Bull Kagoshima Univ Fac Agric 438-41

147 Mallick E H and N G Hajra 1977 Response of rice plant to defoliation treatments at different developmental stages Indian Agric 2151-56

148 Mansingh A 1971 Physiological classification of dormancies in insects Canl Entomol 103983-10)9

149 Martell C G 1974 Catalogue of plant feeding insects of Mexico [in Spanish] Fitofilo 27(69)1-176

150 Martinez 0 J 1982 Principal rice pests in Ilanos orientalis [in Spanish] P~ages 80-94 in Curso deact nalization de conocimientos en arroz Villaniencis Instituto (olonmbiano Agropecuario

151 Martins J F da Silva F Ferrcira A S Prabhu F 1 P Zimmermann 1980 Use of pesticides for the control of the main subterranean upland rice pests esqui Agropecu Bras 1553-u

152 Martins I F da Silva F Fe eira rod BdamSilveira Pinheiro I Q2 Damage simulation caused by defoliators of upland rice [in Portuguese with English summary] Pesqui Agropecu Bras 171113-1119

153 Martins 1F da Silva N(iyuen van Tan and 1Ida Silveira Pinheiro 1981 Stem borer resistance in upland rice associated with plant morphological characters [in Portuuce Iesqui Agropecu Bras 16187-192

154 Masaki S 1980 Sumnmer diapause Ann Rev Entomiol 251-25 155 Masuzawa T 11 Suwa and F Nakasoji 1983 l)ifkleences of

oviposition preferencc and survival rate of two skipper butterflies Parnara guttata and PIchlpidai iathiais (Iepidoptera HIesperiidac) on rice plant and cogon grass New Entomol 3247-56

156 Maurya D M and C P Vaish 1984 Upland rice in India Pages 69-92 in An overview oftpland rice research Proceedings ofthe 1982 Bouakamp Ivory Coast upland rice workshop International Rice Research Institute IO Box 933 Mnila Philippines

157 Menon M G R R N Katiyai S Kuar and S S Misra 1970 Termites attacking the paddy crop Sci Cult 36294-295

158 Miyahara Y and 1 Nishikubo 1967 Behavior of young nymphs of the red rice root aphid Iholahsipiun ri liabhdontinalisSasaki in relation to movement from the leaf to the roots of upland rice [in Japanese] Pioc Assoc Plant Prot Kyushu 1363-66

159 Mo ljoa Tjien 1952 White grub of upland rice (Holotrichiahelleri Brsk) [in Indonesian with English summary] Contrib Agric Res Stn Bogor No 13134 p

160 Mochida 0 and T Suryana 1979 The occurrence of the brown planthopper and ragged stunt on lowland and upland rice ill Central Java and logyakarta in tile late of January 1979 Entomnolgy Survey Report No 15 Central Research Institute of Agriculture (CRIA) Sukamandi Indonesia I February 1979 9 p

161Monet C 1979 Varietal resistance to upland rice pests [inI French] Pages 35-39 in Annual report 1977 Institut des Savanes Department des Cultures Vivrieres Bouak6 Ivory Coast

162 Monte 0 1942 A caterpillar ofrice-fields[in Portuguese] Biol6gico (Sao Paulo) 8161-163 (Rev Appl Entomol A31414)

163 Munroe I) D 1979 Pest management strategy for hill rice in four resettlement areas in Sarawak Malaysia Pages 89-102 i1 Guidelines for integrated control of rice insect pests Food and Agriculture Organization Plant Production and Protection Paper 14 Rome Italy 115 p

40 IRPS No 123 January 1987

164 Nakasuji F M Ishii and Jen-Zon Ito 1984 Rice skippers in Taiwan and their life histories ZZrC (Taipei) Tech Bull 82 10 p

165 Navas I) 1976 Fall armyworum in rice Pages 99-106 il Ecological animal control by habitat management Proc Tall Timbers Con-ference 28 Feb-I Mar 1974 Fall Timbers Res Stn Misc Publ 6 Tallahassee Horida USA

166 Nickel 1 1 1961 Biological control of rice stem borer A feasibility study Tcch Bull 2 International Rice Rcarch Institute PO Box 933 Manila Philippines IIl p

167 Nigan P M and R A Vernia 1985 Insect pests of uplaild rice in Uttar Pradesh Int Rice Res Newsl 10(6)22

168 Nilakhe S S 1985 -cological observations on spittle bugs with emphasis on their occirence in rice Pcsqtui Agropecu Bras 20407-414

169 Nixon G EJ 195 1 lie association of ats with aphids and coccids Commonwealth Iusti lcte of Entonology Iondon UK 36 p

170 Nurdin F an Ishak Manti 1985 Study oit seedling fly of upland rice Interlational Upland Rice Moitoring Tour Report 26 lebruary-2 MarLI Sukaranti West Sumatra

171 Oka 1 N 1979 Ciltural conttol of the brown planthopper Pages 357-369 in Brown planthopper threat to rice production in Asia International Rice Research Institutc 1p0 Box 933 Manila Philippines

172 Ohni M and A Vilnaii 1975 Rice insect pests of Italy biology of Sipha grcria (Kaltenbach) and its control [in Italian] Riso (Milan) 2459-71

173 Otanes F Q atnd 1 I Sison 194 1 Pests of rice Philipp J Agric 12211-259

174 Painter R I 1951 Insect icsistance in crop plants University Press of Kansits Lawrence Kansas USA 520 p

175 Pantua P C and 1A litsinger 1981 Arthropod abundance in continuous vs biannual rice cropping systeis Int Rice Res Ncwsl 6(2)20-21

176 Pathak M ) and V A l)yck 1975 Sttudics ott insect pests of upland rice Pages 186-191 in Major research in upland rice International Rice Research Institute PO Box 933 Ilanila Philippines

177 Patil B R B M IHasabe and 1) S Ajri 198 I Chemical control of white gritb lIditriihinhorraia Fabr intfesting groundntut paddy atnd sorghum Indian 1Entomol 43232-236

178 Pawar A 1) 1974 (ollection and identification of insect pests of rice rlerminal repirt lInternational Rice Research Institute PO Box 933 Manila Philippines 72 p (tnimco)

179 IerrajU A aid A S Reedy 1965 Observations ott diapausc itt tie steti borer ohrice Rice News Teller (lai-Mar)9-13

180 Pont A C 1972 A review of the Oriental species of theri ona Rondani (I)iptera Muscidac) of econtomic importance Pages 27-102 in Control of sorglium shoot fly M G lomtani and W R Young eds Oxford atd IH1 Ptthl Co New I)clhi India 324 p

181 Prasad S C 1PChand aid M 1PSingh 1983 Incidence of rice gall midge in the upland plateau region of Hihar Oryza 20235-236

182 Rai P S 198 0 Nisaga simpex damage to rice in the hill tracts of South India Int Rice kles News[ 5(I)12

183 Rathorc Y S 1969 Gonocplhahli dcprssunt Fabr as itpredator of (tilo zonwIlhis (Swinhoc) and other moths in field cage at Patnagar Uttar Pradesh Indian 1Entotnol 3127

184 Reed W 1983 Crop losses causcd by insect pists in the developing world Proceedings 10th Internatial Congress of Plant Protection lrighton UK 174-80

185 Reedy M V 1981 Ateactogasterinhens(Walker)a new root-grub pest of tipland paddy and inaize Curr Sci 501076

186 Reveno M A 1968 Catalogue of the insect pests of economic crops in Colombia [in Spaiish] Asociacion Latino Americana de Entomo-logia (AIAE) 129-33

187 Reyes G M 1957 Rice dwarf disease in the Philippines Food and Agriculture Organiiation Plant Prot Bull 617-19

188 Rice S E A A Grigarick and M 0 Way 1982 Effect of leaf and panicle feeding byarmyworm (Lepidoptera Noctuidae) larvae on rice grain yield Environ Entomol 7593-595

189 Risch S J 1) Andow and M A Altieri 1983 Agioccosystem diversity arnd pest control data tentative conclusions and nizw research dirctions Environ Entomol 12625-629

190 Rivera C T 1956 Three organic insecticides in thecontrol of insects affecting upland rice Philipp Agric 39465-472

191 Roffrey 1 1972 Locusta outbreaks in the Philippines Acrida 1177-188

192 Rojas R 1) 1941 The effect of pruning at varying ages upon the yield of Ramairio Phipp Agric 29851-860

193 Roonwal M i 1979 Ternite life and termite control in tropical South Asia Scientilic Pnbl odhpr India 139 p

194 Rosetto C1 F da Silva Martins N C Schmidt and L EAzzini 1978 Damage caused by pasturespittle bugs 4Dvoi)lavopictaandDJ schach) on rice lBragantia Nota 6 (Matcli)25-27

195 Rosztto C1- S S Neto I- Amantc I) Iink 1Garcia-Vieira D M de SoLza N V Bansa t and A M de Olivcira 1971 Rice pests of Brazil [in Portuguese] Pages 149-238 inrFA()-International Rice Commission 2nd meeting of the Rice Committee for the Americas 6-11 l)ecember 1971 Pelotas Rio Grande do Sul Brazil

196 Rothschild G H L 1969 Obseratiois on the armyworm Spodoptera matritia acronrctoiiles Gn (Lep Noctuidae) Bull Entomol Res 59143-160

197 Rothschild G 1- L 1974 Parasites of the armworm Spolopiera mauritiaacroncitoihes in Malaysia Fntornopliaga 19293-299

198 Sachan i N S K Gangwar and 1N Katujar 1980 Field crickets damaging cages in North Eastern IHill Region Indian J Entomol 42526-527

199 Sands 1) P A 1977 The biology and ecology of Lepocorisa (Hemiptera Alydidac) in Papua New Guinea Res Bull 18 Departshyment of Primary Industry Port Moresby Papua New Guinea 104 p

200 Sarawak 1968 titomology division Pages 97-106 in Report of the research branch of the Department of Altriculture 1966 Kuching (Rev AIppl Eintomol A56603)

201 Satpathy JM M S Das and K Naik 1977 Effect of multiple and mixed cropping on the incidence of scine important pestsI Entomol Res 178-85

202 Sauer IHF G 1939 Notes Ott Elasmpnlpualtsligniosellths Zeller (Lep Pyr) a serious pest ofcereals in Sao Paolo [in Portuguese] Arq Inst Biol 10199-206 (Rev Appl Entotnol A28378)

203 Satinders 1L A 1 SKing and C L Vargas S 1983 Crop pests of Central Anerica with t bibliography [in Spanish] Centro Agroshynotnico Tropical de Investigacion y Ensenanza Turrialba Costa Rica (CATIE) Tec Bol 9 90 p

204 Sauphanor B 1985 Some factors of upland rice tolerttce to stem borers in West Africa Inseu Sci Appl 6429434

205 Sen P atnd S Chakravorty 1976 Stentflics in Haritighata Nadia West Bengal India [it Rice Res Newsl 1(12) 15

206 Senapai B and 1M Satpathy 1983 The seedling fly Atherigona orrzae Malloch is a pest of rice in Orissa (India) J Entomol Res 777-79

207 Shhjahan M S Alatn M Ah nad and BN Islam 1983 A new scarabaeid beetle pest of dryland rice in Bangladesh Int Rice Res Ncwsl 8(I)16-17

208 Sharma G L S C Modgol and R C Gautam 1979 Effect of depth of submergence ott incidence of bacterial blight and white grub infestation in transplanted rice Int Rice Res Newsl 4(6)12-13

209 Sharma S 1985 Scarabaeid beetle outbreak itt upland rice Int Rice Res Newsl 1(1(l)24

210 Shiraki T 1917 The paddy borer Schoenobhits incertulas Walker [achaku Agric Exp Stn Formosa 256 p

211 Silva A de Brito and 11P Magalhaes 1981 Insect pests of rice [in Portuguese] Circular Tecnica No 22 EMBRAPA Centro de Pesquisa t ecuaria de Tropico Unido (CPAJU) Belem Brazil 14 p

212 Singh S A 1979 Emergence of rice leaf butterfly as a rice pest in Manipur India int Rice Res Newsl 4(4)16-17

213 SinghS A 1983 Changing trends of insect pestsof rice in Manipur Int Rice Res Newsl 8(f)12

214 Siwi S S and P H van Doesburg 1984 Lenovorisa Latreille in Indonesia (Heteroptera Coreidae Alydinae) Zool Meded (Leiden) 58117-129

215 Soejitno J 1977 Reaction of some upland rice varieties toseedling fly eltherikonaexigua Stein [in Indonesian] Siri Hana I Penyakit 630-35 Lembga I osat Fenclilian Pertanian Bogor Indonesia

216 Socnardi 1967 Insect pests of rice in Indonesia Pages675-683 in The major inse pests of thu rice plant Proceedings of a symposium at the Internatital Rice Rec Wi itt S pternber 1964 The Johns Hopkins Press Baltimore Maryland USA

217 Soto P E A T Petcz and I W Buddenhagen 1976 Survey of insect pests and diseases of rice in different ecological zones in Nigeria Rice Entomol Ncwsl 435-46

218 Soto 11E and Z Ziddiqi 1976 Screening for resistance to African rice insect pests Varietal improsetnent sCmi n r Bouak Ivory Coast 13-17 Sept 1976 West Africa Rice l)evelopment Association Monrovia Iiberia

219 Squire F A 1972 Entomological problems in Bolivia PANS (PestArtic News Sumr) 18249-268

220 Stapely 1 II 1978 The rice leaf roller in the Solomon Islands Int Rice Res Newsl 3(l)13

221 Suenaga H 1937 On aphids attacking the roots of rice Oyo-I)oubts Zasshi 9150-152 [in Japanese] (Rev Appl Entomol A25745)

222 Suenaga H 1952 On the seasonal abundance of three subterranean aphids on upland rice Bull Kyushu Agric Exp Sti 1249-262

223 Sutynso Yas and Dandi Soekaria 1964 Beberapa hana yang ttenperang tana man padi tanah kering [in Indonesian] Rapat kerja Padi Tanah Kering 6 sd 8 Juily 1964 Kaliurang 9 p

224 Tallamy I) W and R F Denno 1983 Alternative life history patterns in risky environments in example from lace bugs Pages 129-147 in Insect life history patterns R F l)cnio and If Dingle eds Spring-Verlag 225 p

225 Tamura 1 1939 Observations on some insect pests of field crops [in Japanese] 1 Plant Prot 26414-420 (Rev Appl EntomolA27-556)

226 Tamura and Suzuki 1964 Influences of the early-transplanting practice in rice culture upon development of the rice stem borer Chilo siql)ressalisWalker Bull Ilokuriku Natil Agric Exp Stn 761-94

227 Tan J P 1924 The rice root aphid Drowhi ihirsoia Philipp Agric 13277-288

228 Tanaka 1 1957 Taxonomy and distribution of some subterranean aphids injurious to upland rice in Japan with description of a new species [in Japanese English summary] Botyu-Kagaku 22168-176

229 Tanaka T 1961 The rice root aphids their ecology and control Spec [full College of Agriculture Utsunomiya University Japan F3 p

230 Tauber M IC A Tauber R Nechols and 1Obrycki 1983 Seasonal activity of parasitoids control by external internal and genetic factors Pages 87-108 itt I)iapause and life cycle strategies in insects V K Brown and I Ilodek eds )r W Junk The Hague Netherlands

231 Taylor W F 1972 Effects of artificial defoliation (simulating pest damage) on varieties of upland rice Exp Agric 879-83

232 feotia T P S and S Natud 1966 Biotoomics of the rice skipper Parnaraniathiis Fabricius (Lepidoptera Hesperiidac) Indian J Entomol 28181-186

233 Teran F 0 1971 Rice pests of Santa Cruz [in Spanish Ministerio de Agricultura Sita Cruz Bolivia Bull 45 8 p

234 Than Itun 1976 Population dynamics of the yellow rice borer Tryporza itcertulas (Walker) and its damage to the rice plant MS thesis University of the Philippines at Ios Bafios Laguna Philippines 92 p

235 Uichanco L B 1911 Pests Pages 55-74 it Rice Philipp Agric 11-200

IRPS No 123 January 1987 41

236 van der Laan P A 195 1 Life history of the rice seedling fly Contrib Gen Agric Res Stn Boor Indonesia 118 15 p

237 van der Laan 11A 1959 Correlation between rainfall in the dry season and the occurrence of the white rice borer (Scirpolhaga innotala WIk) in lava Entomol Fxp Appl 212-20

238 Van Driesche R G 1983 Meaning of percent parasitismin studies of insect parasitism Fnviron Entomol 121 I1- 122

239 Venugopal M S and A litsingcr 1984 Fffect ofcarbofuran on rice growth Prot Ecol 7313-317

240 Vietnam Directorate of Rural Affairs 1962 Annual work progress report on crop improvement program of rice sugarcane vegetable and field crops (lor the period from July 1961 to June 1962) (11) 299 p (Rev Appl Entomol A51526-527)

241 Wan M T K 1972 Observations on rice leaf- and planthoppers in Sarawak (Malaysian Borneo) Malay Agric 148308-335

242 West Africa Rice I)esclopnient Association 1978 Annual research report Vol 2 Statistics Monrovia Liberia

243 West Africa Rice Development Association 1979 Annual research report p47 Monrovia Liberia

244 Williams ) F and CS lofgren 1981 Fli I illy l-468 anew bait toxicant for control of the red imported lire at Fla Entomol 64472477

245 Williams I) 1 1970 The mealybugs OFlomoptera Coccoidea Iscudococcidac) of sugar-cane rice and sorghum Bull Entomol Res 0109-188

246 Williams D) 1 1982 Pulinaria icervi (Signoret) and its allies ott sugar cane and other grasses Btll Entoinol Res 72111-117

247 Wirjosuiardjo S 1976 Study ofchenmical cont rot of insect pests ott upland rice Imiu Pertanian (Jogjakarta) 1323-329

248 Wirjosuhardjo S and N Wirarta 1976 Chemical control of the white grub tIolorithiatlltriin upland rice in M uneng Irobolinggo Indonesia) [in Indonesian] Imut Pertanian (Jogjakarta) 1330-332

249 Yano K T Miyake and V F Eastop 1983 The biology and economic importance of rice aphids (Henuiptera Aphid idae) a review Bull Entomol Res 73539-566

250 Yelp D and P Rhodes 19(4 lhe success of drilled dryland rice iii the Fiji Islands International Rice Commission Working Party on Rice Production and Protection Manila Philippines 3-10 March1964 Food and Agriculture Organization Rome Italy 13 p

251 Young W R 1981 Fifty-five years of research on the sorghum shootfly Insect Sci Appi 23-9

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No 101The economics of hybrid rice production in (hina No 113 Boron toxicity in rice No 102 Rice ratooning No 114 Energy analysis rice production systems and rice research No 103 Growth and development of the (Jeep water rice plant No 115 Production risk and optimal fertilizer rates an application of the

No 104 Faridpur acomputer-assisted instruction nmodel forrainfed lowland random coefficient model No 116 Consumer demand for rice grain quality in Thailand Indonesia andrice

No 105 A readingand listening comprehcnsion test inEnglish for non native the Philippines speakers applying for tiraining at IRIRI No 117 Morphological changes in rice panicle development a review of

No 106 Rice grassy stunt virus 2 a new strain of rice grassy stunt in tire literature

Philippines No 118 IRRI-Korca collaborative project for the development of cold-No 107 Physical losses and quality deterioration in rice plstprodnCtion tolerant lines through anthei culture

systemis No 119 Problen soils as potenial areas for adverse soils-tolerant rice No 108 (olptblication ofIRRI materials a survey of translators and varieties in South and Ssuthcasi Asia

publishers No 120 Changes isinall-farm iLtireliing techii)hgy in Thailand and [he No 109 Classification of Philippine rainfall patterns Philippines No 110 Contributions of modern rice varieties to rutrition in Asia No 121 Landforms and modern rice varieties No Ill Changes in rice breeding in 10 Asian countries 1965-84 No 122 Yield stability and modern rice technology No 112 Design parameters affecting the performance of the IRRI-designed

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IRPS No 123 January 1987 37

41 Cendcafia S M and F B Caora 1967 Insect pests of rice in the 62 Ene 1 C 1981 I)iiorldhol)ierusihilis Slater (licteroplera Philippines Pages 591-616 in The najor insect pests ofthe rice plant lygacidue Illissinac) a new rice pest front Nigeria lit Rice Res Proceedings of a symposium at the International Rice Research Newsl N3)15 Institute September 1964 The Johns Hopkins Press Baltimore 63 Fugenio T S 1965 rhe effect oi e-vCgetation on rice production Maryland USA 729 p and the possibility of overcoming it by leaf pruning MS thesis

42 Centro Ihternacional de Agricklittira Tropical 1984 Upland rice in University ol the Philippines at Los Baijos ILaguna Ihilippins 69 p Iatin Anerica Pages 93-119 hi An os ervies of uplaind rice research 04 Farrow R A 1975 The Alrican nmigratory locust ii is main Proceedings of the 1982 Boinakamp Ivory Coast upland rice workshop outbreak area of the middle Niger qlinlitatise studies of solitary International Rice Research Institute P0 Blox 933 Manila populations in relation to environmental factors Locusla No II Philippines 198 p

43 Chandra G 1979 Taxonomy ad hn riononlics ofthe imect parasites of 65 Fennal R G 1943 Food crop investigation Windward and rice Icalloppers and planthoppers in the Pihilippines and their Ieeward Islands First report Report onl work dunring the periodiiiportairce iii natural biol)gical cointrol Philipp Fntoltol 4119-139 April 1942-April 1943 Imperial College of Tropical Agriculture

44 Chang T T 1976 The originii colut ion enlIivation dissemination Trinidad 16 p (Rev of Appl Fintornol A3222-25) and diersificition of Asian and African rices -uphytica 25425-44 1 66 Ferreira F 1983 Integrated pest ritaageiret of upland rice [ill

45 Chatierjee P B 1979 Rice leaffolder attacks in India lit Rice Res Portuguese English summary] Pages 323-341 in Upland riceculttre Newsl 4(3)21 factors affecting production E Ferreira T Yamada and F-

46 Cheng Cbien-pan 19(4 Upland rice in Taiwan International Rice Malmolta eds Institutoda Potassae Foslato Institnto Internacional Cnmmission Wiorking Party oit Rice Production ard Protection I()th da Potassa Piracicaha Sao Paulo lraid Meeting 3-10 March 1964 Manila Philippines Food and Agricul- 67 Ferreira E arid F dl Silva Martins 1979 Occurrence of ture Organiaion Rome Italy 5 p Sogatohs orvziola (Muir) inupland rice in Goias Ilrail lit Rice

47 Chcrrett 1 M and 1) 1Peregrine 1976 A review of the status of Rcs Ncwssl 4(4)18 leaf-cutting ants and their control Ann Appl Biol 84124-128 68 Ferreira E and 1F da Silva Martiris 1984 Insect pestof rice ill

48 Chipeta F M 1974 Pests of rice in Malawi Pages 90-95 in Brazil and their control Jill Portugucsej FMIlRAtPA-CNPAF Proceedings Sith eastern Africa cereals research conference 10-15 Goiania Brmiil 67 p March 1974 DRB Mialda ed Ministry of Agriculture lilongwe 69 Ferreira FI F ca Silha Miartins S S Nero and 1 It P de Malawi Carvallio 1982 M tltiplc regression analysis todeterimine the effect of

49 Chu Yau-i 1979 The d atiaging period of the rice seedling fly insect populatiois ott upland rice yield [inIPortuguese Pesqui (Atlterigotiavxigu Stcin) (NItiscide Diptera) oil cori) at Fast Java Agropecu Bras 17671-675 Indonesia Plant Prot Bill 21403413 70 Ferreira F 1F da Silva Martins and F 1PZimmermann 1979

50 Conklin H C 1957 IHaninoo agriculture a report on aln integral Rice varietal resistance to tle lesser corn stalk borer [in Port uguese] systent of shi fti rig cult ivationi in the Pltilippincs Food and Agriculture Pesqui Agropccu Biras 14317-321 Organization Rome Italy 209 p 71 Fleming W F 1968 Biological control of lie hapanese beetle U S

51 Cramer H II 1967 Plant proection and world crop production Dep Agric RLs Sers [ech Bull 1383 78 p Pilaniensclhutz-Nachr 201-524 72 Food and Agriculture Organiation of the United Nations 1982

52 l)ani R C and Majunidar 1978 Prelininary observations on tie Country report surmimary Pages 27-37 in Report on the first incidence of root aphids on differeit rice cultnrcs Sci Cult 4488-89 FAOIUN)IPhail and regional training coiuirse in iniproved ctltural

53 Davidson R I 1979 lffects of root feeding on foliage yield Pages practices for upland rice 5-23 Oct 1981 Northern Region Agrculturd117-120 in T K Ci osb and R B Pottinger eds Proceedings of the Developririt Center Chiang Mai Food and Agricuilture Organiiashy2nd Australian conlerence on grassland invertebrate ecology Govt tion Rome Italy 37 n Printer Wellington New Zealand 73 Franco do Ariaral S 195 1 Residual action of insecticides in turfy

54 l)ean G 1W 1976 Rice insect pests inlaos Int Rice Res Newsl soil--the results of a field experiment [in Portuguese] Biol6gico (Sao 1(2)15 Paulo) 17209-214 (Rev Appl Entomol A41423)

55 I)ean G J W 1978 Insect pests of rice in Laos PANS 24(3)280-289 74 Freeman ) 1970 Report onithe Iban london School of -coroiics 56 )ingle H 1984 Behavior genes and lifc histories conplex adapta- Monographs on Social Anthropology No 41 Athlane Press

tions to uncertain environments Pages 169-194 in A new ecology London UK 317 p novel approaches to interactive systems P W Price C N Slobod- 75 Fuchs T W1 A Harding and 1W Smithlr 1979 Induction and chikoff and W S Gaud cdsl ohn Wiley New York USA termination oldiapause in the sugarcane borer Ann Flitonlol Soc

57 Dobelmann-JP and M Falais 1968 Upland rice cultivation ill tie Am 72271-274 North-western part of Madagascar in 1966-1967 [in French English 76 (hbriel B1) 1975 A review tf the major insect pests of some tpland summary] Trop Agron 2313-21 crops in tlc Philippines Pages 86-111 in Ad Htoc panel of experts on

58 Doniiciano N L 1979 Chemical control of tie lesser corn stalk pest disease arnd weed problems in some rainfed crops Bangkok (lolsonno~paloIgnowslhts) (Zeller 1848) (I-epid(ptera Ihycitid ae) Thailand Food aid Agriculture Organization Roire Italy

on rice in Ponta Gressa Parara 1977-1978 [in Portuguese] Pages 77 Garg D K and N K Shal 1983 White grub outbreak on rainfed 291-303 in Anais 1 Reuniao de tecnicos cmiirizicultura do estado de dryland rice in Uttar Pradeslh Int Rice Res Ncwsl 81(4)16-17 Sao Paulo-Campinas Cati Brazil 78 Garg I) K and IP Tandon 1982 Major insect pests of rice on hilly

59 Dunsmore 1 R 1970 Investigations on tlie varieties pests and tracts of Uttar Pradesh India lilt Rice Res Newsl 7(11)11-12diseases of upland rice (hill padi) in Sarawak Malaysia Int Rice 79 Garrity 1) P 1983 Upland rice research an international bibliograpy Comm Newsl 1929-35 1975-1982 Iternational Rice Research hnstitnte PO Box 933

60 Elias 1R1965 The new decline of rice seedlings is not caused by Manila Philippines 121 p parasites [in Portuguese] Biol6gico (Sao Paulo) 313-6 80 Ghauri M S K 1982 A new subspecies of Iinorh)rone bull

61 Empresa Brasiliera de Pesquisa Agropecuaria 1984 Upland rice itt cornlus Slater (Hemiptera Iygaeidae) front Papua New Guinea on Brazil Pages 121-134 it Av overview of upland rice research rice and carpet grass Ihll lEntoniol Res 72133-137 Proceedings ofthe 1982 Blouak Ivory Coast Upland rice workshop 81 Gonzales F 0 PArregoces R I ihernandef and OT Parada International Rice Research Institute PO Box 933 Manila 1983 Insect and imite pests of rice arid their control in Latin America Philippines [in Spanish] la Federacion Nacional de Arrocerts (Fedearroz)

Bogota Colombia 60 n

38 IRPS No 123 January 1987

82 Grist D H and R J A W Lever 1969 Pests of rice Longmans London UK520 p

83 Hale P R 1979 Two subterranean pests of upland rice in Papua New Guirnca IntRice Res Newsl 4(2)16-17

84 Hale P R and J T Hale 1975 Insect pests of rice in the New Guinea Islands Rice Entomol Newsl 34

85 Harris W V 1969 Termites as pests of crops and trees Common-wealth Institute of Entomology London UK 41 p

86 Hattori 1 1968 A method to distinguish between grass leaf roller (Ctaplha rocis medinalisGuenee) and the rice leaf roller (Susurtia exigua Butler) [in Japanese] Plant Prot (Japan) 23167-170

87 Henry I E and 1A Onsager 1984 Experimental control of the mormon cricket Iahrosshnph by NJsena locustae(Aficrospora Microsporidae) a protozoan parasite of grasshoppers (Ort Acri-didae) Entomophaga 27197-201

88 Hidaka 1 P Vungsilabulr and SKadkao 1974 Studies on ecology and control of the rice gall midge in Thailand Tech Bull 6 TropicalAgricultural Research Center Tokyo Japan 113 p

89 Hincklcy A 1) 1963 Ecology and control of rice planthoppers in Fiji Bull Entomrol Res 53467 481

90 Hirao J 1982 Ecology and chtInical control of the rice leaffolder Jpn Pestic Inf 4114-17

91 Ho DT 1984 Sten borer incifience in rice ecosystems in KenyaEast Africa Pages 307-320 in A overview of upland rice research Proceedings of the 1982 Bouac Ivory Coast upland rice workshop International Rice Research Institute PO Box 933 Manila Philippines

92 Ho )T J 1Njokah and J GKibuka 1983 Studies on rice sten borers in Kenya with emphasis on ifaliarpiaipiraellRag Insect Sci Appl 465-73

93 Ichihara I T Ito A NIatsuo and M Sawada 1971 Chemical control of the rice striped stem borer in upland rice grown in plastic mulch [iii Japanese] Proc Kanto-Tosan Plant Prot Soc 1882

94 IclhiharaIand A Matsuo 1969 Damage caused by rice striped stett borer to upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1686-87

95 Ichhara I atid A MIatsun11970 Conside ration oft lie extraordinary prevalence of rice striped stem borer in upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1778-79

96 Institut de Recherches Agrolnliqes Iropicale et des Cultures Vivrieres 1978 Insect resistance Pages 12-13 in Upland rice 1978 atnual report Bouak Ivory Coast 27 p

97 International Institute for Tropical Agriculture 1983 Aphids itt rice roots I ITA Res Briefs 4(1-3)4-5

98 International Institite fi Tropical Agriculture 1983 Hopperburn in rice caused by brown plathopper IITA Res Briefs 4(l-3)6-7

99 International Rice Research Institute 1975 Annual report for 1974 PO Box 933 Manila Philippines p 216

100 International Rice Restarch Institute 1976 Annual repiort for 1975 PO Box 933 Manila Philippines p 228

101 International Rice Research Institute 1979 Cropping systems program Component technology development and evaluation Page 404 i Annual report for 1978 11O Box 933 Manila Philippines

102 International Rice Research Institute Department of Entomology 1981) Insecticid evaluation 1979 Tables 73 74 PO Box 933Manila Philipes

103 International Rice Research Institute Department of Entonmology 1981 Insecticide evalomtion 1980 Table 81 POBox 933 Manila Philippines

104 International Rice Research Institute 1981 Report of a meeting of Asian rice-based cropping systems entomologists Eleventh cropping systems working group meeting 18-22 May 1981 PO Box 933 Manila Philippines p 132 133 144

105 International Rice Research Institute 1984 Cropping systems program Pages 368-373 in Annual report for 1983 PO Box 933 Manila Philippines

106 International Rice Research Institutc 1984 Upland rice in Asia Pages 45-68 in An overview of upland rice research Proceedings of the 1982 Bouakamp Ivory Coast upland rice workiiop loternational Rice Research Institute 11O Box 933 Manila Philippines

107 Irshad M 1977 Grasshoppers in rice in Pakistan lit Rice Res Newsl 2(3)8-9

108 Ito R and TOno 1964 Upland lice its culture and varieties in Japan lIternational Rice Cormmission loth Meeting of the Working Party on Rice Production and Protection 3-10 March 1964 Manila Philippines Food and Agriculture Organization Rome Italy 12 p

109 Jeffrey M I and V A Dyck 1983 Effect of moonlight on lightshytrap catches of brown planthopper lit Rice Res Newsl 8(6) 13

110 Jennings P R and T Alicia Pineda 1983 Management ofthe rice plnthoppers Sogatodsorivi(coIl in Latin America Presented at the meeting on stable plant resistance FAO and lIndocsian I)inectorate of Food Crops Protection Bali lndonesia 29 Novcniber-2 l)ccemshyber 9 p

IllJimenez Gomez J A 1979 Rice insect pests i lla ios ()riCntales lil Spanish] Pages 99-103 in Rice ciltivation Instituto (olotibiatto Agropecuario Compndio No 29

112 Jongeleen F JJ Soesanto and A Mockidjo 1979 Studies oti pathogens of two species of Scarabacidac froinIndonesia Z Angew Enlomnol 87412-417

113 Jouvenaz 1) PC S lofgren and W A Baiks 1981 Biological control of imported fire antsa review of cirrent knowledge Bull Entomol Soc Am 27203-208

114 Kainoh Y K Shiniiu S MarUani Y Taiaki 1980 lost finding behavior of tie rice bu lplotori+JchinvittjI)allas (liciniptera Corcidac) with special reference to diet pattern ol aggregation and feeding on the rice plant Appl Entomol Zool 15225-233

115 Kakkar K 0 P Sharma and P I Sharma 1977 Note on Ihteronyrchus liodresRedt (Coleopicra I)ynastidae) as asporadic pest of paddy and maie inHimachal Pradesh Corr Agric 156-57

116 Kalode M131 79 Major insect pest problems and their control in upland rice (Unpublishcd mineogriph CWI oJNaaiat lni Chandaratat (198 1)original not seen)

117 Kalshoven L i 98 I The pests of crops in I ldoniesia (translated by P A vatn der Laan) 1 T Ichtiar B3aru-vati Iloeve lakarta Indonesia 701 p

118 Kamal N Q 1981 Suppression of whitebacked planto ner Sogatella fircaifra (Horvath) aLId rice lealfolder (nhal -cis licdb lis (Guenee) populations by natural cnclmie hlI) tiesis Grcgorio Araneta University Foundation Victoncta Park Metto Manila Philippines 179 Ip

119 Karim A N M Reaul 1975 A species ofliealy bug Trioni i70s ceres new oii rice in the Philippines Rice Entoniol Newsl 247

120 Katanyukul Wand C Chandaratat 1981 Insect pestsof uplaid rice and their control Report firstFAOUNlFi Tliailand training course on improved cultural practices for tplad rice Chiang Mai Thailand 5-23 Oct 1981 Food and Agriculture Organiation Rome Italy 22 p

121 Kern F 1955 lhletwioa(I)uninemts) bhhit Biturm(Coleoptera) a new pest of sugarcane inVenezuela [inSpanish with Etnglish sunmary] Agron Trop (Maracay) 527-29 (Rev Appl Eitoniol A4572)

122 Khan K M and S Ghai 1974 White grubs ald their control in India Pesticides (1Bombay) 819-25

123 Kiritani K 1975 Integrated pest management of rice pests Pages 66-70 itt Proceedings platning workshop ott cooperative hIeld research in pest managementP SMotooka ed East-West Center Honololu Hawaii USA

124 Kisimoto R 1967 Ecology of insect vectors forecasting and chemical control Pages 243-255 in The virus diseases of tle rice plant Proceedings of a symposium atthe International Rice Research Institute April 1967 The Johns Hopkins Press altimore Maryshyland USA

125 Kisimoto R and V A Dyck 1976 Climate and rice insects Pages 367-391 in Climate and rice International Rice Research Institute PO Box 933 Manila Philippines

126 Kureha S S Kobayashi SAsari and T Ishihara 1977 Life history and control of the mole cricket Grrllotalpa africana Palisot de Beauvois [in Japanese English translation 7520 1RRI library] Proc Kanto-Tosan Plant Prot Soc 21192-197

127 Kushwaha K S 1981 The black beetle as a pest of rice in Haryana India Int Rice Res Newsl 6(4)14

128 Lawani S M 1982 A review of tile effects of various agronomic practices on cereal stem borers Trop Pest Manage 28266-276

129 Leuck D B 1966 Biology of the lesser cornstalk borer in South Georgia J Econ Entomol 59 797-801

130 lever R J A W 1955 Insect pests of the rice plant in Malaya Int Rice Comm Newsl 1612-21

131 Li Ching-Sing and Shin-Foon Chiu 1951 The study ofthe rice gall midge Iachydiplosis orvza Wood Mason [in Chinese with English summary] 1Taiwan Agric Res 21-13

132 Iitsinger J A 1984 Assesament of the need-based application for rice Proceedings TechnologyTransfer Workshop 15-16 March 1984 International Rice Research Institute 1PO Box 933 Manila Philippines 6 p

133 Litsinger J A 1984 Environmental diversity and insect pest abundance with reference to the Pacific Pages 285-307 in Proceedings sub-regional training course on methods of controlling diseases insects and other pests of plants in the South Pacific 4-20 Oct 1982 Vaini Kingdom of Tonga Nukualofa

134 Litsinger J A RF Apostol and M BObusan 1983 White grub Leucopholis irrorata (Coleoptera Scarabaeidae) pest status popula-tion dynamics and chemical conllol in arice-maize cropping pattern in the Philippines J Econ Entoinol 761133-1138

135 Litsinger J A C de la Cruz F Raymundo A T Barrion M D Lumaban J P Bandong M S Venugopal F Paragna and E Balete 1982 Insect pests and insecticide reponse of dry-seeded bunded rice Pages 179-193 in Report of workshop on cropping systems research in Asia International Rice Research Institute Los Bafos Philippine

136 Litsinger J A M 1) Luinaban J P Bandong A T Barrion W Katanyukul and N Panda 1978 Insect pest management in rainfcd lowland rice agroecosystms of tropical Asia International Rice Research Conference 17-21 April 1978 International Rice Research Institute Los Bafios Philippines

137 Litsinger J A and K Moody 1976 Integrated pest management in multiple cropping systems Pages 293-317 in R I Papendick P A Sanchez and G B Triplett eds Multiple cropping American Society of Agronomy Madison Wisconsin USA

138 Loevinsohn M E 1984 The ecology and control of rice pests in relation to the intensity and synchrony of cultivation Ph D thesis University of London 36) p

139 Loevinsohn M E J A Litsinger N Panda M A Austria R F Apostol C Andrion and J Bandong 1982 A basis for predicting rice insect pest abundance in cropping systems Paper presented at the 1982 International rice research conference on rice and cropping systems 19-23 April 1982 International Rice Research Institute Los Bafios Iaguna Philippines 25 p

140 Lopez A W 1930 Repot of the Entomology Department Pages 145-172 in Annual report of the Philippine Sugar Association 1929-30 Manila

141 Lopez AW 1931 Entomology Department Pages 227-266 in Annual report of the director or research 1930-31 Philippine Sugar Association Manila

142 Lopez A W 1932 Entomology Deparment Pages 252-279 in Annual report of the Philippine Sugar Association 1931-32 Manila

143 Lungibill P and G G Ainslee 1917 The lessercornstalk borer US Dep Agric Agric Res Serv Tech Bull 539 27 p

IRPS No 123 January 1987 39

144 McGuire J U Jr and BS Crandall 1967 Survey of insect pests and plant diseases ofselected food crops of Mexico Central America and Panama U SDepartment of Agriculture International Agriculshytural Development Service Agricultuial Researrh Service October 1967 11 p

145 McNeill Sand T R ESouthwood 1978 The role of nitrogen in the development of insectplant relations Pages 77-9 in lliochemical aspects of plant and animal coevolution J B Ilarbornu ed Academic Press London UK

146 Maebara H 1955 Spatial distribution of lastinstiarlanaeof purplish stem borer (Sesamia iqferens Walker) hibernating in an upland rice field Bull Kagoshima Univ Fac Agric 438-41

147 Mallick E H and N G Hajra 1977 Response of rice plant to defoliation treatments at different developmental stages Indian Agric 2151-56

148 Mansingh A 1971 Physiological classification of dormancies in insects Canl Entomol 103983-10)9

149 Martell C G 1974 Catalogue of plant feeding insects of Mexico [in Spanish] Fitofilo 27(69)1-176

150 Martinez 0 J 1982 Principal rice pests in Ilanos orientalis [in Spanish] P~ages 80-94 in Curso deact nalization de conocimientos en arroz Villaniencis Instituto (olonmbiano Agropecuario

151 Martins J F da Silva F Ferrcira A S Prabhu F 1 P Zimmermann 1980 Use of pesticides for the control of the main subterranean upland rice pests esqui Agropecu Bras 1553-u

152 Martins I F da Silva F Fe eira rod BdamSilveira Pinheiro I Q2 Damage simulation caused by defoliators of upland rice [in Portuguese with English summary] Pesqui Agropecu Bras 171113-1119

153 Martins 1F da Silva N(iyuen van Tan and 1Ida Silveira Pinheiro 1981 Stem borer resistance in upland rice associated with plant morphological characters [in Portuuce Iesqui Agropecu Bras 16187-192

154 Masaki S 1980 Sumnmer diapause Ann Rev Entomiol 251-25 155 Masuzawa T 11 Suwa and F Nakasoji 1983 l)ifkleences of

oviposition preferencc and survival rate of two skipper butterflies Parnara guttata and PIchlpidai iathiais (Iepidoptera HIesperiidac) on rice plant and cogon grass New Entomol 3247-56

156 Maurya D M and C P Vaish 1984 Upland rice in India Pages 69-92 in An overview oftpland rice research Proceedings ofthe 1982 Bouakamp Ivory Coast upland rice workshop International Rice Research Institute IO Box 933 Mnila Philippines

157 Menon M G R R N Katiyai S Kuar and S S Misra 1970 Termites attacking the paddy crop Sci Cult 36294-295

158 Miyahara Y and 1 Nishikubo 1967 Behavior of young nymphs of the red rice root aphid Iholahsipiun ri liabhdontinalisSasaki in relation to movement from the leaf to the roots of upland rice [in Japanese] Pioc Assoc Plant Prot Kyushu 1363-66

159 Mo ljoa Tjien 1952 White grub of upland rice (Holotrichiahelleri Brsk) [in Indonesian with English summary] Contrib Agric Res Stn Bogor No 13134 p

160 Mochida 0 and T Suryana 1979 The occurrence of the brown planthopper and ragged stunt on lowland and upland rice ill Central Java and logyakarta in tile late of January 1979 Entomnolgy Survey Report No 15 Central Research Institute of Agriculture (CRIA) Sukamandi Indonesia I February 1979 9 p

161Monet C 1979 Varietal resistance to upland rice pests [inI French] Pages 35-39 in Annual report 1977 Institut des Savanes Department des Cultures Vivrieres Bouak6 Ivory Coast

162 Monte 0 1942 A caterpillar ofrice-fields[in Portuguese] Biol6gico (Sao Paulo) 8161-163 (Rev Appl Entomol A31414)

163 Munroe I) D 1979 Pest management strategy for hill rice in four resettlement areas in Sarawak Malaysia Pages 89-102 i1 Guidelines for integrated control of rice insect pests Food and Agriculture Organization Plant Production and Protection Paper 14 Rome Italy 115 p

40 IRPS No 123 January 1987

164 Nakasuji F M Ishii and Jen-Zon Ito 1984 Rice skippers in Taiwan and their life histories ZZrC (Taipei) Tech Bull 82 10 p

165 Navas I) 1976 Fall armyworum in rice Pages 99-106 il Ecological animal control by habitat management Proc Tall Timbers Con-ference 28 Feb-I Mar 1974 Fall Timbers Res Stn Misc Publ 6 Tallahassee Horida USA

166 Nickel 1 1 1961 Biological control of rice stem borer A feasibility study Tcch Bull 2 International Rice Rcarch Institute PO Box 933 Manila Philippines IIl p

167 Nigan P M and R A Vernia 1985 Insect pests of uplaild rice in Uttar Pradesh Int Rice Res Newsl 10(6)22

168 Nilakhe S S 1985 -cological observations on spittle bugs with emphasis on their occirence in rice Pcsqtui Agropecu Bras 20407-414

169 Nixon G EJ 195 1 lie association of ats with aphids and coccids Commonwealth Iusti lcte of Entonology Iondon UK 36 p

170 Nurdin F an Ishak Manti 1985 Study oit seedling fly of upland rice Interlational Upland Rice Moitoring Tour Report 26 lebruary-2 MarLI Sukaranti West Sumatra

171 Oka 1 N 1979 Ciltural conttol of the brown planthopper Pages 357-369 in Brown planthopper threat to rice production in Asia International Rice Research Institutc 1p0 Box 933 Manila Philippines

172 Ohni M and A Vilnaii 1975 Rice insect pests of Italy biology of Sipha grcria (Kaltenbach) and its control [in Italian] Riso (Milan) 2459-71

173 Otanes F Q atnd 1 I Sison 194 1 Pests of rice Philipp J Agric 12211-259

174 Painter R I 1951 Insect icsistance in crop plants University Press of Kansits Lawrence Kansas USA 520 p

175 Pantua P C and 1A litsinger 1981 Arthropod abundance in continuous vs biannual rice cropping systeis Int Rice Res Ncwsl 6(2)20-21

176 Pathak M ) and V A l)yck 1975 Sttudics ott insect pests of upland rice Pages 186-191 in Major research in upland rice International Rice Research Institute PO Box 933 Ilanila Philippines

177 Patil B R B M IHasabe and 1) S Ajri 198 I Chemical control of white gritb lIditriihinhorraia Fabr intfesting groundntut paddy atnd sorghum Indian 1Entomol 43232-236

178 Pawar A 1) 1974 (ollection and identification of insect pests of rice rlerminal repirt lInternational Rice Research Institute PO Box 933 Manila Philippines 72 p (tnimco)

179 IerrajU A aid A S Reedy 1965 Observations ott diapausc itt tie steti borer ohrice Rice News Teller (lai-Mar)9-13

180 Pont A C 1972 A review of the Oriental species of theri ona Rondani (I)iptera Muscidac) of econtomic importance Pages 27-102 in Control of sorglium shoot fly M G lomtani and W R Young eds Oxford atd IH1 Ptthl Co New I)clhi India 324 p

181 Prasad S C 1PChand aid M 1PSingh 1983 Incidence of rice gall midge in the upland plateau region of Hihar Oryza 20235-236

182 Rai P S 198 0 Nisaga simpex damage to rice in the hill tracts of South India Int Rice kles News[ 5(I)12

183 Rathorc Y S 1969 Gonocplhahli dcprssunt Fabr as itpredator of (tilo zonwIlhis (Swinhoc) and other moths in field cage at Patnagar Uttar Pradesh Indian 1Entotnol 3127

184 Reed W 1983 Crop losses causcd by insect pists in the developing world Proceedings 10th Internatial Congress of Plant Protection lrighton UK 174-80

185 Reedy M V 1981 Ateactogasterinhens(Walker)a new root-grub pest of tipland paddy and inaize Curr Sci 501076

186 Reveno M A 1968 Catalogue of the insect pests of economic crops in Colombia [in Spaiish] Asociacion Latino Americana de Entomo-logia (AIAE) 129-33

187 Reyes G M 1957 Rice dwarf disease in the Philippines Food and Agriculture Organiiation Plant Prot Bull 617-19

188 Rice S E A A Grigarick and M 0 Way 1982 Effect of leaf and panicle feeding byarmyworm (Lepidoptera Noctuidae) larvae on rice grain yield Environ Entomol 7593-595

189 Risch S J 1) Andow and M A Altieri 1983 Agioccosystem diversity arnd pest control data tentative conclusions and nizw research dirctions Environ Entomol 12625-629

190 Rivera C T 1956 Three organic insecticides in thecontrol of insects affecting upland rice Philipp Agric 39465-472

191 Roffrey 1 1972 Locusta outbreaks in the Philippines Acrida 1177-188

192 Rojas R 1) 1941 The effect of pruning at varying ages upon the yield of Ramairio Phipp Agric 29851-860

193 Roonwal M i 1979 Ternite life and termite control in tropical South Asia Scientilic Pnbl odhpr India 139 p

194 Rosetto C1 F da Silva Martins N C Schmidt and L EAzzini 1978 Damage caused by pasturespittle bugs 4Dvoi)lavopictaandDJ schach) on rice lBragantia Nota 6 (Matcli)25-27

195 Rosztto C1- S S Neto I- Amantc I) Iink 1Garcia-Vieira D M de SoLza N V Bansa t and A M de Olivcira 1971 Rice pests of Brazil [in Portuguese] Pages 149-238 inrFA()-International Rice Commission 2nd meeting of the Rice Committee for the Americas 6-11 l)ecember 1971 Pelotas Rio Grande do Sul Brazil

196 Rothschild G H L 1969 Obseratiois on the armyworm Spodoptera matritia acronrctoiiles Gn (Lep Noctuidae) Bull Entomol Res 59143-160

197 Rothschild G 1- L 1974 Parasites of the armworm Spolopiera mauritiaacroncitoihes in Malaysia Fntornopliaga 19293-299

198 Sachan i N S K Gangwar and 1N Katujar 1980 Field crickets damaging cages in North Eastern IHill Region Indian J Entomol 42526-527

199 Sands 1) P A 1977 The biology and ecology of Lepocorisa (Hemiptera Alydidac) in Papua New Guinea Res Bull 18 Departshyment of Primary Industry Port Moresby Papua New Guinea 104 p

200 Sarawak 1968 titomology division Pages 97-106 in Report of the research branch of the Department of Altriculture 1966 Kuching (Rev AIppl Eintomol A56603)

201 Satpathy JM M S Das and K Naik 1977 Effect of multiple and mixed cropping on the incidence of scine important pestsI Entomol Res 178-85

202 Sauer IHF G 1939 Notes Ott Elasmpnlpualtsligniosellths Zeller (Lep Pyr) a serious pest ofcereals in Sao Paolo [in Portuguese] Arq Inst Biol 10199-206 (Rev Appl Entotnol A28378)

203 Satinders 1L A 1 SKing and C L Vargas S 1983 Crop pests of Central Anerica with t bibliography [in Spanish] Centro Agroshynotnico Tropical de Investigacion y Ensenanza Turrialba Costa Rica (CATIE) Tec Bol 9 90 p

204 Sauphanor B 1985 Some factors of upland rice tolerttce to stem borers in West Africa Inseu Sci Appl 6429434

205 Sen P atnd S Chakravorty 1976 Stentflics in Haritighata Nadia West Bengal India [it Rice Res Newsl 1(12) 15

206 Senapai B and 1M Satpathy 1983 The seedling fly Atherigona orrzae Malloch is a pest of rice in Orissa (India) J Entomol Res 777-79

207 Shhjahan M S Alatn M Ah nad and BN Islam 1983 A new scarabaeid beetle pest of dryland rice in Bangladesh Int Rice Res Ncwsl 8(I)16-17

208 Sharma G L S C Modgol and R C Gautam 1979 Effect of depth of submergence ott incidence of bacterial blight and white grub infestation in transplanted rice Int Rice Res Newsl 4(6)12-13

209 Sharma S 1985 Scarabaeid beetle outbreak itt upland rice Int Rice Res Newsl 1(1(l)24

210 Shiraki T 1917 The paddy borer Schoenobhits incertulas Walker [achaku Agric Exp Stn Formosa 256 p

211 Silva A de Brito and 11P Magalhaes 1981 Insect pests of rice [in Portuguese] Circular Tecnica No 22 EMBRAPA Centro de Pesquisa t ecuaria de Tropico Unido (CPAJU) Belem Brazil 14 p

212 Singh S A 1979 Emergence of rice leaf butterfly as a rice pest in Manipur India int Rice Res Newsl 4(4)16-17

213 SinghS A 1983 Changing trends of insect pestsof rice in Manipur Int Rice Res Newsl 8(f)12

214 Siwi S S and P H van Doesburg 1984 Lenovorisa Latreille in Indonesia (Heteroptera Coreidae Alydinae) Zool Meded (Leiden) 58117-129

215 Soejitno J 1977 Reaction of some upland rice varieties toseedling fly eltherikonaexigua Stein [in Indonesian] Siri Hana I Penyakit 630-35 Lembga I osat Fenclilian Pertanian Bogor Indonesia

216 Socnardi 1967 Insect pests of rice in Indonesia Pages675-683 in The major inse pests of thu rice plant Proceedings of a symposium at the Internatital Rice Rec Wi itt S pternber 1964 The Johns Hopkins Press Baltimore Maryland USA

217 Soto P E A T Petcz and I W Buddenhagen 1976 Survey of insect pests and diseases of rice in different ecological zones in Nigeria Rice Entomol Ncwsl 435-46

218 Soto 11E and Z Ziddiqi 1976 Screening for resistance to African rice insect pests Varietal improsetnent sCmi n r Bouak Ivory Coast 13-17 Sept 1976 West Africa Rice l)evelopment Association Monrovia Iiberia

219 Squire F A 1972 Entomological problems in Bolivia PANS (PestArtic News Sumr) 18249-268

220 Stapely 1 II 1978 The rice leaf roller in the Solomon Islands Int Rice Res Newsl 3(l)13

221 Suenaga H 1937 On aphids attacking the roots of rice Oyo-I)oubts Zasshi 9150-152 [in Japanese] (Rev Appl Entomol A25745)

222 Suenaga H 1952 On the seasonal abundance of three subterranean aphids on upland rice Bull Kyushu Agric Exp Sti 1249-262

223 Sutynso Yas and Dandi Soekaria 1964 Beberapa hana yang ttenperang tana man padi tanah kering [in Indonesian] Rapat kerja Padi Tanah Kering 6 sd 8 Juily 1964 Kaliurang 9 p

224 Tallamy I) W and R F Denno 1983 Alternative life history patterns in risky environments in example from lace bugs Pages 129-147 in Insect life history patterns R F l)cnio and If Dingle eds Spring-Verlag 225 p

225 Tamura 1 1939 Observations on some insect pests of field crops [in Japanese] 1 Plant Prot 26414-420 (Rev Appl EntomolA27-556)

226 Tamura and Suzuki 1964 Influences of the early-transplanting practice in rice culture upon development of the rice stem borer Chilo siql)ressalisWalker Bull Ilokuriku Natil Agric Exp Stn 761-94

227 Tan J P 1924 The rice root aphid Drowhi ihirsoia Philipp Agric 13277-288

228 Tanaka 1 1957 Taxonomy and distribution of some subterranean aphids injurious to upland rice in Japan with description of a new species [in Japanese English summary] Botyu-Kagaku 22168-176

229 Tanaka T 1961 The rice root aphids their ecology and control Spec [full College of Agriculture Utsunomiya University Japan F3 p

230 Tauber M IC A Tauber R Nechols and 1Obrycki 1983 Seasonal activity of parasitoids control by external internal and genetic factors Pages 87-108 itt I)iapause and life cycle strategies in insects V K Brown and I Ilodek eds )r W Junk The Hague Netherlands

231 Taylor W F 1972 Effects of artificial defoliation (simulating pest damage) on varieties of upland rice Exp Agric 879-83

232 feotia T P S and S Natud 1966 Biotoomics of the rice skipper Parnaraniathiis Fabricius (Lepidoptera Hesperiidac) Indian J Entomol 28181-186

233 Teran F 0 1971 Rice pests of Santa Cruz [in Spanish Ministerio de Agricultura Sita Cruz Bolivia Bull 45 8 p

234 Than Itun 1976 Population dynamics of the yellow rice borer Tryporza itcertulas (Walker) and its damage to the rice plant MS thesis University of the Philippines at Ios Bafios Laguna Philippines 92 p

235 Uichanco L B 1911 Pests Pages 55-74 it Rice Philipp Agric 11-200

IRPS No 123 January 1987 41

236 van der Laan P A 195 1 Life history of the rice seedling fly Contrib Gen Agric Res Stn Boor Indonesia 118 15 p

237 van der Laan 11A 1959 Correlation between rainfall in the dry season and the occurrence of the white rice borer (Scirpolhaga innotala WIk) in lava Entomol Fxp Appl 212-20

238 Van Driesche R G 1983 Meaning of percent parasitismin studies of insect parasitism Fnviron Entomol 121 I1- 122

239 Venugopal M S and A litsingcr 1984 Fffect ofcarbofuran on rice growth Prot Ecol 7313-317

240 Vietnam Directorate of Rural Affairs 1962 Annual work progress report on crop improvement program of rice sugarcane vegetable and field crops (lor the period from July 1961 to June 1962) (11) 299 p (Rev Appl Entomol A51526-527)

241 Wan M T K 1972 Observations on rice leaf- and planthoppers in Sarawak (Malaysian Borneo) Malay Agric 148308-335

242 West Africa Rice I)esclopnient Association 1978 Annual research report Vol 2 Statistics Monrovia Liberia

243 West Africa Rice Development Association 1979 Annual research report p47 Monrovia Liberia

244 Williams ) F and CS lofgren 1981 Fli I illy l-468 anew bait toxicant for control of the red imported lire at Fla Entomol 64472477

245 Williams I) 1 1970 The mealybugs OFlomoptera Coccoidea Iscudococcidac) of sugar-cane rice and sorghum Bull Entomol Res 0109-188

246 Williams D) 1 1982 Pulinaria icervi (Signoret) and its allies ott sugar cane and other grasses Btll Entoinol Res 72111-117

247 Wirjosuiardjo S 1976 Study ofchenmical cont rot of insect pests ott upland rice Imiu Pertanian (Jogjakarta) 1323-329

248 Wirjosuhardjo S and N Wirarta 1976 Chemical control of the white grub tIolorithiatlltriin upland rice in M uneng Irobolinggo Indonesia) [in Indonesian] Imut Pertanian (Jogjakarta) 1330-332

249 Yano K T Miyake and V F Eastop 1983 The biology and economic importance of rice aphids (Henuiptera Aphid idae) a review Bull Entomol Res 73539-566

250 Yelp D and P Rhodes 19(4 lhe success of drilled dryland rice iii the Fiji Islands International Rice Commission Working Party on Rice Production and Protection Manila Philippines 3-10 March1964 Food and Agriculture Organization Rome Italy 13 p

251 Young W R 1981 Fifty-five years of research on the sorghum shootfly Insect Sci Appi 23-9

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No 101The economics of hybrid rice production in (hina No 113 Boron toxicity in rice No 102 Rice ratooning No 114 Energy analysis rice production systems and rice research No 103 Growth and development of the (Jeep water rice plant No 115 Production risk and optimal fertilizer rates an application of the

No 104 Faridpur acomputer-assisted instruction nmodel forrainfed lowland random coefficient model No 116 Consumer demand for rice grain quality in Thailand Indonesia andrice

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No 106 Rice grassy stunt virus 2 a new strain of rice grassy stunt in tire literature

Philippines No 118 IRRI-Korca collaborative project for the development of cold-No 107 Physical losses and quality deterioration in rice plstprodnCtion tolerant lines through anthei culture

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publishers No 120 Changes isinall-farm iLtireliing techii)hgy in Thailand and [he No 109 Classification of Philippine rainfall patterns Philippines No 110 Contributions of modern rice varieties to rutrition in Asia No 121 Landforms and modern rice varieties No Ill Changes in rice breeding in 10 Asian countries 1965-84 No 122 Yield stability and modern rice technology No 112 Design parameters affecting the performance of the IRRI-designed

axial-flow pump

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38 IRPS No 123 January 1987

82 Grist D H and R J A W Lever 1969 Pests of rice Longmans London UK520 p

83 Hale P R 1979 Two subterranean pests of upland rice in Papua New Guirnca IntRice Res Newsl 4(2)16-17

84 Hale P R and J T Hale 1975 Insect pests of rice in the New Guinea Islands Rice Entomol Newsl 34

85 Harris W V 1969 Termites as pests of crops and trees Common-wealth Institute of Entomology London UK 41 p

86 Hattori 1 1968 A method to distinguish between grass leaf roller (Ctaplha rocis medinalisGuenee) and the rice leaf roller (Susurtia exigua Butler) [in Japanese] Plant Prot (Japan) 23167-170

87 Henry I E and 1A Onsager 1984 Experimental control of the mormon cricket Iahrosshnph by NJsena locustae(Aficrospora Microsporidae) a protozoan parasite of grasshoppers (Ort Acri-didae) Entomophaga 27197-201

88 Hidaka 1 P Vungsilabulr and SKadkao 1974 Studies on ecology and control of the rice gall midge in Thailand Tech Bull 6 TropicalAgricultural Research Center Tokyo Japan 113 p

89 Hincklcy A 1) 1963 Ecology and control of rice planthoppers in Fiji Bull Entomrol Res 53467 481

90 Hirao J 1982 Ecology and chtInical control of the rice leaffolder Jpn Pestic Inf 4114-17

91 Ho DT 1984 Sten borer incifience in rice ecosystems in KenyaEast Africa Pages 307-320 in A overview of upland rice research Proceedings of the 1982 Bouac Ivory Coast upland rice workshop International Rice Research Institute PO Box 933 Manila Philippines

92 Ho )T J 1Njokah and J GKibuka 1983 Studies on rice sten borers in Kenya with emphasis on ifaliarpiaipiraellRag Insect Sci Appl 465-73

93 Ichihara I T Ito A NIatsuo and M Sawada 1971 Chemical control of the rice striped stem borer in upland rice grown in plastic mulch [iii Japanese] Proc Kanto-Tosan Plant Prot Soc 1882

94 IclhiharaIand A Matsuo 1969 Damage caused by rice striped stett borer to upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1686-87

95 Ichhara I atid A MIatsun11970 Conside ration oft lie extraordinary prevalence of rice striped stem borer in upland rice grown with plastic mulch [in lapanese] Proc Kanto-Tosan Plant Prot Soc 1778-79

96 Institut de Recherches Agrolnliqes Iropicale et des Cultures Vivrieres 1978 Insect resistance Pages 12-13 in Upland rice 1978 atnual report Bouak Ivory Coast 27 p

97 International Institute for Tropical Agriculture 1983 Aphids itt rice roots I ITA Res Briefs 4(1-3)4-5

98 International Institite fi Tropical Agriculture 1983 Hopperburn in rice caused by brown plathopper IITA Res Briefs 4(l-3)6-7

99 International Rice Research Institute 1975 Annual report for 1974 PO Box 933 Manila Philippines p 216

100 International Rice Restarch Institute 1976 Annual repiort for 1975 PO Box 933 Manila Philippines p 228

101 International Rice Research Institute 1979 Cropping systems program Component technology development and evaluation Page 404 i Annual report for 1978 11O Box 933 Manila Philippines

102 International Rice Research Institute Department of Entomology 1981) Insecticid evaluation 1979 Tables 73 74 PO Box 933Manila Philipes

103 International Rice Research Institute Department of Entonmology 1981 Insecticide evalomtion 1980 Table 81 POBox 933 Manila Philippines

104 International Rice Research Institute 1981 Report of a meeting of Asian rice-based cropping systems entomologists Eleventh cropping systems working group meeting 18-22 May 1981 PO Box 933 Manila Philippines p 132 133 144

105 International Rice Research Institute 1984 Cropping systems program Pages 368-373 in Annual report for 1983 PO Box 933 Manila Philippines

106 International Rice Research Institutc 1984 Upland rice in Asia Pages 45-68 in An overview of upland rice research Proceedings of the 1982 Bouakamp Ivory Coast upland rice workiiop loternational Rice Research Institute 11O Box 933 Manila Philippines

107 Irshad M 1977 Grasshoppers in rice in Pakistan lit Rice Res Newsl 2(3)8-9

108 Ito R and TOno 1964 Upland lice its culture and varieties in Japan lIternational Rice Cormmission loth Meeting of the Working Party on Rice Production and Protection 3-10 March 1964 Manila Philippines Food and Agriculture Organization Rome Italy 12 p

109 Jeffrey M I and V A Dyck 1983 Effect of moonlight on lightshytrap catches of brown planthopper lit Rice Res Newsl 8(6) 13

110 Jennings P R and T Alicia Pineda 1983 Management ofthe rice plnthoppers Sogatodsorivi(coIl in Latin America Presented at the meeting on stable plant resistance FAO and lIndocsian I)inectorate of Food Crops Protection Bali lndonesia 29 Novcniber-2 l)ccemshyber 9 p

IllJimenez Gomez J A 1979 Rice insect pests i lla ios ()riCntales lil Spanish] Pages 99-103 in Rice ciltivation Instituto (olotibiatto Agropecuario Compndio No 29

112 Jongeleen F JJ Soesanto and A Mockidjo 1979 Studies oti pathogens of two species of Scarabacidac froinIndonesia Z Angew Enlomnol 87412-417

113 Jouvenaz 1) PC S lofgren and W A Baiks 1981 Biological control of imported fire antsa review of cirrent knowledge Bull Entomol Soc Am 27203-208

114 Kainoh Y K Shiniiu S MarUani Y Taiaki 1980 lost finding behavior of tie rice bu lplotori+JchinvittjI)allas (liciniptera Corcidac) with special reference to diet pattern ol aggregation and feeding on the rice plant Appl Entomol Zool 15225-233

115 Kakkar K 0 P Sharma and P I Sharma 1977 Note on Ihteronyrchus liodresRedt (Coleopicra I)ynastidae) as asporadic pest of paddy and maie inHimachal Pradesh Corr Agric 156-57

116 Kalode M131 79 Major insect pest problems and their control in upland rice (Unpublishcd mineogriph CWI oJNaaiat lni Chandaratat (198 1)original not seen)

117 Kalshoven L i 98 I The pests of crops in I ldoniesia (translated by P A vatn der Laan) 1 T Ichtiar B3aru-vati Iloeve lakarta Indonesia 701 p

118 Kamal N Q 1981 Suppression of whitebacked planto ner Sogatella fircaifra (Horvath) aLId rice lealfolder (nhal -cis licdb lis (Guenee) populations by natural cnclmie hlI) tiesis Grcgorio Araneta University Foundation Victoncta Park Metto Manila Philippines 179 Ip

119 Karim A N M Reaul 1975 A species ofliealy bug Trioni i70s ceres new oii rice in the Philippines Rice Entoniol Newsl 247

120 Katanyukul Wand C Chandaratat 1981 Insect pestsof uplaid rice and their control Report firstFAOUNlFi Tliailand training course on improved cultural practices for tplad rice Chiang Mai Thailand 5-23 Oct 1981 Food and Agriculture Organiation Rome Italy 22 p

121 Kern F 1955 lhletwioa(I)uninemts) bhhit Biturm(Coleoptera) a new pest of sugarcane inVenezuela [inSpanish with Etnglish sunmary] Agron Trop (Maracay) 527-29 (Rev Appl Eitoniol A4572)

122 Khan K M and S Ghai 1974 White grubs ald their control in India Pesticides (1Bombay) 819-25

123 Kiritani K 1975 Integrated pest management of rice pests Pages 66-70 itt Proceedings platning workshop ott cooperative hIeld research in pest managementP SMotooka ed East-West Center Honololu Hawaii USA

124 Kisimoto R 1967 Ecology of insect vectors forecasting and chemical control Pages 243-255 in The virus diseases of tle rice plant Proceedings of a symposium atthe International Rice Research Institute April 1967 The Johns Hopkins Press altimore Maryshyland USA

125 Kisimoto R and V A Dyck 1976 Climate and rice insects Pages 367-391 in Climate and rice International Rice Research Institute PO Box 933 Manila Philippines

126 Kureha S S Kobayashi SAsari and T Ishihara 1977 Life history and control of the mole cricket Grrllotalpa africana Palisot de Beauvois [in Japanese English translation 7520 1RRI library] Proc Kanto-Tosan Plant Prot Soc 21192-197

127 Kushwaha K S 1981 The black beetle as a pest of rice in Haryana India Int Rice Res Newsl 6(4)14

128 Lawani S M 1982 A review of tile effects of various agronomic practices on cereal stem borers Trop Pest Manage 28266-276

129 Leuck D B 1966 Biology of the lesser cornstalk borer in South Georgia J Econ Entomol 59 797-801

130 lever R J A W 1955 Insect pests of the rice plant in Malaya Int Rice Comm Newsl 1612-21

131 Li Ching-Sing and Shin-Foon Chiu 1951 The study ofthe rice gall midge Iachydiplosis orvza Wood Mason [in Chinese with English summary] 1Taiwan Agric Res 21-13

132 Iitsinger J A 1984 Assesament of the need-based application for rice Proceedings TechnologyTransfer Workshop 15-16 March 1984 International Rice Research Institute 1PO Box 933 Manila Philippines 6 p

133 Litsinger J A 1984 Environmental diversity and insect pest abundance with reference to the Pacific Pages 285-307 in Proceedings sub-regional training course on methods of controlling diseases insects and other pests of plants in the South Pacific 4-20 Oct 1982 Vaini Kingdom of Tonga Nukualofa

134 Litsinger J A RF Apostol and M BObusan 1983 White grub Leucopholis irrorata (Coleoptera Scarabaeidae) pest status popula-tion dynamics and chemical conllol in arice-maize cropping pattern in the Philippines J Econ Entoinol 761133-1138

135 Litsinger J A C de la Cruz F Raymundo A T Barrion M D Lumaban J P Bandong M S Venugopal F Paragna and E Balete 1982 Insect pests and insecticide reponse of dry-seeded bunded rice Pages 179-193 in Report of workshop on cropping systems research in Asia International Rice Research Institute Los Bafos Philippine

136 Litsinger J A M 1) Luinaban J P Bandong A T Barrion W Katanyukul and N Panda 1978 Insect pest management in rainfcd lowland rice agroecosystms of tropical Asia International Rice Research Conference 17-21 April 1978 International Rice Research Institute Los Bafios Philippines

137 Litsinger J A and K Moody 1976 Integrated pest management in multiple cropping systems Pages 293-317 in R I Papendick P A Sanchez and G B Triplett eds Multiple cropping American Society of Agronomy Madison Wisconsin USA

138 Loevinsohn M E 1984 The ecology and control of rice pests in relation to the intensity and synchrony of cultivation Ph D thesis University of London 36) p

139 Loevinsohn M E J A Litsinger N Panda M A Austria R F Apostol C Andrion and J Bandong 1982 A basis for predicting rice insect pest abundance in cropping systems Paper presented at the 1982 International rice research conference on rice and cropping systems 19-23 April 1982 International Rice Research Institute Los Bafios Iaguna Philippines 25 p

140 Lopez A W 1930 Repot of the Entomology Department Pages 145-172 in Annual report of the Philippine Sugar Association 1929-30 Manila

141 Lopez AW 1931 Entomology Department Pages 227-266 in Annual report of the director or research 1930-31 Philippine Sugar Association Manila

142 Lopez A W 1932 Entomology Deparment Pages 252-279 in Annual report of the Philippine Sugar Association 1931-32 Manila

143 Lungibill P and G G Ainslee 1917 The lessercornstalk borer US Dep Agric Agric Res Serv Tech Bull 539 27 p

IRPS No 123 January 1987 39

144 McGuire J U Jr and BS Crandall 1967 Survey of insect pests and plant diseases ofselected food crops of Mexico Central America and Panama U SDepartment of Agriculture International Agriculshytural Development Service Agricultuial Researrh Service October 1967 11 p

145 McNeill Sand T R ESouthwood 1978 The role of nitrogen in the development of insectplant relations Pages 77-9 in lliochemical aspects of plant and animal coevolution J B Ilarbornu ed Academic Press London UK

146 Maebara H 1955 Spatial distribution of lastinstiarlanaeof purplish stem borer (Sesamia iqferens Walker) hibernating in an upland rice field Bull Kagoshima Univ Fac Agric 438-41

147 Mallick E H and N G Hajra 1977 Response of rice plant to defoliation treatments at different developmental stages Indian Agric 2151-56

148 Mansingh A 1971 Physiological classification of dormancies in insects Canl Entomol 103983-10)9

149 Martell C G 1974 Catalogue of plant feeding insects of Mexico [in Spanish] Fitofilo 27(69)1-176

150 Martinez 0 J 1982 Principal rice pests in Ilanos orientalis [in Spanish] P~ages 80-94 in Curso deact nalization de conocimientos en arroz Villaniencis Instituto (olonmbiano Agropecuario

151 Martins J F da Silva F Ferrcira A S Prabhu F 1 P Zimmermann 1980 Use of pesticides for the control of the main subterranean upland rice pests esqui Agropecu Bras 1553-u

152 Martins I F da Silva F Fe eira rod BdamSilveira Pinheiro I Q2 Damage simulation caused by defoliators of upland rice [in Portuguese with English summary] Pesqui Agropecu Bras 171113-1119

153 Martins 1F da Silva N(iyuen van Tan and 1Ida Silveira Pinheiro 1981 Stem borer resistance in upland rice associated with plant morphological characters [in Portuuce Iesqui Agropecu Bras 16187-192

154 Masaki S 1980 Sumnmer diapause Ann Rev Entomiol 251-25 155 Masuzawa T 11 Suwa and F Nakasoji 1983 l)ifkleences of

oviposition preferencc and survival rate of two skipper butterflies Parnara guttata and PIchlpidai iathiais (Iepidoptera HIesperiidac) on rice plant and cogon grass New Entomol 3247-56

156 Maurya D M and C P Vaish 1984 Upland rice in India Pages 69-92 in An overview oftpland rice research Proceedings ofthe 1982 Bouakamp Ivory Coast upland rice workshop International Rice Research Institute IO Box 933 Mnila Philippines

157 Menon M G R R N Katiyai S Kuar and S S Misra 1970 Termites attacking the paddy crop Sci Cult 36294-295

158 Miyahara Y and 1 Nishikubo 1967 Behavior of young nymphs of the red rice root aphid Iholahsipiun ri liabhdontinalisSasaki in relation to movement from the leaf to the roots of upland rice [in Japanese] Pioc Assoc Plant Prot Kyushu 1363-66

159 Mo ljoa Tjien 1952 White grub of upland rice (Holotrichiahelleri Brsk) [in Indonesian with English summary] Contrib Agric Res Stn Bogor No 13134 p

160 Mochida 0 and T Suryana 1979 The occurrence of the brown planthopper and ragged stunt on lowland and upland rice ill Central Java and logyakarta in tile late of January 1979 Entomnolgy Survey Report No 15 Central Research Institute of Agriculture (CRIA) Sukamandi Indonesia I February 1979 9 p

161Monet C 1979 Varietal resistance to upland rice pests [inI French] Pages 35-39 in Annual report 1977 Institut des Savanes Department des Cultures Vivrieres Bouak6 Ivory Coast

162 Monte 0 1942 A caterpillar ofrice-fields[in Portuguese] Biol6gico (Sao Paulo) 8161-163 (Rev Appl Entomol A31414)

163 Munroe I) D 1979 Pest management strategy for hill rice in four resettlement areas in Sarawak Malaysia Pages 89-102 i1 Guidelines for integrated control of rice insect pests Food and Agriculture Organization Plant Production and Protection Paper 14 Rome Italy 115 p

40 IRPS No 123 January 1987

164 Nakasuji F M Ishii and Jen-Zon Ito 1984 Rice skippers in Taiwan and their life histories ZZrC (Taipei) Tech Bull 82 10 p

165 Navas I) 1976 Fall armyworum in rice Pages 99-106 il Ecological animal control by habitat management Proc Tall Timbers Con-ference 28 Feb-I Mar 1974 Fall Timbers Res Stn Misc Publ 6 Tallahassee Horida USA

166 Nickel 1 1 1961 Biological control of rice stem borer A feasibility study Tcch Bull 2 International Rice Rcarch Institute PO Box 933 Manila Philippines IIl p

167 Nigan P M and R A Vernia 1985 Insect pests of uplaild rice in Uttar Pradesh Int Rice Res Newsl 10(6)22

168 Nilakhe S S 1985 -cological observations on spittle bugs with emphasis on their occirence in rice Pcsqtui Agropecu Bras 20407-414

169 Nixon G EJ 195 1 lie association of ats with aphids and coccids Commonwealth Iusti lcte of Entonology Iondon UK 36 p

170 Nurdin F an Ishak Manti 1985 Study oit seedling fly of upland rice Interlational Upland Rice Moitoring Tour Report 26 lebruary-2 MarLI Sukaranti West Sumatra

171 Oka 1 N 1979 Ciltural conttol of the brown planthopper Pages 357-369 in Brown planthopper threat to rice production in Asia International Rice Research Institutc 1p0 Box 933 Manila Philippines

172 Ohni M and A Vilnaii 1975 Rice insect pests of Italy biology of Sipha grcria (Kaltenbach) and its control [in Italian] Riso (Milan) 2459-71

173 Otanes F Q atnd 1 I Sison 194 1 Pests of rice Philipp J Agric 12211-259

174 Painter R I 1951 Insect icsistance in crop plants University Press of Kansits Lawrence Kansas USA 520 p

175 Pantua P C and 1A litsinger 1981 Arthropod abundance in continuous vs biannual rice cropping systeis Int Rice Res Ncwsl 6(2)20-21

176 Pathak M ) and V A l)yck 1975 Sttudics ott insect pests of upland rice Pages 186-191 in Major research in upland rice International Rice Research Institute PO Box 933 Ilanila Philippines

177 Patil B R B M IHasabe and 1) S Ajri 198 I Chemical control of white gritb lIditriihinhorraia Fabr intfesting groundntut paddy atnd sorghum Indian 1Entomol 43232-236

178 Pawar A 1) 1974 (ollection and identification of insect pests of rice rlerminal repirt lInternational Rice Research Institute PO Box 933 Manila Philippines 72 p (tnimco)

179 IerrajU A aid A S Reedy 1965 Observations ott diapausc itt tie steti borer ohrice Rice News Teller (lai-Mar)9-13

180 Pont A C 1972 A review of the Oriental species of theri ona Rondani (I)iptera Muscidac) of econtomic importance Pages 27-102 in Control of sorglium shoot fly M G lomtani and W R Young eds Oxford atd IH1 Ptthl Co New I)clhi India 324 p

181 Prasad S C 1PChand aid M 1PSingh 1983 Incidence of rice gall midge in the upland plateau region of Hihar Oryza 20235-236

182 Rai P S 198 0 Nisaga simpex damage to rice in the hill tracts of South India Int Rice kles News[ 5(I)12

183 Rathorc Y S 1969 Gonocplhahli dcprssunt Fabr as itpredator of (tilo zonwIlhis (Swinhoc) and other moths in field cage at Patnagar Uttar Pradesh Indian 1Entotnol 3127

184 Reed W 1983 Crop losses causcd by insect pists in the developing world Proceedings 10th Internatial Congress of Plant Protection lrighton UK 174-80

185 Reedy M V 1981 Ateactogasterinhens(Walker)a new root-grub pest of tipland paddy and inaize Curr Sci 501076

186 Reveno M A 1968 Catalogue of the insect pests of economic crops in Colombia [in Spaiish] Asociacion Latino Americana de Entomo-logia (AIAE) 129-33

187 Reyes G M 1957 Rice dwarf disease in the Philippines Food and Agriculture Organiiation Plant Prot Bull 617-19

188 Rice S E A A Grigarick and M 0 Way 1982 Effect of leaf and panicle feeding byarmyworm (Lepidoptera Noctuidae) larvae on rice grain yield Environ Entomol 7593-595

189 Risch S J 1) Andow and M A Altieri 1983 Agioccosystem diversity arnd pest control data tentative conclusions and nizw research dirctions Environ Entomol 12625-629

190 Rivera C T 1956 Three organic insecticides in thecontrol of insects affecting upland rice Philipp Agric 39465-472

191 Roffrey 1 1972 Locusta outbreaks in the Philippines Acrida 1177-188

192 Rojas R 1) 1941 The effect of pruning at varying ages upon the yield of Ramairio Phipp Agric 29851-860

193 Roonwal M i 1979 Ternite life and termite control in tropical South Asia Scientilic Pnbl odhpr India 139 p

194 Rosetto C1 F da Silva Martins N C Schmidt and L EAzzini 1978 Damage caused by pasturespittle bugs 4Dvoi)lavopictaandDJ schach) on rice lBragantia Nota 6 (Matcli)25-27

195 Rosztto C1- S S Neto I- Amantc I) Iink 1Garcia-Vieira D M de SoLza N V Bansa t and A M de Olivcira 1971 Rice pests of Brazil [in Portuguese] Pages 149-238 inrFA()-International Rice Commission 2nd meeting of the Rice Committee for the Americas 6-11 l)ecember 1971 Pelotas Rio Grande do Sul Brazil

196 Rothschild G H L 1969 Obseratiois on the armyworm Spodoptera matritia acronrctoiiles Gn (Lep Noctuidae) Bull Entomol Res 59143-160

197 Rothschild G 1- L 1974 Parasites of the armworm Spolopiera mauritiaacroncitoihes in Malaysia Fntornopliaga 19293-299

198 Sachan i N S K Gangwar and 1N Katujar 1980 Field crickets damaging cages in North Eastern IHill Region Indian J Entomol 42526-527

199 Sands 1) P A 1977 The biology and ecology of Lepocorisa (Hemiptera Alydidac) in Papua New Guinea Res Bull 18 Departshyment of Primary Industry Port Moresby Papua New Guinea 104 p

200 Sarawak 1968 titomology division Pages 97-106 in Report of the research branch of the Department of Altriculture 1966 Kuching (Rev AIppl Eintomol A56603)

201 Satpathy JM M S Das and K Naik 1977 Effect of multiple and mixed cropping on the incidence of scine important pestsI Entomol Res 178-85

202 Sauer IHF G 1939 Notes Ott Elasmpnlpualtsligniosellths Zeller (Lep Pyr) a serious pest ofcereals in Sao Paolo [in Portuguese] Arq Inst Biol 10199-206 (Rev Appl Entotnol A28378)

203 Satinders 1L A 1 SKing and C L Vargas S 1983 Crop pests of Central Anerica with t bibliography [in Spanish] Centro Agroshynotnico Tropical de Investigacion y Ensenanza Turrialba Costa Rica (CATIE) Tec Bol 9 90 p

204 Sauphanor B 1985 Some factors of upland rice tolerttce to stem borers in West Africa Inseu Sci Appl 6429434

205 Sen P atnd S Chakravorty 1976 Stentflics in Haritighata Nadia West Bengal India [it Rice Res Newsl 1(12) 15

206 Senapai B and 1M Satpathy 1983 The seedling fly Atherigona orrzae Malloch is a pest of rice in Orissa (India) J Entomol Res 777-79

207 Shhjahan M S Alatn M Ah nad and BN Islam 1983 A new scarabaeid beetle pest of dryland rice in Bangladesh Int Rice Res Ncwsl 8(I)16-17

208 Sharma G L S C Modgol and R C Gautam 1979 Effect of depth of submergence ott incidence of bacterial blight and white grub infestation in transplanted rice Int Rice Res Newsl 4(6)12-13

209 Sharma S 1985 Scarabaeid beetle outbreak itt upland rice Int Rice Res Newsl 1(1(l)24

210 Shiraki T 1917 The paddy borer Schoenobhits incertulas Walker [achaku Agric Exp Stn Formosa 256 p

211 Silva A de Brito and 11P Magalhaes 1981 Insect pests of rice [in Portuguese] Circular Tecnica No 22 EMBRAPA Centro de Pesquisa t ecuaria de Tropico Unido (CPAJU) Belem Brazil 14 p

212 Singh S A 1979 Emergence of rice leaf butterfly as a rice pest in Manipur India int Rice Res Newsl 4(4)16-17

213 SinghS A 1983 Changing trends of insect pestsof rice in Manipur Int Rice Res Newsl 8(f)12

214 Siwi S S and P H van Doesburg 1984 Lenovorisa Latreille in Indonesia (Heteroptera Coreidae Alydinae) Zool Meded (Leiden) 58117-129

215 Soejitno J 1977 Reaction of some upland rice varieties toseedling fly eltherikonaexigua Stein [in Indonesian] Siri Hana I Penyakit 630-35 Lembga I osat Fenclilian Pertanian Bogor Indonesia

216 Socnardi 1967 Insect pests of rice in Indonesia Pages675-683 in The major inse pests of thu rice plant Proceedings of a symposium at the Internatital Rice Rec Wi itt S pternber 1964 The Johns Hopkins Press Baltimore Maryland USA

217 Soto P E A T Petcz and I W Buddenhagen 1976 Survey of insect pests and diseases of rice in different ecological zones in Nigeria Rice Entomol Ncwsl 435-46

218 Soto 11E and Z Ziddiqi 1976 Screening for resistance to African rice insect pests Varietal improsetnent sCmi n r Bouak Ivory Coast 13-17 Sept 1976 West Africa Rice l)evelopment Association Monrovia Iiberia

219 Squire F A 1972 Entomological problems in Bolivia PANS (PestArtic News Sumr) 18249-268

220 Stapely 1 II 1978 The rice leaf roller in the Solomon Islands Int Rice Res Newsl 3(l)13

221 Suenaga H 1937 On aphids attacking the roots of rice Oyo-I)oubts Zasshi 9150-152 [in Japanese] (Rev Appl Entomol A25745)

222 Suenaga H 1952 On the seasonal abundance of three subterranean aphids on upland rice Bull Kyushu Agric Exp Sti 1249-262

223 Sutynso Yas and Dandi Soekaria 1964 Beberapa hana yang ttenperang tana man padi tanah kering [in Indonesian] Rapat kerja Padi Tanah Kering 6 sd 8 Juily 1964 Kaliurang 9 p

224 Tallamy I) W and R F Denno 1983 Alternative life history patterns in risky environments in example from lace bugs Pages 129-147 in Insect life history patterns R F l)cnio and If Dingle eds Spring-Verlag 225 p

225 Tamura 1 1939 Observations on some insect pests of field crops [in Japanese] 1 Plant Prot 26414-420 (Rev Appl EntomolA27-556)

226 Tamura and Suzuki 1964 Influences of the early-transplanting practice in rice culture upon development of the rice stem borer Chilo siql)ressalisWalker Bull Ilokuriku Natil Agric Exp Stn 761-94

227 Tan J P 1924 The rice root aphid Drowhi ihirsoia Philipp Agric 13277-288

228 Tanaka 1 1957 Taxonomy and distribution of some subterranean aphids injurious to upland rice in Japan with description of a new species [in Japanese English summary] Botyu-Kagaku 22168-176

229 Tanaka T 1961 The rice root aphids their ecology and control Spec [full College of Agriculture Utsunomiya University Japan F3 p

230 Tauber M IC A Tauber R Nechols and 1Obrycki 1983 Seasonal activity of parasitoids control by external internal and genetic factors Pages 87-108 itt I)iapause and life cycle strategies in insects V K Brown and I Ilodek eds )r W Junk The Hague Netherlands

231 Taylor W F 1972 Effects of artificial defoliation (simulating pest damage) on varieties of upland rice Exp Agric 879-83

232 feotia T P S and S Natud 1966 Biotoomics of the rice skipper Parnaraniathiis Fabricius (Lepidoptera Hesperiidac) Indian J Entomol 28181-186

233 Teran F 0 1971 Rice pests of Santa Cruz [in Spanish Ministerio de Agricultura Sita Cruz Bolivia Bull 45 8 p

234 Than Itun 1976 Population dynamics of the yellow rice borer Tryporza itcertulas (Walker) and its damage to the rice plant MS thesis University of the Philippines at Ios Bafios Laguna Philippines 92 p

235 Uichanco L B 1911 Pests Pages 55-74 it Rice Philipp Agric 11-200

IRPS No 123 January 1987 41

236 van der Laan P A 195 1 Life history of the rice seedling fly Contrib Gen Agric Res Stn Boor Indonesia 118 15 p

237 van der Laan 11A 1959 Correlation between rainfall in the dry season and the occurrence of the white rice borer (Scirpolhaga innotala WIk) in lava Entomol Fxp Appl 212-20

238 Van Driesche R G 1983 Meaning of percent parasitismin studies of insect parasitism Fnviron Entomol 121 I1- 122

239 Venugopal M S and A litsingcr 1984 Fffect ofcarbofuran on rice growth Prot Ecol 7313-317

240 Vietnam Directorate of Rural Affairs 1962 Annual work progress report on crop improvement program of rice sugarcane vegetable and field crops (lor the period from July 1961 to June 1962) (11) 299 p (Rev Appl Entomol A51526-527)

241 Wan M T K 1972 Observations on rice leaf- and planthoppers in Sarawak (Malaysian Borneo) Malay Agric 148308-335

242 West Africa Rice I)esclopnient Association 1978 Annual research report Vol 2 Statistics Monrovia Liberia

243 West Africa Rice Development Association 1979 Annual research report p47 Monrovia Liberia

244 Williams ) F and CS lofgren 1981 Fli I illy l-468 anew bait toxicant for control of the red imported lire at Fla Entomol 64472477

245 Williams I) 1 1970 The mealybugs OFlomoptera Coccoidea Iscudococcidac) of sugar-cane rice and sorghum Bull Entomol Res 0109-188

246 Williams D) 1 1982 Pulinaria icervi (Signoret) and its allies ott sugar cane and other grasses Btll Entoinol Res 72111-117

247 Wirjosuiardjo S 1976 Study ofchenmical cont rot of insect pests ott upland rice Imiu Pertanian (Jogjakarta) 1323-329

248 Wirjosuhardjo S and N Wirarta 1976 Chemical control of the white grub tIolorithiatlltriin upland rice in M uneng Irobolinggo Indonesia) [in Indonesian] Imut Pertanian (Jogjakarta) 1330-332

249 Yano K T Miyake and V F Eastop 1983 The biology and economic importance of rice aphids (Henuiptera Aphid idae) a review Bull Entomol Res 73539-566

250 Yelp D and P Rhodes 19(4 lhe success of drilled dryland rice iii the Fiji Islands International Rice Commission Working Party on Rice Production and Protection Manila Philippines 3-10 March1964 Food and Agriculture Organization Rome Italy 13 p

251 Young W R 1981 Fifty-five years of research on the sorghum shootfly Insect Sci Appi 23-9

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No 104 Faridpur acomputer-assisted instruction nmodel forrainfed lowland random coefficient model No 116 Consumer demand for rice grain quality in Thailand Indonesia andrice

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No 106 Rice grassy stunt virus 2 a new strain of rice grassy stunt in tire literature

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publishers No 120 Changes isinall-farm iLtireliing techii)hgy in Thailand and [he No 109 Classification of Philippine rainfall patterns Philippines No 110 Contributions of modern rice varieties to rutrition in Asia No 121 Landforms and modern rice varieties No Ill Changes in rice breeding in 10 Asian countries 1965-84 No 122 Yield stability and modern rice technology No 112 Design parameters affecting the performance of the IRRI-designed

axial-flow pump

Page 39: UPLANDRICE. INSECTl--1i M T -E!R.EOLOU t RTANCE ,AND …delphacid.s3.amazonaws.com › 6958.pdf · The significance of abiotic factors in upland rice insect ecology will be apparent

125 Kisimoto R and V A Dyck 1976 Climate and rice insects Pages 367-391 in Climate and rice International Rice Research Institute PO Box 933 Manila Philippines

126 Kureha S S Kobayashi SAsari and T Ishihara 1977 Life history and control of the mole cricket Grrllotalpa africana Palisot de Beauvois [in Japanese English translation 7520 1RRI library] Proc Kanto-Tosan Plant Prot Soc 21192-197

127 Kushwaha K S 1981 The black beetle as a pest of rice in Haryana India Int Rice Res Newsl 6(4)14

128 Lawani S M 1982 A review of tile effects of various agronomic practices on cereal stem borers Trop Pest Manage 28266-276

129 Leuck D B 1966 Biology of the lesser cornstalk borer in South Georgia J Econ Entomol 59 797-801

130 lever R J A W 1955 Insect pests of the rice plant in Malaya Int Rice Comm Newsl 1612-21

131 Li Ching-Sing and Shin-Foon Chiu 1951 The study ofthe rice gall midge Iachydiplosis orvza Wood Mason [in Chinese with English summary] 1Taiwan Agric Res 21-13

132 Iitsinger J A 1984 Assesament of the need-based application for rice Proceedings TechnologyTransfer Workshop 15-16 March 1984 International Rice Research Institute 1PO Box 933 Manila Philippines 6 p

133 Litsinger J A 1984 Environmental diversity and insect pest abundance with reference to the Pacific Pages 285-307 in Proceedings sub-regional training course on methods of controlling diseases insects and other pests of plants in the South Pacific 4-20 Oct 1982 Vaini Kingdom of Tonga Nukualofa

134 Litsinger J A RF Apostol and M BObusan 1983 White grub Leucopholis irrorata (Coleoptera Scarabaeidae) pest status popula-tion dynamics and chemical conllol in arice-maize cropping pattern in the Philippines J Econ Entoinol 761133-1138

135 Litsinger J A C de la Cruz F Raymundo A T Barrion M D Lumaban J P Bandong M S Venugopal F Paragna and E Balete 1982 Insect pests and insecticide reponse of dry-seeded bunded rice Pages 179-193 in Report of workshop on cropping systems research in Asia International Rice Research Institute Los Bafos Philippine

136 Litsinger J A M 1) Luinaban J P Bandong A T Barrion W Katanyukul and N Panda 1978 Insect pest management in rainfcd lowland rice agroecosystms of tropical Asia International Rice Research Conference 17-21 April 1978 International Rice Research Institute Los Bafios Philippines

137 Litsinger J A and K Moody 1976 Integrated pest management in multiple cropping systems Pages 293-317 in R I Papendick P A Sanchez and G B Triplett eds Multiple cropping American Society of Agronomy Madison Wisconsin USA

138 Loevinsohn M E 1984 The ecology and control of rice pests in relation to the intensity and synchrony of cultivation Ph D thesis University of London 36) p

139 Loevinsohn M E J A Litsinger N Panda M A Austria R F Apostol C Andrion and J Bandong 1982 A basis for predicting rice insect pest abundance in cropping systems Paper presented at the 1982 International rice research conference on rice and cropping systems 19-23 April 1982 International Rice Research Institute Los Bafios Iaguna Philippines 25 p

140 Lopez A W 1930 Repot of the Entomology Department Pages 145-172 in Annual report of the Philippine Sugar Association 1929-30 Manila

141 Lopez AW 1931 Entomology Department Pages 227-266 in Annual report of the director or research 1930-31 Philippine Sugar Association Manila

142 Lopez A W 1932 Entomology Deparment Pages 252-279 in Annual report of the Philippine Sugar Association 1931-32 Manila

143 Lungibill P and G G Ainslee 1917 The lessercornstalk borer US Dep Agric Agric Res Serv Tech Bull 539 27 p

IRPS No 123 January 1987 39

144 McGuire J U Jr and BS Crandall 1967 Survey of insect pests and plant diseases ofselected food crops of Mexico Central America and Panama U SDepartment of Agriculture International Agriculshytural Development Service Agricultuial Researrh Service October 1967 11 p

145 McNeill Sand T R ESouthwood 1978 The role of nitrogen in the development of insectplant relations Pages 77-9 in lliochemical aspects of plant and animal coevolution J B Ilarbornu ed Academic Press London UK

146 Maebara H 1955 Spatial distribution of lastinstiarlanaeof purplish stem borer (Sesamia iqferens Walker) hibernating in an upland rice field Bull Kagoshima Univ Fac Agric 438-41

147 Mallick E H and N G Hajra 1977 Response of rice plant to defoliation treatments at different developmental stages Indian Agric 2151-56

148 Mansingh A 1971 Physiological classification of dormancies in insects Canl Entomol 103983-10)9

149 Martell C G 1974 Catalogue of plant feeding insects of Mexico [in Spanish] Fitofilo 27(69)1-176

150 Martinez 0 J 1982 Principal rice pests in Ilanos orientalis [in Spanish] P~ages 80-94 in Curso deact nalization de conocimientos en arroz Villaniencis Instituto (olonmbiano Agropecuario

151 Martins J F da Silva F Ferrcira A S Prabhu F 1 P Zimmermann 1980 Use of pesticides for the control of the main subterranean upland rice pests esqui Agropecu Bras 1553-u

152 Martins I F da Silva F Fe eira rod BdamSilveira Pinheiro I Q2 Damage simulation caused by defoliators of upland rice [in Portuguese with English summary] Pesqui Agropecu Bras 171113-1119

153 Martins 1F da Silva N(iyuen van Tan and 1Ida Silveira Pinheiro 1981 Stem borer resistance in upland rice associated with plant morphological characters [in Portuuce Iesqui Agropecu Bras 16187-192

154 Masaki S 1980 Sumnmer diapause Ann Rev Entomiol 251-25 155 Masuzawa T 11 Suwa and F Nakasoji 1983 l)ifkleences of

oviposition preferencc and survival rate of two skipper butterflies Parnara guttata and PIchlpidai iathiais (Iepidoptera HIesperiidac) on rice plant and cogon grass New Entomol 3247-56

156 Maurya D M and C P Vaish 1984 Upland rice in India Pages 69-92 in An overview oftpland rice research Proceedings ofthe 1982 Bouakamp Ivory Coast upland rice workshop International Rice Research Institute IO Box 933 Mnila Philippines

157 Menon M G R R N Katiyai S Kuar and S S Misra 1970 Termites attacking the paddy crop Sci Cult 36294-295

158 Miyahara Y and 1 Nishikubo 1967 Behavior of young nymphs of the red rice root aphid Iholahsipiun ri liabhdontinalisSasaki in relation to movement from the leaf to the roots of upland rice [in Japanese] Pioc Assoc Plant Prot Kyushu 1363-66

159 Mo ljoa Tjien 1952 White grub of upland rice (Holotrichiahelleri Brsk) [in Indonesian with English summary] Contrib Agric Res Stn Bogor No 13134 p

160 Mochida 0 and T Suryana 1979 The occurrence of the brown planthopper and ragged stunt on lowland and upland rice ill Central Java and logyakarta in tile late of January 1979 Entomnolgy Survey Report No 15 Central Research Institute of Agriculture (CRIA) Sukamandi Indonesia I February 1979 9 p

161Monet C 1979 Varietal resistance to upland rice pests [inI French] Pages 35-39 in Annual report 1977 Institut des Savanes Department des Cultures Vivrieres Bouak6 Ivory Coast

162 Monte 0 1942 A caterpillar ofrice-fields[in Portuguese] Biol6gico (Sao Paulo) 8161-163 (Rev Appl Entomol A31414)

163 Munroe I) D 1979 Pest management strategy for hill rice in four resettlement areas in Sarawak Malaysia Pages 89-102 i1 Guidelines for integrated control of rice insect pests Food and Agriculture Organization Plant Production and Protection Paper 14 Rome Italy 115 p

40 IRPS No 123 January 1987

164 Nakasuji F M Ishii and Jen-Zon Ito 1984 Rice skippers in Taiwan and their life histories ZZrC (Taipei) Tech Bull 82 10 p

165 Navas I) 1976 Fall armyworum in rice Pages 99-106 il Ecological animal control by habitat management Proc Tall Timbers Con-ference 28 Feb-I Mar 1974 Fall Timbers Res Stn Misc Publ 6 Tallahassee Horida USA

166 Nickel 1 1 1961 Biological control of rice stem borer A feasibility study Tcch Bull 2 International Rice Rcarch Institute PO Box 933 Manila Philippines IIl p

167 Nigan P M and R A Vernia 1985 Insect pests of uplaild rice in Uttar Pradesh Int Rice Res Newsl 10(6)22

168 Nilakhe S S 1985 -cological observations on spittle bugs with emphasis on their occirence in rice Pcsqtui Agropecu Bras 20407-414

169 Nixon G EJ 195 1 lie association of ats with aphids and coccids Commonwealth Iusti lcte of Entonology Iondon UK 36 p

170 Nurdin F an Ishak Manti 1985 Study oit seedling fly of upland rice Interlational Upland Rice Moitoring Tour Report 26 lebruary-2 MarLI Sukaranti West Sumatra

171 Oka 1 N 1979 Ciltural conttol of the brown planthopper Pages 357-369 in Brown planthopper threat to rice production in Asia International Rice Research Institutc 1p0 Box 933 Manila Philippines

172 Ohni M and A Vilnaii 1975 Rice insect pests of Italy biology of Sipha grcria (Kaltenbach) and its control [in Italian] Riso (Milan) 2459-71

173 Otanes F Q atnd 1 I Sison 194 1 Pests of rice Philipp J Agric 12211-259

174 Painter R I 1951 Insect icsistance in crop plants University Press of Kansits Lawrence Kansas USA 520 p

175 Pantua P C and 1A litsinger 1981 Arthropod abundance in continuous vs biannual rice cropping systeis Int Rice Res Ncwsl 6(2)20-21

176 Pathak M ) and V A l)yck 1975 Sttudics ott insect pests of upland rice Pages 186-191 in Major research in upland rice International Rice Research Institute PO Box 933 Ilanila Philippines

177 Patil B R B M IHasabe and 1) S Ajri 198 I Chemical control of white gritb lIditriihinhorraia Fabr intfesting groundntut paddy atnd sorghum Indian 1Entomol 43232-236

178 Pawar A 1) 1974 (ollection and identification of insect pests of rice rlerminal repirt lInternational Rice Research Institute PO Box 933 Manila Philippines 72 p (tnimco)

179 IerrajU A aid A S Reedy 1965 Observations ott diapausc itt tie steti borer ohrice Rice News Teller (lai-Mar)9-13

180 Pont A C 1972 A review of the Oriental species of theri ona Rondani (I)iptera Muscidac) of econtomic importance Pages 27-102 in Control of sorglium shoot fly M G lomtani and W R Young eds Oxford atd IH1 Ptthl Co New I)clhi India 324 p

181 Prasad S C 1PChand aid M 1PSingh 1983 Incidence of rice gall midge in the upland plateau region of Hihar Oryza 20235-236

182 Rai P S 198 0 Nisaga simpex damage to rice in the hill tracts of South India Int Rice kles News[ 5(I)12

183 Rathorc Y S 1969 Gonocplhahli dcprssunt Fabr as itpredator of (tilo zonwIlhis (Swinhoc) and other moths in field cage at Patnagar Uttar Pradesh Indian 1Entotnol 3127

184 Reed W 1983 Crop losses causcd by insect pists in the developing world Proceedings 10th Internatial Congress of Plant Protection lrighton UK 174-80

185 Reedy M V 1981 Ateactogasterinhens(Walker)a new root-grub pest of tipland paddy and inaize Curr Sci 501076

186 Reveno M A 1968 Catalogue of the insect pests of economic crops in Colombia [in Spaiish] Asociacion Latino Americana de Entomo-logia (AIAE) 129-33

187 Reyes G M 1957 Rice dwarf disease in the Philippines Food and Agriculture Organiiation Plant Prot Bull 617-19

188 Rice S E A A Grigarick and M 0 Way 1982 Effect of leaf and panicle feeding byarmyworm (Lepidoptera Noctuidae) larvae on rice grain yield Environ Entomol 7593-595

189 Risch S J 1) Andow and M A Altieri 1983 Agioccosystem diversity arnd pest control data tentative conclusions and nizw research dirctions Environ Entomol 12625-629

190 Rivera C T 1956 Three organic insecticides in thecontrol of insects affecting upland rice Philipp Agric 39465-472

191 Roffrey 1 1972 Locusta outbreaks in the Philippines Acrida 1177-188

192 Rojas R 1) 1941 The effect of pruning at varying ages upon the yield of Ramairio Phipp Agric 29851-860

193 Roonwal M i 1979 Ternite life and termite control in tropical South Asia Scientilic Pnbl odhpr India 139 p

194 Rosetto C1 F da Silva Martins N C Schmidt and L EAzzini 1978 Damage caused by pasturespittle bugs 4Dvoi)lavopictaandDJ schach) on rice lBragantia Nota 6 (Matcli)25-27

195 Rosztto C1- S S Neto I- Amantc I) Iink 1Garcia-Vieira D M de SoLza N V Bansa t and A M de Olivcira 1971 Rice pests of Brazil [in Portuguese] Pages 149-238 inrFA()-International Rice Commission 2nd meeting of the Rice Committee for the Americas 6-11 l)ecember 1971 Pelotas Rio Grande do Sul Brazil

196 Rothschild G H L 1969 Obseratiois on the armyworm Spodoptera matritia acronrctoiiles Gn (Lep Noctuidae) Bull Entomol Res 59143-160

197 Rothschild G 1- L 1974 Parasites of the armworm Spolopiera mauritiaacroncitoihes in Malaysia Fntornopliaga 19293-299

198 Sachan i N S K Gangwar and 1N Katujar 1980 Field crickets damaging cages in North Eastern IHill Region Indian J Entomol 42526-527

199 Sands 1) P A 1977 The biology and ecology of Lepocorisa (Hemiptera Alydidac) in Papua New Guinea Res Bull 18 Departshyment of Primary Industry Port Moresby Papua New Guinea 104 p

200 Sarawak 1968 titomology division Pages 97-106 in Report of the research branch of the Department of Altriculture 1966 Kuching (Rev AIppl Eintomol A56603)

201 Satpathy JM M S Das and K Naik 1977 Effect of multiple and mixed cropping on the incidence of scine important pestsI Entomol Res 178-85

202 Sauer IHF G 1939 Notes Ott Elasmpnlpualtsligniosellths Zeller (Lep Pyr) a serious pest ofcereals in Sao Paolo [in Portuguese] Arq Inst Biol 10199-206 (Rev Appl Entotnol A28378)

203 Satinders 1L A 1 SKing and C L Vargas S 1983 Crop pests of Central Anerica with t bibliography [in Spanish] Centro Agroshynotnico Tropical de Investigacion y Ensenanza Turrialba Costa Rica (CATIE) Tec Bol 9 90 p

204 Sauphanor B 1985 Some factors of upland rice tolerttce to stem borers in West Africa Inseu Sci Appl 6429434

205 Sen P atnd S Chakravorty 1976 Stentflics in Haritighata Nadia West Bengal India [it Rice Res Newsl 1(12) 15

206 Senapai B and 1M Satpathy 1983 The seedling fly Atherigona orrzae Malloch is a pest of rice in Orissa (India) J Entomol Res 777-79

207 Shhjahan M S Alatn M Ah nad and BN Islam 1983 A new scarabaeid beetle pest of dryland rice in Bangladesh Int Rice Res Ncwsl 8(I)16-17

208 Sharma G L S C Modgol and R C Gautam 1979 Effect of depth of submergence ott incidence of bacterial blight and white grub infestation in transplanted rice Int Rice Res Newsl 4(6)12-13

209 Sharma S 1985 Scarabaeid beetle outbreak itt upland rice Int Rice Res Newsl 1(1(l)24

210 Shiraki T 1917 The paddy borer Schoenobhits incertulas Walker [achaku Agric Exp Stn Formosa 256 p

211 Silva A de Brito and 11P Magalhaes 1981 Insect pests of rice [in Portuguese] Circular Tecnica No 22 EMBRAPA Centro de Pesquisa t ecuaria de Tropico Unido (CPAJU) Belem Brazil 14 p

212 Singh S A 1979 Emergence of rice leaf butterfly as a rice pest in Manipur India int Rice Res Newsl 4(4)16-17

213 SinghS A 1983 Changing trends of insect pestsof rice in Manipur Int Rice Res Newsl 8(f)12

214 Siwi S S and P H van Doesburg 1984 Lenovorisa Latreille in Indonesia (Heteroptera Coreidae Alydinae) Zool Meded (Leiden) 58117-129

215 Soejitno J 1977 Reaction of some upland rice varieties toseedling fly eltherikonaexigua Stein [in Indonesian] Siri Hana I Penyakit 630-35 Lembga I osat Fenclilian Pertanian Bogor Indonesia

216 Socnardi 1967 Insect pests of rice in Indonesia Pages675-683 in The major inse pests of thu rice plant Proceedings of a symposium at the Internatital Rice Rec Wi itt S pternber 1964 The Johns Hopkins Press Baltimore Maryland USA

217 Soto P E A T Petcz and I W Buddenhagen 1976 Survey of insect pests and diseases of rice in different ecological zones in Nigeria Rice Entomol Ncwsl 435-46

218 Soto 11E and Z Ziddiqi 1976 Screening for resistance to African rice insect pests Varietal improsetnent sCmi n r Bouak Ivory Coast 13-17 Sept 1976 West Africa Rice l)evelopment Association Monrovia Iiberia

219 Squire F A 1972 Entomological problems in Bolivia PANS (PestArtic News Sumr) 18249-268

220 Stapely 1 II 1978 The rice leaf roller in the Solomon Islands Int Rice Res Newsl 3(l)13

221 Suenaga H 1937 On aphids attacking the roots of rice Oyo-I)oubts Zasshi 9150-152 [in Japanese] (Rev Appl Entomol A25745)

222 Suenaga H 1952 On the seasonal abundance of three subterranean aphids on upland rice Bull Kyushu Agric Exp Sti 1249-262

223 Sutynso Yas and Dandi Soekaria 1964 Beberapa hana yang ttenperang tana man padi tanah kering [in Indonesian] Rapat kerja Padi Tanah Kering 6 sd 8 Juily 1964 Kaliurang 9 p

224 Tallamy I) W and R F Denno 1983 Alternative life history patterns in risky environments in example from lace bugs Pages 129-147 in Insect life history patterns R F l)cnio and If Dingle eds Spring-Verlag 225 p

225 Tamura 1 1939 Observations on some insect pests of field crops [in Japanese] 1 Plant Prot 26414-420 (Rev Appl EntomolA27-556)

226 Tamura and Suzuki 1964 Influences of the early-transplanting practice in rice culture upon development of the rice stem borer Chilo siql)ressalisWalker Bull Ilokuriku Natil Agric Exp Stn 761-94

227 Tan J P 1924 The rice root aphid Drowhi ihirsoia Philipp Agric 13277-288

228 Tanaka 1 1957 Taxonomy and distribution of some subterranean aphids injurious to upland rice in Japan with description of a new species [in Japanese English summary] Botyu-Kagaku 22168-176

229 Tanaka T 1961 The rice root aphids their ecology and control Spec [full College of Agriculture Utsunomiya University Japan F3 p

230 Tauber M IC A Tauber R Nechols and 1Obrycki 1983 Seasonal activity of parasitoids control by external internal and genetic factors Pages 87-108 itt I)iapause and life cycle strategies in insects V K Brown and I Ilodek eds )r W Junk The Hague Netherlands

231 Taylor W F 1972 Effects of artificial defoliation (simulating pest damage) on varieties of upland rice Exp Agric 879-83

232 feotia T P S and S Natud 1966 Biotoomics of the rice skipper Parnaraniathiis Fabricius (Lepidoptera Hesperiidac) Indian J Entomol 28181-186

233 Teran F 0 1971 Rice pests of Santa Cruz [in Spanish Ministerio de Agricultura Sita Cruz Bolivia Bull 45 8 p

234 Than Itun 1976 Population dynamics of the yellow rice borer Tryporza itcertulas (Walker) and its damage to the rice plant MS thesis University of the Philippines at Ios Bafios Laguna Philippines 92 p

235 Uichanco L B 1911 Pests Pages 55-74 it Rice Philipp Agric 11-200

IRPS No 123 January 1987 41

236 van der Laan P A 195 1 Life history of the rice seedling fly Contrib Gen Agric Res Stn Boor Indonesia 118 15 p

237 van der Laan 11A 1959 Correlation between rainfall in the dry season and the occurrence of the white rice borer (Scirpolhaga innotala WIk) in lava Entomol Fxp Appl 212-20

238 Van Driesche R G 1983 Meaning of percent parasitismin studies of insect parasitism Fnviron Entomol 121 I1- 122

239 Venugopal M S and A litsingcr 1984 Fffect ofcarbofuran on rice growth Prot Ecol 7313-317

240 Vietnam Directorate of Rural Affairs 1962 Annual work progress report on crop improvement program of rice sugarcane vegetable and field crops (lor the period from July 1961 to June 1962) (11) 299 p (Rev Appl Entomol A51526-527)

241 Wan M T K 1972 Observations on rice leaf- and planthoppers in Sarawak (Malaysian Borneo) Malay Agric 148308-335

242 West Africa Rice I)esclopnient Association 1978 Annual research report Vol 2 Statistics Monrovia Liberia

243 West Africa Rice Development Association 1979 Annual research report p47 Monrovia Liberia

244 Williams ) F and CS lofgren 1981 Fli I illy l-468 anew bait toxicant for control of the red imported lire at Fla Entomol 64472477

245 Williams I) 1 1970 The mealybugs OFlomoptera Coccoidea Iscudococcidac) of sugar-cane rice and sorghum Bull Entomol Res 0109-188

246 Williams D) 1 1982 Pulinaria icervi (Signoret) and its allies ott sugar cane and other grasses Btll Entoinol Res 72111-117

247 Wirjosuiardjo S 1976 Study ofchenmical cont rot of insect pests ott upland rice Imiu Pertanian (Jogjakarta) 1323-329

248 Wirjosuhardjo S and N Wirarta 1976 Chemical control of the white grub tIolorithiatlltriin upland rice in M uneng Irobolinggo Indonesia) [in Indonesian] Imut Pertanian (Jogjakarta) 1330-332

249 Yano K T Miyake and V F Eastop 1983 The biology and economic importance of rice aphids (Henuiptera Aphid idae) a review Bull Entomol Res 73539-566

250 Yelp D and P Rhodes 19(4 lhe success of drilled dryland rice iii the Fiji Islands International Rice Commission Working Party on Rice Production and Protection Manila Philippines 3-10 March1964 Food and Agriculture Organization Rome Italy 13 p

251 Young W R 1981 Fifty-five years of research on the sorghum shootfly Insect Sci Appi 23-9

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No 101The economics of hybrid rice production in (hina No 113 Boron toxicity in rice No 102 Rice ratooning No 114 Energy analysis rice production systems and rice research No 103 Growth and development of the (Jeep water rice plant No 115 Production risk and optimal fertilizer rates an application of the

No 104 Faridpur acomputer-assisted instruction nmodel forrainfed lowland random coefficient model No 116 Consumer demand for rice grain quality in Thailand Indonesia andrice

No 105 A readingand listening comprehcnsion test inEnglish for non native the Philippines speakers applying for tiraining at IRIRI No 117 Morphological changes in rice panicle development a review of

No 106 Rice grassy stunt virus 2 a new strain of rice grassy stunt in tire literature

Philippines No 118 IRRI-Korca collaborative project for the development of cold-No 107 Physical losses and quality deterioration in rice plstprodnCtion tolerant lines through anthei culture

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publishers No 120 Changes isinall-farm iLtireliing techii)hgy in Thailand and [he No 109 Classification of Philippine rainfall patterns Philippines No 110 Contributions of modern rice varieties to rutrition in Asia No 121 Landforms and modern rice varieties No Ill Changes in rice breeding in 10 Asian countries 1965-84 No 122 Yield stability and modern rice technology No 112 Design parameters affecting the performance of the IRRI-designed

axial-flow pump

Page 40: UPLANDRICE. INSECTl--1i M T -E!R.EOLOU t RTANCE ,AND …delphacid.s3.amazonaws.com › 6958.pdf · The significance of abiotic factors in upland rice insect ecology will be apparent

40 IRPS No 123 January 1987

164 Nakasuji F M Ishii and Jen-Zon Ito 1984 Rice skippers in Taiwan and their life histories ZZrC (Taipei) Tech Bull 82 10 p

165 Navas I) 1976 Fall armyworum in rice Pages 99-106 il Ecological animal control by habitat management Proc Tall Timbers Con-ference 28 Feb-I Mar 1974 Fall Timbers Res Stn Misc Publ 6 Tallahassee Horida USA

166 Nickel 1 1 1961 Biological control of rice stem borer A feasibility study Tcch Bull 2 International Rice Rcarch Institute PO Box 933 Manila Philippines IIl p

167 Nigan P M and R A Vernia 1985 Insect pests of uplaild rice in Uttar Pradesh Int Rice Res Newsl 10(6)22

168 Nilakhe S S 1985 -cological observations on spittle bugs with emphasis on their occirence in rice Pcsqtui Agropecu Bras 20407-414

169 Nixon G EJ 195 1 lie association of ats with aphids and coccids Commonwealth Iusti lcte of Entonology Iondon UK 36 p

170 Nurdin F an Ishak Manti 1985 Study oit seedling fly of upland rice Interlational Upland Rice Moitoring Tour Report 26 lebruary-2 MarLI Sukaranti West Sumatra

171 Oka 1 N 1979 Ciltural conttol of the brown planthopper Pages 357-369 in Brown planthopper threat to rice production in Asia International Rice Research Institutc 1p0 Box 933 Manila Philippines

172 Ohni M and A Vilnaii 1975 Rice insect pests of Italy biology of Sipha grcria (Kaltenbach) and its control [in Italian] Riso (Milan) 2459-71

173 Otanes F Q atnd 1 I Sison 194 1 Pests of rice Philipp J Agric 12211-259

174 Painter R I 1951 Insect icsistance in crop plants University Press of Kansits Lawrence Kansas USA 520 p

175 Pantua P C and 1A litsinger 1981 Arthropod abundance in continuous vs biannual rice cropping systeis Int Rice Res Ncwsl 6(2)20-21

176 Pathak M ) and V A l)yck 1975 Sttudics ott insect pests of upland rice Pages 186-191 in Major research in upland rice International Rice Research Institute PO Box 933 Ilanila Philippines

177 Patil B R B M IHasabe and 1) S Ajri 198 I Chemical control of white gritb lIditriihinhorraia Fabr intfesting groundntut paddy atnd sorghum Indian 1Entomol 43232-236

178 Pawar A 1) 1974 (ollection and identification of insect pests of rice rlerminal repirt lInternational Rice Research Institute PO Box 933 Manila Philippines 72 p (tnimco)

179 IerrajU A aid A S Reedy 1965 Observations ott diapausc itt tie steti borer ohrice Rice News Teller (lai-Mar)9-13

180 Pont A C 1972 A review of the Oriental species of theri ona Rondani (I)iptera Muscidac) of econtomic importance Pages 27-102 in Control of sorglium shoot fly M G lomtani and W R Young eds Oxford atd IH1 Ptthl Co New I)clhi India 324 p

181 Prasad S C 1PChand aid M 1PSingh 1983 Incidence of rice gall midge in the upland plateau region of Hihar Oryza 20235-236

182 Rai P S 198 0 Nisaga simpex damage to rice in the hill tracts of South India Int Rice kles News[ 5(I)12

183 Rathorc Y S 1969 Gonocplhahli dcprssunt Fabr as itpredator of (tilo zonwIlhis (Swinhoc) and other moths in field cage at Patnagar Uttar Pradesh Indian 1Entotnol 3127

184 Reed W 1983 Crop losses causcd by insect pists in the developing world Proceedings 10th Internatial Congress of Plant Protection lrighton UK 174-80

185 Reedy M V 1981 Ateactogasterinhens(Walker)a new root-grub pest of tipland paddy and inaize Curr Sci 501076

186 Reveno M A 1968 Catalogue of the insect pests of economic crops in Colombia [in Spaiish] Asociacion Latino Americana de Entomo-logia (AIAE) 129-33

187 Reyes G M 1957 Rice dwarf disease in the Philippines Food and Agriculture Organiiation Plant Prot Bull 617-19

188 Rice S E A A Grigarick and M 0 Way 1982 Effect of leaf and panicle feeding byarmyworm (Lepidoptera Noctuidae) larvae on rice grain yield Environ Entomol 7593-595

189 Risch S J 1) Andow and M A Altieri 1983 Agioccosystem diversity arnd pest control data tentative conclusions and nizw research dirctions Environ Entomol 12625-629

190 Rivera C T 1956 Three organic insecticides in thecontrol of insects affecting upland rice Philipp Agric 39465-472

191 Roffrey 1 1972 Locusta outbreaks in the Philippines Acrida 1177-188

192 Rojas R 1) 1941 The effect of pruning at varying ages upon the yield of Ramairio Phipp Agric 29851-860

193 Roonwal M i 1979 Ternite life and termite control in tropical South Asia Scientilic Pnbl odhpr India 139 p

194 Rosetto C1 F da Silva Martins N C Schmidt and L EAzzini 1978 Damage caused by pasturespittle bugs 4Dvoi)lavopictaandDJ schach) on rice lBragantia Nota 6 (Matcli)25-27

195 Rosztto C1- S S Neto I- Amantc I) Iink 1Garcia-Vieira D M de SoLza N V Bansa t and A M de Olivcira 1971 Rice pests of Brazil [in Portuguese] Pages 149-238 inrFA()-International Rice Commission 2nd meeting of the Rice Committee for the Americas 6-11 l)ecember 1971 Pelotas Rio Grande do Sul Brazil

196 Rothschild G H L 1969 Obseratiois on the armyworm Spodoptera matritia acronrctoiiles Gn (Lep Noctuidae) Bull Entomol Res 59143-160

197 Rothschild G 1- L 1974 Parasites of the armworm Spolopiera mauritiaacroncitoihes in Malaysia Fntornopliaga 19293-299

198 Sachan i N S K Gangwar and 1N Katujar 1980 Field crickets damaging cages in North Eastern IHill Region Indian J Entomol 42526-527

199 Sands 1) P A 1977 The biology and ecology of Lepocorisa (Hemiptera Alydidac) in Papua New Guinea Res Bull 18 Departshyment of Primary Industry Port Moresby Papua New Guinea 104 p

200 Sarawak 1968 titomology division Pages 97-106 in Report of the research branch of the Department of Altriculture 1966 Kuching (Rev AIppl Eintomol A56603)

201 Satpathy JM M S Das and K Naik 1977 Effect of multiple and mixed cropping on the incidence of scine important pestsI Entomol Res 178-85

202 Sauer IHF G 1939 Notes Ott Elasmpnlpualtsligniosellths Zeller (Lep Pyr) a serious pest ofcereals in Sao Paolo [in Portuguese] Arq Inst Biol 10199-206 (Rev Appl Entotnol A28378)

203 Satinders 1L A 1 SKing and C L Vargas S 1983 Crop pests of Central Anerica with t bibliography [in Spanish] Centro Agroshynotnico Tropical de Investigacion y Ensenanza Turrialba Costa Rica (CATIE) Tec Bol 9 90 p

204 Sauphanor B 1985 Some factors of upland rice tolerttce to stem borers in West Africa Inseu Sci Appl 6429434

205 Sen P atnd S Chakravorty 1976 Stentflics in Haritighata Nadia West Bengal India [it Rice Res Newsl 1(12) 15

206 Senapai B and 1M Satpathy 1983 The seedling fly Atherigona orrzae Malloch is a pest of rice in Orissa (India) J Entomol Res 777-79

207 Shhjahan M S Alatn M Ah nad and BN Islam 1983 A new scarabaeid beetle pest of dryland rice in Bangladesh Int Rice Res Ncwsl 8(I)16-17

208 Sharma G L S C Modgol and R C Gautam 1979 Effect of depth of submergence ott incidence of bacterial blight and white grub infestation in transplanted rice Int Rice Res Newsl 4(6)12-13

209 Sharma S 1985 Scarabaeid beetle outbreak itt upland rice Int Rice Res Newsl 1(1(l)24

210 Shiraki T 1917 The paddy borer Schoenobhits incertulas Walker [achaku Agric Exp Stn Formosa 256 p

211 Silva A de Brito and 11P Magalhaes 1981 Insect pests of rice [in Portuguese] Circular Tecnica No 22 EMBRAPA Centro de Pesquisa t ecuaria de Tropico Unido (CPAJU) Belem Brazil 14 p

212 Singh S A 1979 Emergence of rice leaf butterfly as a rice pest in Manipur India int Rice Res Newsl 4(4)16-17

213 SinghS A 1983 Changing trends of insect pestsof rice in Manipur Int Rice Res Newsl 8(f)12

214 Siwi S S and P H van Doesburg 1984 Lenovorisa Latreille in Indonesia (Heteroptera Coreidae Alydinae) Zool Meded (Leiden) 58117-129

215 Soejitno J 1977 Reaction of some upland rice varieties toseedling fly eltherikonaexigua Stein [in Indonesian] Siri Hana I Penyakit 630-35 Lembga I osat Fenclilian Pertanian Bogor Indonesia

216 Socnardi 1967 Insect pests of rice in Indonesia Pages675-683 in The major inse pests of thu rice plant Proceedings of a symposium at the Internatital Rice Rec Wi itt S pternber 1964 The Johns Hopkins Press Baltimore Maryland USA

217 Soto P E A T Petcz and I W Buddenhagen 1976 Survey of insect pests and diseases of rice in different ecological zones in Nigeria Rice Entomol Ncwsl 435-46

218 Soto 11E and Z Ziddiqi 1976 Screening for resistance to African rice insect pests Varietal improsetnent sCmi n r Bouak Ivory Coast 13-17 Sept 1976 West Africa Rice l)evelopment Association Monrovia Iiberia

219 Squire F A 1972 Entomological problems in Bolivia PANS (PestArtic News Sumr) 18249-268

220 Stapely 1 II 1978 The rice leaf roller in the Solomon Islands Int Rice Res Newsl 3(l)13

221 Suenaga H 1937 On aphids attacking the roots of rice Oyo-I)oubts Zasshi 9150-152 [in Japanese] (Rev Appl Entomol A25745)

222 Suenaga H 1952 On the seasonal abundance of three subterranean aphids on upland rice Bull Kyushu Agric Exp Sti 1249-262

223 Sutynso Yas and Dandi Soekaria 1964 Beberapa hana yang ttenperang tana man padi tanah kering [in Indonesian] Rapat kerja Padi Tanah Kering 6 sd 8 Juily 1964 Kaliurang 9 p

224 Tallamy I) W and R F Denno 1983 Alternative life history patterns in risky environments in example from lace bugs Pages 129-147 in Insect life history patterns R F l)cnio and If Dingle eds Spring-Verlag 225 p

225 Tamura 1 1939 Observations on some insect pests of field crops [in Japanese] 1 Plant Prot 26414-420 (Rev Appl EntomolA27-556)

226 Tamura and Suzuki 1964 Influences of the early-transplanting practice in rice culture upon development of the rice stem borer Chilo siql)ressalisWalker Bull Ilokuriku Natil Agric Exp Stn 761-94

227 Tan J P 1924 The rice root aphid Drowhi ihirsoia Philipp Agric 13277-288

228 Tanaka 1 1957 Taxonomy and distribution of some subterranean aphids injurious to upland rice in Japan with description of a new species [in Japanese English summary] Botyu-Kagaku 22168-176

229 Tanaka T 1961 The rice root aphids their ecology and control Spec [full College of Agriculture Utsunomiya University Japan F3 p

230 Tauber M IC A Tauber R Nechols and 1Obrycki 1983 Seasonal activity of parasitoids control by external internal and genetic factors Pages 87-108 itt I)iapause and life cycle strategies in insects V K Brown and I Ilodek eds )r W Junk The Hague Netherlands

231 Taylor W F 1972 Effects of artificial defoliation (simulating pest damage) on varieties of upland rice Exp Agric 879-83

232 feotia T P S and S Natud 1966 Biotoomics of the rice skipper Parnaraniathiis Fabricius (Lepidoptera Hesperiidac) Indian J Entomol 28181-186

233 Teran F 0 1971 Rice pests of Santa Cruz [in Spanish Ministerio de Agricultura Sita Cruz Bolivia Bull 45 8 p

234 Than Itun 1976 Population dynamics of the yellow rice borer Tryporza itcertulas (Walker) and its damage to the rice plant MS thesis University of the Philippines at Ios Bafios Laguna Philippines 92 p

235 Uichanco L B 1911 Pests Pages 55-74 it Rice Philipp Agric 11-200

IRPS No 123 January 1987 41

236 van der Laan P A 195 1 Life history of the rice seedling fly Contrib Gen Agric Res Stn Boor Indonesia 118 15 p

237 van der Laan 11A 1959 Correlation between rainfall in the dry season and the occurrence of the white rice borer (Scirpolhaga innotala WIk) in lava Entomol Fxp Appl 212-20

238 Van Driesche R G 1983 Meaning of percent parasitismin studies of insect parasitism Fnviron Entomol 121 I1- 122

239 Venugopal M S and A litsingcr 1984 Fffect ofcarbofuran on rice growth Prot Ecol 7313-317

240 Vietnam Directorate of Rural Affairs 1962 Annual work progress report on crop improvement program of rice sugarcane vegetable and field crops (lor the period from July 1961 to June 1962) (11) 299 p (Rev Appl Entomol A51526-527)

241 Wan M T K 1972 Observations on rice leaf- and planthoppers in Sarawak (Malaysian Borneo) Malay Agric 148308-335

242 West Africa Rice I)esclopnient Association 1978 Annual research report Vol 2 Statistics Monrovia Liberia

243 West Africa Rice Development Association 1979 Annual research report p47 Monrovia Liberia

244 Williams ) F and CS lofgren 1981 Fli I illy l-468 anew bait toxicant for control of the red imported lire at Fla Entomol 64472477

245 Williams I) 1 1970 The mealybugs OFlomoptera Coccoidea Iscudococcidac) of sugar-cane rice and sorghum Bull Entomol Res 0109-188

246 Williams D) 1 1982 Pulinaria icervi (Signoret) and its allies ott sugar cane and other grasses Btll Entoinol Res 72111-117

247 Wirjosuiardjo S 1976 Study ofchenmical cont rot of insect pests ott upland rice Imiu Pertanian (Jogjakarta) 1323-329

248 Wirjosuhardjo S and N Wirarta 1976 Chemical control of the white grub tIolorithiatlltriin upland rice in M uneng Irobolinggo Indonesia) [in Indonesian] Imut Pertanian (Jogjakarta) 1330-332

249 Yano K T Miyake and V F Eastop 1983 The biology and economic importance of rice aphids (Henuiptera Aphid idae) a review Bull Entomol Res 73539-566

250 Yelp D and P Rhodes 19(4 lhe success of drilled dryland rice iii the Fiji Islands International Rice Commission Working Party on Rice Production and Protection Manila Philippines 3-10 March1964 Food and Agriculture Organization Rome Italy 13 p

251 Young W R 1981 Fifty-five years of research on the sorghum shootfly Insect Sci Appi 23-9

IRR Irch Institute Co EN CAS DE NON REMISE RENYOVER A

KLM-PUBLICATION DISTRIBUTION SERVICE Stamp PO BOX 10000 2130 CA HOOFODORP

H0 L L A N D

1286114960 us 1002

LIBRARY DEVELOPMENT RESOURCES

CENT

RM 209 SA-2MPMTDPCT DCAID WASHINGTON

20523USA

ISSN 05-3862Other papers in this series TITLES OF NUMBERS 1-45 ARE LISTED ON THE LAST PAGE OF NO46 THOSE OF NUMBERS 46-70 ARE ON THE LAST PAGE OF NO 71-80AND THOSE OF NUMBERS 71-1O0ARE ON

THE LAST PAGE OF NO 101-122

No 101The economics of hybrid rice production in (hina No 113 Boron toxicity in rice No 102 Rice ratooning No 114 Energy analysis rice production systems and rice research No 103 Growth and development of the (Jeep water rice plant No 115 Production risk and optimal fertilizer rates an application of the

No 104 Faridpur acomputer-assisted instruction nmodel forrainfed lowland random coefficient model No 116 Consumer demand for rice grain quality in Thailand Indonesia andrice

No 105 A readingand listening comprehcnsion test inEnglish for non native the Philippines speakers applying for tiraining at IRIRI No 117 Morphological changes in rice panicle development a review of

No 106 Rice grassy stunt virus 2 a new strain of rice grassy stunt in tire literature

Philippines No 118 IRRI-Korca collaborative project for the development of cold-No 107 Physical losses and quality deterioration in rice plstprodnCtion tolerant lines through anthei culture

systemis No 119 Problen soils as potenial areas for adverse soils-tolerant rice No 108 (olptblication ofIRRI materials a survey of translators and varieties in South and Ssuthcasi Asia

publishers No 120 Changes isinall-farm iLtireliing techii)hgy in Thailand and [he No 109 Classification of Philippine rainfall patterns Philippines No 110 Contributions of modern rice varieties to rutrition in Asia No 121 Landforms and modern rice varieties No Ill Changes in rice breeding in 10 Asian countries 1965-84 No 122 Yield stability and modern rice technology No 112 Design parameters affecting the performance of the IRRI-designed

axial-flow pump

Page 41: UPLANDRICE. INSECTl--1i M T -E!R.EOLOU t RTANCE ,AND …delphacid.s3.amazonaws.com › 6958.pdf · The significance of abiotic factors in upland rice insect ecology will be apparent

212 Singh S A 1979 Emergence of rice leaf butterfly as a rice pest in Manipur India int Rice Res Newsl 4(4)16-17

213 SinghS A 1983 Changing trends of insect pestsof rice in Manipur Int Rice Res Newsl 8(f)12

214 Siwi S S and P H van Doesburg 1984 Lenovorisa Latreille in Indonesia (Heteroptera Coreidae Alydinae) Zool Meded (Leiden) 58117-129

215 Soejitno J 1977 Reaction of some upland rice varieties toseedling fly eltherikonaexigua Stein [in Indonesian] Siri Hana I Penyakit 630-35 Lembga I osat Fenclilian Pertanian Bogor Indonesia

216 Socnardi 1967 Insect pests of rice in Indonesia Pages675-683 in The major inse pests of thu rice plant Proceedings of a symposium at the Internatital Rice Rec Wi itt S pternber 1964 The Johns Hopkins Press Baltimore Maryland USA

217 Soto P E A T Petcz and I W Buddenhagen 1976 Survey of insect pests and diseases of rice in different ecological zones in Nigeria Rice Entomol Ncwsl 435-46

218 Soto 11E and Z Ziddiqi 1976 Screening for resistance to African rice insect pests Varietal improsetnent sCmi n r Bouak Ivory Coast 13-17 Sept 1976 West Africa Rice l)evelopment Association Monrovia Iiberia

219 Squire F A 1972 Entomological problems in Bolivia PANS (PestArtic News Sumr) 18249-268

220 Stapely 1 II 1978 The rice leaf roller in the Solomon Islands Int Rice Res Newsl 3(l)13

221 Suenaga H 1937 On aphids attacking the roots of rice Oyo-I)oubts Zasshi 9150-152 [in Japanese] (Rev Appl Entomol A25745)

222 Suenaga H 1952 On the seasonal abundance of three subterranean aphids on upland rice Bull Kyushu Agric Exp Sti 1249-262

223 Sutynso Yas and Dandi Soekaria 1964 Beberapa hana yang ttenperang tana man padi tanah kering [in Indonesian] Rapat kerja Padi Tanah Kering 6 sd 8 Juily 1964 Kaliurang 9 p

224 Tallamy I) W and R F Denno 1983 Alternative life history patterns in risky environments in example from lace bugs Pages 129-147 in Insect life history patterns R F l)cnio and If Dingle eds Spring-Verlag 225 p

225 Tamura 1 1939 Observations on some insect pests of field crops [in Japanese] 1 Plant Prot 26414-420 (Rev Appl EntomolA27-556)

226 Tamura and Suzuki 1964 Influences of the early-transplanting practice in rice culture upon development of the rice stem borer Chilo siql)ressalisWalker Bull Ilokuriku Natil Agric Exp Stn 761-94

227 Tan J P 1924 The rice root aphid Drowhi ihirsoia Philipp Agric 13277-288

228 Tanaka 1 1957 Taxonomy and distribution of some subterranean aphids injurious to upland rice in Japan with description of a new species [in Japanese English summary] Botyu-Kagaku 22168-176

229 Tanaka T 1961 The rice root aphids their ecology and control Spec [full College of Agriculture Utsunomiya University Japan F3 p

230 Tauber M IC A Tauber R Nechols and 1Obrycki 1983 Seasonal activity of parasitoids control by external internal and genetic factors Pages 87-108 itt I)iapause and life cycle strategies in insects V K Brown and I Ilodek eds )r W Junk The Hague Netherlands

231 Taylor W F 1972 Effects of artificial defoliation (simulating pest damage) on varieties of upland rice Exp Agric 879-83

232 feotia T P S and S Natud 1966 Biotoomics of the rice skipper Parnaraniathiis Fabricius (Lepidoptera Hesperiidac) Indian J Entomol 28181-186

233 Teran F 0 1971 Rice pests of Santa Cruz [in Spanish Ministerio de Agricultura Sita Cruz Bolivia Bull 45 8 p

234 Than Itun 1976 Population dynamics of the yellow rice borer Tryporza itcertulas (Walker) and its damage to the rice plant MS thesis University of the Philippines at Ios Bafios Laguna Philippines 92 p

235 Uichanco L B 1911 Pests Pages 55-74 it Rice Philipp Agric 11-200

IRPS No 123 January 1987 41

236 van der Laan P A 195 1 Life history of the rice seedling fly Contrib Gen Agric Res Stn Boor Indonesia 118 15 p

237 van der Laan 11A 1959 Correlation between rainfall in the dry season and the occurrence of the white rice borer (Scirpolhaga innotala WIk) in lava Entomol Fxp Appl 212-20

238 Van Driesche R G 1983 Meaning of percent parasitismin studies of insect parasitism Fnviron Entomol 121 I1- 122

239 Venugopal M S and A litsingcr 1984 Fffect ofcarbofuran on rice growth Prot Ecol 7313-317

240 Vietnam Directorate of Rural Affairs 1962 Annual work progress report on crop improvement program of rice sugarcane vegetable and field crops (lor the period from July 1961 to June 1962) (11) 299 p (Rev Appl Entomol A51526-527)

241 Wan M T K 1972 Observations on rice leaf- and planthoppers in Sarawak (Malaysian Borneo) Malay Agric 148308-335

242 West Africa Rice I)esclopnient Association 1978 Annual research report Vol 2 Statistics Monrovia Liberia

243 West Africa Rice Development Association 1979 Annual research report p47 Monrovia Liberia

244 Williams ) F and CS lofgren 1981 Fli I illy l-468 anew bait toxicant for control of the red imported lire at Fla Entomol 64472477

245 Williams I) 1 1970 The mealybugs OFlomoptera Coccoidea Iscudococcidac) of sugar-cane rice and sorghum Bull Entomol Res 0109-188

246 Williams D) 1 1982 Pulinaria icervi (Signoret) and its allies ott sugar cane and other grasses Btll Entoinol Res 72111-117

247 Wirjosuiardjo S 1976 Study ofchenmical cont rot of insect pests ott upland rice Imiu Pertanian (Jogjakarta) 1323-329

248 Wirjosuhardjo S and N Wirarta 1976 Chemical control of the white grub tIolorithiatlltriin upland rice in M uneng Irobolinggo Indonesia) [in Indonesian] Imut Pertanian (Jogjakarta) 1330-332

249 Yano K T Miyake and V F Eastop 1983 The biology and economic importance of rice aphids (Henuiptera Aphid idae) a review Bull Entomol Res 73539-566

250 Yelp D and P Rhodes 19(4 lhe success of drilled dryland rice iii the Fiji Islands International Rice Commission Working Party on Rice Production and Protection Manila Philippines 3-10 March1964 Food and Agriculture Organization Rome Italy 13 p

251 Young W R 1981 Fifty-five years of research on the sorghum shootfly Insect Sci Appi 23-9

IRR Irch Institute Co EN CAS DE NON REMISE RENYOVER A

KLM-PUBLICATION DISTRIBUTION SERVICE Stamp PO BOX 10000 2130 CA HOOFODORP

H0 L L A N D

1286114960 us 1002

LIBRARY DEVELOPMENT RESOURCES

CENT

RM 209 SA-2MPMTDPCT DCAID WASHINGTON

20523USA

ISSN 05-3862Other papers in this series TITLES OF NUMBERS 1-45 ARE LISTED ON THE LAST PAGE OF NO46 THOSE OF NUMBERS 46-70 ARE ON THE LAST PAGE OF NO 71-80AND THOSE OF NUMBERS 71-1O0ARE ON

THE LAST PAGE OF NO 101-122

No 101The economics of hybrid rice production in (hina No 113 Boron toxicity in rice No 102 Rice ratooning No 114 Energy analysis rice production systems and rice research No 103 Growth and development of the (Jeep water rice plant No 115 Production risk and optimal fertilizer rates an application of the

No 104 Faridpur acomputer-assisted instruction nmodel forrainfed lowland random coefficient model No 116 Consumer demand for rice grain quality in Thailand Indonesia andrice

No 105 A readingand listening comprehcnsion test inEnglish for non native the Philippines speakers applying for tiraining at IRIRI No 117 Morphological changes in rice panicle development a review of

No 106 Rice grassy stunt virus 2 a new strain of rice grassy stunt in tire literature

Philippines No 118 IRRI-Korca collaborative project for the development of cold-No 107 Physical losses and quality deterioration in rice plstprodnCtion tolerant lines through anthei culture

systemis No 119 Problen soils as potenial areas for adverse soils-tolerant rice No 108 (olptblication ofIRRI materials a survey of translators and varieties in South and Ssuthcasi Asia

publishers No 120 Changes isinall-farm iLtireliing techii)hgy in Thailand and [he No 109 Classification of Philippine rainfall patterns Philippines No 110 Contributions of modern rice varieties to rutrition in Asia No 121 Landforms and modern rice varieties No Ill Changes in rice breeding in 10 Asian countries 1965-84 No 122 Yield stability and modern rice technology No 112 Design parameters affecting the performance of the IRRI-designed

axial-flow pump

Page 42: UPLANDRICE. INSECTl--1i M T -E!R.EOLOU t RTANCE ,AND …delphacid.s3.amazonaws.com › 6958.pdf · The significance of abiotic factors in upland rice insect ecology will be apparent

IRR Irch Institute Co EN CAS DE NON REMISE RENYOVER A

KLM-PUBLICATION DISTRIBUTION SERVICE Stamp PO BOX 10000 2130 CA HOOFODORP

H0 L L A N D

1286114960 us 1002

LIBRARY DEVELOPMENT RESOURCES

CENT

RM 209 SA-2MPMTDPCT DCAID WASHINGTON

20523USA

ISSN 05-3862Other papers in this series TITLES OF NUMBERS 1-45 ARE LISTED ON THE LAST PAGE OF NO46 THOSE OF NUMBERS 46-70 ARE ON THE LAST PAGE OF NO 71-80AND THOSE OF NUMBERS 71-1O0ARE ON

THE LAST PAGE OF NO 101-122

No 101The economics of hybrid rice production in (hina No 113 Boron toxicity in rice No 102 Rice ratooning No 114 Energy analysis rice production systems and rice research No 103 Growth and development of the (Jeep water rice plant No 115 Production risk and optimal fertilizer rates an application of the

No 104 Faridpur acomputer-assisted instruction nmodel forrainfed lowland random coefficient model No 116 Consumer demand for rice grain quality in Thailand Indonesia andrice

No 105 A readingand listening comprehcnsion test inEnglish for non native the Philippines speakers applying for tiraining at IRIRI No 117 Morphological changes in rice panicle development a review of

No 106 Rice grassy stunt virus 2 a new strain of rice grassy stunt in tire literature

Philippines No 118 IRRI-Korca collaborative project for the development of cold-No 107 Physical losses and quality deterioration in rice plstprodnCtion tolerant lines through anthei culture

systemis No 119 Problen soils as potenial areas for adverse soils-tolerant rice No 108 (olptblication ofIRRI materials a survey of translators and varieties in South and Ssuthcasi Asia

publishers No 120 Changes isinall-farm iLtireliing techii)hgy in Thailand and [he No 109 Classification of Philippine rainfall patterns Philippines No 110 Contributions of modern rice varieties to rutrition in Asia No 121 Landforms and modern rice varieties No Ill Changes in rice breeding in 10 Asian countries 1965-84 No 122 Yield stability and modern rice technology No 112 Design parameters affecting the performance of the IRRI-designed

axial-flow pump