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Systematic Entomology (2012), 37, 125–171 DOI: 10.1111/j.1365-3113.2011.00601.x Generic revision and phylogeny of Microweiseinae (Coleoptera: Coccinellidae) HERMES E. ESCALONA 1,2 and A D A M ´ SLIPI ´ NSKI 2 1 Museo del Instituto de Zoología Agrícola, FAGRO-Universidad Central de Venezuela, Maracay, Venezuela and 2 CSIRO-Ecosystem Sciences, Canberra, Australia Abstract. The monophyly and phylogenetic relationships of the subfamily Microwei- seinae were investigated. Twenty-three in-group taxa, representing all known genera of Microweiseinae (except for Microcapillata Gordon) were included in a cladis- tic analysis, based on 45 adult morphological characters. The parsimony analysis of the resulting data matrix supported the monophyly of Microweiseinae, Carinodulini, Serangiini and Microweiseini (inclusive of Sukunahikonini). The recognition of Suku- nahikonini renders Microweiseini paraphyletic, and consequently both tribes were synonymized, retaining Microweiseini as a senior family-group name (syn.n.). Gen- era and tribes of Microweiseinae are diagnosed thoroughly, illustrated and keys to their identification are provided. The following nomenclatural changes were made: Hikonasukuna Sasaji and Orculus Sicard are synonymized with Scymnomorphus Weise (syn.n.); Gnathoweisea Gordon is synonymized with Microweisea Cockerell (syn.n.); Hikonasukuna monticola Sasaji and Orculus castaneus Sicard are transferred to Scymnomorphus (comb.n.); Smilia planiceps Casey, Gnathoweisea hageni Gordon, Gnathoweisea ferox Gordon, Gnathoweisea micula Gordon and Gnathoweisea texana Gordon are transferred to Microweisea (comb.n). Three new genera are described: Allenius gen.n. for Allenius californianus sp.n. (type species) from Mexico and Alle- nius iviei sp.n. from U.S.A.; Gordoneus gen.n. (type species Gnathoweisea schwarzi Gordon from U.S.A.); and Cathedrana gen.n. (type species Cathedrana natalen- sis sp.n. from South Africa). The first African member of Carinodulini, Carinodulina ruwenzorii sp.n. is described. The genera Microweisea, Coccidophilus, Serangium and Delphastus are well-known predators of sternorrynchous Hemiptera, such as scale insects (Diaspididae) and whiteflies (Aleyrodidae), and play a significant role in agricultural ecosystems as biocontrol agents. Host data and biological records are summarized for each genus. Introduction The family Coccinellidae is a relatively large, widely dis- tributed group of beetles, containing many well-known species that are predators of Hemiptera and other invertebrates. It is also a family of economic importance as it includes many species that are either biological control agents or serious agricultural pests, damaging tomato, potato, cucurbits and bean Correspondence: Hermes E. Escalona, CSIRO-Ecosystem Sci- ences, GPO Box 1700, Canberra ACT 2601, Australia. E-mail: [email protected] crops in tropical and subtropical regions (Hodek & Honˇ ek, 1996; Obrycki & Kring, 1998). Members of the subfamily Microweiseinae are among the most bizarre, smallest and least recognizable Coccinellidae (Figs 2, 3). The small, pubescent body forms of Microwei- seinae more closely resemble some Endomychidae (e.g. Anamorphinae and Mycetaeinae) and Corylophidae than ‘true’ lady beetles. The Microweiseinae often have the head with- drawn into the prothorax or deflexed and held closely against the ventral side, and also lack the familiar aposematic colouring of common garden coccinellids. © 2011 CSIRO Systematic Entomology © 2011 The Royal Entomological Society 125

Generic revision and phylogeny of Microweiseinae (Coleoptera: Coccinellidae)

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Page 1: Generic revision and phylogeny of Microweiseinae (Coleoptera: Coccinellidae)

Systematic Entomology (2012), 37, 125–171 DOI: 10.1111/j.1365-3113.2011.00601.x

Generic revision and phylogeny of Microweiseinae(Coleoptera: Coccinellidae)

H E R M E S E . E S C A L O N A1,2 and A D A M S L I P I N S K I2

1Museo del Instituto de Zoología Agrícola, FAGRO-Universidad Central de Venezuela, Maracay, Venezuela and2CSIRO-Ecosystem Sciences, Canberra, Australia

Abstract. The monophyly and phylogenetic relationships of the subfamily Microwei-seinae were investigated. Twenty-three in-group taxa, representing all known generaof Microweiseinae (except for Microcapillata Gordon) were included in a cladis-tic analysis, based on 45 adult morphological characters. The parsimony analysis ofthe resulting data matrix supported the monophyly of Microweiseinae, Carinodulini,Serangiini and Microweiseini (inclusive of Sukunahikonini). The recognition of Suku-nahikonini renders Microweiseini paraphyletic, and consequently both tribes weresynonymized, retaining Microweiseini as a senior family-group name (syn.n.). Gen-era and tribes of Microweiseinae are diagnosed thoroughly, illustrated and keys totheir identification are provided. The following nomenclatural changes were made:Hikonasukuna Sasaji and Orculus Sicard are synonymized with ScymnomorphusWeise (syn.n.); Gnathoweisea Gordon is synonymized with Microweisea Cockerell(syn.n.); Hikonasukuna monticola Sasaji and Orculus castaneus Sicard are transferredto Scymnomorphus (comb.n.); Smilia planiceps Casey, Gnathoweisea hageni Gordon,Gnathoweisea ferox Gordon, Gnathoweisea micula Gordon and Gnathoweisea texanaGordon are transferred to Microweisea (comb.n). Three new genera are described:Allenius gen.n. for Allenius californianus sp.n. (type species) from Mexico and Alle-nius iviei sp.n. from U.S.A.; Gordoneus gen.n. (type species Gnathoweisea schwarziGordon from U.S.A.); and Cathedrana gen.n. (type species Cathedrana natalen-sis sp.n. from South Africa). The first African member of Carinodulini, Carinodulinaruwenzorii sp.n. is described. The genera Microweisea, Coccidophilus, Serangiumand Delphastus are well-known predators of sternorrynchous Hemiptera, such asscale insects (Diaspididae) and whiteflies (Aleyrodidae), and play a significant rolein agricultural ecosystems as biocontrol agents. Host data and biological records aresummarized for each genus.

Introduction

The family Coccinellidae is a relatively large, widely dis-tributed group of beetles, containing many well-known speciesthat are predators of Hemiptera and other invertebrates. It isalso a family of economic importance as it includes manyspecies that are either biological control agents or seriousagricultural pests, damaging tomato, potato, cucurbits and bean

Correspondence: Hermes E. Escalona, CSIRO-Ecosystem Sci-ences, GPO Box 1700, Canberra ACT 2601, Australia. E-mail:[email protected]

crops in tropical and subtropical regions (Hodek & Honek,1996; Obrycki & Kring, 1998).

Members of the subfamily Microweiseinae are among themost bizarre, smallest and least recognizable Coccinellidae(Figs 2, 3). The small, pubescent body forms of Microwei-seinae more closely resemble some Endomychidae (e.g.Anamorphinae and Mycetaeinae) and Corylophidae than ‘true’lady beetles. The Microweiseinae often have the head with-drawn into the prothorax or deflexed and held closely againstthe ventral side, and also lack the familiar aposematic colouringof common garden coccinellids.

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126 H. E. Escalona and A. Slipinski

Despite their small body size, some species of Microwei-seinae are important biological control agents. Serangiini(species of Serangium Blackburn and Delphastus Casey) arewell-known predators of Bemisia spp. whiteflies (Aleyrodi-dae), and are used against whiteflies in different crops, partic-ularly in glasshouses (Legaspi et al., 1996; Obrycki & Kring,1998; Ellis et al., 2001; Al–Zyoud & Sengonca, 2004; Sim-mons et al., 2008). Microweiseini and Sukunahikonini havebeen recorded as predators of various scale insects (Pope, 1962;Chapin, 1965; Chazeau, 1975, 1979; Gordon, 1977, 1985;Vazirani, 1982; Silva et al., 2004).

Historically, following landmark research on lady beetlemorphology by Hirouki Sasaji (1968b, 1971), small and‘atypical’ Coccinellidae (Pseudococcinellidae of Sicard, 1907)were placed in the subfamily Sticholotidinae. Sticholotidinaesensu Sasaji (1968b) included Serangiini, Sukunahikoniniand Sticholotini (including Microweisea), and was definedprimarily by the presence of a narrow and apically pointedterminal maxillary palpomere and a narrow junction betweenmentum and submentum. This diagnosis has been blurred overthe years by the subsequent inclusion of taxa like Shirozuellini,Limnichopharini, Argentipilosini, Cephaloscymnini, Plotininior Carinodulini, in which the terminal palpomere is parallelsided, distally expanded or even knifelike (cultriform).

Subsequent attempts to organize the classification of Sti-cholotidinae have only added to the confusion. Gordon(1977) revised the New World genera of Sticholotidinaeand for the first time recognized Microweiseini in the mod-ern sense, in addition to Sukunahikonini, Serangiini andSticholotidini. Gordon’s (1977) phylogenetic reconstruction(Sukunahikonini + (Microweiseini + (Sticholotidini + Seran-giini))) was questioned by Gordon et al. (1989), who pointed toa natural grouping (Carinodulini + Sukunahikonini + Micro-weiseini + Serangiini) excluding heterogeneous Sticholoti-dini, based on characters of male and female genitalia.In a more recent and comprehensive classification of Coc-cinellidae (Kovar, 1996), Sticholotidinae was divided intoten tribes representing two series, distinguished by char-acters on the metendosternite, pronotum and genitalia ofboth sexes. Kovar (1996) provided no basis for the mono-phyly of Sticholotidinae, and listed only selected char-acters to support some of his groupings (Vandenberg &Perez-Gelabert, 2007). He provided a dendrogram depictinghypothetical phylogenetic relationships among the recog-nized tribes in Sticholotidinae, with the clade (Carin-odulini + (Sikunahikonini + (Microweiseini + Serangiini)))as a sister to the remaining six tribes of Sticholotidinae.This dendrogram from Kovar (1996) was used by Duverger(2003) to revise the classification of Coccinellidae, includingScotoscymninae as a new name for Microweiseinae withoutapplying principles of the International Code of ZoologicalNomenclature. Irrespective of nomenclature and recognizedtribal groupings, the branching dendrograms of Kovar (1996)and Duverger (2003) postulated that Sticholotidinae sensuKovar (1996) was a sister group to the remaining Coccinel-lidae. That relationship was challenged by Slipinski (2007)who segregated Microweiseinae containing Sukunahikonini,

Microweiseini and Serangiini as a monophyletic group, basedon adult, larval and pupal synapomorphies, and placed remain-ing tribes of the former Sticholotidinae (Shirozuellini, Limni-chopharini, Argentopilosini, Cephaloscymnini, Plotinini andSticholotidini) in a redefined subfamily Coccinellinae. Carin-odulini were included only tentatively in Microweiseinaeby Slipinski (2007) because some adult characters (11-segmented antenna and double mandibular apex) were tran-sitional between Microweiseinae and Coccinellinae. The basalsplit of Coccinellidae into Microweiseinae and Coccinellinaeproposed by Slipinski (2007) has been supported by molecularanalyses (Robertson et al., 2008; Giorgi et al., 2009), and bycombined molecular and morphological analyses (Seago et al.,2011), and was implemented by Slipinski & Tomaszewska(2010) in a comprehensive synopsis of Coccinellidae.

The present study was inspired by the discovery of unusualnew genera of Microweiseini, expanding the morphologicaland geographical limits of the tribe. Building on the findingsof Seago et al. (2011) (monophyletic Microweiseinae andCarinodulini; paraphyletic Sukunahikonini with Serangiini andMicroweiseini nested within), we present the first worldgeneric revision and phylogenetic analysis of the group.The main objective of our cladistic analyses was to testthe monophyly of the tribes within Microweiseinae and toinvestigate the placement of the three new genera describedhere. Our study provides a phylogenetic framework for theproposed synonymies, and attempts to reconstruct phylogeneticrelationships within all genera of Microweiseinae.

As currently defined, Microweiseinae contains three tribes,23 genera and around 150 described species, representingabout 3.5% of the Coccinellidae species, distributed mostlyin tropical and subtropical ecosystems.

Material and methods

Material examined and type specimens are deposited in thefollowing collections: AAC, Albert Allen Personal Collection,Boise, Idaho, U.S.A.; ANIC, Australian National InsectCollection, Canberra, Australia; FMNH, Field Museum ofNatural History, Chicago, U.S.A.; MHNG, Museum d’histoirenaturelle de Geneve, Switzerland; MRAC, Mussee Royalde l’Afrique Centrale, Tervuren, Belgium; MTEC, MontanaEntomology Collection, to be placed on permanent loan atCalifornia Academy of Sciences; NHML, Natural HistoryMuseum, London, U.K.; NMNH, National Museum of NaturalHistory, Smithsonian Institution, Washington D.C., U.S.A.

To allow for a thorough examination of morphologicalcharacters, entire specimens or body parts were cleared incold 10% KOH and disarticulated, then examined and digitallyphotographed in glycerol on slides, using a JVC digital cameraattached to a Leitz compound microscope. Subsequently someparts were gold coated to produce SEM images with aZeiss EvoLS15 (CSIRO). Habitus pictures were generatedwith a JVC camera mounted on a Leica M205C dissectingmicroscope, and on the Visionary Digital BK Plus Lab System(Richmond, VA, U.S.A.). Measurements and composite digital

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Generic revision and phylogeny of Microweiseinae 127

images were obtained with automontage (Syncroscopy), andwere enhanced in Adobe photoshop.

Morphological terms follow Lawrence & Britton (1994),Lawrence et al. (2010, 2011) and Slipinski (2007); theCoccinellidae classification follows Slipinski & Tomaszewska(2010) and Seago et al. (2011). The arrangement of the genericdescriptions follows the results of our phylogenetic hypothesis.

Phylogenetic analysis

Taxon selection

Exemplar taxa for phylogenetic analyses were selected fromavailable adult specimens only, as the larvae or larval descrip-tions were available for just four of the 23 genera. Exemplarsincluded all recognized genera of Microweiseinae (Microwei-seini, Sukunahikonini, Serangiini and Carinodulini) except forMicrocapillata Gordon (which could not be scored based onits single, incomplete holotype). In most cases both sexeswere used, if available, for dissection. Although we madeevery effort to score the type species of each genus, ifthat was impossible, our specimen was compared carefullywith the type species to ensure consistency of critical mor-phological characters. Two taxa of Coccinellinae (RhyzobiusStephens and Sticholotis Crotch) and one of Corylophidae,Peltinodinae (Holopsis Broun) were used as out-groups for theanalyses.

In-group taxa

Microweiseinae

Carinodulini.– Carinodulina burakowskii Slipinski & Jadwiszczak (♂, ♀),

Thailand.– Carinodulinka baja Slipinski & Tomaszewska (♂, ♀), Baja

California, Mexico.– Carinodula campbelli Gordon et al. (♂), 5 miles south-

west of Teopisca, Mexico.

Serangiini.– Serangium mysticum Blackburn (♂, ♀), Tasmania, Aus-

tralia.– Delphastus sp. (♂, ♀), Great Smoky Mountains, U.S.A.– Serangiella sp. (♂, ♀), Danbulla, Sri Lanka.

Sukunahikonini.– Paraphellus rostratus (Lea) (♂, ♀), NSW, Australia.– Pharellus popei Slipinski & Tomaszewska (♂, ♀), NT,

Australia.– Orculus castaneus Sicard (♂, ♀), Ivory Coast.– Scymnomorphus rotundatus Weise (♂), Kikuyu, Gatamayu,

Kenya.– Hikonasukuna sp. (♂, ♀), Alishan, Taiwan.

Microweiseini.– Paracoelopterus inconspicuus (Pope) (♂, ♀) South Africa.– Microfreudea cyclica Fursh (♂, ♀), Transvaal, South Africa.

– Hong glorius Slipinski (♀), South Queensland, Australia.– Gnathoweisea planiceps (Casey) (♂, ♀) California, U.S.A.– Gnathoweisea schwarzi Gordon (♂, ♀) Arizona, U.S.A.– Coccidophilus citricola Brethes (♂, ♀), Rio de Janeiro,

Brazil.– Microweisea misella (LeConte) (♀), Tehuacan, Mexico;

Microweisea sp. (♂) Idaho, U.S.A.– Stictospilus darwini Brethes (♂), Chiloe Prov., Chile.– Parasidis sp. (♂), Nuble Prov., Chile.– Nipus biplagiatus Casey (♂), California, U.S.A.

New taxa– Cathedrana gen.n. (♂), Natal, South Africa.– Allenius gen.n. (♂, ♀), Baja California, Mexico.

Out-group taxa– Corylophidae, Corylophinae, Peltinodini. Holopsis sp.

(♂, ♀), Australia.

Coccinellidae, Coccinellinae.– Sticholotidini, Sticholotis ruficeps Weise (♂, ♀), Christmas

Island, Australia.– Coccidulini, Rhizobius ventralis (Erichson) (♂, ♀), ACT,

Australia.

Character coding

Forty-five discrete morphological characters were selectedand coded using delta editor (Dallwitz, 1980; Dallwitzet al., 1999). Unknown character states were coded with ‘?’,and a single inapplicable character was coded with ‘–’. Mostcharacters were based on our own observations and those inSlipinski (2007).

The following characters and character states were used inthe cladistic analysis of the Microweiseinae. All characterswere unordered and unweighted.

1. Anterior edge of clypeus: (1) smooth; (2) margined. Thisrefers to a relatively deep groove along the anterior edgeof the clypeus that continues along the lateral edges andinner margins of the eyes in Serangiini (Fig. 6A).

2. Frontoclypeus in front of eyes: (1) straight or weaklyemarginate (Fig. 4A); (2) distinctly emarginate (Figs16A, 20A).

3. Distance between antennal insertions: (1) less than thatbetween inner eye margins; (2) equal or greater than thatbetween inner eye margins.

4. Posterior margin of antennal socket: (1) distinctly in frontof eyes (Fig. 6A); (2) adjacent to anterior eye margin orin between eyes. In Microweiseinae the antennal socketsare visible dorsally and located in front and between theeyes (Figs 4A, 6A, 20A), as opposed to a location withinthe area of eye emargination in most Coccinellinae.

5. Supraorbital sulcus: (1) absent (Fig. 6A); (2) present(Fig. 4A). This refers to a deep groove extending fromabout antennal insertions along inner margins of eyes

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128 H. E. Escalona and A. Slipinski

seen in Carinodulini and Sticholotis. The similarly posi-tioned line or carina (Fig. 6A) is coded as supraorbital line(character 6).

6. Supraorbital marginal line: (1) absent (Fig. 4A); (2) pre-sent (Fig. 6A).

7. Occipital endocarina: (1) present (Fig. 3G); (2) absent(Fig. 3I). The occipital endocarina is an internal cuticularstructure of variable length, found in few Microweiseini.It is not visible externally and has to be examined on acleared specimen.

8. Ligula: (1) sclerotized (Fig. 4B); (2) membranous anteri-orly (Figs 6D, 12B).

9. Distance between insertions of labial palps: (1) at least asgreat as width of palpifer (Fig. 5E); (2) less than width ofpalpifer (Fig. 15B) or palps contiguous (Fig. 6D).

10. Apical margin of mentum: (1) truncate (Figs 6D, 12B);(2) shallowly emarginate (Fig. 4B); (3) deeply emarginate(Fig. 16C).

11. Maxillary galea: (1) absent; (2) present. Maxilla withoutgalea occurs only in Corylophidae (Holopsis).

12. Maxillary stipes: (1) divided into basistipes and medio-stipes (Fig. 6C); (2) fused into single, elongate structure(Fig. 24G).

13. Maxillary palpifer: (1) convex, not receiving maxillarypalp in repose (Fig. 24G); (2) foveate externally to receivemaxillary palp in repose (Fig. 6C). In the Microweiseinae,maxillary palps are long and geniculate with palpomere 2often being received by foveate palpifer or sometimescardo and palpifer.

14. Shape of terminal maxillary palpomere: (1) knife-like(cultriform) (Figs 4H, 15A); (2) oval and broadenedmedially (Figs 6C, 26A); (3) conical or parallel sided(Figs 12A, 16B). The character states are not clearlydefined, as there is a gradual transition from one state toanother. Maxillary palpomere cultriform (14, 1) has beencoded for Carinodulini and Cathedrana (Microweiseini)that approach this condition.

15. External border of mandible dorsally: (1) normal, broadlyrounded (Fig. 6F); (2) projected into a process (Fig. 10F).This peculiar modification of the mandible in Serangiinihas not been observed before. It appears that theprominent dorsal process forms a ‘false condyle’ that isdorsally articulating with a corresponding clypeal structure(Fig. 10B).

16. Mandibular apex: (1) bidentate; (2) unidentate (Fig. 24D);(3) broadly rounded, reduced. All Microweiseinae, exceptfor Carinodulini, have mandibular apex unidentate. Holop-sis mandibles are blunt to apex, and somewhat reduced;they are coded as a third state.

17. Mandibular mola: (1) developed but smooth, withoutgrinding surfaces or molar tooth (Fig. 6F); (2) withgrinding surfaces or molar tooth. The typical mola inCoccinellidae is smooth, without grinding surfaces, butwith a single or double sclerotized tooth (Slipinski, 2007).The molar tooth is absent in Microweiseinae, althoughthe mola is somewhat angulate in Carinodulini. Grindingmolar surfaces are present only in Holopsis in our dataset.

18. Gena ventrally: (1) extending forwards and forming aframe for mouthparts, approaching clypeus from below(Fig. 10B); (2) extending forwards and forming a framefor mouthparts, and entirely fused to clypeus in front ofantennal insertions (Fig. 24E); (3) not extending forwardsand clearly separated from clypeus. This character isdifficult to observe without scanning electron microscope(SEM) images showing head laterally with antennaremoved (as in Fig. 24E). All Microweiseinae have thegena extending forwards, caused by somewhat rostratehead and strongly retracted ventral mouthparts.

19. Glandular structures or glandular openings on subgenalarea: (1) well delimited and separated from the mouthcavity (Fig. 15B); (2) not well delimited and closelyadjacent to the mouth cavity (Fig. 6D); (3) absent. Itappears that in all members of Microweiseinae the subgenabears a pit or distinctive patch of cuticle consistingof rough granules and micro-openings that could be ofglandular nature. This structure has not been reportedbefore for in Coccinellidae, and its function remainsunknown.

20. Number of antennal segments: (1) 7–10; (2) 11. Thenumber of antennal segments often varies within agenus in some Microweiseinae (e.g. Scymnomorphus andSerangium).

21. Antennal club: (1) one-segmented and distinctly flat-tened (Figs 6H, 10E); (2) one-segmented and roundedin cross section (Fig. 12H); (3) two- to four-segmented(Figs 15C, 26D).

22. Anterior edge of pronotum: (1) medially emarginate(Fig. 23G); (2) arcuate (Fig. 21G).

23. Sublateral carina on pronotum: (1) absent; (2) at leastpartially visible (Fig. 5D). The sublateral carina is adistinct structure, present in Carinodulini only, separatefrom the pronotal carina, which forms the lateral edge ofthe prothorax.

24. Anterior corners of pronotum: (1) separated from discby line joined to lateral and anterior margins (Fig. 8D);(2) separated from disc by a line joined to anterior marginonly (Fig. 18D); (3) not separated from disc. Most generaof Sukunahikonini and Microweiseini have the pronotaledges margined by continuous or fragmented marginallines. These structures may vary slightly and appear as aline, carina, shallow sulcus or a combination of these. Themarginal line near anterior corners often cuts across thedisc (Fig. 23G), thus separating corners from the pronotaldisc.

25. Pits on pronotum: (1) absent (Fig. 7B); (2) present(Fig. 4G).

26. Pits on prosternum: (1) absent; (2) present.27. Prosternum in front of coxa: (1) strongly reduced

(Fig. 16F); (2) well developed and always longer than halfof coxa length (Figs 4I, 21E). The length of the proster-num and corresponding length of the coxa is measuredat the middle of the procoxal cavity, and not along themidline of the prosternum that often includes a prominentchinpiece (Fig. 21E).

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28. Prosternum: (1) entirely raised, forming a triangularplate (Fig. 7A); (2) with anteromedian prominence (chin-piece as defined by Lawrence et al., 2011) (Fig. 21E);(3) straight anteriorly, without chinpiece (Fig. 4I); (4) ant-erior margin broadly arcuate, emarginate laterally toreceive antennal funicle in repose (Fig. 12D).

29. Prosternal process: (1) broad and extending behind coxa(Fig. 4I); (2) narrow, carina-like and often incompleteposteriorly (Fig. 17C).

30. Procoxal cavities: (1) with lateral slits (Fig. 7A); (2) with-out slits (Fig. 4I).

31. Prosternal rest: (1) absent (Fig. 22J); (2) present (Fig. 8C).32. Anterior margin of mesoventrite: (1) about same level as

mesometaventral junction (Fig. 4D); (2) on much lowerlevel than the mesometaventral junction (Fig. 7F).

33. Mesometaventral junction: (1) narrow, as wide or lessthan a coxal diameter (Figs 4D, 7F); (2) broad, distinctlybroader than a coxal diameter (Fig. 11C).

34. Metaventral postcoxal lines: (1) joined medially (Fig. 4D);(2) separate medially (Fig. 11C).

35. Metaventral postcoxal lines: (1) V-shaped (Fig. 4D);(2) descending laterally (Fig. 13B); (3) recurved(Fig. 14E).

36. Metaventral postcoxal lines and associated crural impres-sions: (1) reaching metanepisternum (Fig. 7F); (2) limitedto metaventrite only (Fig. 12I). The ventral foveae (cru-ral impressions) to accommodate legs are present only inSerangiini in our dataset. Metanepisternum is used insteadof the traditional metepisternum following a revisednomenclature of beetle ventral sclerites by Lawrenceet al. (2011).

37. Number of abdominal ventrites: (1) 5 (Fig. 11A); (2) 6(Fig. 16E). The number of abdominal ventrites is noteasy to code consistently because ventrite 6 is sometimesvisible apically as a result of sexual dimorphism ordistortion. In taxa coded here as having abdomen with sixventrites, ventrite 5 is truncate apically and the ventrite 6is at the same level as ventrite 5.

38. Abdominal postcoxal lines: (1) V-shaped (Fig. 4F);(2) descending and incomplete, not reaching lateral edge(Fig. 13I); (3) descending and complete, reaching lateraledge (Fig. 17H); (4) recurved (Fig. 7E).

39. Accessory postcoxal line: (1) absent (Fig. 12J); (2) present(Fig. 15H). The abdominal postcoxal region in Coccinel-lidae is very variable and often difficult to code. InMicroweiseinae the postcoxal lines are often accom-panied by a shorter, oblique accessory line extendingfrom the coxal cavity, and by various pores or deeppunctures.

40. Elytral epipleuron: (1) without foveae receiving legs inrepose (Fig. 17G); (2) with foveae receiving legs in repose(Fig. 11G).

41. Number of tarsomeres: (1) three; (2) four. This charactervaries within genera because of partial fusion of the lasttwo tarsomeres.

42. Pretarsal claws: (1) simple; (2) appendiculate.

43. Male genitalia with tegmen (at least phallobase): (1) asym-metrical (Fig. 28B, G); (2) symmetrical. All Microwei-seinae are characterized by asymmetrical phallobase inmale genitalia, this is often further pronounced byasymmetry in parameres (Fig. 28G).

44. Number of abdominal spiracles: (1) 7; (2) 5. All Coccinel-lidae have five pairs of functional spiracles but seven pairsare usually found in Corylophidae (Slipinski et al., 2009),including Holopsis.

45. Penis base: (1) without capsule (Fig. 28J); (2) with dis-tinct capsule (Fig. 14L).

Analysis

Tree searches were conducted in nona (Goloboff, 1999) runwithin winclada 1.00.08 (Nixon, 2002) and in tnt (Goloboffet al., 2003). Initial parsimony analyses were performedusing the Parsimony Ratchet (Nixon, 1999), under thefollowing settings: 1000 iterations, ten trees held, six charactersresampled and ambiguity set to polytomy. Unsupported nodeswere collapsed in winclada and only the ‘best trees’ werefurther investigated. Bremer support values (Bremer, 1994)were calculated in tnt for suboptimal trees set to ten stepslonger. To reduce the effects of homoplasy in our dataset,further parsimony searches with implied weighting wereconducted using tnt. Under the implied weight criterion theexisting character conflicts in the dataset are resolved in favourof the characters with lower homoplasy by searching trees witha maximum total fit. We repeated this analysis with concavity(K) values of 3–15.

Results and discussion

The resulting data matrix (see Table S1) comprised 26exemplar species and 45 morphological characters from adultbeetles. All characters were considered to be of equal weight,and multistate characters were treated as unordered.

Parsimony analysis using the ratchet yielded 28 mostparsimonious trees (tree length, L = 110 steps; consistencyindex, CI = 0.51; retention index, RI = 0.73) with rather vari-able crown topology. The resulting trees provided conflict-ing support for the position of Serangiini in Microweiseinae.Some reconstructions show Serangiini as sister to the cladethat includes Microweiseini + Sukunahikonini (e.g. Fig. 1A),whereas others (e.g. Fig. 1B) place it well embedded withinthe clade that now includes Microweiseini + Sukunahikonini+ Serangiini. This conflict resulted in the large polytomy in thestrict consensus tree (Fig. 1C) and the lack of convincing Bre-mer support for tribal relationships in Microweiseinae, exceptfor Carinodulini. No parsimony reconstruction supporteda monophyletic Microweiseini (sensu Fursch, 1985), butmonophyletic Sukunahikonini was always embedded betweenbasal Old World clades (Microfreudea + Paracoleopterus) +Hong and the clade containing all the New World genera.

Further parsimony analyses under implied weighting intnt with the concavity constant (K) set in the range 3–15

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130 H. E. Escalona and A. Slipinski

A

B

C

Fig. 1. Phylogenetic hypothesis for Microweiseinae genera. A, strict consensus of the two shortest trees obtained using implied weighting intnt with K = 10 (tree length, L = 111 steps; consistency index, CI = 51; retention index, RI = 72), with characters mapped on branches usingunambiguous character changes in winclada (•, non-homoplasious changes; ©, homoplasious changes; the numbers above and below each circlecorrespond to the character number and character state, respectively); the boldface numbers above the branches indicate the Bremer support values.B, one of the 28 most parsimonious trees found by Ratchet in winclada, with Serangiini in terminal position. C, strict consensus tree of 28 mostparsimonious trees found by Ratchet in winclada.

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A B C

D E F

G H I

Fig. 2. A, Stictospilus darwini ; B, Cathedrana natalensis; C, Carinodula sp.; D, Parasidis sp.; E, Scymnomorphus castaneus; F, Carinodulinkabaja; G, Microfreudea cyclica; H, Coccidophilus sp.; I, Carinodulina burakowskii.

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132 H. E. Escalona and A. Slipinski

A

E F G

H I J

B C D

Fig. 3. A, B, Allenius californianus: (A) dorsal; (B) lateral. C, D, Gordoneus schwarzi : (C) dorsal; (D) lateral. E, F, Hong glorius: (E) lateral;(F) dorsal. G, Gnathoweisea sp., head. H, Paracoelopterus sp. I, Cathedrana natalensis, head. J, Gordoneus schwarzi, head.

found very few minimum length trees, often three or four,but always returned Serangiini as a sister to the cladeMicroweiseini + Sukunahikonini. Increasing K from three toten yielded little change in the tree topology, with the exceptionof Parasidis, which moved from an unresolved position nextto Cathedrana (on trees with K = 3) to the crown group ofNew World taxa (K = 10 or 15).

As a preferred hypothesis for the relationships for Microwei-seinae, we have chosen the results from implied weightinganalyses with K = 10 (Fig. 1A). These results are consistentwith the majority-rule consensus of most parsimonious treesfound by nona, but with much better resolution for the genericrelationships within Microweiseinae. The tree illustrated fordiscussion (Fig. 1A) is a strict consensus of two trees found bytnt with concavity (K) set to ten. The characters and characterstates were mapped on the cladogram using unambiguousoptimization in winclada.

Phylogenetic assessment of tribes of Microweiseinae

Microweiseinae. The monophyly of Microweiseinae (includ-ing Carinodulini) is supported by four unambiguous synapo-morphies. Of these, the position of antennal insertions (4, 1)and asymmetrical tegmen (at least phallobase) in the malegenitalia (43, 1) are distinctive modifications not found inCoccinellinae (with very few exceptions that are apparentlyof secondary origin, c.f. Seago et al., 2011) or Corylophi-dae. Tegminal asymmetry is found in ‘higher’ Endomychidae(Tomaszewska, 2000), but it involves a substantial reductionof the phallobase and parameres. The placement of antennalinsertions well before the anterior margins of eyes is unique tothis group of Coccinellidae; it may be correlated with a gen-eral elongation of the head, but is also pronounced in the veryshort, transverse head capsule of Serangiini.

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Carinodulini. The monophyly of Carinodulini is clearlysupported in this study by the peculiar sublateral carinae onpronotum (23, 2) often associated with dorsal pits (absent inCarinodula) and the peculiar V-shaped metaventral postcoxallines (35, 1).

Serangiini. Morphologically this is the best-defined group inMicroweiseinae, with several unique apomorphies. The mostapparent are: a mandible with a long dorsal ‘condyle’ (15, 2);the large and flattened one-segmented antennal club, (21, 1);the prosternum raised and forming a triangular plate (28, 1)with anterior margin meeting the clypeus in repose, and theventral side with distinct crural impressions accommodatingthe legs (36, 1; 40, 2). Our hypothesis favours a separatetribe Serangiini; further research is necessary to determine

the relationships of Serangiini and Microweiseini, because ofexisting conflicting hypotheses (Fig. 1A, B).

Sukunahikonini. Five genera of this tribe form a mono-phyletic group based on two homoplasious characters: therelatively reduced prosternum (27, 1) and narrowly separatedcoxae (29, 2). The origin of this lineage well within a broadlydefined Microweiseini prevents recognition of this clade as avalid tribe (Seago et al., 2011)

Microweiseini. All of the New World members of tradi-tional Microweiseini (sensu Gordon, 1977) form a mono-phyletic group supported by the prominent prosternal chinpiece(28, 2). However, the chinpiece is absent in North Ameri-can Gordoneus (former Gnathoweisea schwarzi Gordon), butis present in the Australian Hong. The clade that includes

A

D

G H I

F

E

C

B

Fig. 4. Carinodulina ruwenzorii : (A) head, dorsal; (B) mouthparts; (C) head, ventral; (D) meso–metaventrite; (E) antenna; (F) abdomen;(G) pronotum; (H) terminal maxillary palpomere; (I) prothorax, ventral.

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134 H. E. Escalona and A. Slipinski

Microweiseini and Sukunahikonini can be recognized as a sis-ter group to Serangiini based on the shared presence of uniqueglandular structures on the ventral side of the mouth cavity (19,1) and the homoplastic form of the abdominal postcoxal lines(38, 2). The African–Mediterranean clade of Microfreudea +Paracoleopterus is postulated here as a sister group to theremaining Microweiseini. These genera show some similarities(one-segmented antennal club, large and anteriorly prominentprosternum and abdomen occasionally with five ventrites) toSerangiini, and always occupy an intermediate position on thecladograms.

The generic relationships of Microweiseini are rela-tively well resolved here and most generic groups arestrongly supported and recognizable by morphological diag-nostic characters. Three genera of former Sukunahikonini −Scymnomorphus, Orculus and Hikonasukuna (Fig. 1A) forma monophyletic group supported by a narrow prosternal pro-cess and multisegmented antennal club. Examination of almostall world species of the large and variable Scymnomorphusrevealed clear transitions in diagnostic characters (develop-ment of wings, body shape, epipleural width). In the absenceof generic diagnostic characters and a study of most speciesworldwide we propose to include Orculus and Hikonasukunain Scymnomorphus. Two species of Gnathoweisea are placedvery far apart on our cladograms, with the type species(G. planiceps) being closely allied to Microweisea (and heresynonymized with that genus because of the absence of sepa-rate diagnostic characters), and G. schwarzi nested outside thatclade (described as a separate genus, Gordoneus). The poly-tomy formed by Coccidophilus, Stictospilus, Microweisea andParasidis cannot be resolved yet.

This is the first comprehensive phylogenetic and genericstudy of the subfamily Microweiseinae. Its monophyly wascorroborated as suggested by previous studies (Slipinski, 2007;Robertson et al., 2008; Giorgi et al., 2009; Seago et al., 2011),the internal phylogenetic relationships between its genera wererecognized and congruent with those of Seago et al. (2011),with the addition of several new morphological charactersthat define major clades and more clearly diagnose eachgroup.

Classification

Consistent with our phylogenetic hypotheses, we proposethe following classification:

Family Coccinellidae Latreille, 1807Subfamily Microweiseinae Leng, 1920Tribe Carinodulini Gordon et al., 1989

– Carinodula Gordon et al., 1989– Carinodulina Slipinski & Jadwiszczak, 1995– Carinodulinka Slipinski & Tomaszewska, 2002

Tribe Serangiini Pope, 1962– Serangium Blackburn, 1889– Delphastus Casey, 1899– Serangiella Chapin, 1940

Tribe Microweiseini Leng, 1920= Sukunahikonini Kamiya, 1960 syn.n.

– Paracoelopterus Normand, 1936– Microfreudea Fursh, 1985– Hong Slipinski, 2007– Cathedrana gen.n.– Gordoneus gen.n.– Coccidophilus Brethes, 1905– Microweisea Cockerell, 1903

= Gnathoweisea Gordon, 1970 syn.n.– Microcapillata Gordon, 1977– Stictospilus Brethes, 1925– Parasidis Brethes, 1915– Nipus Casey, 1899– Allenius gen.n.– Paraphellus Chazeau, 1981– Pharellus Sicard, 1928– Scymnomorphus Weise, 1897

= Orculus Sicard, 1931 syn.n.= Hikonasukuna Sasaji, 1967 syn.n.

Microweiseinae Leng

Diagnosis. Adult. Maximum length 3.5 mm. Antenna inser-ted in front of eyes and often separated from eyes anteriorly(Figs 4A, 6A, 20A); antennal insertions exposed and relativelyclose together (Fig. 20A); frontoclypeus well developed andemarginate around antennal insertions (Figs 6A, 19D); sub-gena with glandular openings (Figs 6D, 8F, 15A); mandiblewith well-developed mola but without molar tooth (Figs 6F,25C). Ventral mouthparts retracted causing unusual projectionof genae into a frame (Figs 4C, 8B, 21B) enclosing stronglyelongate maxillae and labium. Male genitalia with asymmetri-cal tegmen, including at least phallobase (Fig. 28B), and oftenpenis guide and parameres.

Larva. Mandible triangular with single apical tooth and bluntnon-projected molar area; abdomen without gland openings;tibiotarsus with a pair of flattened, spatulate setae.

Pupa without urogomphi.

Distribution. Worldwide, but most of the species occurin tropical and subtropical regions; not recorded from NewZealand.

Key to tribes of Microweiseinae

1. Mandibular apex bidentate; prothorax and mesoventrite usu-ally with deep pits; pronotum with sublateral carina well sep-arated from lateral margin (Figs 4G, 5D); terminal maxillarypalpomere knife-like (cultriform) (Fig. 4H) . . . . . Carinodulini– Mandibular apex unidentate (Fig. 24D); prothorax andmesoventrite without pits; pronotum without sublateral cari-nae (Figs 13F, 21G); terminal maxillary palpomere variable(Figs 6C, 10A, 12A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2 (1) Entire prosternum elevated, forming large triangular plate(Figs 7A, 8E, 10C), its anterior margin closes with anterior

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margin of clypeus in repose; ventral side, including epipleuradeeply foveate, receiving folded legs (Figs 7F, 9D); abdomenfive-segmented with ventrite 5 as long as ventrites 2–4combined (Fig. 7E); antenna eight- or nine-segmented withflattened one-segmented antennal club (Fig. 6H) . . . Serangiini– Prosternum variable, flat, strongly reduced or variouslylobed anteriorly (Figs 12D, 14D, 17C, 20D, 21E, 22B),partially or almost completely concealing mouthparts; ventralside, including epipleura, without foveae receiving foldedlegs (Figs 17G, 25E); abdomen five- or six-segmented withventrite 5 almost always short; antenna eight- to ten-segmentedwith one- to three-segmented antennal club that is circular incross section (Figs 12H, 14C, 24C) . . . . . . . . . . Microweiseini

Review of tribes and genera

Carinodulini Gordon, Pakaluk & Slipinski

Carinodulini Gordon et al., 1989: 360. Type genus Carin-odula Gordon et al., 1989.

Diagnosis. Wingless. Eyes reduced, consists of coarse facets(Fig. 4A); head with deep supraorbital sulcus along inner mar-gin of eye, extending anteriorly (Fig. 4A); mandibular apexbidentate, molar part angulate; pronotum with sublateral carina(Fig. 4G); prosternum flat, long in front of procoxae; prosternalprocess extending well beyond procoxae (Fig. 4I). Mesoven-trite long and flat; mesometaventral process narrow; metaven-tral postcoxal lines deep, V-shaped (Fig. 4D). Abdominal post-coxal lines V-shaped (Fig. 4F) or shallow and close to coxa.

Distribution. Known species occur in relictual isolatedmountainous areas of North and Central America, Africa andAsia.

Comments. Contrary to the well-known general tendency ofshortening of the thoracic ventrites in wingless beetles, allCarinodulini have long metaventrite.

Key to genera of Carinodulini

1. Pronotum and prosternum without pits; tarsi four-segmented.Mexico . . . . . . . . . . . Carinodula Gordon, Pakaluk & Slipinski– Pronotum and prosternum with dorsal (Fig. 4G) and/orventral pits; tarsi three-segmented (Fig. 5H) . . . . . . . . . . . . . . 2

2 (1) Antenna 11-segmented (Fig. 4E); prothorax transverse(Fig. 4G); pronotum with two pairs of pits and often one pairof pits on prosternum; mesoventrite without pits (Fig. 4D);abdominal postcoxal lines extending to posterior margin ofventrite (Fig. 4F). Thailand, D.R. Congo, India. . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . Carinodulina Slipinski & Jadwiszczak– Antenna ten-segmented; prothorax elongate; pronotum andprosternum each with two pairs of pits; mesoventrite with a pairof pits along mesoventral suture; abdominal postcoxal linesshallow and close to coxal cavity. Mexico, U.S.A. . . . . . . . . .. . . . . . . . . . . . . . . . . . . Carinodulinka Slipinski & Tomaszewska

Carinodula Gordon, Pakaluk & Slipinski(Fig. 2C)

Carinodula Gordon, Pakaluk & Slipinski, 1989: 360. Typespecies Carinodula campbelli Gordon et al., 1989 (originaldesignation).

Diagnosis. Carinodula and Carinodulina combine11-segmented antennae (ten-segmented antennae in Carin-odulinka) with a complete sublateral carina on pronotum.Carinodula can be easily separated from Carinodulina by itsfour-segmented tarsi and prothorax without deep pits.

Description. Length 1.4 mm. Elongate and flattened; dorsalsurfaces covered with uniform long setae (Fig. 2C). Colour yel-lowish brown to light brown. Head transverse; eye consists of17–19 coarse facets. Frontoclypeus as long as antennal socketlength, lateral edges margined, slightly emarginate aroundantennal insertions. Subantennal grooves short. Subgenal glandopenings present. Gula elongate, surface smooth with few api-cal setae, gular sutures distinct. Antenna 11-segmented withtwo-segmented club (Fig. 4E); antennomeres 8 and 9 trans-verse. Mouthparts: mandibular apex bidentate, molar surfacenarrow and smooth, with a weakly developed tooth, prosthecareduced; terminal maxillary palpomere knife-like (cultriform)(Fig. 4H); submentum broad, mentum rectangular, as wide assubmentum; ligula slightly shorter than labial palp and wellsclerotized, apex fringed with setae. Prothorax transverse, with-out pits. Pronotal disc convex, lateral edge finely crenulate,sublateral carina complete. Prosternum in front of coxae about1.3 times coxa length. Prosternal process parallel-sided, apexrounded. Elytra irregularly punctate; epipleura broad reach-ing abdominal ventrite 2, without foveae. Scutellum triangularand transverse. Mesoventrite transverse, flat, without pits, pro-coxal rest present, mesometaventral junction at midline slightlynarrower than coxa. Metaventrite transverse, shorter thanabdominal ventrite 1; postcoxal lines V-shaped. Abdomen withsix ventrites: ventrite 1 longer than ventrites 2–4 combined;postcoxal lines V-shaped (Fig. 4F). Legs slender, tarsi four-segmented; pretarsal claws simple. Male genitalia: parameresnot observed; penis very long and coiled. Ovipositor triangular,styli elongate with simple apical setae; spermatheca constrictednear middle and globular at sides.

Distribution. South Mexico–Central America.

Species included. Carinodula campbelli Gordon et al.

Material examined. Carinodula campbelli and undescribedspecies from Central America.

Comments. The type series of C. campbelli was collectedsifting litter in Quercus and Pinus forest; gut contents includedfractions of sternorrhynchous Hemiptera (Gordon et al., 1989).

Carinodulina Slipinski & Jadwiszczak(Figs 2I, 4A–I, 5A–I, 28J, K)

Carinodulina Slipinski & Jadwiszczak, 1995: 344. Typespecies Carinodulina burakowskii Slipinski & Jadwiszczak,1995 (original designation).

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136 H. E. Escalona and A. Slipinski

A

C

F

G

H

I

D

E

B

Fig. 5. Carinodulina spp.: (A) C. burakowskii, prosternum detail; (B) C. ruwenzorii, prosternum detail; (C–E) C. burakowskii (C) head, ventral;(D) pronotum detail; (E) mouthparts detail; (F) C. burakowskii, antenna; (G) C. ruwenzorii, middle leg; (H) C. ruwenzorii, middle tarsus;(I) C. burakowskii, epipleura.

Diagnosis. The 11-segmented antennae and complete prono-tal sublateral carina are reminiscent of Carinodula, but Carin-odulina has deep pronotal pits, three-segmented tarsi and well-developed parameres in male genitalia.

Description. Length 1.4 mm. Elongate oval (Fig. 2I), dor-sal surfaces covered with uniform long setae. Colour lightbrown. Head longer than wide (Fig. 4A); eyes consist of 14–22coarse facets with few fine setae. Frontoclypeus (Fig. 4A)shallowly emarginate around antennal insertions, lateral edgesmargined. Subantennal grooves short. Subgenal gland openingsas Fig. 4B. Gula transverse, surface smooth with few scatteredsetae, gular sutures parallel and distinct. Antenna 11-segmented(Figs 4E, 5F) with two-segmented club; antennomeres 7–9transverse. Mouthparts (Figs 4B, 5C, E): mandibular apexbidentate, molar surface smooth, prostheca well developed;maxillary palpifer foveate, terminal maxillary palpomereelongate and knife-like (cultriform) (Fig. 4H); submentum

broad, mentum rectangular, as wide as submentum; ligulashorter than labial palp, apex fringed with setae; labial palpsseparated by slightly less than palpomere 1 diameter. Protho-rax transverse (Fig. 4G, I), with three pairs of pits, two onpronotum and one on prosternum in front of coxae. Pronotum(Fig. 4G) with lateral edge crenulate, disc convex; sublateralcarina extends from anterior pit to posterior edge (Figs 4G,5D). Prosternum (Fig. 4I) in front of coxae about 1.5 coxalength, not raised; prosternal process subparallel, 0.3–0.5 coxawidth. Elytra with irregular punctation; epipleura incomplete,setose (Fig. 5I). Scutellum transverse, broadly rounded poste-riorly and slightly constricted at base. Mesoventrite (Fig. 4D)flat, procoxal rest well extended, without pits, in front of coxa,as long as coxa. Metaventrite (Fig. 4D) well developed andtransverse, about as long as abdominal ventrite 1; postcoxallines recurved and complete or reduced. Abdomen (Fig. 4F)with six ventrites: ventrite 1 longer than ventrites 2 and 3

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combined; postcoxal lines V-shaped. Legs slender; tarsi three-segmented. Male genitalia (Figs 28J, K): penis guide veryshort; parameres articulated as long as tegmen, slightly asym-metrical with setose apex; penis curved, without basal capsule.Ovipositor elongate, styli cylindrical and terminal with simpleapical setae, spermatheca constricted at centre, globular andelongated.

Distribution. Mountains in Thailand, southern India andDemocratic Republic of Congo.

Species included. Carinodulina burakowskii Slipinski &Jadwiszczak; Carinodulina ruwenzorii sp.n.

Material examined. Carinodulina burakowskii ; C. ruwen-zorii sp.n. and undescribed species from India.

Comments. The single male specimen from India (AnamalaiHills, MHNG) differs from the two named species by lackingprosternal pits.

Carinodulina ruwenzorii sp.n.(Figs 4A–I, 5B, G, H, 28J, K)

Diagnosis. This species is very similar to C. burakowskii ;apart from differences in the male genitalia it differs from thatspecies by sublateral carina (Fig. 4G) very close to lateral edgeof prothorax, and prosternum lateral edge at the same level(Fig. 5B) as compared with prosternum lateral edge descendingbefore apex in C. burakowskii (Fig. 5A).

Description. Length 1.4 mm; width 0.5 mm. Colour light todark brown. Dorsum covered with dense, fine setae; pronotumand elytra finely punctate, punctures more than five diametersapart (Fig. 4G). Prosternal process 0.4 times the width ofcoxa (Fig. 4I). Metaventral postcoxal lines deeply recurvedand complete (Fig. 4D). Abdominal postcoxal lines reachingventrite 2 (Fig. 4F). Male genitalia as Figs 28J, K.

Etymology. Named after the type locality, the astonishingRuwenzori Mountains (currently Rwenzori Mountains) inCentral Africa.

Type material. Holotype, ♂, CONGO BELGE: PNA, MassifRuwenzori, Mont Vuhonga, vallee de la Lume, 2200 m.,14.328–29, 22–viii–1955, (Vanschuytbroeck ) (MRAC).

Paratypes, as holotype (1, MRAC); as holotype but14.280–83, Massif Ruwenzori, Kalonge, 2480 m., (etagebamboos) Riv. Nyamwamba–Ihongero (1, MRAC; 1, NHML);as holotype but 14.372–80, Kibombole (lieu–dit), pisteKalonge–Mahungu, 2700 m., terreau (2, MRAC).

Carinodulinka Slipinski & Tomaszewska(Fig. 2F)

Carinodulinka Slipinski & Tomaszewska, 2002: 489. Typespecies Carinodulinka baja Slipinski & Tomaszewska, 2002(original designation).

Diagnosis. Carinodulinka differs from Carinodula andCarinodulina in having elongate, flattened and weakly sclero-tized body, ten-segmented antenna and incomplete sublateralcarina on pronotal disc.

Description. Length 0.9–1.0 mm. Body elongate and flat-tened (Fig. 2F), lightly sclerotized, dorsal surfaces coveredwith short setae. Colour yellowish to reddish brown. Headelongate, eyes consist of five or six coarse facets, glabrous.Frontoclypeus shorter than antennal socket length, weaklyemarginate around antennal insertions. Subantennal groovesabsent. Subgenal gland openings present. Gula elongate,gular sutures reduced and converging. Antenna ten-segmentedwith two-segmented club, antennomere 8 transverse. Mouth-parts: mandibular apex bidentate, molar surface broad andsmooth, prostheca reduced; maxillary palpifer foveate; terminalmaxillary palp large and knife-like (cultriform); submentumand mentum narrow, mentum slightly expanded to apex, ligulavery short, labial palps separated by about palpomere 1 diam-eter. Prothorax subquadrate, with four pairs of pits: two pairson pronotum and two pairs on prosternum (in front of coxaalong the notosternal suture). Pronotum narrow at base, ante-rior angles obtuse, posterior angles well-defined, lateral edgefinely crenulate, disc flattened, sublateral carinae weak andjoining anterior and posterior pits. Prosternum twice coxallength. Prosternal process 0.2–0.3 times as broad as coxa.Elytra exposing apex of pygidium, apices obtuse; surfaces withconfuse punctation, epipleura narrow, reaching abdominal ven-trite 1. Scutellum triangular and glabrous. Mesoventrite withpair of pits along the mesoventral suture; coxae separated byless than coxal width. Metaventrite longer than abdominal ven-trite 1; postcoxal lines incomplete and V-shaped. Abdomenwith six ventrites: ventrite 6 partially visible; ventrite 1 longerthan two or three combined; postcoxal lines arcuate and veryclose to coxal cavity. Femora and tibia slender, tarsi three-segmented. Male genitalia: penis guide as long as parameres,acuminate apically; parameres slightly asymetrical and setose;penis very thin and coiled, without basal capsule, spinuloseand membranous at apex. Ovipositor broad and elongate; stylismall and terminal, with simple apical setae; spermathecasimple.

Distribution. Mexico, Baja California; U.S.A., California.

Species included. Carinodulinka baja Slipinski & Tomas-zewska.

Material examined. Carinodulinka baja from Mexico andone unnamed specimens from U.S.A. (CA, Los Angeles Co.Santa Monica, Mts. Topanga SP, Prunus litter, M.S. Caterino& K.J. Hopp, 3–iii–2009, ANIC).

Comments. The type series of C. baja was collected withpitfall traps (Slipinski & Tomaszewska, 2002).

Serangiini Pope

Serangiini Pope, 1962: 627. Type genus Serangium Black-burn, 1889. Sasaji, 1967: 7. Pakaluk et al., 1994: 250.Slipinski & Burckhardt, 2006: 38.

Diagnosis. Body broadly oval and convex; dorsum glabrous,rarely sparsely setose; ventral side and elytral epipleura foveateto receive legs in repose (Fig. 9D); head transverse withlarge, finely facetted eyes (Fig. 6A); anterior border of clypeus

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margined (Fig. 6A); mandible apex simple, antenna with large,flattened one-segmented club (Fig. 6H) in repose, receivedin hypomeral cavity; prosternum elevated and forms large,triangular plate (Fig. 7A); femora distinctly flattened (Fig. 7D);abdomen with five ventrites (Fig. 7E), last ventrite long androunded apically.

Distribution. Delphastus is restricted to the New World andis more diverse in tropical areas, with a few species reachingtemperate or subtropical environments in northern U.S.A. andArgentina. Serangium is a large and diverse genus distributedfrom Africa, Madagascar, through India, the Far East Russiaand Japan to Australia and Oceania, it appears that thereare relatively few species in Africa, with a large number ofdescribed and undescribed species from Asia and Australia.Serangiella is so far known from India, Sri Lanka, China,Vietnam and Japan, and seems to be a specialized predator,as indicated by the peculiar modification of the mouthparts(Fig. 10F). Some species of Delphastus or Serangium havebeen introduced to different countries in biological controlprogrammes.

Key to genera of Serangiini

1 Mandible reduced with external border deeply emarginate(Fig. 10F); antennomere 3 short and asymmetrical (Fig. 10E).Tropical Asia, Japan . . . . . . . . . . . . . . . . . . . Serangiella Chapin– Mandible not reduced, external border broadly curved(Fig. 6F); antennomere 3 elongate, symmetrical (Fig. 6H) orslightly asymmetrical . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2 (1) Pronotum with lateral edges distinctly margined (Fig. 8D);middle and hind tibia angulate externally (Fig. 9A, G); tarsithree-segmented. New World . . . . . . . . . . . . .Delphastus Casey– Pronotum with lateral edges narrowly margined (Fig. 7C);middle and hind tibiae not or slightly angulate externally; tarsithree- or four-segmented (Fig. 7D). Africa, Asia, Australia . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Serangium Blackburn

Serangium Blackburn(Figs 6A–H, 7A–H, 14G–I)

Serangium Blackburn, 1889: 209. Type species Serangiummysticum Blackburn, 1889 (monotypy). – Sicard, 1909:151; Chapin, 1940: 268; Sasaji, 1967: 8, 1971: 52;Miyatake, 1994: 239; Slipinski & Burckhardt, 2006: 39;Slipinski, 2007: 53.

Semichnoodes Weise, 1892: 15. Type species Semichnoodeskunowi Weise, 1892 (monotypy). Synonymized byWeise, 1908: 13.

Catana Chapin, 1940: 266. Type species Catana clauseniChapin, 1940 (original designation). Sasaji, 1967: 8;Gordon, 1977: 209; Miyatake, 1994: 240. Synonymizedby Slipinski & Burckhardt, 2006: 39.

Catanella Miyatake, 1961: 136. Type species Catanella for-mosana Miyatake, 1961 (original designation). Sasaji,1967: 10; Miyatake, 1994: 242. Synonymized bySlipinski & Burckhardt, 2006: 39.

Diagnosis. The Old World Serangium can be distinguishedfrom Serangiella in having normally developed mandibles(Fig. 6F) and antennomere 3 long and often parallel sided(Fig. 6H).

Description. Length 1.0–3.5 mm. Hemispherical, convexand compact; clypeus meets anterior margin of prosternum inrepose; dorsal surface with sparse to dense and uniform finesetae often associated with micropunctures (Fig. 7G). Colouryellowish to black, elytra rarely with colour pattern. Winged.Head transverse, eyes large, glabrous and partially margineddorsally (Fig. 6A). Frontoclypeus (Fig. 6A) as long as anten-nal socket, weakly emarginate around antennal insertions, apexmargined. Subantennal grooves present, extending behind eyelevel (Fig. 6B). Subgenal gland openings not well defined,as shown in Figs 6D, 7H. Gula transverse, surface setose(Fig. 6B). Antenna (Fig. 6E, H) eight- or nine-segmentedwith one-segmented club; antennomere 3 often symmetrical,cylindrical and moderately to strongly elongated, in somespecies asymmetrical, resembling that of Serangiella. Mouth-parts (Fig. 6B): mandible as in Fig. 6F, prostheca present,mola reduced; maxillary palpifer foveate, last maxillary palpelongate conical or barrel shaped (Fig. 6C); submentum andmentum broad, labial palps contiguous, ligula well developed.Pronotum transverse (Fig. 7B), lateral edges smooth with verynarrow marginal lines. Prosternum (Fig. 7A) strongly elevatedand prominent anteriorly, forming a broad triangular plate con-cealing mouthparts from below, anteriorly truncate and deeplynotched laterally to accommodate antennal club. Elytra sur-face smooth, sutural line rarely present; epipleura reaching atleast apical third, with foveae to accommodate apices of midand hind femora. Scutellum triangular (Fig. 6G). Mesoventrite(Fig. 7F) with mesoventral process very short and transverse,prosternal and procoxal rest present. Metaventrite (Fig. 7F)transverse, longer than ventrites 1–4; postcoxal lines com-plete and extending to metanepisternum. Abdomen (Fig. 7E)with five ventrites: ventrites 1 and 5 much longer than ven-tries 2 and 3 combined, ventrite 5 sometimes finely crenulate;postcoxal lines incomplete, reaching lateral margin of ventrite,without accessory lines or pores. Femora (Fig. 7D), particu-larly anterior femur, broad, flattened and closely fitting intodepression on meso- and metaventrite; tibia external borderslightly curved; tarsi three- or four-segmented (Fig. 7D); pre-tarsal claws simple. Male genitalia: parameres short or dis-tinctly reduced with setose apex; penis elongated and curved,with basal capsule weak to absent. Ovipositor triangular, elon-gate with short styli; spermatheca globular with at least onechamber, sometimes with slightly sclerotized basis; membra-nous infundibulum rarely present.

Distribution. Old World.Species included. Serangium adustum (Weise); Serangium

baculum Xiao; Serangium bakeri Chapin; Serangium bellumSlipinski & Burckhardt; Serangium buettikeri Fursch; Seran-gium chapini (Kapur); Serangium cinctum (Weise); Serangiumclauseni (Chapin); Serangium comperi Chapin; Serangiumcoxale Weise; Serangium decorsei Sicard; Serangium dimidia-tum (Chapin); Serangium drepnicum Xiao; Serangium for-mosanum (Miyatake); Serangium giffardi Grandi; Serangium

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A

C E

F HG

B

D

Fig. 6. Serangium sp. (Queensland, Australia): (A) head, dorsal; (B) head, ventral; (C) maxilla; (D) mouthparts detail; (E) antenna detail;(F) mandible, ventral; (G) scutellum; (H) antenna.

glorius Slipinski & Burckhardt; Serangium haleemae Afroze& Haider; Serangium hanamense Hoang; Serangium hirtuosumSlipinski & Burckhardt; Serangium howdenorum Slipinski &Burckhardt; Serangium japonicum Chapin; Serangium kunowi(Weise); Serangium luzonicum Chapin; Serangium lygaeumIablokoff–Khnzorian; Serangium maculigerum Blackburn;Serangium magnomaculatum Fursch; Serangium magnumSlipinski & Burckhardt; Serangium metasternale Chapin;Serangium microscopicum Lea; Serangium montazerii Fursch;Serangium monteithi Slipinski & Burckhardt; Serangium mon-ticola Sicard; Serangium mysticum Blackburn; Serangiumnigrum Slipinski & Burckhardt; Serangium nitidum Slipinski& Burckhardt; Serangium parcesetosum Sicard; Serangiumpunctatum Weise; Serangium punctum Miyatake; Serangiumruficolle Kamiya; Serangium ryukyuense Kamiya; Serangiumsabahense (Sasaji); Serangium sculptum Slipinski &

Burckhardt; Serangium serratum Poorani; Serangium spilo-tum Weise; Serangium toamasinae Sicard; Serangium yamSlipinski & Burckhardt; Serangium yasumatsui (Sasaji).

Material examined. Serangium japonicum; S. spilotum;S. dimidiatum; S. clauseni ; S. parcesetosum; S. kunowi ;S. maculigerum; S. luzonicum; S. bakery ; S. metasternale;S. bellum; S. howdenorum; S. glorius; S. magnum; S. mon-teithi ; S. nitidum; S. sculptum; S. yam; S. hirtuosum;S. nigrum.

Comments. The larva of Serangium has been described byTimofeeva & Hoang (1978) and Slipinski (2007). Poorani(1998) provided a key to species occurring in India. Serangiumclauseni was introduced to Cuba from Indonesia to controlthe citrus blackfly Aleurocanthus woglumi Ashby (Chapin,1940; Gordon, 1977). Serangium spp. is used as a biological

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140 H. E. Escalona and A. Slipinski

A B

C

E

G

D

F

H

Fig. 7. Serangium sp. (Queensland, Australia): (A) prothorax, ventral; (B) pronotum; (C) pronotum detail; (D) middle leg; (E) abdomen;(F) meso–metaventrite; (G) frontoclypeus surface, detail; (H) subgenal gland openings, detail.

control agent of the whitefly Bemisia spp. (Legaspi et al.,1996; Obrycki & Kring, 1998; Poorani, 1998; Ellis et al., 2001;Al–Zyoud & Sengonca, 2004; Al-Zyoud, 2008; Yao et al.,2011).

Delphastus Casey(Figs 8A–F, 9A–G)

Delphastus Casey, 1899: 111. Type species Oeneis pusillusLeConte, 1852 (subsequent designation by Korschefsky,1931). Chapin, 1940: 264; Gordon, 1970c: 357; 1977:209; 1985: 61; 1994: 73; Vandenberg, 2002: 382.

Lioscymnus Champion, 1913: 125. Type species Lioscymnusdiversipes Champion, 1913 (monotypy). Synonymizedby Chapin, 1940: 263.

Diagnosis. Delphastus can be distinguished from Serangiumand Serangiella by its angulate middle and hind tibiae(Fig. 9A, G), three-segmented tarsi and distinctly marginedlateral pronotal edges (Fig. 8D).

Description. Length 0.9–1.6 mm. Hemispherical, convexand compact; clypeus meets anterior margin of prosternumin repose; dorsal surface polished, glabrous or covered withsparse long fine setae, often associated with micropunc-tures. Colour black to light brown, elytra rarely maculate.Winged. Head (Fig. 8A, B) transverse, eyes large, glabrousand margined dorsally. Frontoclypeus (Fig. 8A) as long asantennal socket length, emarginate around antennal inser-tions, lateral sides and apex margined. Subantennal groovespresent. Subgenal gland openings not well defined, closeto mouth cavity as Fig. 8F. Gula (Fig. 8B) transverse and

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well developed, gular sutures parallel, surface setose. Antennanine-segmented with one-segmented club; antennomere 3 oftensymmetrical, cylindrical and moderately to strongly elon-gate. Mouthparts (Fig. 8B, F): mandible triangular, pros-theca present, mola reduced; maxillary palpifer not foveate,last maxillary palpomere subparallel and diagonally truncatedto apex; submentum very narrow, mentum and prementumbroader; labial palps subcontiguous, ligula very short. Prono-tum (Fig. 8D) transverse, disc convex with complete marginallines, lateral edges smooth. Prosternum (Fig. 8E) stronglyelevated and prominent anteriorly, forming a broad trian-gular plate concealing mouthparts from below, deeply andbroadly notched laterally to accommodate antennal funicle andclub. Elytra (Fig. 9C–E) convex, sutural line absent; epipleurareaching apical third, with cavities to accommodate apicesof middle and hind femur. Scutellum triangular. Mesoventrite

(Fig. 8C) with mesoventral process transverse and very short,procoxal rest present. Metaventrite (Fig. 8C) transverse, longerthan abdominal ventrites 1–3; postcoxal lines complete andextending to metanepisternum. Abdomen (Fig. 9B) with fiveventrites; ventrites 1 and 5 each longer than ventrites 2 and 3,respectively, ventrite 5 smooth apically; postcoxal lines arcu-ate and complete, without accessory lines or pores. Femora,especially anterior one, broad, flattened and closely fittinginto depression on meso- and metaventrite; tibiae (Fig. 9A,F, G): mid and hind tibiae externally angulate medially(Fig. 9A, G); tarsi four-segmented; pretarsal claws toothed.Male genitalia: parameres, apex setose, in some species theparameres are strongly reduced. Ovipositor weakly sclerotizedand triangular, styli with at least three apical setae; sper-matheca variable with one or more chambers and sclerotizedinfundibulum.

A B

C

E

D

F

Fig. 8. Delphastus sp. (Alabama, U.S.A.): (A) head, dorsal; (B) head, ventral; (C) meso–metaventrite; (D) pronotum; (E) prothorax, ventral;(F) mouthparts detail.

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142 H. E. Escalona and A. Slipinski

A B

C

F G

D E

Fig. 9. Delphastus sp. (Alabama, U.S.A.): (A) middle leg; (B) abdomen; (C) elytron, internal; (D) elytron, ventral; (E) elytron, lateral; (F) anteriorleg; (G) hindleg.

Distribution. New World.

Species included. Delphastus abditus Gordon; Delphastusamazonensis Gordon; Delphastus anthracinus Gordon; Del-phastus argentinicus Nunenmacher; Delphastus bechyneorumGordon; Delphastus berryi Gordon; Delphastus boothi Gor-don; Delphastus catalinae (Horn); Delphastus celatus Gor-don; Delphastus chapini Gordon; Delphastus collaris Chapin;Delphastus davidsoni Gordon; Delphastus dejavu Gordon;Delphastus diversipes (Champion); Delphastus dubitalis Gor-don; Delphastus ecuadorensis Gordon; Delphastus flavicollisGordon; Delphastus hirtulus (Kirsch); Delphastus huahuaiGordon; Delphastus mexicanus Gordon; Delphastus minu-tus Gordon; Delphastus nebulosus Chapin; Delphastus occi-dentalis Juarez & Zaragoza; Delphastus pallidus (LeConte);Delphastus punctum Gordon; Delphastus pusillus (LeConte);Delphastus quinculus Gordon; Delphastus sonoricus Casey.

Material examined. Delphastus chapini ; D. boothi ;D. ecuadorensis; D. collaris; D. flavicollis; D. anthracinus;D. catalinae; D. nebulosus; D. pallidus; D. pusillus;D. dejavu; D. sonoricus; D. minutus; D. amazonensis;D. davidsoni ; D. mexicanus; D. argentinicus; D. quinculus;D. diversipes.

Comments. Gordon (1994) revised Delphastus and foundthe genus very diverse in the New World tropics, with a fewspecies occurring in U.S.A. and Argentina. Recently, Shock-ley & Vandenberg (2011) transferred the species Bystus hir-tula (Kirsch) (Anamorphinae, Endomychidae) to Delphastus.Delphastus larva was included in a key by Rees et al. (1994)and described by Liu & Stansly (1996).

The following host whiteflies have been recorded forDelphastus: Aleurotrachelus sp., Aleurothrixus sp., Peliussp., Dialeurodes sp., Aleurocanthus sp., Orthezia sp.,

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A

C

E F

H

D

G

B

Fig. 10. Serangiella sp. (Sri Lanka): (A) head, ventral; (B) head, frontal; (C) prothorax, ventral; (D) prosternum; (E) antenna; (F) mandible,ventral; (G) middle tarsi; (H) mouthparts detail.

Trialeurodes sp., Aleurodes sp., Bemisia sp. and Paraley-rodes sp. (Quezada, 1974; Watve & Clower, 1976; Cherry& Dowell, 1979; Meyerdirk et al., 1980; Dowell & Cherry,1981; Hoelmer et al., 1993; Gordon, 1994; Lucas et al., 2004;Simmons & Legaspi, 2004; Liu, 2005; Legaspi et al., 2006;Legaspi et al., 2008; Simmons et al., 2008; Hodek & Honek,2009).

Delphastus catalinae, previously confused with D. pusillus,has been cultured and sold worldwide as a predator ofwhiteflies, particularly Bemisia tabaci (Hoelmer & Pickett,2003; Hoelmer et al., 1993). Zang & Liu (2007) investigatedintraguild interactions involving D. catalinae, and Liu &Stansly (1999) investigated its feeding behaviour.

Serangiella Chapin(Figs 10A–H, 11A–H)

Serangiella Chapin, 1940: 271. Type species Serangiellaflavescens Chapin, 1940 (subsequent designation byHoang, 1977). Miyatake, 1994: 241; Slipinski & Bur-ckhardt, 2006: 49.

Microserangium Miyatake, 1961: 37. Type species Micro-serangium shikokense Miyatake, 1961. Sasaji, 1971: 61;Miyatake, 1994: 241; Poorani, 2000: 45. Synonymizedby Slipinski & Burckhardt, 2006: 49.

Diagnosis. Serangiella can be distinguished from Serangiumand Delphastus by its reduced mandibles (Fig. 10F) and short,asymmetrical antennomere 3 (Fig. 10E).

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144 H. E. Escalona and A. Slipinski

A

C

F G H

D E

B

Fig. 11. Serangiella sp. (Sri Lanka): (A) abdomen; (B) anterior leg; (C) meso–metaventrite; (D) elytron, ventral; (E) middle leg; (F) elytron,lateral; (G) elytron, ventral; (H) scutellum.

Description. Length about 1.0 mm. Hemispherical, convexand compact; clypeus meets anterior margin of prosternum inrepose; dorsal surface polished with sparse long setae, oftenassociated with micropunctures, concentrated anteriorly on ely-tra. Colour dark to light brown. Winged. Head transverse(Fig. 10A), eyes large and glabrous, not margined. Fronto-clypeus emarginate around antennal insertions, lateral sidesmargined. Subantennal grooves present, extending behind eyes.Subgenal gland openings not well defined. Gula (Fig. 10A)transverse and short, surface with scattered setae. Antenna(Fig. 10E) nine-segmented with one-segmented club, anten-nomere 3 strongly asymmetrical, triangular and bearing setaeon its asymmetrical angle, lateral border of last anten-nomere diagonally truncated, apex subtruncated. Mouthparts(Fig. 10A, H): mandible (Fig. 10F) narrow and pointed api-cally, external border deeply emarginate, prostheca present,mola indistinct; maxillary palpifer not foveate, last maxillary

palpomere ovoid; submentum very short, about as wide asmentum; prementum and ligula very short, labial palps con-tiguous. Pronotum (Fig. 10D) transverse, lateral edge smooth,disc convex, lateral sides narrowly margined, line separatinganterior corners joining anterior and lateral margins. Proster-num (Fig. 10C) strongly prominent medially, forming trian-gular plate that conceals mouthparts from below, narrow anddeeply notched laterally to accommodate antennal club. Ely-tra (Fig. 11F, G) usually smooth and polished, sutural lineabsent, epipleura reaching apical third, with cavities to accom-modate apices of middle and hind femora. Scutellum triangular(Fig. 11H). Mesoventral process strongly transverse (Fig. 11C)and very short, prosternal and procoxal rest present. Metaven-trite transverse, longer than abdominal ventrites 1–4 com-bined; postcoxal lines complete, extending to metanepisternum.Abdomen (Fig. 11A) with five ventrites: ventrite 1 or 5 muchlonger than ventrites 2 and 3 combined; ventrite 5 smooth

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posteriorly; postcoxal lines complete, without accessory linesor pores. Femora (Fig. 11B, E), especially anterior femur,flattened and closely fitting into depressions on meso- andmetaventrite; middle and hind tibia (Fig. 11E) angulate exter-nally; tarsi three-segmented or with a fourth partially fusedtarsomere; pretarsal claws simple. Male genitalia: parameresshort with setose apex. Ovipositor weakly sclerotized and nar-rowing to apex, styli strongly reduced; spermatheca with twoor three chambers and sclerotized infundibulum.

Distribution. Vietnam, India, Sri Lanka, China and Japan.

Species included. Serangiella bacthaiensis (Hoang); Serang-iella brunneonigra (Poorani); Serangiella ceylonica (Weise);Serangiella coconiensis (Duverger); Serangiella flavescensChapin; Serangiella hainanensis (Miyatake); Serangiella lat-eralis (Motschulsky); Serangiella okinawensis (Miyatake);Serangiella shikokensis (Miyatake); Serangiella vietnamicaHoang.

Material examined. Serangiella flavescens; Serangiella sp.(Sri Lanka).

Microweiseini Leng

Microweiseini Leng, 1920: 213. Type genus MicroweiseaCockerell, 1903. Gage, 1920: 42 (larva); Korschefsky,1931: 209; Gordon, 1977: 200, 1978: 205; Vandenberg,2002: 382; Slipinski, 2007: 49.

Microweisini Gordon, 1985: 36 (misspelling).

Pharini Casey, 1899: 110. – Pope, 1962: 267 (in part).

Sukunahikonini Kamiya, 1960: 24. Type genus SukunahikonaKamiya, 1960. Sasaji, 1967: 4, 1971: 47; Gordon,1977: 188; Miyatake, 1994: 233; Slipinski & Tomas-zewska, 2005: 370. syn.n.

Diagnosis. Body elongate to oval (Figs 2A, B, 3A–F), con-vex, almost always distinctly pubescent; ventral side (Fig. 13B)and elytral epipleura (Fig. 13D) without foveae to receive legsin repose; mandibular apex simple; antenna seven- to ten-segmented, club with one or more segments, not flattened andnot received on prothoracic hypomeron; prosternum promi-nent anteriorly (Figs 13E, 21E), flat (Fig. 15E) or stronglyreduced in front of coxae (Fig. 16F); abdomen with five orsix ventrites, ventrite 5 not markedly longer than ventrite 4;abdominal postcoxal lines rarely recurved; legs slender; tarsithree- or four-segmented.

Distribution. Most of the genera and species of Microwei-seini occur in the New World. Three genera and three specieshave been described from southern Africa; one of them, Para-coleopterus, is also distributed in the Middle East and theMediterranean region; a monotypic genus Hong has beendescribed from subtropical Australia.

Comments. Microweiseini has been credited historically toCasey (1899), under the preoccupied name Pharini (Pope,1962; Gordon, 1977), but actually Casey’s definition of Phar-ini corresponds to Sticholotidini (Slipinski, 2004, 2007). Casey(1899) studied two genera of current Microweiseini but placed

them in Cranophorini (Nipus Casey) and Scymnini (SmiliaWeise = Microweisea Cockerell). Gordon (1977) found thatMicroweisea and related genera form a natural group, andintroduced Microweiseini into modern Coccinellidae system-atics. This concept has been followed by Fursch (1985),Vandenberg (2002) and Slipinski (2007).

Key to genera of Microweiseini

1 Pronotum arcuate anteriorly covering head from above,anterior angles indistinct (Figs 21G, 22C) . . . . . . . . . . . . . . . . 2– Pronotum emarginate anteriorly, at most covering part ofhead from above, anterior angles distinct (Fig. 23G) . . . . . . .3

2 (1) Prosternum in front of coxa at least four times coxallongitudinal diameter (Fig. 22B); head very long (Fig. 22A),in repose entirely retracted into prothorax (Fig. 3B). Mexico,U.S.A. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Allenius gen.n.– Prosternum in front of coxa much less than four timescoxa length (Fig. 21E); head short (Fig. 21A), in repose notretracted into prothorax. U.S.A. . . . . . . . . . . . . . . . Nipus Casey

3 (1) Prosternum without chin piece, strongly reduced infront of coxae (Figs 16F, 19F), its length less than 0.5 coxallongitudinal diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4– Prosternum often with chin piece, variable but always aboutas long as coxal longitudinal diameter (Fig. 21E) . . . . . . . . . 7

4 (3) Pronotum without line separating anterior corner; anten-nal club three-segmented (Fig. 20B); mesoventrite anteriorlyraised and folding backwards (Fig. 20C). U.S.A. . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gordoneus gen.n.

– Pronotum often with line separating anterior corner(Fig. 19B); antenna variable; mesoventrite flat . . . . . . . . . . . . 5

5 (4) Antenna eight-segmented, with one-segmented antennalclub (Fig. 14J). Australian, Pacific . . . . . Paraphellus Chazeau– Antenna nine- or ten-segmented, with two- or three-segmented antennal club . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

6 (5) Submentum and mentum equally broad (Fig. 19C);mentum not emarginate apically (Fig. 19C); maxillary cardovery narrow, partially visible. Tropical Asia, Australia . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Pharellus Sicard– Submentum notably narrower than mentum (Fig. 16C);mentum deeply emarginate apically (Fig. 16C); maxillarycardo broad and clearly visible. Pantropical . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Scymnomorphus Weise7 (3) Antennal club one-segmented (Fig. 12H); prosternumbroadly prominent anteriorly, with narrow slits laterally toaccommodate antenna (Figs 12D, 13E) . . . . . . . . . . . . . . . . . . . 8– Antennal club multisegmented (Figs 14C, 15C); prosternumflattened (Figs 15E, 20D) or forming a chinpiece with definedlateral sides (Fig. 21E) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

8 (7) Pronotum with line separating anterior corner (Fig. 12G);prosternal carinae present, incomplete (Fig. 12D); submentumpoorly developed, not clearly visible ventrally (Fig. 12A);antenna eight-segmented; abdominal postcoxal lines complete(Fig. 12J). South Africa . . . . . . . . . . . . . . . . Microfreudea Fursh

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146 H. E. Escalona and A. Slipinski

– Pronotum without line separating anterior corner (Fig. 13F);prosternal carinae absent (Fig. 13E); submentum welldeveloped and visible ventrally (Fig. 13A); antenna seven-segmented; abdominal postcoxal lines incomplete (Fig. 13I).South Africa, Mediterranean, Pakistan . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paracoelopterus Normand

9 (7) Prosternum with anterior margin straight, not prominentinto a chinpiece (Figs 14D, 15E) . . . . . . . . . . . . . . . . . . . . . . . 10– Prosternum anterior margin with distinct chinpiece (Figs 21E,22B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

10 (9) Terminal maxillary palp knife-like (cultriform)(Fig. 15A); antennal club two-segmented (Fig. 15C); proster-num well developed in front of coxa without posterior chin-piece (Fig. 15E); pronotal and elytral margins not serrate.South Africa . . . . . . . . . . . . . . . . . . . . . . . . . . . Cathedrana gen.n.– Terminal maxillary palpomere elongate and conical(Fig. 14B); antennal club three-segmented (14C); prosternumwith chinpiece posterior to prosternal margin, forming a ver-tical shelf (Fig. 14D); pronotal (Fig. 14D) and elytral marginsfinely serrate. Australia . . . . . . . . . . . . . . . . . . . . .Hong Slipinski

11 (9) Tarsal formula 4-4-4; prementum relatively broad(Fig. 26A). Chile . . . . . . . . . . . . . . . . . . . . . . . . Parasidis Brethes– Tarsal formula 3-3-3; prementum very narrow (Fig. 24B) . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

12 (11) Antenna nine-segmented, antennal club two-segmented(Fig. 24F). New World . . . . . . . . . . . . . . Coccidophilus Brethes– Antenna ten-segmented, antennal club three-segmented(Fig. 27C) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

13 (12) Dorsum at most with few scattered short to minutesetae (Fig. 2H). U.S.A. and Mexico . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Microweisea Cockerell– Dorsum covered with long fine setae (Fig. 2A) . . . . . . . . 14

14 (13) Maxillary palp slightly broader at middle (Fig. 27B).Chile . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stictospilus Brethes– Maxillary palp conical. Intercepted in U.S.A. . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Microcapillata Gordon

Microfreudea Fursch(Figs 2G, 12A–J)

Microfreudea Fursch, 1985: 292. Type species Microfreudeacyclica Fursch, 1985 (original designation).

Diagnosis. Paracoleopterus and Microfreudea are verysimilar and share setose, compact body, short antenna withone-segmented club and the prosternum projected anteriorly(Fig. 12D), but not forming a chinpiece. Microfreudea canbe distinguished from Paracoleopterus in having anteriorpronotal corners separated from the disc by a line (Fig. 12G),well-developed prosternal carinae (Fig. 12D) and metaventralpostcoxal lines recurved (Fig. 12I).

Description. Length 1.0–1.2 mm. Body ovoid and con-vex, dorsal surfaces finely and densely punctured and setose.Colour black. Winged. Head transverse; eyes large, glabrous

not margined. Frontoclypeus about 1.5× as long as antennalsocket length, lateral sides margined, well emarginate aroundantennal insertions. Subantennal grooves extending behindeye level. Subgenal gland openings as Fig. 12C. Gula short,transverse, gular sutures reduced, surface setose. Antenna(Fig. 12H) eight-segmented, club large and one-segmented.Mouthparts (Figs 12A,B): mandible with prostheca, molareduced; maxillary palpifer foveate, terminal maxillary palpconical, apically pointed; submentum strongly reduced, men-tum shield-like, prementum well developed, labial palps nar-rowly separated, ligula large, broadly rounded and setose.Pronotum (Fig. 12G) transverse, lateral sides narrowly margi-ned, disc convex with line separating anterior corners frompronotal disc; lateral edges smooth. Prosternum (Fig. 12D)prominent, forming a broadly rounded anterior lobe that con-ceals mouthparts from below, notched laterally to accommo-date antennal club; prosternal process slightly narrower thancoxa; prosternal carinae diverging anteriorly reaching mid-dle of prosternum. Elytral surfaces with dense fine punctures(Fig. 12E), sutural line absent; epipleura reaching apical third.Scutellum triangular. Mesoventrite (Fig. 12I), mesoventral pro-cess twice as wide as mesocoxa; procoxal and prosternal restpresent. Metaventrite (Fig. 12I) transverse, 1.2× longer thanabdominal ventrite 1; postcoxal lines recurved and complete;meso–metaventral junction straight. Abdomen (Fig. 12J) withfive ventrites, in some specimens ventrite 6 barely visible, sur-face with coarse punctures; ventrite 1 as long as ventrites 2–4combined, ventrite 5 longer than ventrite 4, rounded apically;postcoxal lines arcuate and complete, without accessory line.Legs slender, tarsi three-segmented; pretarsal claws weaklytoothed. Male genitalia: penis guide long, triangular; parameresshort and strongly asymmetrical; penis with basal capsule.

Distribution. Cape Province, South Africa.

Species included. Microfreudea cyclica Fursch.

Material examined. Microfreudea cyclica.

Paracoelopterus Normand(Figs 3H, 13A–I)

Paracoelopterus Normand, 1936: 97. Type species Para-coelopterus rufinus Normand, 1936 (monotypy).

Diloponis Pope, 1962: 633. Type species Diloponis incon-spicuus Pope, 1962 (original designation). Fursch1985: 289; Ahmadi & Yazdani 1993: 30. Synonymizedby Kovar, 2007: 72.

Diagnosis. Paracoleopterus is very similar to Microfreudea,but can be distinguished from that genus in having seven-segmented antenna, anterior pronotal corners not separatedfrom the disc by a line (Fig. 13F), prosternal carinae absent(Fig. 13E) and metaventral postcoxal lines distinctly descend-ing (Fig. 13B).

Description. Length 1.0–1.3 mm. Body ovoid to rounded,convex, dorsal surfaces setose with associated irregular microp-unctures and punctures. Colour light to dark brown. Winged.Head (Fig. 13A, G) transverse, eyes large, with fine interfacetal

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Generic revision and phylogeny of Microweiseinae 147

A

D

G

I

J

H

EF

B C

Fig. 12. Microfreudea sp. (South Africa): (A) head, ventral; (B) mouthparts detail; (C) head detail; (D) prothorax, ventral; (E) elytron dorsalsurface, detail; (F) antenna detail; (G) pronotum; (H) antenna; (I) meso–metaventrite; (J) abdomen.

setae, not margined dorsally. Frontoclypeus (Fig. 13G) as longas antennal socket length, emarginate around antennal inser-tions, lateral sides weakly margined. Subantennal grooves welldeveloped. Subgenal gland openings present. Gula (Fig. 13A)short, transverse and triangular, surface with few sparse setae.Antenna seven-segmented, with one-segmented club. Mouth-parts: mandibles narrow and elongated, prostheca present, molareduced; maxillary palpifer at least partially foveolate, terminalmaxillary palpomere parallel to conical (Fig. 13A); submen-tum broad, mentum elongate, prementum very short, labialpalps narrowly separated, ligula well developed, setose toapex. Pronotum (Fig. 13F) transverse, disc convex with lat-eral sides deeply margined, lateral edges smooth; anterior cor-ners not separated from disc. Prosternum (Fig. 13E) promi-nent, forming a large and broadly rounded plate, marginedand deeply notched laterally; prosternal process about 0.6×as wide as coxa; hypomeron with weak or deep C-shapedgroove near anterior corners. Elytral (Fig. 13C, D) surface

smooth and densely punctured, sutural line absent; epipleura(Fig. 13D) reaching apical fourth and setose. Scutellum trian-gular. Mesoventral process transverse (Fig. 13B), wider thancoxa, anterior edge raised; prosternal and procoxal rests present.Metaventrite (Fig. 13B) transverse, 1.2× longer than abdom-inal ventrite 1, mesometaventral junction partially fused andstraight, postcoxal lines separated at middle, descending andcontinuing along lateral margin. Abdomen (Fig. 13I) with fiveventrites, in male ventrite 6 barely visible, ventrite 1 as longas ventrites 2–3 combined; postcoxal lines arcuate and incom-plete, accessory line absent. Legs slender; tarsi four-segmented;pretarsal claws simple. Male genitalia: parameres distinct, oftenslightly asymmetrical; penis curved, basal capsule weakly dif-ferentiate. Female genitalia: bursa copulatrix with infundibu-lum; spermatheca sides rounded.

Distribution. Mediterranean region, Afghanistan, Iran andSouth Africa.

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148 H. E. Escalona and A. Slipinski

Species included. Paracoleopterus rufinus Normand;Paracoleopterus berytensis (Weise); Paracoleopterus fuerschiAhmadi & Yazdani; Paracoleopterus inconspicuus (Pope).

Material examined. Paracoleopterus berytensis ; Paracol-eopterus inconspicuus.

Comments. Paracoleopterus inconspicuus and Paracol-eopterus fuerschi feed on citrus scales (Brettell, 1964; Ahmadi& Yazdani, 1993).

Hong Slipinski(Figs 3E, F, 14A–E)

Hong Slipinski, 2007: 50. Type species Hong gloriousSlipinski, 2007 (original designation).

Diagnosis. This genus is externally similar to Cathedranafrom South Africa, but can be easily recognized by its crenulate

pronotal and elytral edges, three-segmented antennal club(Fig. 14C), shelf-like prosternal chinpiece (Fig. 14D) and four-segmented tarsi.

Description. Length about 1.2 mm. Body elongate, oval andconvex (Fig. 3E, F), setose; dorsal surface polished, clothedwith long fine setae. Colour brown. Winged. Head elongate(Fig. 14B); eyes with coarse facets, glabrous, without dorsalmarginal line. Frontoclypeus (Fig. 14B) more than three timesas long as antennal socket length, emarginate around anten-nal insertions, lateral sides margined. Subantennal groovesindistinct. Subgenal gland openings present. Gula longer thanwide, with few apical setae. Antenna ten-segmented with three-segmented club (Fig. 14C). Mouthparts: maxillary palpifernot foveate, terminal maxillary palpomere elongate and nar-rowing apically (Fig. 14B); submentum and mentum verynarrow, mentum longer than submentum and prementum;

A

C

G H I

D

E

F

B

Fig. 13. Paracoelopterus sp. (Turkey): (A) head, ventral; (B) meso–metaventrite; (C) elytron, lateral; (D) elytron, ventral; (E) prothorax, ventral;(F) pronotum; (G) head, dorsal; (H) abdominal postcoxal line detail; (I) abdomen.

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Generic revision and phylogeny of Microweiseinae 149

A BC

D

G

FE

H I

J

K

L

M

Fig. 14. Hong glorius, ♀: (A) abdomen; (B) head, dorsal; (C) antenna; (D) prothorax, ventral; (E) meso–metaventrite; (F) abdominal postcoxalline; (G) Serangium maculigerum, spermatheca; (H) Serangium magnum, tegmen; (I) S. magnum, penis; (J) Paraphellus rostratus, antenna;(K) Scymnomorphus sp. (Australia), spermatheca; (L) Scymnomorphus luteus, penis; (M) Scymnomorphus sp. (Australia), tegmen.

labial palps narrowly separated, ligula not observed. Pronotum(Fig. 3F) transverse; anterior margin arcuate, without lineseparating anterior corners from pronotal disc, lateral edgesweakly crenulate. Prosternum (Fig. 14D) with shelf-like, pos-teriorly located chinpiece; prosternal process about 0.6× aswide as coxa; prosternal carinae absent. Elytra (Fig. 3F) withcoarse irregular punctures, interspaces smooth, lateral sidesfinely crenulate; epipleura reaching apical third. Scutellum tri-angular. Mesoventrite (Fig. 14E) transverse and very short,about 0.8× as wide as mesocoxa; procoxal and proster-nal rest present; mesometaventral junction arcuate anteriorly.

Metaventrite (Fig. 14E) transverse, about as long as abdominalventrite 1; postcoxal lines recurved and complete. Abdomen(Fig. 14A, F) with five ventrites, ventrite 1 longer than ven-trites 2 and 3 combined; postcoxal lines short descending withaccessory oblique line. Legs slender, tarsi four-segmented;pretarsal claws simple. Female genitalia: coxites triangularand elongate, styli terminal; infundibulum absent; spermathecamulticameral and sclerotized. Male unknown.

Distribution. South Queensland, Australia.

Species included. Hong glorious Slipinski.

Material examined. Hong glorious (holotype, ♀).

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150 H. E. Escalona and A. Slipinski

AB

C D

F

E

G H I

Fig. 15. Cathedrana natalensis, ♂ (South Africa): (A) head, ventral; (B) mouthparts detail; (C) antenna; (D) elytron, ventral; (E) prothorax, ventral;(F) meso–metaventrite; (G) abdomen; (H) abdominal postcoxal line; (I) abdominal surface, detail.

Cathedrana gen.n.(Figs 2B, 3I, 15A–I, 28L–N)

Type species. Cathedrana natalensis sp.n.

Diagnosis. Cathedrana gen.n. is a very distinctive memberof Microweiseini, resembling the genus Hong from Australiaand an unnamed taxon from Chile, with a relatively longprosternum not forming a chinpiece (Fig. 15E) and a knife-like(cultriform) maxillary palp.

Description. Length about 1.1 mm. Colour orange to yel-lowish. Body 1.6× longer than wide, sides subparallel(Fig. 2B). Dorsum covered with fine long setae and irregularcoarse punctures; venter with coarse punctures and associatedfine setae (Fig. 15I). Head about as long as wide, rostrate(Fig. 15A). Subantennal grooves short. Subgenal gland open-ings as Fig. 15A, B. Gula (Fig. 15A) well developed, about

as long as wide, gular sutures short. Eyes reduced, coarselyfacetted and glabrous. Frontoclypeus, twice or more the lengthof antennal socket, emarginate around antennal insertions, lat-eral sides weakly margined. Temple longer than eye length.Occipital region without endocarina. Antenna (Fig. 15C) nine-segmented, with two-segmented club. Mandible not exam-ined. Maxilla with palpifer foveate; last palpomere elongateand knife-like (cultriform) (Fig. 15A). Submentum about asbroad as mentum, mentum long and broad (Fig. 15B); pre-mentum subquadrate, labial palps narrowly separated, ligulabroadly rounded and setose. Pronotum transverse, anteriorangles obtuse, posterior angles acute; disc convex withoutline separating anterior corners from pronotal disc. Prosternumin front of coxae slightly shorter than coxa; prosternal pro-cess (Fig. 15E) broad and subparallel, apex rounded. Meso-and metaventrite (Fig. 15F) not foveate for legs reception.Scutellum abruptly elevated, longer than wide, apex rounded.

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Generic revision and phylogeny of Microweiseinae 151

Mesoventrite more or less at same level, at middle longer thancoxa; procoxal and prosternal rest present; mesoventral pro-cess flattened, about as wide as coxa. Elytra subparallel withapical third, widest near middle; lateral margins mostly visiblefrom above; epipleura narrow (Fig. 15D), reaching abdominalventrite 4 and setose. Wings present, veins reduced. Metaven-trite (Fig. 15F) transverse, slightly shorter than abdominal ven-trite 1; postcoxal lines descending and complete. Abdomen(Fig. 15G) with six ventrites, ventrite 6 visible apically; ven-trite 1 slightly longer than ventrites 2–4 combined; postcoxallines short, arcuate and incomplete (Fig. 15H), with accessoryline and pores. Legs slender; tarsi four-segmented; pretarsalclaws simple. Male terminalia: spiculum gastrale Y-shaped,with doubled rod at base. Genitalia: tegmen as Fig. 28M,N; parameres well developed, longer than penis guide, withdisperse setae (Fig. 28N); penis curved, basal capsule barelydifferentiate (Fig. 28L).

Distribution. KwaZulu Natal, South Africa.

Etymology. Named after the type species locality, CathedralPeak State Forest, South Africa. Gender feminine.

Species included. Cathedrana natalensis sp.n.

Comments. The types were collected in Podocarpus forest.

Cathedrana natalensis sp.n.(Figs 2B, 3I, 15A–I, 28L–N)

Description. Length: 1.1 mm. Width: 0.6 mm. Colour(Fig. 2B) light orange to yellowish; dorsal surfaces clothedwith long, dense, fine setae and coarse punctures. Head(Fig. 15A) as long as wide; dorsal surface mostly microretic-ulate. Eye reduced to about 50 coarse facets, dorsally about0.2× as long as head capsule length; interocular distance about3.6× eye diameter; temple about 2× eye length. Maxillary

A

C

E

B

D

F

G

Fig. 16. Scymnomorphus sp., ♀ (Cameroon): (A) head, dorsal; (B) head, ventral; (C) mouthparts detail; (D) meso–metaventrite; (E) abdomen;(F) prothorax, ventral; (G) female coxites, detail.

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152 H. E. Escalona and A. Slipinski

A

C

F G H

D

E

B

Fig. 17. Scymnomorphus castaneus: (A) meso–metaventrite; (B) head ventral; (C) prothorax, ventral; (D) anterior leg; (E) mouthparts detail;(F) elytron, lateral view; (G) elytron, ventral view; (H) abdomen.

palpomere 4 about 3.4× longer than palpomere 3 (Fig. 15A).Pronotum transverse (Fig. 15E), 0.6× as long as wide, widestat base; lateral edges smooth; disc densely punctured, punc-tures smaller than those on elytra; prosternum with coarseirregular punctures. Elytral surfaces smooth and polishedwith dense punctures, coarse, sparse and irregular. Metaven-trite with coarse punctures, smaller on disc. Abdominal ven-trite 1 about 2.6× as long as ventrite 2 (Fig. 15G, H), withcoarse punctures between postcoxal lines. Male genitalia asin Fig. 28L–N; parameres conical, in lateral view narrow-ing to apex and setose, penis guide in lateral view nar-rowing to apex. Penis as Fig. 28L. Female genitalia: notexamined.

Etymology. Named after KwaZulu Natal Province, SouthAfrica.

Type material. Holotype: ♀, SOUTH AFRICA, Natal, 75 kmWSW Estcourt, Cathedral Peaks For. Sta., 7–31-xii-1979,Rainbow Gorge, Podocarp For. 1600 m, malaise traps, 8–13-xii-1979 (S. & J. Peck ) (NHML). (Specimen bears two labels.)

Paratypes: as holotype, except 1 ♀, 1500 m, malaise trough,7–31-xii-1979 (NHML), 1 ♂, in part coated for SEM (ANIC);as holotype, except 1 ♀, 1500 m, veg. sweeping, 17-xii-1979 (ANIC).

Scymnomorphus Weise(Figs 2E, 14K–M, 16A–G, 17A–H, 18A, B, D, E)

Scymnomorphus Weise, 1897: 303. Type species Scymnomor-phus rotundatus Weise, 1897 (subsequent designationby Pope, 1962). Slipinski & Tomaszewska, 2005: 380.Fursch, 2006: 116.

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Generic revision and phylogeny of Microweiseinae 153

A

B

D

C

E

F

G H

Fig. 18. A, B, Scymnomorphus hirtus: (A) meso–metaventrite; (B) mouthparts detail. C, Paraphellus rostratus: abdominal postcoxal lines.D, Symnomorphus fulvus: head and pronotum lateral view. E, Scymnomorphus hirtus: prosternal process. F–H, Paraphellus rostratus: (F) prosternalprocess; (G) habitus; (H) head dorsal view.

Scotoscymnus Weise, 1901: 458 (unnecessary replacementname). – Gordon, 1977: 189; Fursch, 1985: 283; Miy-atake, 1994: 235.

Sukunahikona Kamiya, 1960: 22. Type species Sukunahikonajaponica Kamiya, 1960 (original designation). Syn-onymized by Fursch, 1985: 283.

Hikonasukuna Sasaji, 1967: 4. Type species Hikonasukunamonticola Sasaji, 1967. Fursch, 1985: 287 syn.n.

Orculus Sicard, 1931: 233. Type species Orculus castaneusSicard, 1931 (monotypy). Fursch, 1985: 289 syn.n.

Diagnosis. Scymnomorphus, Pharellus and Paraphellusshare a strongly reduced prosternum, but Scymnomorphusdiffers from those genera by submentum distinctly narrowerthan the mentum (Fig. 16B), mentum deeply emarginate

apically (Figs 17E, 18B), maxillary cardo broad and clearlyvisible externally and multisegmented antennal club.

Description. Maximum length 1.5 mm. Convex, lateralsides of elytra explanate in some species, dorsal surfaceswith long setae or mixed short setae. Colour from lightbrown to black. Usually winged. Head transverse (Figs 16A,17B), often rostrate; eyes large and glabrous. Frontoclypeus(Fig. 16A) variable in length usually longer than one antennalsocket length, emarginate around antennal insertions, lateralsides partially margined. Subantennal grooves short. Subgenalgland openings as in Fig. 17E. Gula (Figs 16B, 17B) aslong as mouthparts cavity, subtriangular, wider than longwith sparse setae. Antenna nine- or ten-segmented, withtwo- or three-segmented club, antennomere 7 sometimesconsiderably smaller. Mouthparts (Figs 16B, 17B, E, 18B):

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154 H. E. Escalona and A. Slipinski

mandibular mola reduced, mesal edge with prominence nearapical third, prostheca well developed; maxillary palpiferpartially excavated for maxillary palp reception, last maxillarypalpomere elongate and conical, apex diagonally truncate(Figs 16B, 17B); submentum very narrow (Fig. 16B) to absent(Fig. 17B), mentum broader and longer than submentum, apexdeeply emarginate to accommodate prementum (Figs 17E,18B); prementum short, labial palps narrowly separated, ligulawith minute setae. Pronotum transverse, lateral edge smooth,disc convex with distinct line or ridge separating anteriorcorners from pronotal disc (Fig. 18D), sometimes extendingalong lateral edge. Prosternum (Figs 16F, 17C) reduced infront of coxae; prosternal process narrow, often forms carinabetween coxae (Figs 16F, 17C, 18E). Elytra (Fig. 17F, G):punctures at least forming an irregular row along suture;epipleura narrow or very broad and incomplete; lateral part ofelytron often with a submarginal carina from humeral angle,parallel or very close to lateral margin and extending to apicalthird. Scutellum triangular. Mesoventral process (Figs 16D,17A, 18A) transverse, short and elevated anteriorly; mesocoxaseparated by more or less a coxa width; procoxal and prosternalrest weakly differentiate. Metaventrite transverse, longer thanabdominal ventrite 1; postcoxal lines descending, completeor incomplete. Abdomen (Figs 16E, 17H) with six ventrites,ventrite 6 partially visible; ventrites 1 and 2 completelyto partially fused, ventrite 1 as long as ventrites 2 and 3combined, ventrite 5 with apex slightly truncate; postcoxallines arcuate, incomplete or complete, with accessory line oftenpresent. Femora slender to robust, tibia slender (Fig. 17D),tarsi three- or four-segmented; pretarsal claws barely toothed.Male genitalia (Figs 14L,M): parameres reduced, penis guideasymmetrical and elongated, penis long, slender and curved,basal capsule evident. Ovipositor cylindrical, flattened, stylielongated with apical setae (Fig. 16G), spermatheca bulb-like,constricted medially.

Distribution. Pantropical with large number of species stillundescribed.

Species included. Scymnomorphus castaneus (Sicard)comb.n.; Scymnomorphus monticola (Sasaji) comb.n.; Scym-nomorphus elongatus (Gordon); Scymnomorphus globosus(Gordon); Scymnomorphus orchidion (Gordon); Scymnomor-phus facetus (Gordon); Scymnomorphus apterus (Gordon);Scymnomorphus pygmaeus (Gordon); Scymnomorphus step-hensi (Gordon); Scymnomorphus perpusillus (Gordon);Scymnomorphus colombianus (Gordon); Scymnomorphusrotundatus Weise; Scymnomorphus punctipennis Weise; Scym-nomorphus parvulus Weise; Scymnomorphus niger Weise;Scymnomorphus fulvus Slipinski & Tomaszewska; Scym-nomorphus luteus Slipinski & Tomaszewska; Scymnomor-phus hirtus Slipinski & Tomaszewska; Scymnomorphus storeyiSlipinski & Tomaszewska; Scymnomorphus ker Slipinski &Tomaszewska; Scymnomorphus australis (Chazeau); Scym-nomorphus zeijsti (Chazeau); Scymnomorphus prapawan(Chazeau); Scymnomorphus japonicus (Reitter); Scymnomor-phus bicolor (Kamiya); Scymnomorphus japonicus (Kamiya);Scymnomorphus popei (Vazirani); Scymnomorphus minutus

(Fursch); Scymnomorphus glabripilosus (Fursch); Scymnomor-phus maximus (Fursch); Scymnomorphus punctatissimus(Fursch); Scymnomorphus wagneri (Fursch); Scymnomorphustriangularis (Gordon & Almeida).

Material examined. Scymnomorphus pygmaeus; S. fulvus;S. luteus; S. hirtus; S. storeyi ; S. ker ; S. japonicus; S. bicolor ;S. castaneus; S. stephensi ; S. colombianus.

Comments. Scymnomorphus larva have been described byKamiya (1965), Sasaji (1968a) and commented by Slipinski(2007).

Gordon (1977) described several Neotropical species(S. apterous, S. facetus, S. orchidion, S. globosus andS. elongatus) with nine-segmented antenna and one-segmentedantennal club, suggesting relationships with Paraphellus. How-ever, detailed observations of antenna of S. elongatus revealedtwo-segmented club but with the penultimate segment dis-proportionally small (as illustrated in Gordon, 1977). Sicard(1931) and Fursch (1985) erroneously recorded 11-segmentedantennae for S. castaneus.

Host records include Aulacaspis sp., Chrysomphalus aonid-ium (Linnaeus), Aspidotus sp., Pseudaulacaspis sp., Unaspissp. and other Diaspididae (Kamiya, 1960, 1966; Chazeau,1975, 1979; Vazirani, 1982; Slipinski, 2007).

Pharellus Sicard(Figs 19A–G, 20G)

Pharellus Sicard, 1928: 300. Type species Pharellus minuti-simus Sicard, 1928 (monotypy). Chapin, 1965: 228; Chazeau,1977: 216; Hoang, 1982: 79; Miyatake, 1994: 237; Slipinski& Tomaszewska, 2005: 370.

Diagnosis. Pharellus differs from Scymnomorphus in hav-ing submentum and mentum equally broad, mentum notemarginate anteriorly and strongly reduced externally max-illary cardo (Figs 19C, 20G), and from Paraphellus by themultisegmented antennal club.

Description. Maximum length 1.3 mm. Convex; head inrepose strongly deflexed; dorsal vestiture double, consistingof sparse long setae mixed with fine minute hairs. Colourusually dark brown. Winged. Head transverse (Fig. 19D) orabout as long as wide, often rostrate; eyes large and glabrous.Frontoclypeus (Fig. 19D) up to four times antennal socketlength, slightly emarginate around antennal insertions, weaklymargined laterally. Subantennal grooves short. Subgenal glandopenings as Fig. 19C. Gula as long as the mouth cavity andabout as long as wide. Antenna nine- (P. glabratus) or ten-segmented with three-segmented club. Mouthparts (Figs 19C,20G): mandibular molar surface reduced, prostheca welldeveloped; maxillary cardo very small (Fig. 19C); maxillarypalpifer not foveate for maxillary palp reception, terminalmaxillary palpomere elongate and conical, apex narrow anddiagonally truncate; mentum broad and long, as wide assubmentum, submentum short, ligula about as long as labialpalps. Pronotum transverse, lateral edge smooth, disc convex;anterior corners separated from pronotal disc by distinctline or ridge. Prosternum strongly reduced and incomplete,

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AB

DC

E F G

Fig. 19. A–D, Pharellus popei : (A) lateral view; (B) head and pronotum detail; (C) head ventral; (D) head dorsal. E–G, Pharellus glabratus:(E) abdomen; (F) prosternal process; (G) meso–metaventrite.

prosternal process parallel or somewhat triangular (Fig. 19F).Elytra with irregular setose punctuation; epipleura narrowand incomplete. Scutellum triangular. Mesoventrite transverse,anterior margin raised; mesocoxa separated by more or lessa coxa width; mesometaventral junction (Fig. 19G) arcuatedanteriorly, procoxal and prosternal rest present. Metaventritetransverse, longer than ventrites 1–2; postcoxal lines complete,recurved. Abdomen (Fig. 19E) with six ventrites, ventrite 6partially visible; ventrite 1 as long as ventrites 2 and 3combined; postcoxal lines arcuated reaching ventrite apex, andvery close to lateral edge, with or without short accessory line.Femora slightly robust, tibiae slender, tarsi four-segmented.Male genitalia: parameres absent to highly reduced; penis guideasymmetrical, long and narrowing to apex; penis long, slenderwith evident basal capsule. Ovipositor triangular, styli elongatewith simple apical setae; spermatheca worm like.

Distribution. Oriental and Australian.

Species included. Pharellus thanhhoaensis Hoang; Pharel-lus minutisimus Sicard; Pharellus popei Slipinski & Tomas-zewska; Pharellus glabratus Slipinski & Tomaszewska;Pharellus villosus Chapin; Pharellus setosus Chapin; Pharellusyapensis Chapin; Pharellus calvus Chapin; Pharellus ponapen-sis Chapin; Pharellus dybasi Chapin; Pharellus palauensisChapin; Pharellus guamencis Chapin.

Material examined. Pharellus popei ; P. glabratus;P. villosus; P. setosus; P. yapensis; P. calvus; P. ponapensis;P. dybasi ; P. palauensis; P. guamencis.

Comments. Pharellus was synonymized with Scotoscymnusby Gordon (1977), but it was recognized as valid by Chazeau(1977) and Slipinski & Tomaszewska (2005).

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156 H. E. Escalona and A. Slipinski

A

B C

ED

G H

F

Fig. 20. A–F, Gordoneus schwarzi : (A) head, dorsal; (B) antenna; (C) meso–metaventrite; (D) prothorax, ventral; (E) elytron, lateral;(F) abdomen. G, Pharellus glabratus, head ventral. H, Paraphellus rostratus, pronotum detail.

Paraphellus Chazeau(Figs 14J, 18C, F–H, 20H)

Paraphellus Chazeau, 1981: 119. Type species Paraphel-lus pacificus Chazeau, 1981 (original designation). Fursch,1985: 288; Miyatake, 1994: 237; Slipinski & Tomaszewska,2005: 379.

Diagnosis. In size and general appearance, Paraphellusresembles Scymnomorphus and Pharellus, but is distinguishedeasily from these genera by its eight-segmented antenna withlarge one-segmented antennal club (Fig. 14J).

Description. Maximum length 1.5 mm. Convex (Fig. 18G),dorsal surfaces with uniform sparse setae. Colour usually darkbrown. Winged or wingless. Head (Fig. 18H) slightly elongateand rostrate; eyes large and glabrous. Frontoclypeus (Fig. 18H)about twice as long as antennal socket length, emarginatearound antennal insertions, lateral sides weakly margined. Sub-antennal grooves shallow. Subgenal gland openings present.

Gula longer than wide. Antenna eight-segmented with largeone-segmented club (Fig. 14J). Mouthparts: mandible withmolar surface reduced, prostheca well developed; maxillarypalpifer not foveate for maxillary palp reception, last maxillarypalpomere elongate and conical; submentum about as broad asmentum, labial palps narrowly separated, ligula well devel-oped. Pronotum transverse, lateral edge smooth, disc convex;without distinct line separating anterior corners from pronotaldisc. Prosternum strongly reduced and incomplete, prosternalprocess (Fig. 18F) narrow and parallel between coxa. Elytra(Fig. 18G) with irregular and dense punctuation, without lateralcarina; epipleura narrow and incomplete. Scutellum triangu-lar. Mesoventrite transverse, anterior margin raised; mesocoxaseparated by more or less a coxa width; mesometaventraljunction truncated anteriorly, procoxal rest present, prosternalrest weakly demarcate. Metaventrite transverse, longer thanabdominal ventrite 1, postcoxal lines incomplete, very close tolateral margins. Abdomen with six ventrites, ventrite 6 partially

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visible, ventrite 1 longer than ventrites 2 and 3 combined;postcoxal lines arcuated and incomplete, with accessory addi-tional lines and pores (Fig. 18C). Femora slightly robust, tibiaeslender, tarsi four-segmented. Male genitalia highly variable,parameres well developed or reduced, penis guide asymmet-rical and narrow to apex; penis very long and slender, withbasal capsule evident. Ovipositor triangular, styli elongate withsimple apical setae; spermatheca worm like.

Distribution. Pacific, Australian.

Species included. Paraphellus magnopunctatus Slipinski& Tomaszewska; Paraphellus rostratus (Lea); Paraphelluspacificus Chazeau.

Material examined. Paraphellus magnopunctatus; P. rost-ratus.

Gordoneus gen.n.(Figs 3C, D, J, 20A–F)

Type species. Gnathoweisea schwarzi Gordon, 1970a.

Diagnosis. The single species of this genus resemblesrostrate species of Microweisea but differs in having anteriormargin of prosternum without chinpiece (Fig. 20D), thepronotum without line separating anterior corners and theanterior margin of mesoventrite raised and folded backwards(Fig. 20C).

Description. Length about 1.0 mm. Body elongate, ovoidand convex (Fig. 3C, D); glabrous. Colour dark to light brown.Winged. Head about 1.3× as long as wide, rostrate, eyes large,glabrous, not margined dorsally. Frontoclypeus (Fig. 20A) aslong as 2.5× antennal socket length, lateral sides marginated,emarginate around antennal insertions. Subantennal groovesshort and shallow. Subgenal gland openings present. Gulalonger than wide, surface setose. Antenna nine-segmentedwith three-segmented club (Fig. 20B). Mouthparts: mandiblenarrow, prostheca well developed, mola reduced; maxillarypalpifer not foveate, terminal maxillary palpomere elongate,apex diagonally truncate; submentum very narrow, carinalike, mentum elongate, prementum as long as labial palps;labial palps narrowly separated, ligula membranous and setose.Pronotum transverse, disc convex without line separating ante-rior corners, lateral edges smooth. Prosternum (Fig. 20D) veryshort in front of coxae, about 0.4× coxa length, without chin-piece; prosternal process about 0.3× as wide as coxa surfaceflattened. Elytral (Fig. 20E) surface smooth and polished, withdense irregular punctures, sutural line absent; epipleura reach-ing apical third. Scutellum triangular. Mesoventrite (Fig. 20C)transverse and very short, anterior margin raised and foldingbackwards forming a shelf; mesometaventral junction trun-cate anteriorly, procoxal and prosternal rest present. Metaven-trite (Fig. 20C) transverse, about as long as abdominal ven-trite 1; postcoxal lines descending, extending to apical fourthof metaventrite. Abdomen with six ventrites (Fig. 20F), ven-trite 6 partially visible; ventrite 1 as long as ventrites 2 and3 combined; postcoxal line diagonal and incomplete; acces-sory line oblique and close to coxa. Legs slender, tarsi three-segmented; pretarsal claws simple. Male genitalia: penis guide

longer than parameres; parameres strongly asymmetrical; penislong with basal capsule.

Distribution. U.S.A., Arizona.

Species included. Gordoneus schwarzi (Gordon) comb.n.

Material examined. Gordoneus schwarzi, type series.

Etymology. We dedicate this genus to Dr. Robert Gordon(U.S.A.) in recognition for his pioneering work on Microwei-seinae systematics. Gender masculine.

Nipus Casey(Fig. 21A–G)

Nipus Casey, 1899: 132. Type species Nipus biplagia-tus Casey, 1899 (subsequent designation by Korschefsky,1931). Gordon, 1970d: 72, 1977: 208, 1985: 56. Vanden-berg, 2002: 382.

Diagnosis. The anteriorly prominent pronotum (Fig. 21G)with rounded and not margined anterior edge is a characteristicof this genus shared only with Allenius. Nipus is distinguishedfrom Allenius by a much shorter prosternum bearing a distinctchinpiece anteriorly (Fig. 21E), and by the head not retractedinto the prothorax when in repose.

Description. Length 1.0–1.5 mm. Body elongate, oval,compact and convex, smooth, dorsum glabrous or with sparseminute setae. Colour dark to light brown, in some specieswith yellowish spots on elytra. Winged. Head (Fig. 21A)slightly elongate, in repose dorsally covered by pronotum;eyes large, glabrous, not margined dorsally. Frontoclypeus(Fig. 21A) about 2.4× as long as antennal socket length,emarginate around antennal insertions, lateral sides clearlymargined. Subantennal grooves short. Subgenal gland openingsas Fig. 21B. Gula (Fig. 21B) elongate, as long as mouthpartscavity, gular sutures short, narrowly separated, surface withsparse setae. Antenna ten-segmented with three-segmentedclub (Fig. 21C). Mouthparts (Fig. 21B): prostheca present,mola reduced; maxillary palpifer not foveate, terminal max-illary palpomere subparallel and diagonally truncate apically;submentum reduced to a carina, mentum elongate divergentto apex with two long setae on disc, prementum subtrian-gular, labial palps narrowly separated, ligula well developed,setose and broadly rounded. Pronotum (Fig. 21G) transverse,with anterior margin arcuate; disc convex without marginal lineseparating anterior corners. Prosternum (Fig. 21E) with large,broadly rounded chinpiece; prosternal process 0.6× as wideas coxa; prosternal carinae present. Elytral surface smooth andpolished, sutural line absent; epipleura reaching apical third.Scutellum triangular. Mesoventral (Fig. 21F) process trans-verse, slightly broader than coxa; with prosternal and procoxalrest. Metaventrite (Fig. 21F) transverse, slightly longer thanabdominal ventrite 1; postcoxal lines descending and incom-plete laterally; mesometaventral junction arcuate anteriorly.Abdomen (Fig. 21D) with six ventrites, ventrite 1 slightlylonger than ventrites 2 and 3 combined, postcoxal lines diag-onal and incomplete with accessory line very close to post-coxal line. Legs slender, tarsi three-segmented; pretarsal claws

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A B

C

ED

F G

Fig. 21. Nipus sp. (U.S.A.): (A) head, dorsal; (B) head, ventral; (C) antenna detail; (D) abdomen; (E) prothorax, ventral; (F) meso–metaventrite;(G) pronotum.

simple. Male genitalia: penis guide long and rotated asymmet-rically; parameres strongly asymmetrical; penis long with basalcapsule.

Distribution. Western U.S.A.

Species included. Nipus biplagiatus Casey; Nipus occiduusGordon; Nipus niger Casey; Nipus planatus Gordon.

Material examined. Nipus planatus; N. occiduus; N. niger ;N. biplagiatus and unnamed species from New Mexico,U.S.A.

Comments. Originally Casey placed Nipus in Cranophorini,but Gordon (1970d) transferred it to Sticholotidini, and then in1977 to Microweiseini. The only known host for this genusis Ehrhornia cupressi (Ehrhorn) (Pseudococcidae) (Gordon,1985).

Allenius gen.n.(Figs 3A, B, 22A–J, 28A–I)

Type species. Allenius californianus sp.n.

Diagnosis. Allenius is similar to Nipus in having a rela-tively flat body, anteriorly prominent pronotum (Fig. 22C) withanterior and lateral edges not margined; however, the proster-num in Allenius (Fig. 22B) is strongly elongate in front ofcoxae and does not bear typical chinpiece present in Nipus.

Description. Length about 1.4 mm. Body ovoid and elon-gate (Fig. 3A, B); pronotum projected anteriorly and flat-tened, elytra convex. Winged. Head (Fig. 22A) flattened, abouttwo times longer than wide, in repose retracted into pro-thorax; subantennal grooves absent. Subgenal gland openingspresent. Gula longer than wide, slightly divergent to apex.

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A

D

H I

J

F G

E

B C

Fig. 22. Allenius californianus (Mexico): (A) head, dorsal; (B) prothorax, ventral; (C) pronotum; (D) antenna; (E) maxilla; (F) middle leg;(G) abdomen; (H) abdominal postcoxal line; (I) elytron, lateral; (J) meso–metaventrite.

Eyes (Fig. 22A) reduced to about 14 coarse facets, dorsallyabout 0.1× as long as head capsule; interocular distance about6.0× an eye diameter; interfacetal setae indistinct; temple morethan 3.0× eye length. Frontoclypeus (Fig. 22A) emarginatearound antennal insertions, lateral sides margined. Occipitalregion without endocarina. Antenna (Fig. 22D) ten-segmentedwith compact four-segmented club; antennomere 4 distinctlylonger than antennomere 3, antennomere 5 very short andtransverse. Maxillary palpifer not foveate; terminal maxillarypalpomere elongate and conical (Fig. 22E). Mandible withmolar area broad, prostheca reduced. Submentum and men-tum very narrow; prementum slightly triangular; labial palpsnarrowly separated; ligula membranous, longer than labialpalpomere 1. Pronotum (Fig. 22C) about as long as wide,widest at base, slightly constricted medially, anterior angles

obtuse, apex broadly arcuate anteriorly, without line separat-ing anterior corners; disc convex, especially along median andbasal parts, anterolaterally sloping towards lateral margins; sur-face with microsculpture. Prosternum (Fig. 22B) in front ofcoxae about four times coxa length, carina extending fromprosternal process. Prosternal process (Fig. 22B) about 0.5×as wide as coxa and subparallel. Scutellum abruptly raised andtriangular. Mesoventrite transverse (Fig. 22J); procoxal andprosternal rests absent, mesometaventral junction at midlineslightly wider than coxa. Elytra (Fig. 22I) widest near mid-dle; lateral margins visible from above; surface smooth andsetose; epipleura narrow and incomplete. Wing with reducedvenation, edges fringed with fine setae; anal lobe well devel-oped. Metaventrite (Fig. 22J) 0.5× as long as wide, longerthan abdominal ventrite 1; postcoxal lines descending, close

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160 H. E. Escalona and A. Slipinski

to lateral margin. Abdomen (Fig. 22G) with six ventrites,ventrite 6 partially visible; ventrite 1 about 1.5× as long asventrite 2; postcoxal lines arcuate and incomplete (Fig. 22H),without accessory line. Legs slender (Fig. 22F); tarsi three-segmented; pretarsal claws simple. Male terminalia: spiculumgastrale asymmetrical, Y-shaped; male genitalia: tegminal strutas long as ventrites 1–5, parameres apex setose (Fig. 28C),penis long and curved. Female genitalia: coxites sclerotized,about twice longer than abdominal ventrite 6; triangular, withfew apical setae; stylus reduced and apical; infundibulum short;spermatheca sclerotized and rounded.

Distribution. Mexico, U.S.A.

Species included. Allenius californianus sp.n.; Allenius ivieisp.n.

Etymology. Dedicated to the well-known beetle enthusiastAlbert Allen (Idaho, U.S.A.), in recognition of his help withthis paper and his insatiable passion for beetles. Gendermasculine.

Allenius californianus sp.n.(Figs 3A, B, 22A–J, 28A–D)

Description. Length: 1.3 mm; width: 0.7 mm. Colour(Fig. 3A) reddish to yellowish, lighter on anterior half ofpronotum and on median spots on elytra; surface with sparsefine setae denser on elytra. Elytral disc with sparse punc-tures. Male genitalia (Fig. 28A–D): tegmen as Fig. 28B–C,parameres as Fig. 28B; penis apex as Fig. 28A (only apicalpart of penis was recovered in dissection). Female genitalia,spermatheca globular narrow and pointed to apex.

Etymology. Named after the type locality, Baja California,Mexico.

Type material. Holotype: ♂, MEXICO, Baja California,6 km E. San Agustin, 29◦59′N, 114◦56′W, 580 m, ethyleneglycol pitfall trap, 16-vii-1991–27-v-992 (Clark, Blom, Clark,Ward Jr.) (NMNH).

Paratypes: MEXICO, Baja California, Mesa Palmarito,29◦47′N, 114◦44′W, 840 m, ethylene glycol pitfall trap, 27-iv – 1991–21-vi-1990 (Clark, Clark, Ward Jr.) (one specimen,AAC). Mesa, ex. pitfall trap, 30-viii-1989 (Clark ) onespecimen (ANIC, on slide or coated for SEM).

Allenius iviei sp.n.(Fig. 28E–I)

Diagnosis. This species can be easily distinguished fromA. californianus by the form of the male genitalia.

Description. Length: 1.4 mm; width: 0.7 mm. Differs fromA. californianus in the elytra with punctures more dense, uni-form and male genitalia (Fig. 28F–I): tegmen as Fig. 28G–H,penis guide broad in lateral view (Fig. 28G); penis apex asshown in Fig. 28I.

Etymology. Named after Dr Michael A. Ivie (Montana StateUniversity) who provided us with the only known malespecimen of this interesting species.

Type material. Holotype: ♂, U.S.A., Montana, BeaverheadCo., Centennial Valley, Pitfall, 44◦41.878′N, 111◦48.558′′W,WS-15A AUG/24-05/SEP/2009 (Winton). Voucher: Winton2010, MS Thesis, Centennial Sandhills (MTEC).

Paratype, ♀, U.S.A., Idaho, Blaine CO, Butte CO, 5 Mi E.Arco, 4-viii-1981 (Allen) (AAC).

Microweisea Cockerell(Figs 3G, 23A–G)

Microweisea Cockerell, 1903: 38 (replacement name). Typespecies Smilia felschei Weise, 1891 (monotypy). Pope,1962: 637; Gordon, 1970b: 209, 1977: 203, 1985: 37;Gordon & Hilburn 1990: 267; Belicek, 1976: 296;Vandenberg, 2002: 382.

Smilia Weise, 1891: 288 (not Germar, 1833) (junior homo-nym).

Epismilia Cockerell, 1900: 606 (not Fromental, 1861) (juniorhomonym).

Pseudoweisea Schwarz, 1904: 118 (lapsus calami).Gnathoweisea Gordon, 1970a: 47. Type species Smilia plan-

iceps Casey, 1899 (original designation). Gordon,1977: 204, 1985: 49. Vandenberg, 2002: 382 syn.n.

Diagnosis. Microweisea is similar and difficult to dis-tinguish from Coccidophilus. The only reliable diagnosticcharacter that separates these genera is antenna not seg-mented with two-segmented club in Coccidophilus (Fig. 24F),and the ten-segmented antenna with three-segmented club inMicroweisea.

Description. Length up to 1.4 mm. Body elongate, ovaland convex, dorsal surfaces smooth or sparsely setose, setaeshort and commonly associated with micropunctures, in somespecies pronotum microsculptured. Colour black to lightbrown. Winged. Head slightly transverse to elongate; eyeslarge, often with fine interfacetal setae not margined dor-sally. Occiput often with distinct endocarina (Fig. 3G) (notvisible externally). Frontoclypeus (Fig. 23A) two times ormore the length of antennal socket, emarginate around anten-nal insertions, lateral sides clearly margined. Subantennalgrooves shallow. Subgenal gland openings as Fig. 23B. Gula(Fig. 23B) transverse to elongated; gular sutures parallel, sur-face not well sclerotized with few apical setae. Antennaten-segmented with three-segmented club (in some speciesthe antennal club can be interpreted as four-segmented).Mouthparts (Fig. 23B, C): mandible, prostheca present, molareduced; maxillary palpifer not foveate, terminal maxillarypalpomere elongate and subparallel, apex diagonally trun-cate; submentum very reduced, carina like; mentum elon-gate and narrow, prementum wider than mentum and welldeveloped, labial palps narrowly separated, ligula broadlyrounded and setose. Pronotum (Fig. 23G) transverse, disc con-vex, lateral edges smooth, with complete line separating ante-rior corners from pronotal disc, lateral sides not margined.Prosternum (Fig. 23F) with chinpiece; prosternal process about0.6× as wide as a coxa. Elytral surface smooth and pol-ished, sutural line absent; epipleura narrow, reaching apical

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A

D

F G

E

B

C

Fig. 23. Microweisea sp. (U.S.A.): (A) head, dorsal; (B) head, ventral; (C) mouthparts detail; (D) meso–metaventrite; (E) abdomen detail;(F) prothorax, ventral; (G) pronotum.

third. Scutellum triangular. Mesoventrite (Fig. 23D) transverseprocoxal and prosternal rest present; mesometaventral junc-tion at midline slightly broader than coxa; mesometaventraljunction arcuate anteriorly. Metaventrite (Fig. 23D) transverse,longer than abdominal ventrites 1–2 combined, postcoxal linesdescending. Abdomen (Fig. 23E) with six ventrites; ventrite 6partially visible; ventrite 1 slightly longer than ventrite 2, post-coxal line arcuate, incomplete; accessory line, oblique and veryshort. Legs slender, tarsi three-segmented, pretarsal claws sim-ple. Male genitalia: penis guide long and rotated; parameresstrongly asymmetrical; penis with evident basal capsule.Female genitalia: bursa copulatrix with long tubular infundibu-lum; spermatheca forming a bulb with elongated apex.

Distribution. New World.Species included. Microweisea planiceps (Casey) comb.n.;

Microweisea hageni (Gordon) comb.n.; Microweisea ferox

(Gordon) comb.n.; Microweisea micula (Gordon) comb.n.;Microweisea texana (Gordon) comb.n.; Microweisea suturalis(Schwarz); Microweisea minuta (Casey); Microweisea misella(LeConte); Microweisea coccidivora (Ashmead); Microweiseaovalis (LeConte).

Material examined. Microweisea coccidivora; M. minuta;M. misella; M. suturalis; M. planiceps; M. hageni ; M. ferox ;M. micula.

Comments. Gordon (1977, 1985) erroneously interpretedGnathoweisea antenna as nine-segmented. An Microweisealarva was described by Boving (1917) and Gage (1920),illustrated by LeSage (1991) and included in a key by Reeset al. (1994): these authors remarked upon the elongated headcapsule with a large occipital endocarina, an unusual character

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162 H. E. Escalona and A. Slipinski

for Coccinellidae larvae. An M. ovalis pupa was described byPhuoc & Stehr (1974).

Host data includes the Diaspididae, Melanaspis obscura(Comstock), Pseudaonidia duplex (Cockerell), Diaspidiotusperniciosus (Comstock), Lepidosaphes spp., Chionaspis spp.,Diaspidiotus ancylus (Putnam), Nuculacaspis californica(Coleman) (Edmunds, 1973; Belicek, 1976; Gordon,1977; Gordon, 1985; Fondren & McCullough, 2005;Gordon & Davidson, 2008) and the Adelgidae (Aphidoidea,Hemiptera) Pineus strobi (Hartig) (Raske & Hodson, 1964).

Coccidophilus Brethes(Figs 2H, 24A–J, 25A–G)

Coccidophilus Brethes, 1905: 76. Type species Cocci-dophilus citricola Brethes, 1905 (original designation).

Costa Lima, 1941: 409; Pope, 1962: 638; Gordon,1977: 203, 1978: 206, 1985: 44; Duverger, 1986: 220;Vandenberg, 2002: 382.

Pseudosmilia Brethes, 1925: 156. Type species Pseudosmiliaarrowi Brethes, 1925 (monotypy). Pope 1962: 638;Gordon 1977: 204. Synonymized by Gonzalez, 2008: 79.

Cryptoweisea Gordon, 1970b: 213. Type species Pentiliamarginata LeConte, 1878. Synonymized by Gordon,1977: 203.

Diagnosis. Coccidophilus is very similar and difficult todistinguish from Microweisea. It can only be distinguishedfrom Microweisea in having a nine-segmented antenna withtwo-segmented club (Fig. 24F).

Description. Length about 1.1 mm. Body (Fig. 2H) ovaland convex, smooth or sparsely setose. Colour light to

A

E

H I J

F

G

B

C

D

Fig. 24. Coccidophilus sp. (Brazil): (A) head, dorsal; (B) head, ventral; (C) antenna; (D) mandible, mesal view; (E) head, lateral view; (F) antennadetail; (G) maxilla; (H) pronotum; (I) elytron ventral; (J) elytron lateral.

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A

D

F

E

G

B C

Fig. 25. Coccidophilus sp. (Brazil): (A) head, ventral detail; (B) maxillae; (C) mandible, ventral; (D) abdomen; (E) meso–metaventrite;(F) prothorax, ventral; (G) prothorax detail, ventral.

dark brown. Winged. Head transverse (Fig. 24A); eyeslarge, glabrous without margin. Frontoclypeus (Fig. 24A) aslong as 1.5× antennal socket length, emarginate aroundantennal insertions, margined. Subantennal grooves short.Subgenal gland openings as shown in Fig. 24B. Gula(Fig. 24B) short and transverse, with few sparse setae. Antenna(Fig. 24C, F) nine-segmented with two-segmented club,antennomere 7 transverse. Mouthparts (Figs 24B, G, 25B):mandible (Figs 24D, 25C) prostheca present, mola reduced;maxillary palpifer not foveate, terminal maxillary palp elon-gated and conical, apically pointed; submentum (Fig. 24B)very narrow, almost carina like, mentum elongate, longerand broader than prementum with two long setae at middle,prementum well developed, labial palps, narrowly separated,ligula broadly rounded and setose. Pronotum (Fig. 24H) trans-verse, convex, basal and lateral borders margined, anterior

corners clearly separated from pronotal disc, lateral edgessmooth. Prosternum (Fig. 25F, G) with broad chinpiece;prosternal process about 0.6× as wide as coxa; prosternalcarinae extend forwards. Elytral surface smooth and polished(Fig. 24J), sutural line absent; epipleura (Fig. 24I) narrow,reaching apical third. Scutellum triangular. Mesoventrite trans-verse and raised anteriorly, procoxal and prosternal restpresent. Meso–metaventral junction arcuate anteriorly, slightlybroader than coxa; metaventrite (Fig. 25E) transverse, longerthan abdominal ventrites 1 and 2 combined; postcoxal linesweakly recurved but often angulate or weakly descendingnear lateral margins. Abdomen with six ventrites (Fig. 25D);ventrite 6 partially visible; postcoxal lines arcuate and usu-ally incomplete, accessory line very short and distant frompostcoxal line, ventrite 1 longer than ventrites 2 and 3 com-bined. Legs slender, tarsi three-segmented; pretarsal claws

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164 H. E. Escalona and A. Slipinski

simple. Male genitalia: penis guide long and rotated; parameresstrongly asymmetrical; penis long with evident basal capsule.Female genitalia: bursa copulatrix without sclerites but withtubular infundibulum; spermatheca simple, bulb-like.

Distribution. New World.

Species included. Coccidophilus arrowi (Brethes); Cocci-dophilus citricola Brethes; Coccidophilus cariba Gordon;Coccidophilus nigra Duverger; Coccidophilus aimogastaensisGonzalez & Pedemonte; Coccidophilus marginata (LeConte);Coccidophilus atronitens (Casey); Coccidophilus peninsularis(Gordon).

Material examined. Coccidophilus citricola; C. penin-sularis; C. cariba; C. marginata; C. atronitens and unnamedspecimens from South America.

Comments. Adults and larvae of various Coccidophilusspecies feed on a broad range of Diaspididae scales, likeAspidiotus spp., Cornuaspis spp., Chrysomphalus spp., Par-latoria cinerea (Doane & Hadden), Selenaspidus articu-latus (Morgan), Chionaspis spp., Pseudolacopis pentagona(Targioni–Tozzetti), Lepidosaphes beckii (Newman), Saisse-tia oleae (Olivier), Aonidiella aurantii (Maskell), Diaspisechinocacti (Bouche), Unaspis citri (Comstock) (Gordon,1977, 1978; Duverger, 1986; Cooper & Cranshaw, 1999;Arruda Filho & Arruda, 2002; Silva et al., 2003, 2004, 2005;Santos & Gravena, 2005; Gonzalez & Pedemonte, 2010).

Coccidophilus citricola has been found feeding on Diaspi-didae on various crops in South America (Silva et al., 2005).It was also introduced to the U.S.A. (Frank & McCoy, 1993;Peck & Thomas, 1998). The larvae of C. citricola has beenkeyed and illustrated by Rees et al. (1994) and described bySilva et al. (2005).

Parasidis Brethes(Figs 2D, 26A–J)

Parasidis Brethes, 1915: 88. Type species Parasidis porteriBrethes, 1925 (subsequent designation by Brethes,1925). Brethes, 1925: 171; Honour & Gonzalez, 2010:59 (revision).

Sarapidus Gordon, 1977: 202. Type species Sarapidusaustralis Gordon, 1977 (original designation). Syn-onymized by Gonzalez, 2008: 79.

Diagnosis. Parasidis can be distinguished from similar andsympatric Coccidophilus and Stictospilus by four-segmentedtarsi and the relatively broad prementum (Fig. 26A).

Description. Length 1.0–2.0 mm. Body elongate, oval andconvex (Fig. 2D), dorsum smooth or sparsely setose, setae oftenassociated with micropunctures. Colour black to light brown,with red or light-brown large spots on elytra. Winged. Headtransverse (Fig. 26B), eyes large, with few fine interfacetalsetae, not dorsally margined. Frontoclypeus (Fig. 26B) aboutas long as antennal socket length, emarginate around antennalinsertions, lateral sides partially margined. Subantennal groovesextending behind eyes. Subgenal gland openings as shown in

Fig. 26F, J. Gula (Fig. 26A) transverse and well developed,gular sutures parallel and well defined, surface with few setaeanteriorly. Antenna ten-segmented with three-segmented club(Fig. 26D), antenomere 7 transverse and setose. Mouthparts(Fig. 26A): mandible, prostheca present, mola reduced; max-illary palpifer foveate, terminal maxillary palpomere ovoid;submentum narrow, mentum triangular with few sparse setae,apex emarginate to accommodate prementum; prementum veryshort, labial palps clearly separated, ligula short and fringedwith setae. Pronotum (Fig. 26E) transverse; disc with completeline separating anterior corners; anterior edge not margined,lateral edges smooth. Prosternum (Fig. 26G) with prominentchinpiece; prosternal process about 0.7× as wide as coxa. Ely-tral surfaces smooth with fine punctures, sutural line absent;epipleura reaching apical third with fine setae. Scutellum heartlike. Mesoventral process transverse (Fig. 26I), raised ante-riorly; procoxal and mesosternal rest present. Metaventrite(Fig. 26I) transverse, 1.2× longer than abdominal ventrite 1;mesometaventral junction arcuate anteriorly; metanpisternumwith longitudinal sulcus in apical half; postcoxal lines descend-ing, laterally continue along lateral margin. Abdomen with sixventrites; ventrite 6 partially visible, ventrite 1 as long as ven-trites 2 and 3 combined; postcoxal lines arcuate and incomplete(Fig. 26H), accessory line long, often joining postcoxal lineapex. Legs slender (Fig. 26C), tarsi four-segmented; pretarsalclaws barely apendiculate. Male genitalia: penis guide longerthan parameres and rotated; parameres slightly asymmetrical;penis with weak basal capsule. Female genitalia: bursa copu-latrix with infundibulum; spermatheca forming a single bulbwith narrowly elongate apex.

Distribution. Central Chile.

Species included. Parasidis porteri Brethes; Parasidis aus-tralis (Gordon); Parasidis barrigai Honour & Gonzalez; Para-sidis brethesi Honour & Gonzalez; Parasidis elguetai Honour& Gonzalez; Parasidis gordoni Honour & Gonzalez.

Material examined. Parasidis australis; P. brethesi ;P. elguetai.

Comments. The identity of the genus and its synonymy waselaborated upon recently by Gonzalez (2008). Some specieshave been collected in Nothofagus forests but their hostsremain unknown (Honour & Gonzalez, 2010).

Stictospilus Brethes(Figs 2A, 27A–H)

Stictospilus Brethes, 1925: 153. Type species Stictospilusdarwini Brethes, 1925 (monotypy). Pope, 1962: 638;Gordon, 1977: 203; Gonzalez, 2008: 80.

Diagnosis. This monotypic genus is very similar toMicroweisea but can be distinguished by its distinctly setosedorsum (Fig. 2A).

Description. Length 1.4 mm. Body (Fig. 2A) elongate,oval and convex; dorsum clothed with long erect setaemixed with minute hairs. Colour light to dark brown, in

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Generic revision and phylogeny of Microweiseinae 165

A

D

B C

F

H

E

G

I

J

Fig. 26. Parasidis sp. (Chile): (A) head, ventral; (B) head, dorsal; (C) middle leg; (D) antenna; (E) pronotum; (F) head, lateral view; (G) prothorax,ventral view; (H) abdomen detail; (I) meso–metaventrite; (J) head detail, ventral view.

which case the pronotum could be darker. Winged. Headtransverse (Fig. 27A), eyes large, glabrous, not margineddorsally. Frontoclypeus (Fig. 27A) as long as antennal socketlength, lateral sides margined, emarginate around antennalinsertions. Subantennal grooves well developed (Fig. 27B),extending behind eyes. Subgenal gland openings as shown inFig. 27B. Gula (Fig. 27B) transverse, gular sutures short andconverging, widely separated, surface with few apical setae.Antenna ten-segmented, with three-segmented club (Fig. 27C),antennomere 7 shorter than antennomeres 6 or 8. Mouthparts(Fig. 27B): mandible with prostheca present, mola reduced;maxillary palpifer not foveate, terminal maxillary palpomeresomewhat ovoid, apex diagonally truncate; submentum verynarrow and carina like; mentum rectangular, slightly expandedapically; prementum very short, labial palps contiguous, ligulawell developed and setose. Pronotum (Fig. 27D) transverse,disc convex, with complete line separating anterior corners

from pronotal disc; lateral sides weakly margined, lateral edgessmooth. Prosternum (Fig. 27F) with large prominent chinpiece;prosternal process about 0.7× as wide as coxa. Elytralsurface smooth, sutural line absent; epipleura reaching apicalthird. Scutellum triangular. Mesoventral process (Fig. 27E)transverse and short, wider than mesocoxa; prosternal andprocoxal rest present. Metaventrite (Fig. 27E) transverse,longer than abdominal ventrite 1, mesometaventral junctionarcuate anteriorly; postcoxal lines descending and continuealong lateral margin. Abdomen with six ventrites (Fig. 27H);ventrite 1 longer than ventrites 2 and 3 combined; postcoxallines arcuate (Fig. 27G) and incomplete, with very shortaccessory line. Legs slender, tarsi three-segmented, pretarsalclaw simple. Male genitalia: parameres slightly asymmetrical;penis guide broad in dorsal and lateral view, penis basalcapsule well defined.

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166 H. E. Escalona and A. Slipinski

A

D

F

G H

B C

E

Fig. 27. Stictospilus sp. (Chile): (A) head, dorsal; (B) head, ventral; (C) antenna; (D) pronotum; (E) meso–metaventrite; (F) prothorax, ventral;(G) abdominal postcoxal line detail; (H) abdomen.

Distribution. Central and southern Chile.

Species included. Stictospilus darwini Brethes.

Material examined. Stictospilus darwini.

Microcapillata GordonMicrocapillata Gordon, 1977: 201. Type species Micro-

capillata clypealis Gordon, 1977 (original designation).

Diagnosis. This monotypic genus is very similar to Stic-tospilus. Apparently it can be distinguished by its denser andmore even body vestiture and narrower terminal maxillarypalpomere, but more specimens are needed to confirm thatdiagnosis.

Description. Length about 1.1 mm. Body elongate, oval andconvex, covered with uniform erect setae. Colour yellowish

brown. Wings not observed. Head elongate; eyes large andcoarsely facetted. Frontoclypeus as long as 2 to 3× antennalsocket length, emarginate around antennal insertions. Sub-antennal grooves short. Antenna ten-segmented, club elon-gate and three-segmented. Mouthparts: labial palps narrowlyseparated, last maxillary palpomere elongate, subparallel anddiagonally truncate apically. Pronotum transverse, disc con-vex, lateral sides margined, anterior corners separated fromdisc by line, lateral edges smooth. Prosternum with moder-ately large chinpiece; prosternal process parallel sided andmargined; hypomeron not foveate. Elytral surface denselypunctate, sutural line absent; epipleura disappearing in apicalhalf. Mesoventrite slightly broader than coxa, prosternal pro-cess rest shallow. Mesoventrite transverse, mesometaventraljunction arcuate anteriorly; postcoxal lines descending. Legs

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Generic revision and phylogeny of Microweiseinae 167

A

B

C

D

E

G

F

H

I

MKJ

L

N

Fig. 28. A–D, Allenius californianus: (A) penis detail; (B) tegmen dorsal; (C) tegmen lateral; (D) spiculum gastrale. E–I, Allenius iviei :(E) tegminal strut; (F) penis; (G) tegmen, dorsal; (H) tegmen, lateral; (I) penis detail. J–K, Carinodulina ruwenzorii : (J) penis; (K) tegmen, dorsal.L–N, Cathedrana natalensis: (L) penis; (M) tegmen, dorsal; (N) tegmen, lateral.

slender, tarsi three-segmented (four-segmented according tooriginal description). Abdomen (Gordon, 1977) with six ven-trites; postcoxal lines arcuate and incomplete; accessory linevery short and hardly visible near coxal cavity only. Male gen-italia: tegmen asymmetrical; parameres strongly asymmetrical;penis long with apparently reduced basal capsule (Gordon,1977: fig. 71).

Distribution. The distribution is uncertain: the holotype ofM. clypealis Gordon was intercepted at the U.S.A. border, onorchids from Mexico (San Luis de Potosi).

Species included. Microcapillata clypealis Gordon.

Material examined. Microcapillata clypealis, holotype.

Comments. The characters listed above were based on anexamination of the holotype and original description (Gordon,

1977); the abdomen and genitalia were lost after the specieswas described.

Supporting Information

Additional Supporting Information may be found in the onlineversion of this article under the DOI reference:10.1111/j.1365-3113.2011.00601.x

Table S1. Data matrix used in the cladistic analysis.

Please note: Neither the Editors nor Wiley-Blackwellare responsible for the content or functionality of anysupporting materials supplied by the authors. Any queries(other than missing material) should be directed to thecorresponding author for the article.

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168 H. E. Escalona and A. Slipinski

Acknowledgements

We are indebted to: Albert Allen (Idaho, U.S.A.), Ivan Lobland Giulio Cuccodoro (MHNG, Switzerland), Roger Booth(NHML, England), Guillermo Gonzalez (Santiago, Chile),Robert Gordon (U.S.A.), Phillip Perkins (Harvard University,U.S.A.), Michael Caterino (Santa Barbara Museum of Nat-ural History, U.S.A.), Natalia Vandenberg (USDA–NMNH,U.S.A.), Alfred Newton Jr. (Chicago Field Museum, U.S.A.),Michael Ivie (Montana State University, U.S.A.) and NathanLord (University of New Mexico, U.S.A.) for provid-ing specimens; Steve Lingafelter and Natalia Vandenberg(USDA–NMNH, U.S.A.) for support and assistance dur-ing a recent visit that HE made to the Smithsonian. Ains-ley Seago (ANIC-CSIRO, Australia), Wioletta Tomaszewska(Museum and Institute of Zoology–PAS, Poland), JohnLawrence (ANIC-CSIRO, Australia), Natalia Vandenberg andRoger Booth read an early draft of the article and offeredvaluable suggestions and corrections. Lourdes Chamorro(USDA–NMNH, U.S.A.) kindly helped with photographs.The help and patience of Natalie Banks and Mark Talbot(CSIRO, Australia) with SEM sessions are appreciated. JurgCarl DeMarmels (MIZA, Venezuela) verified the compositionof the new Latin names. HE thanks the staff of Entomology,Museo del Instituto de Zoología Agrícola–UCV, Venezuelafor years of support. Last, but not least, thanks to Tom Weir,Cate Lemann and Anne Hasting (ANIC-CSIRO, Australia) fortechnical support.

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Accepted 27 July 2011First published online 20 October 2011

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