Encyclopedia of insects

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Resh & Carde. 2009. Encyclopedia of insects.

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  • 1.Encyclopedia ofINSECTS

2. This page intentionally left blank 3. Encyclopedia ofINSECTSSecond Edition Editors Vincent H. Resh University of California, BerkeleyRing T. Carde University of California, Riverside AMSTERDAM BOSTON LONDON NEW YORK OXFORD PARISSAN DIEGO SAN FRANCISCO SINGAPORE SYDNEY TOKYOAcademic Press is an imprint of Elsevier 4. Cover Art: The jewel scarab, Chrysina cusuquensis, known only from a fragment of forest in northern Guatemala(Photograph courtesy of David Hawks).Letter-Opening Photo Credits:R: Snakey (Raphidioptera) photographed at Nanaimo (Vancouver Island), British Columbia, Canada.(Photograph Copyright Jay Patterson.)Y: Aedes aegypti, Uganda strain (a vector of yellow fever), bloodfeeding from the photographers hand.(Photograph by Leonard E. Munstermann.)Other letter-opening photographs furnished by the authors. (See relevant article for credit.)Copyright Exceptions:Cat Fleas by Nancy C. Hinkle and Michael K. Rust, Cell Culture by Dwight E. Lynn, Extension Entomology byNancy C. Hinkle, Beverly Sparks, Linda J. Mason, and Karen M. Vail, and Nomenclature and Classication, Principlesof by F. Christian Thompson are in the public domain.Embiidina by Edward S. Ross, Figures 14 Copyright Edward S. Ross. Swimming, Lake Insects by Werner Nachtigall,Figures 13 Copyright Werner Nachtigall. Wolbachia by Richard Stouthamer, Figure 1 Copyright Richard Stouthamer.This book is printed on acid-free paper.Academic Press is an imprint of Elsevier30 Corporate Drive, Suite 400, Burlington, MA 01803, USA525 B Street, Suite 1900, San Diego, CA 92101-4495, USA32 Jamestown Road, London NW1 7BY, UKSecond edition 2009Copyright 2009, Elsevier, Inc. All rights reserved.No part of this publication may be reproduced, stored in a retrieval system or transmitted inany form or by any means electronic, mechanical, photocopying, recording or otherwisewithout the prior written permission of the publisherPermissions may be sought directly from Elseviers Science & Technology RightsDepartment in Oxford, UK: phone (44) (0) 1865 843830; fax (44) (0) 1865 853333;email: [email protected]. Alternatively you can submit your request onlineby visiting the Elsevier web site at http://www.elsevier.com/locate/permissions, andselecting: Obtaining permission to use Elsevier materialISBN: 978-0-12-374144-8 For information on all Academic Press publications visit our website at www.elsevierdirect.comTypeset by Macmillan Publishing Solutionswww.macmillansolutions.comPrinted and bound in China09 10 11 12 10 9 8 7 6 5 4 3 2 1 5. CONTENTSCONTENTS BY SUBJECT AREA xvii Anatomy: Head, Thorax,CONTRIBUTORSxxi Abdomen, and Genitalia11GUIDE TO THE ENCYCLOPEDIAxxxDavid H. Headrick and Gordon GordhFOREWORDxxxiiAnopheles Mosquito21PREFACE TO THE SECOND EDITION xxxiv see MosquitoesPREFACE xxxvABOUT THE EDITORS xxxviAnoplura21see PhthirapteraAntennae21A Catherine LoudonAcari1Ants24see Mites; TicksNigel R. FranksAccessory Glands 1Aphids27Diana E. WheelerJohn T. SorensenAcoustic Behavior2Apis Species31see Hearing Eva CraneAedes Mosquito 2Aposematic Coloration 33see MosquitoesMathieu JoronAestivation2Apterygota38Sinzo MasakiRing T. Card and Vincent H. ReshAfricanized Bees 4Aquatic Habitats38see Neotropical African BeesRichard W. Merritt and J. Bruce WallaceArachnida 48Agricultural Entomology4see Daddy-Long-Legs; Mites; Scorpions;Marcos Kogan and Ronald ProkopySpiders; Ticks; VinegaroonsAldery8Archaeognatha 48see Megaloptera Helmut SturmAmber8Arthropoda and Related Groups 50George Poinar Jr. James H. Thorp 6. vi Contents Auchenorrhyncha56Blood 112 Christopher H. Dietrichsee Circulatory System Autohemorrhage 64Blood Sucking 112 Vincent H. ReshM. J. Lehane Autotomy64 Body Size 114 Ring T. CardChrister Bjrkman, Karl Gotthard,and Mats W. PetterssonBoll Weevil 116R. N. Foster BBombyx mori 117 Bedbugs 65 Satoshi Takeda Christiane Weirauch and Alec C. GerryBook Louse119 Bee 66 see Psocoptera see Apis SpeciesBorers119 Beekeeping66 Timothy D. Paine Eva CraneBrain and Optic Lobes 121 Bee Products 71Nicholas J. Strausfeld Eva CraneBristletail 130 Beetle 75see Archaeognatha see ColeopteraBubonic Plague130 Biodiversity75 Irwin W. Sherman Nigel E. Stork Biogenic Amines80 Arnd Baumann, Wolfgang Blenau, and Joachim Erber C Biogeographical Patterns 82Caddisy133 Peter Zwicksee Trichoptera Biological Control of Insect Pests 91Caste 133 M. S. Hoddle and R. G. Van DriescheSean ODonnell Bioluminescence 101Caterpillars135 James E. Lloyd and Erin C. GentryFrederick W. Stehr Biotechnology and Insects 105Cat Fleas 138 Bryony C. BonningNancy C. Hinkle and Michael K. Rust Blattodea 108Cave Insects139 Donald G. CochranFrancis G. Howarth 7. Contents viiCell Culture 144Collection and Preservation201Dwight E. LynnCharles V. Covell Jr.Chemical Defense 145Collembola 206Murray S. BlumKenneth A. Christiansen, Peter Bellingerand Frans JanssensChemoreception 148B. K. MitchellColonies 210Sean ODonnellChiggers and OtherDisease-Causing Mites152Colorado Potato Beetle 212Larry G. Arlian George G. KennedyChitin 156Ephraim Cohen Coloration 213Helen GhiradellaChromosomes158Graham C. WebbCommercialization of Insects andTheir Products 220Chrysalis162Gail E. Kampmeier and Michael E. IrwinFrederick W. StehrConservation 227Cicadas163Tim R. NewMaxwell S. MouldsCrickets 232Circadian Rhythms165Richard D. Alexander and Daniel OtteTerry L. PageCrypsis236Circulatory System 169Paul M. BrakeeldThomas A. Miller and Gnther PassCultural Entomology239Classication173James N. Hoguesee Nomenclature and ClassicationCuticle245Cockroach173Svend O. Andersensee BlattodeaCocoon 173Frederick W. Stehr DCodling Moth 174Daddy-Long-Legs247Stephen C. Welter Gonzalo GiribetCoevolution175Damsely 248Douglas J. Futuymasee OdonataCold/Heat Protection 179Dance Language 248David L. DenlingerP. Kirk VisscherColeoptera 183DDT251Joseph V. McHugh and James K. LiebherrFumio Matsumura 8. viii Contents Defensive Behavior252Drosophila melanogaster 301 Justin O. SchmidtPatrick M. OGrady Delusory Parasitosis257Dung Beetles304 see Ekbom Syndrome James Ridsdill-Smith andLeigh W. Simmons Dengue257 Thomas W. Scott Dermaptera Susan M. Rankin and James O. Palmer 259 EEarwig308 Development, Hormonal Control of261see Dermaptera Michael E. AdamsEcdysis 308 Diapause267see Molting David L. DenlingerEcdysteroids308Michael E. Adams Digestion 271 Walter R. TerraEggs311Diana E. Wheeler Digestive System273 Walter R. Terra and Cllia FerreiraEgg Coverings 312Diana E. Wheeler Diplura 281 Markus KochEkbom Syndrome313Nancy C. Hinkle Diptera 284Embiidina 315 Richard W. Merritt, GregoryEdward S. Ross W. Courtney, and Joe B. KeiperEmbryogenesis 316 Diversity 297Lisa Nagy and Miodrag Grbic see BiodiversityEndangered Insects320 Division of Labor in Insect Societies 297Scott Hoffman Black and Gene E. Robinson Mace Vaughan Dobsony299Endopterygota 324 see MegalopteraRing T. Card Dog Heartworm 299Entomological Societies 324 Edward G. PlatzerAlan I. Kaplan Dormancy300Ephemeroptera 328 Richard E. Lee Jr. John E. Brittain and Michel Sartori Dragony301Estivation334 see Odonatasee Aestivation 9. ContentsixEvolution 334Fossil Record 396see Phylogeny of Insects David GrimaldiExcretion 334Freshwater Habitats 403Timothy J. Bradley see Aquatic HabitatsExopterygota339Ring T. CardFruit Fly 403 see Drosophila melanogasterExoskeleton 339Svend O. AndersenExtension EntomologyNancy C. Hinkle, Beverly Sparks,342GLinda J. Mason, and Karen M. VailGallmaking and Insects404Eyes and Vision 345Katherine N. Schick and Donald L. DahlstenMichael F. Land Genetically Modied Plants406 David A. Andow F Genetic Engineering Peter W. Atkinson and David A. OBrochta410Fat Body356Ephraim CohenGenetic Variation 416 George K. Roderick and Maria NavajasFeeding Behavior357R. F. Chapman Grasshopper 419Fire Ants 362see OrthopteraLes Greenberg Genomics419Flea364Peter Arensburger and Peter W. Atkinsonsee Siphonaptera Grassland Habitats424Flight364Teja Tscharntke, Ingolf Steffan-Michael Dickinson and Robert DudleyDewenter, Andreas Kruess, and Carsten ThiesFly 372see DipteraGreenhouse Gases, Global Warming, and Insects 428Folk Beliefs and Superstitions372 Peter StilingJames N. HogueFood, Insects as376Growth, Individual431Gene R. DeFoliartMartin B. Berg and Richard W. MerrittForensic Entomology 381Grylloblattodea 434M. Lee GoffD. C. F. Rentz and Sigfrid IngrischForest Habitats 386Gypsy Moth435David L. Wood and Andrew J. Storer Joseph S. Elkinton 10. xContentsHymenoptera473 HDonald L. J. QuickeHearing440Hypermetamorphosis 484Ron Hoy and Jayne YackJohn D. PintoHemiptera446Hyperparasitism486see Auchenorrhyncha;Daniel J. SullivanProsorrhyncha; SternorrhynchaHemolymph446Michael R. Kanost IHeteroptera449see Prosorrhyncha Imaginal Discs 489Seth S. BlairHibernation449Richard E. Lee Jr.Immunology 492Nancy E. BeckageHistory of Entomology 449Edward H. Smith and George G. Kennedy Industrial Melanism496Michael E. N. MajerusHomeostasis, Behavioral458P. Kirk VisscherInsecta, Overview501Homoptera459Vincent H. Resh and Ring T. Cardsee Auchenorrhyncha; SternorrhynchaInsecticides 502Honey459Fumio MatsumuraEva Crane and P. Kirk VisscherInsecticide and Acaricide Resistance 505Honey Bee461Gregor J. Devine andsee Apis SpeciesIan DenholmHoneydew 461Angela E. Douglas Insectivorous Plants 511Lewis J. FeldmanHornet 463see Wasps Insectivorous Vertebrates514Charles R. CrumlyHost Seeking, by Parasitoids 463J. Daniel Hare and Ronald M. WeselohInsect Zoos516Leslie Saul-GershenzHost Seeking, for Plants 466Elizabeth A. BernaysIntegrated Pest Management 523House Fly469Ronald Prokopy andGregory A. Dahlem Marcos KoganHuman History, Insects Effect on 471Integument 528James N. HogueSvend O. Andersen 11. Contents xiIntroduced Insects 529Lepidoptera559Daniel Simberloff Jerry A. PowellIsland Biogeography533Lice 587George K. Roderick andsee PhthirapteraRosemary G. GillespieLice, Human587Isoptera 535Terri L. MeinkingVernard R. LewisLocusts589R. F. Chapman JJapanese BeetleDavid W. Held and Daniel A. Potter 539MMagnetic Sense 592June Beetles 540John Klotz and Rudolf JanderDaniel A. Potter and David W. HeldMalaria594Juvenile Hormones541William K. ReisenMichael E. AdamsMantodea 597Lawrence E. Hurd KMantophasmatodea 599Klaus-Dieter Klass andKatydid546Christin Grossmannsee OrthopteraKiller Bees546Marine Insects 600see Neotropical African BeesLanna ChengMating Behaviors 604Darryl T. Gwynne LMechanoreception 610Andrew S. French andLacewing 547Pivi H. Torkkelisee NeuropteraMecoptera611Ladybugs 547George W. ByersMichael E. N. MajerusLarva551Medical Entomology 614Frederick W. StehrJohn D. EdmanLearning 552Medicine, Insects in 618Daniel R. Papaj Ronald A. ShermanLegs 555Megaloptera620Peter H. AdlerNorman H. Anderson 12. xii ContentsMetabolism623Nest Building 688S. N. Thompson and R. K. SuarezRobert L. JeanneMetamorphosis 627Neuropeptides 691Frederick W. Stehr Miriam AltsteinMigration 628Neuroptera695Hugh DingleCatherine A. Tauber, Maurice J. Tauber, and Gilberto S. AlbuquerqueMimicry 633Mathieu JoronNomenclature and Classication, Principles of 707Mites 643F. Christian ThompsonBarry M. OConnor Nutrition 715Molting 649S. N. Thompson and S. J. SimpsonLynn M. RiddifordMonarchs654Lincoln P. Brower OMosquitoes658Ocelli and Stemmata 721Bruce F. EldridgeFrederick W. StehrMoth663Odonata 721see LepidopteraK. J. TennessenMouthparts663Orientation 729R. F. ChapmanRing T. CardMovies, Insects in668Orthoptera732May R. Berenbaum and Sigfrid Ingrisch and D. C. F. RentzRichard J. Leskosky Ovarioles 743Muscle System 675Diana E. WheelerRobert Josephson Oviposition Behavior745Museums and Display Collections 680Marc J. KlowdenGordon M. Nishida P N Parasitoids 748Neosomy 685Nick MillsFrank J. Radovsky Parental Care 751Neotropical African Bees686Michelle Pellissier ScottOrley R. Taylor Parthenogenesis in Insects and Mites753Nervous System688Benjamin B. Normark andsee Brain and Optic LobesLawrence R. Kirkendall 13. ContentsxiiiPathogens of Insects757Praying Mantid 837Brian A. Federicisee MantodeaPhasmida765Predation/Predatory Insects837Erich H. Tilgner Ronald M. Weseloh and J. Daniel HarePheromones766Ring T. Card and Jocelyn G. MillarProsorrhyncha839Phoresy 772Carl W. SchaeferMarilyn A. Houck Protura855Photography of Insects774Markus KochMark W. MoffettPsocoptera 858Phthiraptera777Edward L. MockfordRonald A. Hellenthal and Roger D. PricePterygota860Phylogeny of Insects780Ring T. CardPeter S. Cranston and Penny J. GullanPuddling Behavior860Physical Control of Insect Pests794Scott R. SmedleyCharles Vincent, Phyllis Weintraub,Pupa and Puparium862and Guy HallmanFrederick W. StehrPhytophagous Insects798Elizabeth A. BernaysPhytotoxicityAlexander H. Purcell800 Q Queen863Plant Diseases and Insects802see CasteAlexander H. PurcellPlantInsect Interactions 806J. Mark ScriberRPlecoptera810Raphidioptera864Kenneth W. Stewart Ulrike Aspck and Horst AspckPollination and Pollinators 813Rearing of Insects 866Gordon W. Frankie and Robbin W. ThorpNorman C. LepplaPollution, Insect Response to 819Recruitment Communication869David M. Rosenberg and Vincent H. Resh James F. A. TranielloPolyembryony821Regulatory Entomology877Michael R. StrandRobert V. DowellPopulation Ecology826Reproduction, Female 880Joseph S. Elkinton Diana E. Wheeler 14. xiv ContentsReproduction, Female:Silk Production 921Hormonal Control of 882Frantiek Sehnal and Catherine CraigDiana E. Wheeler Silversh 924Reproduction, Male885see ZygentomaMarc J. Klowden Siphonaptera924Reproduction, Male:Michael W. Hastriter andHormonal Control of 887Michael F. WhitingMarc J. Klowden Snakey 928Research Tools, Insects as888see RaphidiopteraKipling W. WillSociality 928 James E. ZablotnyRespiratory System889Jon F. HarrisonSoil Habitats 935 Patricia J. VittumRiver Blindness 895Vincent H. ReshSound Production939 see Hearing Spermatheca 939SMarc J. KlowdenSalivary Glands 897Spermatophore 940Gregory P. WalkerMarc J. KlowdenScale Insect901Spiders 941see Sternorrhyncha Rosemary G. Gillespie and Joseph C. SpagnaScales and Setae901 Springtail951Shaun L. Winterton see CollembolaScorpions 904Stamps, Insects and 951Stanley C. WilliamsCharles V. Covell Jr.Segmentation909Sterile Insect Technique953Paul Z. Liu and Nipam H. Patel Jorge Hendrichs and Alan RobinsonSericulture 912Sternorrhyncha957Satoshi Takeda Penny J. Gullan and Jon H. MartinSex Determination 914Stoney 967Michael F. Antolin and Adam D. Henksee PlecopteraSexual Selection917Stored Products as Habitats 967Kenneth Y. Kaneshiro Rudy Plarre and Wendell E. BurkholderSilk Moth 921Strepsiptera971see Bombyx moriMichael F. Whiting 15. ContentsxvSwimming, Lake Insects972Touch1011Werner Nachtigallsee MechanoreceptionSwimming and Other Tracheal System1011Movements, Stream Insects 975Jon F. HarrisonBernhard Statzner Trichoptera (Caddisies) 1015Symbionts Aiding Digestion978John C. MorseAndreas Brune Tsetse Fly 1020Symbionts, Bacterial983Stephen G. A. LeakMichael E. N. MajerusSystematics 987see Nomenclature and Classication U Urban Habitats 1025TMichael K. RustTaste988see ChemoreceptionTaxonomy 988 Vsee Nomenclature and Classication Venom1028Teaching Resources988Justin O. SchmidtJohn H. Acorn and Felix A. H. Sperling Veterinary Entomology1031Temperature, Effects onBradley A. MullensDevelopment and Growth990 Oldr ich NedvedVibrational Communication1034 Andrej Cokl and Meta Virant-DoberletTermite 993see Isoptera Vinegaroons1038 Justin O. SchmidtTerrestrial Insects 993see Soil HabitatsVision 1041 see Eyes and VisionThermoregulation993Bernd Heinrich Vitellogenesis 1041 William H. TelferThrips999see ThysanopteraThysanopteraLaurence A. Mound999 WThysanura1003Walking and Jumping1044see Archaeognatha; Zygentoma Roy E. Ritzmann and Sasha N. ZillTicks1003Wasps1049Daniel E. Sonenshine Justin O. Schmidt 16. xvi ContentsWater and Ion Balance,Hormonal Control of1052 ZThomas M. Clark Zoonotic Agents, Arthropod-Borne 1065Wings1055Robert S. LaneRobin J. Wootton Zoraptera1069Wolbachia1061Michael S. EngelRichard Stouthamer Zygentoma1070Worker 1063Helmut Sturmsee Caste GLOSSARY 1073 SUBJECT INDEX1093 YYellow Fever 1064Thomas P. MonathYellowjacket 1065see Wasps 17. CONTENTS BY SUBJECT AREAAnatomy ExcretionAnatomy: Head, Thorax, Abdomen, and Genitalia Eyes and VisionAntennaeFat BodyBrain and Optic Lobes FlightChitinGenetic EngineeringColorationGenomicsCuticle HearingDigestive SystemHemolymphExoskeleton HibernationEyes and Vision Homeostasis, BehavioralIntegumentHoneydewLegsImaginal DiscsMouthpartsImmunologyOcelli and Stemmata InsecticidesPupa and Puparium Juvenile HormonesSalivary Glands Magnetic SenseScales and SetaeMechanoreceptionSegmentationMetabolismTracheal System MoltingWings Muscle SystemNeuropeptidesPhysiologyNutritionAestivation Reproduction, FemaleAutohemorrhageReproduction, Female: Hormonal Control ofAutotomyReproduction, MaleBiogenic Amines Reproduction, Male: Hormonal Control ofBioluminescence Respiratory SystemBiotechnology and Insects Salivary GlandsBody Size SegmentationBrain and Optic Lobes Sex DeterminationCell CultureSilk Production in InsectsChemical DefenseSymbionts Aiding DigestionChemoreceptionThermoregulationChitinTracheal SystemChromosomes VitellogenesisCircadian Rhythms Walking and JumpingCirculatory SystemWater and Ion Balance, Hormonal Control ofCold/Heat ProtectionCuticle BehaviorDDT Aposematic ColorationDevelopment, Hormonal Control ofAutohemorrhageDiapauseAutotomyDigestion BioluminescenceDigestive SystemBlood SuckingDormancyBorersEcdysteroidsCaste 18. xviii Contents by Subject AreaChemical Defense Mating BehaviorsChemoreception OvariolesCircadian RhythmsOviposition BehaviorColonies Parthenogenesis in Insects and MitesCrypsisPolyembryonyDance Language Reproduction, FemaleDefensive Behavior Reproduction, Female: Hormonal Control ofDivision of Labor in Insect SocietiesReproduction, MaleEyes and VisionReproduction, Male: Hormonal Control ofFeeding Behavior SpermathecaFlight SpermatophoreHearingVitellogenesisHibernationHost Seeking, by Parasitoids Development And MetamorphosisHost Seeking, for Plants Body SizeLearning CaterpillarsMagnetic Sense ChrysalisMating Behaviors CocoonMechanoreception Development, Hormonal Control ofMigration EcdysteroidsMimicry Egg CoveringsNest Building Growth, IndividualOrientation HypermetamorphosisOviposition Behavior Imaginal DiscsParental Care Juvenile HormonesPheromones LarvaPhoresy MetamorphosisPredation/Predatory Insects MoltingPuddling Behavior NeosomyRecruitment Communication Temperature, Effects on Development and GrowthSex DeterminationSexual SelectionSocialityMajor Groups And Notable FormsSwimming, Lake Insects AntsSwimming and Other Movements, Stream Insects AphidsThermoregulation Apis SpeciesVibrational CommunicationApterygotaWalking and JumpingArchaeognatha Arthropoda and Related GroupsEvolutionAuchenorrhynchaAmberBedbugsAposematic ColorationBlattodeaBiogeographical Patterns Boll WeevilCoevolutionBombyx moriFossil RecordCat FleasGenetic VariationCicadasIndustrial MelanismCodling MothInsecticide and Acaricide Resistance ColeopteraIsland BiogeographyCollembolaMimicryColorado Potato BeetleNomenclature and Classication, Principles ofCricketsPhylogeny of Insects Daddy-Long-LegsSexual Selection DermapteraSocialityDipluraWolbachiaDiptera Drosophila melanogasterReproduction Dung BeetlesAccessory Glands EmbiidinaEgg CoveringsEndopterygotaEggs EphemeropteraEmbryogenesisExopterygota 19. Contents by Subject Area xixFire Ants Predation/Predatory InsectsGrylloblattodea Symbionts Aiding DigestionGypsy MothSymbionts, BacterialHouse Fly VenomHymenoptera Veterinary EntomologyInsecta, Overview WolbachiaIsopteraJapanese Beetle Interactions With HumansJune BeetlesApis SpeciesLadybugsBedbugsLepidoptera Bee ProductsLice, Human BeekeepingLocusts Blood SuckingMantodeaBombyx moriMantophasmatodeaBubonic PlagueMecoptera Chiggers and Other Disease-Causing MitesMegaloptera Commercialization of Insects and Their ProductsMites Cultural EntomologyMonarchsDDTMosquitoesDengueNeotropical African BeesEkbom SyndromeNeuropteraExtension EntomologyOdonata Folk Beliefs and SuperstitionsOrthopteraFood, Insects asPhasmidaForensic EntomologyPhthirapteraHoneyPlecopteraHuman History, Insects Effect onProsorrhyncha InsecticidesProtura Integrated Pest ManagementPsocopteraLice, HumanPterygota MalariaRaphidioptera Medical EntomologyScorpions Medicine, Insects inSiphonapteraMosquitoesSpiders Museums and Display CollectionsSternorrhynchaRegulatory EntomologyStrepsipteraRiver BlindnessThysanopteraSilk Production in InsectsTicks Tsetse FlyTrichoptera Yellow FeverVinegaroons Zoonotic Agents, Arthropod-BorneWaspsZorapteraZygentoma HabitatsAquatic HabitatsInteractions With Other Organisms Cave InsectsAposematic Coloration Forest HabitatsBlood Sucking Grassland HabitatsCat Fleas Marine InsectsChiggers and Other Disease-Causing MitesSoil HabitatsDefensive BehaviorStored Products as HabitatsDog Heartworm Urban HabitatsFeeding BehaviorHost Seeking, by ParasitoidsHost Seeking, for PlantsEcologyHyperparasitism Agricultural EntomologyMimicry Aposematic ColorationParasitoids BiodiversityPathogens of InsectsBiogeographical PatternsPhoresy Biological Control of Insect PestsPhytophagous InsectsBorers 20. xx Contents by Subject Area Coevolution History And Methodology ConservationAmber Crypsis Biotechnology and Insects DDT Cell Culture Endangered InsectsCollection and Preservation Gallmaking and InsectsCultural Entomology Genetic Variation Entomological Societies Genetically Modied PlantsFolk Beliefs and Superstitions Greenhouse Gases, Global Warming, and Insects Forensic Entomology HoneydewGenetically Modied Plants Hyperparasitism Genetic Engineering InsecticidesGenomics Insectivorous PlantsHistory of Entomology Insectivorous Vertebrates Human History, Insects Effect on Integrated Pest ManagementInsect Zoos Introduced InsectsMedical Entomology Migration Movies, Insects in Mimicry Museums and Display Collections Parasitoids Nomenclature and Classication, Principles of Pathogens of InsectsPhotography of Insects Phoresy Rearing of Insects Physical Control of Insect PestsResearch Tools, Insects as Phytophagous InsectsSericulture Phytotoxicity Stamps, Insects and Plant Diseases and InsectsSterile Insect Technique PlantInsect Interactions Teaching Resources Pollination and Pollinators Veterinary Entomology Pollution, Insect Response to Population Ecology Predation/Predatory Insects Swimming, Lake Insects Swimming and Other Movements, Stream Insects 21. CONTRIBUTORSJohn H. AcornHorst AspckUniversity of Alberta, CanadaMedical University of ViennaTeaching Resources RaphidiopteraMichael E. Adams Ulrike AspckUniversity of California, RiversideNaturhistorisches Museum WienDevelopment, Hormonal Control of RaphidiopteraEcdysteroids Peter W. AtkinsonJuvenile Hormones University of California, RiversidePeter H. Adler Genetic EngineeringClemson University GenomicsLegs Arnd BaumannGilberto S. AlbuquerqueForschungszentrum Jlich, GermanyUniversidade Estadual do Norte Fluminense, BrazilBiogenic AminesNeuroptera Nancy E. BeckageRichard D. Alexander University of California, RiversideUniversity of Michigan ImmunologyCrickets Peter Bellinger*Miriam AltsteinCalifornia State University, NorthridgeAgricultural Research Organization, Volcani Center, Israel CollembolaNeuropeptides May R. BerenbaumSvend O. AndersenUniversity of IllinoisThe Royal Danish Academy of Sciences and Letters Movies, Insects inCuticle Martin B. BergExoskeleton Loyola University ChicagoIntegument Growth, IndividualNorman H. Anderson Elizabeth A. BernaysOregon State University University of ArizonaMegaloptera Host Seeking, for PlantsDavid A. Andow Phytophagous InsectsUniversity of Minnesota, St. Paul Christer BjrkmanGenetically Modied Plants Swedish University of Agricultural Sciences andMichael F. Antolin Stockholm UniversityColorado State UniversityBody SizeSex Determination Scott Hoffman BlackPeter ArensburgerThe Xerces SocietyUniversity of California, RiversideEndangered InsectsGenomics Seth S. BlairLarry G. ArlianUniversity of Wisconsin, MadisonWright State UniversityImaginal DiscsChiggers and Other Disease-Causing Mites Wolfgang Blenau Universitt Potsdam, Germany*Deceased November 20, 2000. Biogenic Amines 22. xxii ContributorsMurray S. BlumDonald G. CochranUniversity of Georgia Virginia Polytechnic Institute and State UniversityChemical DefenseBlattodeaBryony C. Bonning Ephraim CohenIowa State University The Hebrew University of JerusalemBiotechnology and Insects ChitinFat BodyTimothy J. BradleyUniversity of California, Irvine Andrej CoklExcretion National Institute of Biology, SloveniaVibrational CommunicationPaul M. BrakeeldLeiden University, The NetherlandsGregory W. CourtneyCrypsis Iowa State UniversityDipteraJohn E. BrittainNatural History Museum, University of OsloCharles V. Covell, Jr.Ephemeroptera University of LouisvilleCollection and PreservationLincoln P. BrowerStamps, Insects andSweet Briar CollegeMonarchsCatherine CraigHarvard University/Tufts UniversityAndreas BruneSilk Production in InsectsMax Planck Institute for Terrestrial Microbiology,Marburg, GermanyEva CraneSymbionts Aiding DigestionInternational Bee Research AssociationApis SpeciesWendell E. BurkholderBeekeepingUniversity of Wisconsin, MadisonBee ProductsStored Products as HabitatsHoneyGeorge W. ByersPeter S. CranstonUniversity of KansasUniversity of California, DavisMecopteraPhylogeny of InsectsRing T. CardCharles R. CrumlyUniversity of California, RiversideUniversity of California Press, BerkeleyApterygotaInsectivorous VertebratesAutotomyEndopterygota Gregory A. DahlemExopterygotaNorthern Kentucky UniversityInsecta, Overview House FlyOrientation Donald L. DahlstenPheromonesUniversity of California, BerkeleyPterygota Gallmaking and InsectsR. F. Chapman*Gene R. DeFoliartUniversity of Arizona University of Wisconsin, MadisonFeeding BehaviorFood, Insects asLocustsMouthpartsIan DenholmRothamsted ResearchLanna Cheng Insecticide and Acaricide ResistanceScripps Institution of OceanographyMarine InsectsDavid L. DenlingerOhio State UniversityKenneth A. Christiansen Cold/Heat ProtectionGrinnell College, Grinnell, IADiapauseCollembolaGregor J. DevineThomas M. Clark Rothamsted ResearchIndiana University, South BendInsecticide and Acaricide ResistanceWater and Ion Balance, Hormonal Control of Deceased September 6, 2007.*Deceased May 2, 2003. Deceased September 3, 2003. 23. Contributors xxiiiMichael DickinsonAndrew S. FrenchCalifornia Institute of Technology Dalhousie University, Halifax, Nova ScotiaFlight MechanoreceptionChristopher H. DietrichDouglas J. FutuymaIllinois Natural History SurveyState University of New York, Stony BrookAuchenorrhynchaCoevolutionHugh DingleErin C. GentryUniversity of California, DavisUniversity of FloridaMigrationBioluminescenceAngela E. DouglasAlec C. GerryCornell University, USAUniversity of California, RiversideHoneydew BedbugsRobert V. Dowell Helen GhiradellaCalifornia Department of Food and AgricultureState University of New York, AlbanyRegulatory EntomologyColorationRobert DudleyRosemary G. GillespieUniversity of California, Berkeley University of California, BerkeleyFlight Island BiogeographyJohn D. EdmanSpidersUniversity of California, Davis Gonzalo GiribetMedical Entomology Harvard UniversityBruce F. EldridgeDaddy-Long-LegsUniversity of California, Davis M. Lee GoffMosquitoes Chaminade University of HonoluluJoseph S. Elkinton Forensic EntomologyUniversity of MassachusettsGypsy Moth Gordon GordhPopulation Ecology U.S. Department of Agriculture Anatomy: Head, Thorax, Abdomen, and GenitaliaMichael S. EngelUniversity of Kansas Karl GotthardZorapteraSwedish University of Agricultural Sciences and Stockholm UniversityJoachim ErberBody SizeTechnische Universitt Berlin, GermanyBiogenic AminesMiodrag Grbic University of Western Ontario, CanadaBrian A. Federici EmbryogenesisUniversity of California, RiversidePathogens of Insects Les Greenberg University of California, RiversideLewis J. Feldman Fire AntsUniversity of California, BerkeleyInsectivorous Plants David GrimaldiCllia FerreiraAmerican Museum of Natural History, New YorkUniversity of So Paulo, BrazilFossil RecordDigestive System Christin GrossmannR. Nelson Foster Museum fr Tierkunde, DresdenU.S. Department of Agriculture MantophasmatodeaBoll WeevilPenny J. GullanGordon W. FrankieUniversity of California, DavisUniversity of California, Berkeley Phylogeny of InsectsPollination and PollinatorsSternorrhynchaNigel R. FranksDarryl T. GwynneUniversity of BristolUniversity of TorontoAnts Mating Behaviors 24. xxivContributorsGuy HallmanRon HoyUSDA-ARS, Weslaco, Texas Cornell UniversityPhysical Control of Insect Pests HearingJ. Daniel Hare Lawrence E. HurdUniversity of California, RiversideWashington and Lee UniversityHost Seeking, by Parasitoids MantodeaPredation/Predatory Insects Sigfrid IngrischJon F. HarrisonMuseum Koenig, BonnArizona State University GrylloblattodeaRespiratory System OrthopteraTracheal System Michael E. IrwinMichael W. Hastriter University of IllinoisBrigham Young University Commercialization of Insects and Their ProductsSiphonaptera Rudolf JanderDavid H. HeadrickUniversity of Kansas, LawrenceCalifornia Polytechnic State UniversityMagnetic SenseAnatomy: Head, Thorax, Abdomen, and Genitalia Frans JanssensBernd Heinrich University of Antwerp, BelgiumUniversity of VermontCollembolaThermoregulation Robert L. JeanneDavid W. HeldUniversity of WisconsinAuburn UniversityNest BuildingJapanese BeetleJune Beetles Mathieu Joron Musum National dHistoire Naturelle, FranceRonald A. Hellenthal Aposematic ColorationUniversity of Notre Dame MimicryPhthiraptera Robert JosephsonJorge HendrichsUniversity of California, IrvineJFAO/IAE Division, Vienna, Austria Muscle SystemSterile Insect Technique Gail E. KampmeierAdam D. Henk Illinois Natural History SurveyColorado State UniversityCommercialization of Insects and Their ProductsSex Determination Kenneth Y. KaneshiroNancy C. HinkleUniversity of HawaiiUniversity of GeorgiaSexual SelectionCat FleasEkbom Syndrome Michael R. KanostExtension Entomology Kansas State University HemolymphM. S. HoddleUniversity of California, RiversideAlan I. KaplanBiological Control of Insect Pests East Bay Regional Park District, Berkeley, CA Entomological SocietiesJames N. HogueCalifornia State University, NorthridgeJoe B. KeiperCultural EntomologyThe Cleveland Museum of Natural HistoryFolk Beliefs and Superstitions DipteraHuman History, Insects Effect on George G. KennedyMarilyn A. Houck North Carolina State UniversityTexas Tech UniversityColorado Potato BeetlePhoresyHistory of EntomologyFrancis G. Howarth Lawrence R. KirkendallB. P. Bishop Museum, Honolulu, HawaiiUniversity of Bergen, NorwayCave Insects Parthenogenesis in Insects and Mites 25. ContributorsxxvKlaus-Dieter Klass James E. LloydMuseum fr Tierkunde, DresdenUniversity of FloridaMantophasmatodea BioluminescenceJohn Klotz Catherine LoudonUniversity of California, RiversideUniversity of California, IrvineMagnetic Sense AntennaeMarc J. KlowdenDwight E. LynnUniversity of IdahoU.S. Department of AgricultureOviposition Behavior Cell CultureReproduction, MaleReproduction, Male: Hormonal Control ofMichael E. N. Majerus*SpermathecaUniversity of CambridgeSpermatophoreIndustrial Melanism LadybugsMarkus KochSymbionts, BacterialFreie Universitt Berlin, GermanyDipluraJon H. MartinProturaThe Natural History Museum, London SternorrhynchaMarcos KoganOregon State UniversitySinzo MasakiAgricultural EntomologyHirosaki UniversityIntegrated Pest Management AestivationAndreas Kruess Linda J. MasonUniversity of Gttingen, Germany Purdue UniversityGrassland Habitats Extension EntomologyMichael F. LandFumio MatsumuraUniversity of Sussex, Brighton University of California, DavisEyes and VisionDDTRobert S. Lane InsecticidesUniversity of California, Berkeley Joseph V. McHughZoonotic Agents, Arthropod-Borne University of GeorgiaStephen G. A. Leak ColeopteraInternational Trypanotolerance Centre, The GambiaTerri L. MeinkingTsetse Fly Global Health Associates of Miami (GHAM)Richard E. Lee, Jr.Lice, HumanMiami University, Oxford, OH Richard W. MerrittDormancy Michigan State UniversityHibernationAquatic HabitatsM. J. Lehane DipteraUniversity of Wales, BangorGrowth, IndividualBlood SuckingJocelyn G. MillarNorman C. Leppla University of California, RiversideUniversity of FloridaPheromonesRearing of Insects Thomas A. MillerRichard J. LeskoskyUniversity of California, RiversideUniversity of Illinois Circulatory SystemMovies, Insects in Nick MillsVernard R. Lewis University of California, BerkeleyUniversity of California, Berkeley ParasitoidsIsoptera B. K. MitchellJames K. LiebherrUniversity of Alberta, CanadaCornell University ChemoreceptionColeopteraPaul Z. LiuUniversity of California, BerkeleySegmentation *Deceased January 27, 2009. 26. xxviContributorsEdward L. Mockford Sean ODonnellIllinois State UniversityUniversity of WashingtonPsocoptera Caste ColoniesMark W. MoffettUniversity of California, Berkeley Patrick M. OGradyPhotography of Insects University of California, Berkeley Drosophila melanogasterThomas P. MonathAcambis Inc., Cambridge, MADaniel OtteYellow Fever Philadelphia Academy of Natural Sciences CricketsJohn C. MorseClemson University Terry L. PageTrichopteraVanderbilt University Circadian RhythmsMax S. MouldsAustralian Museum, SydneyTimothy D. PaineCicadasUniversity of California, Riverside BorersLaurence A. MoundCSIRO Entomology, Canberra, AustraliaJames O. PalmerThysanoptera Allegheny College DermapteraBradley A. Mullens Daniel R. PapajUniversity of California, Riverside University of ArizonaVeterinary Entomology LearningWerner Nachtigall Gnther PassUniversitt der Saarlandes, Germany University of Vienna, AustriaSwimming, Lake Insects Circulatory SystemLisa Nagy Nipam H. PatelUniversity of Arizona University of California, BerkeleyEmbryogenesis SegmentationMaria Navajas Mats W. PetterssonInstitut National de la Recherche Agronomique (INRA), Swedish University of Agricultural Sciences andCentre de Biologie et Gestion des Populations, Stockholm UniversityMontferrier sur Lez, France Body SizeGenetic Variation John D. PintoOldr ich Nedve d University of California, RiversideUniversity of South Bohemia, and Institute of Entomology,HypermetamorphosisCzech RepublicTemperature, Effects on Development and Growth Rudy Plarre Federal German Institute for Materials Research andTim R. New Testing, GermanyLa Trobe University, Australia Stored Products as HabitatsConservation Edward G. PlatzerGordon M. NishidaUniversity of California, RiversideUniversity of California, Berkeley Dog HeartwormMuseums and Display Collections George Poinar, Jr.Benjamin B. NormarkOregon State UniversityUniversity of Massachusetts, Amherst AmberParthenogenesis in Insects and Mites Daniel A. PotterDavid A. OBrochta University of KentuckyUniversity of Maryland Biotechnology Institute Japanese BeetleGenetic EngineeringJune BeetlesBarry M. OConnor Jerry A. PowellUniversity of Michigan University of California, BerkeleyMitesLepidoptera 27. ContributorsxxviiRoger D. Price George K. RoderickUniversity of MinnesotaUniversity of California, BerkeleyPhthiraptera Genetic Variation Island BiogeographyRonald Prokopy*University of MassachusettsDavid M. RosenbergAgricultural EntomologyFreshwater Institute, Winnipeg, CanadaIntegrated Pest Management Pollution, Insect Response toAlexander H. Purcell Edward S. RossUniversity of California, Berkeley California Academy of SciencesPhytotoxicityEmbiidinaPlant Diseases and Insects Michael K. RustDonald L. J. QuickeUniversity of California, RiversideImperial College, University of London Cat FleasHymenopteraUrban HabitatsFrank J. Radovsky Michel SartoriOregon State University Museum of Zoology, LausanneNeosomy EphemeropteraSusan M. Rankin Leslie Saul-GershenzAllegheny College Center for Ecosystem Survival, San Francisco, CADermaptera Insect ZoosWilliam K. ReisenUniversity of California, DavisCarl W. SchaeferMalariaUniversity of Connecticut ProsorrhynchaD. C. F. RentzCalifornia Academy of Sciences, San FranciscoKatherine N. SchickGrylloblattodeaUniversity of California, BerkeleyOrthoptera Gallmaking and InsectsVincent H. ReshJustin O. SchmidtUniversity of California, Berkeley Southwestern Biological Institute, TucsonApterygota Defensive BehaviorAutohemorrhage VenomInsecta, OverviewVinegaroonsPollution, Insect Response toWaspsPterygotaRiver BlindnessMichelle Pellissier Scott University of New HampshireLynn M. RiddifordParental CareUniversity of WashingtonMoltingThomas W. Scott University of California, DavisJames Ridsdill-Smith DengueUniversity of Western Australia, AustraliaDung Beetles J. Mark Scriber Michigan State UniversityRoy E. Ritzmann PlantInsect InteractionsCase Western Reserve UniversityWalking and JumpingFrantiek SehnalAlan RobinsonBiology Centre ASCRJFAO/IAE Division, Vienna, Austria Silk Production in InsectsSterile Insect Technique Irwin W. ShermanGene E. Robinson University of California, Riverside/The Scripps ResearchUniversity of Illinois, Urbana-Champaign Institute, La JollaDivision of Labor in Insect SocietiesBubonic Plague Ronald A. Sherman University of California, Irvine*Deceased May 14, 2004.Medicine, Insects in 28. xxviiiContributorsDaniel SimberloffNigel E. StorkUniversity of TennesseeJames Cook University, AustraliaIntroduced Insects BiodiversityLeigh W. Simmons Richard StouthamerUniversity of Western Australia, Australia University of California, RiversideDung Beetles WolbachiaS. J. SimpsonMichael R. StrandUniversity of Sydney University of GeorgiaNutritionPolyembryonyScott R. Smedley Nicholas J. StrausfeldTrinity College, Connecticut University of ArizonaPuddling BehaviorBrain and Optic LobesEdward H. SmithHelmut SturmCornell University (Emeritus), Asheville, NC University Hildesheim, GermanyHistory of EntomologyArchaeognathaDaniel E. Sonenshine ZygentomaOld Dominion UniversityR. K. SuarezTicksUniversity of California, Santa BarbaraJohn T. Sorensen MetabolismCalifornia Department of Food and AgricultureDaniel J. SullivanAphids Fordham UniversityJoseph C. Spagna HyperparasitismUniversity of California, Berkeley Satoshi TakedaSpidersNational Institute of Agrobiological Sciences, JapanBeverly Sparks Bombyx moriUniversity of GeorgiaSericultureExtension Entomology Catherine A. TauberFelix A. H. Sperling Cornell UniversityUniversity of Alberta, CanadaNeuropteraTeaching Resources Maurice J. TauberBernhard StatznerCornell UniversityCNRSUniversit Lyon 1, France NeuropteraSwimming and Other Movements, Stream Insects Orley R. TaylorIngolf Steffan-DewenterUniversity of KansasUniversity of Gttingen, Germany Neotropical African BeesGrassland Habitats William H. TelferFrederick W. Stehr University of PennsylvaniaMichigan State UniversityVitellogenesisCaterpillars K. J. TennessenChrysalisFlorida State Collection of Arthropods, Gainesville, FLCocoon OdonataLarvaMetamorphosisWalter R. TerraOcelli and StemmataUniversity of So Paulo, BrazilPupa and PupariumDigestion Digestive SystemKenneth W. StewartUniversity of North TexasCarsten ThiesPlecoptera University of Gttingen, Germany Grassland HabitatsPeter StilingUniversity of South FloridaF. Christian ThompsonGreenhouse Gases, Global Warming, and InsectsU.S. Department of Agriculture Nomenclature and Classication, Principles ofAndrew J. StorerMichigan Technological University, HoughtonS. N. ThompsonForest HabitatsUniversity of California, Riverside 29. Contributors xxixMetabolismPhyllis WeintraubNutrition Agricultural Research Organization, GilatResearch Center, IsraelJames H. ThorpPhysical Control of Insect PestsUniversity of KansasArthropoda and Related Groups Christiane WeirauchUniversity of California, RiversideRobbin W. Thorp BedbugsUniversity of California, DavisPollination and Pollinators Stephen C. WelterUniversity of California, BerkeleyErich H. TilgnerCodling MothUniversity of GeorgiaPhasmidaRonald M. WeselohThe Connecticut Agricultural Experiment StationPivi H. Torkkeli Host Seeking, by ParasitoidsDalhousie University, Halifax, Nova ScotiaPredation/Predatory InsectsMechanoreceptionDiana E. WheelerJames F. A. Traniello University of ArizonaBoston University Accessory GlandsRecruitment Communication EggsTeja Tscharntke Egg CoveringsUniversity of Gttingen, GermanyOvariolesGrassland HabitatsReproduction, FemaleReproduction, Female: Hormonal Control ofKaren M. VailUniversity of Tennessee, KnoxvilleMichael F. WhitingExtension EntomologyBrigham Young UniversitySiphonapteraR. G. Van DriescheStrepsipteraUniversity of Massachusetts, AmherstBiological Control of Insect PestsKipling W. WillUniversity of California, BerkeleyMace VaughanResearch Tools, Insects asThe Xerces SocietyStanley C. WilliamsEndangered InsectsSan Francisco State UniversityCharles Vincent ScorpionsAgriculture and Agri-Food Canada, QuebecShaun L. WintertonPhysical Control of Insect PestsNorth Carolina State UniversityMeta Virant-DoberletScales and SetaeNational Institute of Biology, SloveniaDavid L. WoodVibrational CommunicationUniversity of California, BerkeleyP. Kirk VisscherForest HabitatsUniversity of California, RiversideRobin J. WoottonDance LanguageUniversity of ExeterHomeostasis, BehavioralWingsHoneyJayne YackPatricia J. VittumCarleton UniversityUniversity of Massachusetts HearingSoil HabitatsJames E. ZablotnyGregory P. Walker U.S. Department of AgricultureUniversity of California, Riverside SocialitySalivary GlandsSasha N. ZillJ. Bruce WallaceMarshall UniversityUniversity of Georgia Walking and JumpingAquatic HabitatsPeter ZwickGraham C. WebbMax-Planck-Institut fr Limnologie, GermanyThe University of Adelaide, Australia Biogeographical PatternsChromosomes 30. GUIDE TO THEENCYCLOPEDIAThe Encyclopedia of Insects is a complete source of information ARTICLE FORMATon the subject of insects, contained within a single volume. EachEach article in this Encyclopedia begins with an introductory par-article in the Encyclopedia provides an overview of the selectedagraph that denes the topic being discussed and indicates its signi-topic to inform a broad spectrum of readers, from insect biologists cance. For example, the article Exoskeleton begins as follows:and scientists conducting research in related areas, to students andthe interested general public. The exoskeleton is noncellular material that is located on topIn order that you, the reader, will derive the maximum benetof the epidermal cell layer and constitutes the outermost partfrom the Encyclopedia of Insects, we have provided this Guide. Itof the integument. The local properties and appearance of theexplains how the book is organized and how the information withinexoskeleton are highly variable, and nearly all visible featuresits pages can be located.of an insect result from the exoskeleton. The exoskeleton serves as a barrier between the interior of the insect and the environment, preventing desiccation and the penetration ofSUBJECT AREASmicroorganisms. Muscles governing the insects movements The Encyclopedia of Insects presents 273 separate articles on the are attached to the exoskeleton.entire range of entomological study. Articles in the Encyclopedia fallMajor headings highlight important subtopics that are discussedwithin 12 general subject areas, as follows:in the article. For example, the article Flight includes the follow- Anatomy ing topics: Evolution of Flight; Aerodynamics; Neural Control; PhysiologyEnergetics; Ecology and Diversity. Behavior Evolution CROSS-REFERENCES Reproduction Development and Metamorphosis The Encyclopedia of Insects has an extensive system of cross- Major Groups and Notable Formsreferencing. References to other articles may appear either as mar- Interactions with Other Organisms ginal headings within the AZ topical sequence, or as indications of Interactions with Humansrelated topics at the end of a particular article. HabitatsAs an example of the rst type of reference cited above, the fol- Ecology lowing marginal entry appears in the AZ article list between the History and Methodology entries Bee Products and Biodiversity: Beetle see ColeopteraORGANIZATIONThis reference indicates that the topic of Beetles is discussedelsewhere, under the article title Coleoptera, which is the name ofThe Encyclopedia of Insects is organized to provide the maximumthe order including this group.ease of use for its readers. All of the articles are arranged in a singleAn example of the second type, a cross-reference at the end of analphabetical sequence by title. An alphabetical Table of Contentsarticle, can be found in the entry DDT. This article concludes withfor the articles can be found beginning on p. v of this introductorythe statement:section.As a reader of the Encyclopedia, you can use this alphabeticalTable of Contents by itself to locate a topic. Or you can rst identifySee Also the Following Articlesthe topic in the Contents by Subject Area (p. xvii) and then go to theInsecticides Integrated Pest Management Pollutionalphabetical Table to nd the page location. This reference indicates that these three related articles all pro-In order to identify articles more easily, article titles begin withvide some additional information about DDT.the key word or phrase indicating the topic, with any descrip-tive terms following this. For example, Temperature, Effects onDevelopment and Growth is the title assigned to an article, rather BIBLIOGRAPHYthan Effects of Temperature on Development and Growth,The Bibliography section appears as the last element of an article,because the specic term Temperature is the key word. under the heading Further Reading. This section lists recent 31. Guide to the Encyclopedia xxxisecondary sources that will aid the reader in locating more detailed in this Encyclopedia. The terms were identied by the contributorsor technical information on the topic at hand. Review articles and as helpful to the understanding of their entries, and they have beenresearch papers that are important to a more detailed understandingdened by these authors according to their use in the actual articles.of the topic are also listed here. The Bibliography entries in this Encyclopedia are for the benetINDEXof the reader, to provide references for further reading or additionalresearch on the given topic. Thus they typically consist of a limited The Subject Index for the Encyclopedia of Insects contains morenumber of entries. They are not intended to represent a complete than 7000 entries. Within the entry for a given topic, references tolisting of all the materials consulted by the author(s) in preparing the general coverage of the topic appear rst, such as a complete articlearticle. The Bibliography is in effect an extension of the article itself, on the subject. References to more specic aspects of the topic thenand it represents the authors choice as to the best sources available appear below this in an indented list.for additional information.GLOSSARY The Encyclopedia of Insects presents an additional resource forthe reader, following the AZ text. A comprehensive glossary providesdenitions for more than 750 specialized terms used in the articles 32. FOREWORDI would say that creating an encyclopedia of insects was a her- on, contribute millions more. Insect-based biological control of bothculean task, but I think that sells the enterprise short. After all, insect and weed pests is worth additional millions in reclaimed landHercules only had twelve labors assigned to him, and twelveand crop production, and even insect disposal of dung and otheryears to complete themwith insects, there are over 900,000 differ-waste materials, although decidedly unglamorous, is economicallyent species and many, many more stories to tell. Twelve years from signicant in elds, rangelands, and forests throughout the country.now, there will likely be even more. Why, then, would anyone under-So, for no reason other than economic self-interest, theres rea-take the seemingly impossible task of compiling an encyclopedia of son enough for creating an encyclopedia of insects. But what caninsects? To an entomologist, the answer is obvious. For one thing, be learned from insects that cant be learned from an encyclopediatheres the numbers argumentover 70% of all known species are of any other abundant group of organisms? Basically, the biologyinsects, so if any group merits attention in encyclopedic form, surely of insects is the biology of small size. Small size, which has been inits the one that happens to dominate the planet. Moreover, owinglarge part responsible for the overwhelming success of the taxon, atin large part to their staggering diversity, insects are in more differ- the same time imposes major limits on the taxon. The range in sizeent places in the world than virtually any other organism. There are of living organisms, on earth at least, encompasses some 13 ordersinsects in habitats ranging from the High Arctic to tropical rainfor-of magnitude (from a 100 metric ton blue whale to rotifers weigh-ests to petroleum pools to glaciers to mines a mile below the sur- ing less than 0.01 mg). Insects range over ve orders of magnitudeface to caves to sea lion nostrils and horse intestines. About the onlyfrom 30-g beetles to 0.03-g fairyiesso eight orders of magnitudeplace where insects are conspicuously absent is in the deep oceanare missing in the class Insecta. Problems at the upper limit involve(actually, in deep water in general), an anomaly that has frustrated support, transport, and overcoming inertia, issues clearly not criticalmore than a few entomologists who have grown accustomed to world for organisms, like insects, at the lower end of the range.domination. Then theres the fact that insects have been around forWe humans, in the grand scheme of things, are big creatures andlonger than most other high-prole life-forms. The rst proto-insectsas a consequence we interact with the biological and physical worlddate back some 400 million years; by contrast, mammals have been entirely differently. Rules that constrain human biology often arearound only about 230 million years and humans (depending on how suspended for insects, which operate by a completely different set oftheyre dened) a measly one million years.rules. The constraints and benets of small size are reected in every Probably the best justication for an encyclopedia devoted to aspect of insect biology. They hear, smell, taste, and sense the worldinsects is that insects have a direct and especially economic impact in every other way with abilities that stagger the imagination. Theyon humans. In the United States alone, insects cause billions of dol-are capable of physical feats that seem impossiblemost y, somelars in losses to staple crops, fruit crops, truck crops, greenhouse and glow in the dark, and others control the sex of their offspring andnursery products, forest products, livestock, stored grain and pack- even occasionally engage in virgin birth, to cite a few examples. Theiraged food, clothing, household goods and furniture, and just about generation times are so short and reproductive rates so high thatanything else people try to grow or build for sale or for their ownthey can adapt and evolve at rates that continually surprise (and sty-consumption. Beyond the balance sheet, they cause incalculable mie) us. The environment is patchier to smaller organisms, whichlosses as vectors of human pathogens. Theyre involved in transmis-can divide resources more nely than can large, lumbering species.sion of malaria, yellow fever, typhus, plague, dengue, various forms Thus, they can make a living on resources so rare or so nutrient-poorof encephalitis, relapsing fever, river blindness, lariasis, sleeping that it dees belief, such as nectar, dead bodies, and even dung.sickness, and innumerable other debilitating or even fatal diseases, So theyre profoundly different from humans and other big ani-not just abroad in exotic climes but here in the United States as well.mals, and the study of insects can offer many insights into life onAll told, insects represent a drag on the economy unequaled by any earth that simply couldnt be gained from a study of big creatures.other single class of organisms, a seemingly compelling reason for By the same token, though, they are cut from the same cloththekeeping track of them in encyclopedic form.same basic building blocks of life, same genetic code, and the like In the interests of fairness, however, it should be mentioned thatand their utility as research organisms has provided insights into allinsects also amass economic benets in a magnitude unequaled bylife on the planet.most invertebrates (or even, arguably, by most vertebrates). Insect- The Encyclopedia of Insects contains contributions from somepollinated crops in the United States exceed $9 billion in value annu- of the greatest names in entomology today. Such a work has to beally, and insect products, including honey, wax, lacquer, silk, and so a collective effort because nobody can be an expert in everything 33. Foreword xxxiiientomological. Even writing a foreword for such a wide-ranging vol- activities than perhaps any other class of organisms, to be the ulti-ume is a daunting task. To be such an expert would mean mastering mate authority on insects also means mastering the minutiae of his-every biological science from molecular biology (in which the fruit tory, economics, art, literature, politics, and even popular culture.y Drosophila melanogaster serves as a premier model organism) to Nobody can master all of that informationand thats why this ency-ecosystem ecology (in which insects play an important role in rates clopedia is such a welcome volume.of nutrient turnover and energy ow). But, because insects, throughtheir ubiquity and diversity, have had a greater inuence on human May R. Berenbaum 34. PREFACE TO THE SECOND EDITIONW e are pleased to have had the privilege of continuing as edi- Between publication of the rst and second edition of the tors for the second edition of the Encyclopedia of Insects. Encyclopedia several contributors to the rst edition died, including This edition contains several new entries and updates ofPeter Bellinger, Donald Dahlsten, Reginald Chapman, Eva Crane,almost all of the original entries. Many new illustrations have been Michael Majerus, and Ronald Prokopy. Their substantial contributionsadded and references for further readings have been updated. to entomology will long be remembered. The rst edition of the Encyclopedia of Insects was well received.We thank the staff of Elsevier Press for their assistance on thisAwards garnered include: The Most Outstanding Single-Volume project. Christine Minihane originally proposed the preparation of aReference in Science, presented by The Association of Americansecond edition and Andy Richford shepherded it to completion. PatPublishers for 2003; An Outstanding Academic Title, 2003, by Gonzalez was invaluable in managing the ow of revised manuscripts.CHOICE magazine; and Best of Reference, 2003, by both theStephen Pegg and Mani Prabakaran oversaw the printing process. WeNew York Public Librarians and the American Library Journal 2003.especially thank Alan Kaplan for reading over the nal text for consist-These are a tribute to the quality of contributions to that volume.ency and accuracy. David Hawks provided the cover photograph.We anticipate that this updated, second edition will play the same We are pleased to dedicate our efforts in producing this secondrole in assisting students, teachers, and researchers in the entomo- edition to our mentors and professors, whose inuence we still feellogical and biological sciences, along with interested readers among 40 years later: Stuart Neff, Louis Krumholz, Jack Franclemont andthe general public, in obtaining up-to-date and accurate information Wendell Roelofs.about these fascinating organisms. Vincent H. Resh and Ring T. Card 35. PREFACEI nsects are ever present in human lives. They are at once awecompleted. This is the rst order of insects to be described in over inspiring, fascinating, beautiful, and, at the same time, a scourge 80 years, and we are pleased to be able to include it as an entry, of humans because of food loss and disease. Yet despite their further underscoring that there is much left to learn about insects.negative effects, we depend on insects for pollination and for their Some topics, especially the poster insectsthose well-known taxaproducts. As insects are the largest living group on earth (75% of below the level of orders for which entries are presentedmay notall animal species), any understanding of ecological interactions at cover all that are desired by some readers. Given insect biodiversity,local or global scales depends on our knowledge about them. Givenyour indulgence is requested.the current interest in biodiversity, and its loss, it must be remem-We have gathered over 260 experts worldwide to write on thebered that insects represent the major part of existing biodiversity.entries that we have selected for inclusion. These specialists, ofAesthetically, insect images are often with us as well: early images course, have depended on the contributions of thousands of theirinclude Egyptian amulets of sacred scarabs; modern images includeentomological predecessors. Because the modern study of entomol-dragony jewelry, buttery stationery, and childrens puppets. ogy is interdisciplinary, we enlisted experts ranging from arachnolo-The idea of an Encyclopedia of Insects is new, but the concept ofgists to specialists in zoonotic diseases. Given that the two of us havean encyclopedia is quite old. In 1745, Diderot and DAlembert askedspent over 25 combined years as editors of the Annual Review ofthe best minds of their eraincluding Voltaire and MontesquieuEntomology, many of our contributors were also writers for that peri-to prepare entries that would compile existing human knowledge inodical. We thank our contributors for putting up with our compulsiveone place: the worlds rst encyclopedia. It took over 20 years to n- editing, requests for rewrites, and seemingly endless questions.ish the rst edition, which became one of the worlds rst best-sellingOur intended audience is not entomological specialists but ento-books and a triumph of the Enlightenment.mological generalists, whether they be students, teachers, hobbyists,What do we intend this encyclopedia to be? Our goal is to con- or interested nonscientists. Therefore, to cover the diverse interests ofvey the exciting, dynamic story of what entomology is today. It is this readership, we have included not just purely scientic aspects ofintended to be a concise, integrated summary of current knowl- the study of insects, but cultural (and pop-cultural) aspects as well.edge and historical background on each of the nearly 300 entries We thank the staff of Academic Press for their encouragementpresented. Our intention has been to make the encyclopedia scien-and assistance on this project. Chuck Crumly had the original con-tically uncompromising; it is to be comprehensive but not exhaus- cept for this encyclopedia, convinced us of its merit, and helped ustive. Cross-references point the reader to related topics, and further greatly in dening the format. Chris Morris provided suggestionsreading lists at the end of each article allow readers to go into topics about its development. Jocelyn Lofstrom and Joanna Dinsmorein more detail. The presence of a certain degree of overlap is inten-guided the book through printing. Gail Rice managed the ow oftional, because each article is meant to be self-contained.manuscripts and revisions with skill and grace, and made many valu-The Encyclopedia of Insects also includes organisms that are able suggestions. Julie Todd of Iowa State University provided a cru-related to insects and often included in the purview of entomol- cial nal edit of the completed articles. All these professionals haveogy. Therefore, besides the members of the class Insectathe truehelped make this a rewarding and fascinating endeavor.insectsthe biology of spiders, mites, and related arthropods is We dedicate our efforts in editing the Encyclopedia of Insects toincluded. The core of this encyclopedia consists of the articles on theour wives, Cheryl and Anja; their contributions to our entomologicaltaxonomic groupsthe 30 or so generally accepted orders of insects,and personal lives have been indescribable.the processes that insects depend on for their survival and success,and the range of habitats they occupy. The fact that entomology isa dynamic eld is emphasized by the discovery of a new order ofinsects, the Mantophasmatodea, just as this encyclopedia was beingVincent H. Resh and Ring T. Card 36. ABOUT THE EDITORS Ring Card joined the Department of Entomology of the UniversityVincent Resh is Professor of Entomology and a Curator of the Essig of California, Riverside, in 1996 as Distinguished Professor and holdsMuseum at the University of California, Berkeley, since 1975. He isthe position of A. M. Boyce Chair. He has served as Department Chairthe author of more than 300 articles on insects, mainly on the rolesince 2003. He has authored more than 230 articles on insect chemi-of aquatic insects in the assessment of water pollution and as vectors cal messengers, particularly on moth communication by pheromones,of disease. For 22 years, he was an editor of the Annual Review of and has edited four books on insect chemical ecology and pherom-Entomology and served as an ecological advisor to the United Nations ones. He is a fellow of the American Association for the AdvancementWorld Health Organizations program on the control of river blindnessof Science, the Entomological Society of America, the Entomologicalin West Africa. In 1995 he was elected as a Fellow of the California Society of Canada, and the Royal Entomological Society. In 2009Academy of Sciences and was the recipient of the University of he was awarded the Silver Medal by the International Society ofCalifornia at Berkeleys Distinguished Teaching Award. Chemical Ecology. 37. Sperm can be stored for some length of time in spermathecae,with the record belonging to ant queens that maintain sperm viability AAfor a decade or more. Secretions of spermathecal glands are poorlycharacterized, and how sperm is maintained for such extendedperiods is not known. Spermathecal tissue seems to create a chemi-cal environment that maintains sperm viability, perhaps throughreduced metabolism. A nutritional function is also possible.Transport of sperm out of storage can be facilitated by the secre-tions of the spermathecal gland, which presumably activate qui-escent sperm to move toward the primary reproductive tract. Onepotential function of female accessory glands that has been exploredonly slightly is the production of hormonelike substances that modu-late reproduction functions.Production of Egg CoveringsAcariFemale accessory glands that produce protective coverings for see Mites; Ticks eggs are termed colleterial glands. Colleterial glands have been bestcharacterized in cockroaches, which produce an oothecal case sur-rounding their eggs. Interestingly, the left and right glands are ana-tomically different and have different products. Separation of thechemicals permits reactions to begin only at the time of mixing andootheca formation. Other protective substances produced by glandsAccessory Glandsinclude toxins and antibacterials.Diana E. WheelerNourishment for Embryos or Larvae University of Arizona Viviparous insects use accessory glands to provide nourishmentdirectly to developing offspring. Tsetse flies and sheep keds are dip-The accessory glands of reproductive systems in both femaleterans that retain single larvae within their reproductive tracts andand male insects produce secretions that aid in sperm mainte- provide them with nourishment. They give birth to mature larvaenance, transport, and fertilization. In addition, accessory glandsready to pupate. The gland that produces the nourishing secretion,in females provide protective coatings for eggs. Accessory glands can rich in amino acids and lipids, is known as the milk gland. The Pacificbe organs distinct from the main reproductive tract, or they can be beetle roach, Diploptera punctata, is also viviparous and provides itsspecialized regions of the gonadal ducts (ducts leading from the ova- developing embryos with nourishment secreted by the brood sac, anries or testes). Typically, glandular tissue is composed of two cell types: expanded portion of oviduct.one that is secretory and the other that forms a duct. The interplaybetween male and female secretions from accessory glands is a keyelement in the design of diverse mating systems. ACCESSORY GLANDS OF MALES Accessory glands of the male reproductive tract have diverse func-tions related to sperm delivery and to the design of specific matingACCESSORY GLANDS OF FEMALESsystems. Management of Sperm and Other Male Contributions Sperm management by females involves a wide range of proc-Sperm Deliveryesses, including liberation of sperm from a spermatophore, digestion Males of many insects use spermatophores to transfer sperm toof male secretions and sperm, transport of sperm to and from thefemales. A spermatophore is a bundle of sperm contained in a pro-spermatheca, maintenance of stored sperm, and fertilization.tective packet. Accessory glands secrete the structural proteins nec- Accessory gland secretions can have digestive functions impor-essary for the spermatophores construction. Males of the yellowtant in sperm management. First, digestive breakdown of the sper-mealworm, Tenebrio molitor, have two distinct accessory glands, onematophore can free encapsulated sperm for fertilization and storage.bean-shaped and the other tubular (Fig. 1). Bean-shaped accessorySecond, male contributions can provide an important nutritionalglands contain cells of at least seven types and produce a semisolidbenefit to their mates. Female secretions can digest the secretorymaterial that forms the wall and core of the spermatophore. Tubularcomponents of male seminal fluid to facilitate a nutritive role. Inaccessory glands contain only one type of cell, and it produces a mixaddition, females can digest unwanted sperm to transform it intoof water-soluble proteins of unknown function. Spermatophores arenutrients. Third, female secretions in some species are required tonot absolutely required for sperm transfer in all insects. In manydigest sperm coverings that inhibit fertilization.insects, male secretions create a fluid medium for sperm transfer. Transfer of sperm to and from the spermatheca is generallyaccomplished by a combination of chemical signals and muscularcontractions. Secretions of female accessory glands in some speciesEffects on Sperm Management and on the Femaleincrease sperm motility or appear to attract sperm toward the sper-The effects of male accessory gland secretions on the female aremathecae. Transport of fluid out through the wall of the spermathecabest known for the fruit fly, Drosophila melanogaster, in which themay also create negative pressure that draws in sperm.function of several gene products has been explored at the molecular 38. 2Aestivation level. Since insects have a diversity of mating systems, the specificAfunctions of accessory gland secretions are likely to reflect this variation. In Drosophila, the accessory glands are simple sacs consisting of a single layer of secretory cells around a central lumen (Fig. 2). Genes for more than 80 accessory gland proteins have been identified so far. These genes code for hormonelike substances and enzymes, as well as for many novel proteins. The gene products or their deriva- tives have diverse functions, including an increased egg-laying rate, a reduced inclination of females to mate again, increased effective- ness of sperm transfer to a females spermatheca, and various toxic effects most likely involved in the competition of sperm from differ- ent males. A side effect of this toxicity is a shortened life span for females. Other portions of the reproductive tract contribute secre- tions with diverse roles. For example, the ejaculatory bulb secretes one protein that is a major constituent of the mating plug, and another that has antibacterial activity. See Also the Following ArticlesFIGURE 1 Male reproductive system of T. molitor, showing tes-Egg Coverings Spermatheca Spermatophoretes (T), ejaculatory duct (EJD), tubular accessory gland (TAG), andbean-shaped accessory gland (BAG). [From Dailey, P. D., GadzamaFurther ReadingJ. M., and Happ, G. M. (1980). Cytodifferentiation in the accessory Chen, P. S. (1984). The functional morphology and biochemistry of insectglands of Tenebrio molitor. VI. A congruent map of cells and their male accessory glands and their secretions. Annu. Rev. Entomol. 29,secretions in the layered elastic product of the male bean-shaped233255.accessory gland. J. Morphol. 166, 289322. Reprinted by permission Eberhard, W. G. (1996). Female Control: Sexual Selection by Crypticof Wiley-Liss, Inc., a subsidiary of John Wiley & Sons, Inc.]Female Choice. Princeton University Press, Princeton, NJ. Gillott, C. (1988). ArthropodaInsecta. In Accessory Sex Glands (K. G. Adiyodi, and R. G. Adiyodi, eds.), Vol. 3 of Reproductive Biology of NormalInvertebrates, pp. 319471. Wiley, New York. Happ, G. M. (1992). Maturation of the male reproductive system and its mcendocrine regulation. Annu. Rev. Entomol. 37, 303320. Wolfner, M. F. (2001). The gifts that keep on giving: Physiological func- tions and evolutionary dynamics of male seminal proteins in Drosophila. Heredity 88, 8593.eg ed (A)TransgenicAccessory glandsAcoustic Behavior see Hearing Aedes Mosquitosee Mosquitoes (B)FIGURE 2 Accessory gland of D. melanogaster. (A) The cells in thisnormal accessory gland express b-galactosidase driven by a promoter ofa gene for an accessory gland protein. (B) A transgenic accessory gland,Aestivationcells expressing the gene have been selectively killed after eclosion.These flies were used to explore the function of accessory gland secre- Sinzo Masakitions. In transgenic males, accessory glands are small and translationally Hirosaki Universityinert. [From Kalb, J. M., DiBenedetto, A. J., and Wolfner, M. F. (1993). AProbing the function of Drosophila melanogaster accessory glandsestivation is a dormant state for insects to pass the summerby directed cell ablation. Proc. Natl Acad. Sci. USA 90, 80938097. in either quiescence or diapause. Aestivating, quiescentCopyright 1993, National Academy of Sciences, U.S.A.] insects may be in cryptobiosis and highly tolerant to heat and 39. Aestivation3drought. Diapause for aestivation, or summer diapause, serves notonly to enable the insect to tolerate the rigors of summer but also toensure that the active phase of the life cycle occurs during the favo- Arable time of the year. QUIESCENCEQuiescence for aestivation may be found in arid regions. Forexample, the larvae of the African chironomid midge, Polypedilumvanderplanki, inhabit temporary pools in hollows of rocks andbecome quiescent when the water evaporates. Dry larvae of thismidge can revive when immersed in water, even after years of qui-escence. The quiescent larva is in a state of cryptobiosis and toleratesthe reduction of water content in its body to only 4%, surviving evenbrief exposure to temperatures ranging from 102C to 270C.Moreover, quiescent eggs of the brown locust, Locustana pardalina,survive in the dry soil of South Africa for several years until theirwater content decreases to 40%. When there is adequate rain, theyabsorb water, synchronously resume development, and hatch, result-ing in an outburst of hopper populations. The above-mentionedexamples are dramatic, but available data are so scanty that it is dif-ficult to surmise how many species of insects can aestivate in a stateof quiescence in arid tropical regions.SUMMER DIAPAUSESyndromeThe external conditions that insects must tolerate differ sharplyin summer and winter. Aestivating and hibernating insects may showsimilar diapause syndromes: cessation of growth and development,reduction of metabolic rate, accumulation of nutrients, and increasedprotection by body coverings (hard integument, waxy material,cocoons, etc.), which permit them to endure the long period of FIGURE 1 Bogong moths, Agrotis infusa, aestivating in aggre-dormancy that probably is being mediated by the neuroendocrine gation on the roof of a cave at Mt Gingera, A. C. T., Australia.system.[Photograph from Common, I. (1954). Aust. J. Zool. 2, 223263,Migration to aestivation sites is another component of diapausecourtesy of CSIRO Publishing.]syndrome found in some species of moths, butterflies, beetles, andhemipterans. In southeastern Australia, the adults of the Bogongmoth, Agrotis infusa, emerge in late spring to migrate from theplains to the mountains, where they aestivate, forming huge aggrega-tions in rock crevices and caves (Fig. 1).Seasonal Cues Summer diapause may be induced obligatorily or facultativelyby such seasonal cues as daylength (nightlength) and temperature.When it occurs facultatively, the response to the cues is analogousto that for winter diapause; that is, the cues are received duringthe sensitive stage, which precedes the responsive (diapause) stage.The response pattern is, however, almost a mirror image of that forwinter diapause (Fig. 2). Aestivating insects themselves also mayFIGURE 2 Photoperiodic response in the noctuid M. brassicaebe sensitive to the seasonal cues; a high temperature and a long controlling the pupal diapause at 20C. Note the different ranges ofdaylength (short nightlength) decelerate, and a short daylength (longphotoperiod for the induction of summer diapause (dashed line) andnightlength) and a low temperature accelerate the termination of winter diapause (solid line). [From Furunishi et al. (1982), repro-diapause.duced with permission.] The optimal range of temperature for physiogenesis duringsummer diapause broadly overlaps with that for morphogenesis, or the superficial similarity in their dormancy syndromes, the two typesextends even to a higher range of temperature. Aestivating eggs of of diapause involve basically different physiological processes.the brown locust, L. pardalina, can terminate diapause at 35C andthose of the earth mite, Halotydeus destructor, do this even at 70C.The different thermal requirements for physiogenesis clearly distin- See Also the Following Articlesguish summer diapause from winter diapause, suggesting that despiteCold/Heat Protection Diapause Dormancy Migration 40. 4 Agricultural EntomologyFurther Reading of aggregate losses caused by insects as vectors of pathogens and para-A Common, I. F. B. (1954). A study of the biology of the adult Bogong moth,sites of humans and domestic animals, as agents causing direct damageto dwellings and other articial structures, and as pests of crop plants Agrotis infusa (Boisd.) (Lepidoptera: Noctuidae), with special refer-and farm animals, but the amounts run to probably hundreds of bil- ence to its behaviour during migration and aestivation. Aust. J. Zool. 2, 223263. lions of dollars annually. Losses caused by insects and vertebrate pestsFurunishi, S., Masaki, S., Hashimoto, Y., and Suzuki, M. (1982). Diapause worldwide in the production of only eight principal food and cash response to photoperiod and night interruption in Mamestra brassicae crops (barley, coffee, cotton, maize, potato, rice, soybean, and wheat) (Lepidoptera: Noctuidae). Appl. Entomol. Zool. 17, 398409.between 1988 and 1990 have been estimated at $90.5 billion.Hinton, H. E. (1960). Cryptobiosis in the larva of Polypedilum vanderplanki In the late 1800s and early 1900s, entomology became established Hint (Chironomidae). J. Insect Physiol. 5, 286300.in many academic and research institutions as a discipline equal inMasaki, S. (1980). Summer diapause. Annu. Rev. Entomol. 25, 125. rank with botany and zoology. The diversity of insects and their eco-Matthe, J. J. (1951). The structure and physiology of the egg of Locustana nomic importance was the justication for ranking the study of a class pardalina (Walk.). Union S. Afr. Dep. Agric. Sci. Bull. 316, 183.of animals (Insecta) as being equivalent to the study of two kingdomsTauber, M. J., Tauber, C. A., and Masaki, S. (1986). Seasonal Adaptations ofof organisms (plants and animals other than insects). Through the Insects. Oxford University Press, New York.rst half of the twentieth century, there was a schism between basicand applied (or economic) entomology. Since then, common use ofthe expression economic entomology has declined, being replacedby designations of its principal branches, such as agricultural ento-mology, forest entomology, urban entomology, and medical and vet- Africanized Bees erinary entomology. A detailed historical account is beyond the scopeof this article, but Table I provides a chronology of some landmarkssee Neotropical African Beesin the development of agricultural entomology through the ages.The realm of agricultural entomology includes all basic studiesof benecial and pest insects associated with agricultural crops andfarm animals. This article deals mainly with crops, but the generalprinciples and concepts are equally applicable to farm animals. The Agricultural Entomologystarting point of such studies is a correct identication of the insectspecies, in accordance with the science known as biosystematics. Marcos KoganOregon State UniversityBIOSYSTEMATICSScientic nomenclature is a powerful tool for obtaining informa-tion about the basic biology of closely related species within a genus. Ronald ProkopyWhen systematic studies have been extended beyond the naming of University of Massachusettsspecies (taxonomy) and contain detailed information on geographicdistribution, host records, and biology of one or more species in aT he study of all economically important insects is the object genus, it is often possible to extrapolate the information to other of the subdiscipline economic or applied entomology. closely related species of that genus. Although details of the biology Agricultural entomology, a branch of economic entomology,must be ascertained for each individual species, biosystematics offersis dedicated to the study of insects of interest to agriculture because a blueprint to follow when dealing with a new pest. For example, thethey help increase crop production (e.g., pollinators); produce a genus Cerotoma (Coleoptera: Chrysomelidae) contains 1012 speciescommodity (e.g., honey, silk, lacquer); cause injury leading to eco-distributed from southern Brazil to the northeastern United States. Allnomic losses to plants grown for food, feed, ber, or landscaping;seem to be associated with herbaceous plants in the family Fabaceaecause injury to farm animals; or are natural enemies of agricultural(bean family). The biology of two of the species, C. trifurcata in Northpests and, therefore, considered to be benecial. Study of all fun- America and C. arcuata in South America (Fig. 2), has been studieddamental aspects of the ecology, life history, and behavior of insectsextensively. Based on information for these two species, it is possibleassociated with agricultural crops and farm animals falls within theto infer that the other species in the genus share at least some of therealm of agricultural entomology. These studies provide the founda- following features: eggs are laid in the soil adjacent to growing legumi-tion for the design and implementation of integrated pest manage- nous plants; larvae feed on nitrogen-xing root nodules and pupate inment (IPM) programs (Fig. 1). soil inside pupal cases; rst-generation adults emerge when seedlingsemerge, and second-generation adults emerge when plants are in full ECONOMIC ENTOMOLOGYvegetative growth, feeding rst on foliage and, later on, switching tofeeding on developing pods. The biosystematic information on theInsects are regarded by some as the main competitors of humansgenus allows students of agricultural entomology in South, Central, orfor dominance on the earth. Humans depend on insects for pollinationNorth America to understand, at least in general terms, the role ofof many crops, for production of honey and silk, for the decompositionany other species of Cerotoma within their particular agroecosystem.of organic matter and the recycling of carbon, and for many other vitalThe ip side of this notion is recognition that closely related andecological roles. But it is the negative impact of insect pests that hasmorphologically nearly undistinguishable (sibling) species may havebeen of greatest concern to humans. There are no reliable estimatesmany important biological differences. Examples of the critical needfor reliable biosystematics studies are found in the biological controlDeceased May 14, 2004.literature. The present account is based on studies conducted by Paul 41. Agricultural Entomology 5 System integr s ation and information systemA Biological controlBehavioral control Chemical control Cultural controlPlant resistanceGenetic control Integrated pest management bridge WeedscienceRural sociologyPlant CropCroppathologyprotection Agricultural productionAgricultural economicsentomologyBiological Socialsciencessciences The flowing river of societal needs and demandsFIGURE 1 A bridge metaphor: agricultural entomology is conceived as one of the main pillars, together with plant pathology and weedscience, supporting the integrated pest management bridge. The bridge connects two-way trafc between crop production and cropprotection. The other pillar is provided by the social sciences of economics and sociology. The main tension cables, which are system inte-gration and information systems, hold the vertical lines that together give stability to the bridge; these are the tactical components of IPM.Under the bridge runs the river of ever-shifting societal needs and demands. TABLE ISome Landmarks in the Historical Development of Agricultural Entomology Signicant events Years ago from 2000Date Beginnings of agriculture 10,000 8000 B.C.E. First records of insecticide use4,5002500 B.C.E. First descriptions of insect pests3,5001500 B.C.E. Soaps used to control insects in China9001100 Beginning of scientic nomenclature10th edition of Linnaeus, Systema Naturae 2421758 Burgeoning descriptions of insects10020018th and 19th centuries First record of plant resistance to an insect 1691831 Charles Darwin and Alfred Wallace jointly present paper on the theory of evolution1421858 First successful case of biological control: the cottony cushion scale, on citrus, in California, by1121888 the vedalia beetle First record of widespread damage of cotton in Texas by the cotton boll weevil1061894 First record of an insect resistant to an insecticide 86 1914 First edition of C. L. Metcalf and W. P. Flints Destructive and Useful Insects 72 1928 Discovery of DDT and beginning of the insecticide era 61 1939 First report of insect resistance to DDT54 1946 Term pheromone coined by P. Karlson and P. Butenandt, who identied rst such substance 45 1959 in the silkworm moth First edition of Rachel Carsons Silent Spring48 1962 Expression integrated pest management rst appears in the press 32 1968 Rapid development of molecular biology20 1980s Release of Bt transgenic varieties of cotton, corn, and potato51990s Based in part on Norris et al. (2003).DeBach, one of the leading biological control specialists of the twen-of confusion and missed opportunities because of misidentication oftieth century. The California red scale, Aonidiella aurantii, is a seri-its parasitoids. The red scale parasitoid Aphytis chrysomphali had beenous pest of citrus in California and other citrus-producing areas of theknown to occur in California and was not considered to be a very effec-world. Biological control of the red scale in California had a long history tive control agent. When entomologists discovered parasitized scales 42. 6Agricultural Entomology Yield typeDefining factors: CO2A1 Potential Radiation Temperature Crop genetics -Crop physiology -Crop phenology -Canopyarchitecture Limiting factors: Water 2 AttainableNutrients -Nitrogen -PhosphorusYield-increasing -Potassiummeasures Reducing factors: Insect pests 3 ActualVertebrates Pathogens Yield-protectingWeeds measuresPollutants 15005000 10,000 20,000 Production level (kg ha1)FIGURE 3 Factors impacting the yield potential of a genericcrop. (Adapted from information on a Web site originated at IMI/University of Miami, Summer Institute.)FIGURE 2 Morphological diversity and biological similarities inthe genus Cerotoma: four of the dozen known species are illustrated To assess crop losses and attribute the losses to a specic causeby male and female specimens. The species are clearly distinguish-(e.g., the attack of a pest) requires setting up experiments to isolateable by morphological characters, but they have similar life historiesthe effect of the pest from all other constraints. Methodologies varyand behaviors. (From unpublished drawings by J. Sherrod, Illinois with pest categorywhether the pests are insects, vertebrates, plantNatural History Survey.)pathogens, or weeds, for example. The quantitative relationshipbetween crop losses and pest population levels is the basis for com-puting the economic injury level for the pest. The economic injuryduring foreign exploration, the parasitoids were misidentied as A. level is a fundamental concept in IPM.chrysomphali and therefore were not imported into California. It waslater discovered that the parasitoids were in fact two different species,A. lingnanensis and A. melinus, both more efcient natural enemies of LIFE HISTORY AND HABITSthe California red scale than A. chrysomphali. Now A. lingnanensis and Once the identity and pest status of a species have been wellA. melinus are the principal red scale parasitoids in California. Further established, it becomes essential to extend the informational basebiosystematics studies have shown that what was once thought to beon the life history and habits of the species to the conditions undersingle species, A. chrysomphali, parasitic on the California red scale in which the crop is grown. Economically important life history traitsthe Orient and elsewhere, and accidentally established in California, isinclude information on developmental threshold temperatures andin fact a complex including at least seven species having different bio-temperature-dependent developmental rates. These data are used inlogical adaptations but nearly indistinguishable morphologically. modeling the phenology of the pest. Other essential studies includeKnowledge of the name of a species, however, is not an indication the orientation, feeding, host selection, and sexual behavior of theof its true potential economic impact or pest status. A next impor- species. Many of these studies provide the foundation for strategictant phase in agricultural entomology is, therefore, the assessment ofplanning in IPM and for the development of target-specic controlbenets or losses caused by that species. tactics. For example, the study of sexual behavior can involve thedenition of the role of pheromones in mating and the identication PEST IMPACT ASSESSMENT of those pheromones. These, in turn, may be used for monitoringpest incidence and abundance or in mating disruption, both valuableThe mere occurrence of an insect species in association with acomponents of IPM systems for many crops. The study of host selec-crop or a farm animal does not necessarily mean that the species is tion behavior often leads to the identication of kairomones, equallya pest of that crop or animal. To be a pest it must cause economicimportant in IPM development.losses. The assessment of economic losses from pests is the subjectof studies conducted under conditions that match as closely as pos-sible the conditions under which the crop is grown commerciallyPHENOLOGYor the animals are raised. Much of the methodology used in crop The life cycle of different insect species varies greatly, althoughloss assessment has been established under the sponsorship of the all insects undergo the basic stages of development from egg toFood and Agriculture Organization (FAO) of the United Nations asreproductive adult (or imago). Depending on the length of the lifea means of prioritizing budget allocations and research efforts. Keycycle, there is considerable variation in the number of generationsdata for these studies relate to the determination of the yield poten-per year, a phenomenon called voltinism. A univoltine species hastial of a crop. The genetic makeup of a crop variety determines its one generation per year; a multivoltine species may have many gen-maximum yield in the absence of adverse environmental factors.erations per year. The range of variation in the Insecta is evidentThis is known as the attainable yield. To determine the attainablewhen one considers that the 17-year periodical cicada has one gener-yield, the crop is grown under nearly ideal conditions; the actualation every 17 years, whereas whiteies or mosquitoes may completeyield is what occurs when the crop is grown under normal farminga generation in about 21 days. Under temperate climate conditions,conditions. The difference between attainable and actual yields is agenerations often are discrete, but under warmer subtropical condi-measure of crop loss (Fig. 3).tions, they frequently overlap. The denition of temporal periodicity 43. Agricultural Entomology 7in an organisms developmental cycle is called phenology. The rela- The set of species coexisting in an area and interacting to varyingtionship between the phenology of the crop and the phenologies ofits various pests is of interest in agricultural entomology. Fig. 4 showsdegrees form what is known as an ecological community. In a cropcommunity, the crop plants and the weeds that persist within theAan example of such a relationship for soybean grown under condi-crop eld o