Contents by Subject Area xiii Contributors xvii Guide to the Encyclopedia xxiii Foreword xxv Preface...

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Transcript of Contents by Subject Area xiii Contributors xvii Guide to the Encyclopedia xxiii Foreword xxv Preface...

  • Encyclopedia of

    INSECTS

  • Amsterdam Boston London New York Oxford Paris San Diego San Francisco Singapore Sydney Tokyo

    Encyclopedia of

    INSECTS

    Editors

    VINCENT H. RESHUniversity of California, Berkeley

    RING T. CARDUniversity of California, Riverside

  • Cover Art: Monarch butterflies photographed at an overwintering site in centralMexico. Each fall, eastern North American monarchs migrate by the millions tothese high-altitude Oyamel fir forests. The weight of their dense aggregationscan even cause tree limbs to break away. (Photograph by George D. Lepp, aspecialist in images of the natural world.)

    Frontispiece: An 18th century engraving by I. A. Covinus depicts the Biblicaldescription of the creation of insects. The image appears in Physique sacre, ouhistoire naturelle de la Bible, a massive eight-volume study by the prominentSwiss naturalist Johann Jacob Scheuchzer (16721737). The artist Covinushere combines an accurate rendering of various insect forms with an allegoricaltreatment of the Book of Genesis; this approach reflects the authorScheuchzers lifelong efforts to provide scientific evidence for the literal truthof the Bible. (Courtesy of the History of Science Collections, CornellUniversity Library.)

    Letter Openers:R: Snakefly (Raphidioptera) photographed at Nanaimo (Vancouver Island),British Columbia, Canada. (Photograph Jay Patterson.)Y: Aedes aegypti, Uganda strain (a vector of yellow fever), bloodfeeding fromthe photographers hand. (Photograph by Leonard E. Munstermann.)Other letter opener images furnished by the authors. (See relevant articlefor credit.)

    This book is printed on acid-free paper.

    Copyright 2003, Elsevier Science (USA).

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    8

  • v

    Contents by Subject Area xiii

    Contributors xvii

    Guide to the Encyclopedia xxiii

    Foreword xxv

    Preface xxvii

    Acarisee Mites; Ticks

    Accessory Glands 1Diana E. Wheeler

    Acoustic Behaviorsee Hearing

    Aedes Mosquitosee Mosquitoes

    Aestivation 3Sinzo Masaki

    Africanized Beessee Neotropical African Bees

    Agricultural Entomology 4Marcos Kogan and Ronald Prokopy

    Alderflysee Megaloptera

    Amber 9George Poinar Jr.

    Anatomy: Head, Thorax,Abdomen, and Genitalia 12David H. Headrick and Gordon Gordh

    Anopheles Mosquitosee Mosquitoes

    Anoplurasee Phthiraptera

    Antennae 26Catherine Loudon

    Ants 29Nigel R. Franks

    Aphids 32John T. Sorenson

    Apis Species 37Eva Crane

    Aposematic Coloration 39Mathieu Joron

    Apterygota 45The Editors

    Aquatic Habitats 45Richard W. Merritt and J. Bruce Wallace

    Arachnidasee Scorpions; Spiders

    Archaeognatha 57Helmut Sturm

    Arthropoda and Related Groups 59James H. Thorp

    Auchenorrhyncha 66Christopher H. Dietrich

    Autohemorrhage 74The Editors

    Autotomy 75The Editors

    Beesee Apis Species

    Beekeeping 76Eva Crane

    Bee Products 82Eva Crane

    Beeswax 84Eva Crane

    Beetlesee Coleoptera

    Biodiversity 85Nigel E. Stork

    CONTENTS

  • Biogenic Amines 91Arnd Baumann, Wolfgang Blenau,and Joachim Erber

    Biogeographical Patterns 94Peter Zwick

    Biological Control of Insect Pests 103R. G. Van Driesche and M. S. Hoddle

    Bioluminescence 115James E. Lloyd and Erin C. Gentry

    Biotechnology and Insects 120Bryony C. Bonning

    Blattodea 123Donald G. Cochran

    Bloodsee Circulatory System

    Blood Sucking 127M. J. Lehane

    Body Size 130Christer Bjrkman and Mats W. Pettersson

    Boll Weevil 132R. Nelson Foster

    Bombyx mori 133Satoshi Takeda

    Book Lousesee Psocoptera

    Borers 135Timothy D. Paine

    Brain and Optic Lobes 137Nicholas J. Strausfeld

    Bristletailsee Archaeognatha

    Bubonic Plague 147Irwin W. Sherman

    Bumblebeesee Hymenoptera

    Butterflysee Lepidoptera

    Caddisflysee Trichoptera

    Caste 151Sean ODonnell

    Caterpillars 154Frederick W. Stehr

    Cat Fleas 156Nancy C. Hinkle and Michael K. Rust

    Cave Insects 158Francis G. Howarth

    Cell Culture 163Dwight E. Lynn

    Chemical Defense 165Murray S. Blum

    Chemoreception 169B. K. Mitchell

    Chiggers and Other Disease-Causing Mites 174Larry G. Arlian

    Chitin 179Ephraim Cohen

    Chromosomes 180Graham C. Webb

    Chrysalis 185Frederick W. Stehr

    Cicadas 186Max S. Moulds

    Circadian Rhythms 188Terry L. Page

    Circulatory System 192Thomas A. Miller

    Classificationsee Nomenclature and Classification

    Cockroachsee Blattodea

    Cocoon 197Frederick W. Stehr

    Codling Moth 198Stephen C. Welter

    Coevolution 199Douglas J. Futuyma

    Cold/Heat Protection 204David L. Denlinger

    Coleoptera 209James K. Liebherr and Joseph V. McHugh

    Collection and Preservation 230Charles V. Covell, Jr.

    Collembola 235Kenneth A. Christiansen and Peter Bellinger

    Colonies 239Sean ODonnell

    Colorado Potato Beetle 242George G. Kennedy

    Coloration 244Helen Ghiradella

    Commercial Products from Insects 251Michael E. Irwin and Gail E. Kampmeier

    Conservation 260Tim R. New

    Crickets 265Richard D. Alexander and Daniel Otte

    Crypsis 269Paul M. Brakefield

    Cultural Entomology 273James N. Hogue

    Cuticle 281Svend O. Andersen

    Daddy-Long-Legs 283Gonzalo Giribet

    Damselflysee Odonata

    Dance Language 284P. Kirk Visscher

    vi Content s

  • DDT 288Fumio Matsumura

    Defensive Behavior 289Justin O. Schmidt

    Dengue 295Thomas W. Scott

    Dermaptera 297Susan M. Rankin and James O. Palmer

    Development, Hormonal Control of 300Michael E. Adams

    Diapause 305David L. Denlinger

    Digestion 310Walter R. Terra

    Digestive System 313Walter R. Terra and Cllia Ferreira

    Diplura 323Robert T. Allen

    Diptera 324Richard W. Merritt, Gregory W. Courtney, and Joe B. Keiper

    Diversitysee Biodiversity

    Division of Labor in Insect Societies 340Gene E. Robinson

    Dobsonflysee Megaloptera

    Dog Heartworm 343Edward G. Platzer

    Dormancy 344Richard E. Lee, Jr.

    Dragonflysee Odonata

    Drosophila melanogaster 345Patrick M. OGrady

    Dung Beetles 348James Ridsdill-Smith

    Earwigsee Dermaptera

    Ecdysissee Molting

    Ecdysteroids 352Michael E. Adams

    Eggs 355Diana E. Wheeler

    Egg Coverings 356Diana E. Wheeler

    Embiidina 358Edward S. Ross

    Embryogenesis 359Lisa Nagy and Miodrag Grbic

    Endangered Insects 364Scott Hoffman Black and Mace Vaughan

    Endopterygota 369The Editors

    Entomological Societies 369Alan I. Kaplan

    Ephemeroptera 373John E. Brittain and Michel Sartori

    Estivationsee Aestivation

    Evolutionsee Phylogeny

    Excretion 380Timothy J. Bradley

    Exopterygota 387The Editors

    Exoskeleton 387Svend O. Andersen

    Extension Entomology 390Nancy C. Hinkle, Beverly Sparks, Linda J.Mason, and Karen M. Vail

    Eyes and Vision 393Michael F. Land

    Fat Body 407Ephraim Cohen

    Feeding Behavior 409Reginald F. Chapman

    Fire Ants 414Les Greenberg

    Fleasee Siphonaptera

    Flight 416Michael Dickinson and Robert Dudley

    Flysee Diptera

    Folk Beliefs and Superstitions 426James N. Hogue

    Food, Insects as 431Gene R. DeFoliart

    Forensic Entomology 437M. Lee Goff

    Forest Habitats 442David L. Wood and Andrew J. Storer

    Fossil Record 455David Grimaldi

    Freshwater Habitatssee Aquatic Habitats

    Fruit Flysee Drosophila melanogaster

    Gallmaking and Insects 464Katherine N. Schick and Donald L. Dahlsten

    Genetically Modified Plants 466David A. Andow

    Genetic Engineering 471Peter W. Atkinson and David A. OBrochta

    Content s vii

  • Genetic Variation 478George K. Roderick

    Grasshoppersee Orthoptera

    Grassland Habitats 481Teja Tscharntke, Ingolf Steffan-Dewenter,Andreas Kruess, and Carsten Thies

    Greenhouse Gases, GlobalWarming, and Insects 486Peter Stiling

    Growth, Individual 489Martin B. Berg and Richard W. Merritt

    Grylloblattodea 492D. C. F. Rentz

    Gypsy Moth 493Joseph S. Elkinton

    Hearing 498Jayne Yack and Ron Hoy

    Hemipterasee Auchenorrhyncha; Prosorrhyncha;Sternorrhyncha

    Hemolymph 505Michael R. Kanost

    Heteropterasee Prosorrhyncha

    Hibernation 508Richard E. Lee, Jr.

    History of Entomology 509Edward H. Smith and Janet R. Smith

    Homeostasis, Behavioral 519P. Kirk Visscher

    Homopterasee Auchenorrhyncha; Sternorrhyncha

    Honey 521Eva Crane

    Honey Beesee Apis Species

    Honeydew 523Thomas E. Mittler and Angela E. Douglas

    Hornetsee Wasps

    Host Seeking, by Parasitoids 526Ronald M. Weseloh

    Host Seeking, for Plants 529Elizabeth A. Bernays

    House Fly 532Gregory A. Dahlem

    Hymenoptera 534Donald L. J. Quicke

    Hypermetamorphosis 546John D. Pinto

    Hyperparasitism 549Daniel J. Sullivan

    Imaginal Discs 552Seth S. Blair

    Immunology 555Nancy E. Beckage

    Industrial Melanism 560Michael E. N. Majerus

    Insecta, Overview 564Vincent H. Resh and Ring T. Card

    Insecticides 566Fumio Matsumura

    Insecticide and Acaricide Resistance 569Gregor J. Devine and Ian Denholm

    Insectivorous Plants 577Lewis J. Feldman

    Insectivorous Vertebrates 580The Editors

    Insect Zoos 582Leslie Saul-Gershenz

    Integrated Pest Management 589Ronald Prokopy and Marcos Kogan

    Integument 595Svend O. Andersen

    Introduced Insects 597Daniel Simberloff

    Island Biogeography and Evolution 602George K. Roderick and Rosemary G. Gillespie

    Isoptera 604Vernard R. Lewis

    Japanese Beetle 609David W. Held and Daniel A. Potter

    June Beetles 610Daniel A. Potter and David W. Held

    Juvenile Hormones 611Michael E. Adams

    Katydidsee Orthoptera

    Killer Beessee Neotropical African Bees

    Lacewingsee Neuroptera

    Ladybugs 618Michael E. N. Majerus

    viii Content s

  • Larva 622Frederick W. Stehr

    Learning 624Daniel R. Papaj

    Legs 627Peter H. Adler

    Lepidoptera 631Jerry A. Powell

    Licesee Phthiraptera

    Lice, Human 664Terri L. Meinking

    Locusts 666Reginald F. Chapman

    Magnetic Sense 670John Klotz and Rudolf Jander

    Malaria 672William K. Reisen

    Mallophagasee Phthiraptera

    Mantodea 675Lawrence E. Hurd

    Mantophasmatodea 677Klaus-Dieter Klass

    Marine Insects 679Lanna Cheng

    Mating Behaviors 682Darryl T. Gwynne

    Mayflysee Ephemeroptera

    Mechanoreception 689Andrew S. French and Pivi H. Torkkeli

    Mecoptera 690George W. Byers

    Medical Entomology 693John D. Edman

    Medicine, Insects in 698Ronald A. Sherman

    Megaloptera 700Norman H. Anderson

    Metabolism 703S. Nelson Thompson and R. K. Suarez

    Metamorphosis 707Frederick W. Stehr

    Migration 708Hugh Dingle

    Mimicry 714Mathieu Joron

    Mites 726Barry M. OConnor

    Molting 733Lynn M. Riddiford

    Monarchs 739Lincoln P. Brower

    Mosquitoes 743Bruce F. Eldridge

    Mothsee Lepidoptera

    Mouthparts 750Reginald F. Chapman

    Movies, Insects in 756May R. Berenbaum and Richard J. Leskosky

    Musca domesticasee House Fly

    Muscle System 763Robert Josephson

    Museums and Display Collections 768Gordon M. Nishida

    Neosomy 775Frank J. Radovsky

    Neotropical African Bees 776Orley R. Taylor

    Nervous Systemsee Brain and Optic Lobes

    Nest Building 778Robert L. Jeanne

    Neuropeptides 782Miriam Altstein

    Neuroptera 785Catherine A. Tauber, Maurice J. Tauber, andGilberto S. Albuquerque

    Nomenclature and Classification,Principles of 798F. Christian Thompson

    Nutrition 807S. Nelson Thompson and S. J. Simpson

    Ocelli and Stemmata 814Frederick W. Stehr

    Odonata 814K. J. Tennessen

    Orientation 823Ring T. Card

    Orthoptera 827D. C. F. Rentz and You Ning Su

    Ovarioles 839Diana E. Wheeler

    Oviposition Behavior 841Marc J. Klowden

    Parasitoids 845Nick Mills

    Parental Care 848Michelle Pellissier Scott

    Content s ix

  • Parthenogenesis 851Lawrence R. Kirkendall and Benjamin Normark

    Pathogens of Insects 856Brian A. Federici

    Phasmida 865Erich H. Tilgner

    Pheromones 866Ring T. Card and Jocelyn G. Millar

    Phoresy 873Marilyn A. Houck

    Photography of Insects 875Mark W. Moffett

    Phthiraptera 879Ronald A. Hellenthal and Roger D. Price

    Phylogeny of Insects 882Peter S. Cranston and Penny J. Gullan

    Physical Control of Insect Pests 898Charles Vincent and Bernard Panneton

    Phytophagous Insects 902Elizabeth A. Bernays

    Phytotoxemia 905Alexander H. Purcell

    Plant Diseases and Insects 907Alexander H. Purcell

    PlantInsect Interactions 912J. Mark Scriber

    Plecoptera 915Kenneth W. Stewart

    Pollination and Pollinators 919Gordon W. Frankie and Robbin W. Thorp

    Pollution, Insect Response to 926David M. Rosenberg and Vincent H. Resh

    Polyembryony 928Michael R. Strand

    Population Ecology 933Joseph S. Elkinton

    Praying Mantidsee Mantodea

    Predation/Predatory Insects 944Ronald M. Weseloh

    Prosorrhyncha 947Carl W. Schaefer

    Protura 965Robert T. Allen

    Psocoptera 966Edward L. Mockford

    Pterygota 969The Editors

    Puddling Behavior 969Scott R. Smedley

    Pupa and Puparium 970Frederick W. Stehr

    Queensee Caste

    Raphidioptera 973Ulrike Aspck and Horst Aspck

    Rearing of Insects 975Norman C. Leppla

    Recruitment Communication 980James F. A. Traniello

    Regulatory Entomology 988Robert V. Dowell

    Reproduction, Female 991Diana E. Wheeler

    Reproduction, Female: HormonalControl of 994Diana E. Wheeler

    Reproduction, Male 997Marc J. Klowden

    Reproduction, Male: HormonalControl of 999Marc J. Klowden

    Research Tools, Insects as 1000Kipling W. Will

    Respiratory System 1002Jon F. Harrison

    River Blindness 1007Vincent H. Resh

    Rostrum 1008Reginald F. Chapman

    Royal Jelly 1009Eva Crane

    Salivary Glands 1011Gregory P. Walker

    Scale Insectsee Sternorrhyncha

    Scales and Setae 1017Shaun L. Winterton

    Scorpions 1020Stanley C. Williams

    Segmentation 1025Nipam H. Patel

    Sericulture 1027Satoshi Takeda

    Sex Determination 1029Michael F. Antolin and Adam D. Henk

    Sexual Selection 1032Kenneth Y. Kaneshiro

    x Content s

  • Silk Mothsee Bombyx mori

    Silk Production 1038Catherine L. Craig

    Silverfishsee Zygentoma

    Siphonaptera 1040Michael W. Hastriter and Michael F. Whiting

    Snakeflysee Raphidioptera

    Sociality 1044James E. Zablotny

    Soil Habitats 1053Patricia J. Vittum

    Sound Productionsee Hearing

    Spermatheca 1058Marc J. Klowden

    Spermatophore 1059Marc J. Klowden

    Spiders 1060Rosemary G. Gillespie and Joseph C. Spagna

    Springtailsee Collembola

    Stamps, Insects and 1073Charles V. Covell, Jr.

    Sterile Insect Technique 1074Jorge Hendrichs and Alan Robinson

    Sternorrhyncha 1079Penny J. Gullan and Jon H. Martin

    Stoneflysee Plecoptera

    Stored Products as Habitats 1089Rudy Plarre and Wendell E. Burkholder

    Strepsiptera 1094Michael F. Whiting

    Swimming, Lake Insects 1096Werner Nachtigall

    Swimming, Stream Insects 1098Bernhard Statzner

    Symbionts Aiding Digestion 1102Andreas Brune

    Symbionts, Bacterial 1107Michael E. N. Majerus

    Systematicssee Nomenclature and Classification

    Tastesee Chemoreception

    Taxonomysee Nomenclature and Classification

    Teaching Resources 1113John H. Acorn and Felix A. H. Sperling

    Temperature, Effects onDevelopment and Growth 1116Frantisek Sehnal, Oldrich Nedved, and Vladimir Kostl

    Termitesee Isoptera

    Terrestrial Insectssee Soil Habitats

    Thermoregulation 1119Bernd Heinrich

    Thripssee Thysanoptera

    Thysanoptera 1127Laurence A. Mound

    Thysanurasee Archaeognatha; Zygentoma

    Ticks 1132Daniel E. Sonenshine

    Touchsee Mechanoreception

    Tracheal System 1142Jon F. Harrison

    Trichoptera 1145John C. Morse

    Tsetse Fly 1151Stephen G. A. Leak

    Urban Habitats 1157Michael K. Rust

    Venom 1160Justin O. Schmidt

    Veterinary Entomology 1163Bradley A. Mullens

    VibrationalCommunication 1167Andrej Cokl and Meta Virant-Doberlet

    Visionsee Eyes and Vision

    Vitellogenesis 1171William H. Telfer

    Walking and Jumping 1174Roy E. Ritzmann and Sasha N. Zill

    Walking Stickssee Phasmida

    Content s xi

  • Wasps 1179Justin O. Schmidt

    Water and Ion Balance,Hormonal Control of 1183Thomas M. Clark

    Weevilsee Boll Weevil

    Wings 1186Robin J. Wootton

    Wolbachia 1192Richard Stouthamer

    Workersee Caste

    Yellow Fever 1196Thomas P. Monath

    Yellowjacketsee Wasps

    Zoonoses, Arthropod-Borne 1198Robert S. Lane

    Zoraptera 1201Michael S. Engel

    Zygentoma 1203Helmut Sturm

    Glossary 1207

    Subject Index 1227

    xii Content s

  • xiii

    ANATOMYAnatomy: Head, Thorax,

    Abdomen, and GenitaliaAntennaeBrain and Optic LobesChitinColorationCuticleDigestive SystemExoskeletonEyes and VisionIntegumentLegsMouthpartsOcelli and StemmataPupa and PupariumRostrumSalivary GlandsScales and SetaeSegmentationTracheal SystemWings

    PHYSIOLOGYAestivationAutohemorrhage

    AutotomyBiogenic AminesBioluminescenceBiotechnology and InsectsBody SizeBrain and Optic LobesCell CultureChemical DefenseChemoreceptionChitinChromosomes Circadian RhythmsCirculatory SystemCold/Heat ProtectionCuticleDDTDevelopment, Hormonal

    Control ofDiapauseDigestionDigestive SystemDormancyEcdysteroidsExcretionEyes and VisionFat BodyFlightGenetic EngineeringHearingHemolymph

    HibernationHomeostasis, BehavioralHoneydewImaginal DiscsImmunologyInsecticidesJuvenile HormonesMagnetic SenseMechanoreceptionMetabolism MoltingMuscle SystemNeuropeptidesNutritionReproduction, FemaleReproduction, Female: Hormonal

    Control ofReproduction, MaleReproduction, Male: Hormonal

    Control ofRespiratory SystemSalivary GlandsSegmentationSex DeterminationSilk ProductionSymbionts Aiding DigestionThermoregulationTracheal SystemVitellogenesisWalking and Jumping

    CONTENTS BYSUBJECT AREA

  • Water and Ion Balance, HormonalControl of

    BEHAVIORAposematic ColorationAutohemorrhageAutotomyBioluminescenceBlood SuckingBorersCasteChemical DefenseChemoreceptionCircadian RhythmsColoniesCrypsisDance LanguageDefensive BehaviorDivision of Labor in Insect SocietiesEyes and VisionFeeding BehaviorFlightHearingHibernationHost Seeking, by ParasitoidsHost Seeking, for PlantsLearningMagnetic SenseMating BehaviorsMechanoreceptionMigrationMimicryNest BuildingOrientationOviposition BehaviorParental CarePheromonesPhoresyPredation/Predatory InsectsPuddling BehaviorRecruitment CommunicationSex DeterminationSexual SelectionSociality Swimming, Lake Insects

    Swimming, Stream InsectsThermoregulationVibrational CommunicationWalking and Jumping

    EVOLUTIONAmberAposematic ColorationBiogeographical PatternsCoevolutionFossil RecordGenetic VariationIndustrial MelanismInsecticide and Acaricide

    ResistanceIsland Biogeography and EvolutionMimicryNomenclature and Classification,

    Principles ofPhylogeny of InsectsSexual SelectionSociality Wolbachia

    REPRODUCTIONAccessory GlandsEgg CoveringsEggsEmbryogenesisMating BehaviorsOvariolesOviposition BehaviorParthenogenesisPolyembryonyReproduction, FemaleReproduction, Female: Hormonal

    Control ofReproduction, MaleReproduction, Male: Hormonal

    Control ofSpermathecaSpermatophoreVitellogenesis

    DEVELOPMENT ANDMETAMORPHOSISBody SizeCaterpillarsChrysalisCocoonDevelopment, Hormonal

    Control ofEcdysteroidsEgg CoveringsGrowth, IndividualHypermetamorphosisImaginal DiscsJuvenile HormonesLarvaMetamorphosisMoltingNeosomyRoyal JellyTemperature, Effects on

    Development and Growth

    MAJOR GROUPS ANDNOTABLE FORMSAntsAphidsApterygotaArchaeognathaArthropoda and Related GroupsAuchenorrhynchaBlattodeaBoll WeevilBombyx moriCat FleasCicadasCodling MothColeopteraCollembolaColorado Potato BeetleCrickets Daddy-Long-Legs DermapteraDiplura

    xiv Content s by Subjec t Area

  • Diptera Drosophila melanogasterDung BeetlesEmbiidina EndopterygotaEphemeroptera ExopterygotaFire AntsGrylloblattodea Gypsy MothHouse Fly HymenopteraInsecta, OverviewIsopteraJapanese BeetleJune BeetlesLadybugsLepidopteraLice, HumanLocustsMantodeaMantophasmatodeaMecoptera Megaloptera MitesMonarchsMosquitoesNeotropical African BeesNeuroptera Odonata Orthoptera Phasmida Phthiraptera PlecopteraProsorrhynchaProturaPsocopteraPterygotaRaphidiopteraScorpionsSiphonapteraSpidersSternorrhynchaStrepsipteraThysanopteraTicks Trichoptera

    WaspsZorapteraZygentoma

    INTERACTIONS WITHOTHER ORGANISMSAposematic ColorationBlood SuckingCat FleasChiggers and Other Disease-

    Causing MitesDefensive BehaviorDog HeartwormFeeding BehaviorHost Seeking, by ParasitoidsHost Seeking, for PlantsHyperparasitismMimicryParasitoidsPathogens of InsectsPhoresyPhytophagous InsectsPredation/Predatory InsectsSymbionts Aiding DigestionSymbionts, BacterialVenomVeterinary EntomologyWolbachia

    INTERACTIONS WITHHUMANSApis Species Bee ProductsBeekeepingBeeswaxBlood SuckingBombyx moriBubonic PlagueChiggers and Other Disease-

    Causing MitesCommercial Products from InsectsCultural Entomology

    DDTDengueExtension EntomologyFolk Beliefs and SuperstitionsFood, Insects asForensic EntomologyHoneyInsecticidesIntegrated Pest ManagementLice, Human MalariaMedical EntomologyMedicine, Insects inMosquitoesMuseums and Display CollectionsRegulatory EntomologyRiver BlindnessSilk ProductionTsetse FlyYellow FeverZoonoses, Arthropod-Borne

    HABITATSAquatic HabitatsCave InsectsForest HabitatsGrassland HabitatsMarine InsectsSoil HabitatsStored Products as HabitatsUrban Habitats

    ECOLOGYAgricultural EntomologyAposematic ColorationBiodiversityBiogeographical PatternsBiological Control of Insect PestsBorersCoevolutionConservationCrypsis

    Content s by Subjec t Area xv

  • DDTEndangered InsectsGallmaking and InsectsGenetic VariationGenetically Modified PlantsGreenhouse Gases, Global

    Warming, and InsectsHoneydewHyperparasitismInsecticidesInsectivorous PlantsInsectivorous VertebratesIntegrated Pest ManagementIntroduced InsectsMigrationMimicryParasitoidsPathogens of InsectsPhoresyPhysical Control of Insect Pests

    Phytophagous InsectsPhytotoxemiaPlant Diseases and InsectsPlantInsect InteractionsPollination and PollinatorsPollution, Insect Response toPopulation EcologyPredation/Predatory InsectsSwimming, Lake InsectsSwimming, Stream Insects

    HISTORY ANDMETHODOLOGYAmberBiotechnology and InsectsCell CultureCollection and PreservationCultural Entomology

    Entomological SocietiesFolk Beliefs and SuperstitionsForensic EntomologyGenetically Modified PlantsGenetic EngineeringHistory of EntomologyInsect ZoosMedical EntomologyMovies, Insects inMuseums and Display CollectionsNomenclature and Classification,

    Principles ofPhotography of InsectsRearing of InsectsResearch Tools, Insects asSericultureStamps, Insects andSterile Insect TechniqueTeaching ResourcesVeterinary Entomology

    xvi Content s by Subjec t Area

  • xvii

    JOHN H. ACORNUniversity of Alberta, CanadaTeaching Resources

    MICHAEL E. ADAMSUniversity of California, RiversideDevelopment, Hormonal Control of Ecdysteroids Juvenile Hormones

    PETER H. ADLERClemson UniversityLegs

    GILBERTO S. ALBUQUERQUEUniversidade Estadual do Norte Fluminense,BrazilNeuroptera

    RICHARD D. ALEXANDERUniversity of MichiganCrickets

    ROBERT T. ALLENParis, ArkansasDiplura Protura

    MIRIAM ALTSTEINAgricultural Research Organization, VolcaniCenter, IsraelNeuropeptides

    SVEND O. ANDERSENCopenhagen University, DenmarkCuticle ExoskeletonIntegument

    NORMAN H. ANDERSONOregon State UniversityMegaloptera

    DAVID A. ANDOWUniversity of Minnesota, St. PaulGenetically Modified Plants

    MICHAEL F. ANTOLINColorado State UniversitySex Determination

    LARRY G. ARLIANWright State UniversityChiggers and Other Disease-Causing Mites

    HORST ASPOCKUniversity of Vienna, AustriaRaphidioptera

    ULRIKE ASPOCKNatural History Museum, Vienna, AustriaRaphidioptera

    PETER W. ATKINSONUniversity of California, Riverside Genetic Engineering

    ARND BAUMANN Forschungszentrum Jlich, GermanyBiogenic Amines

    NANCY E. BECKAGEUniversity of California, RiversideImmunology

    PETER BELLINGER

    California State University, NorthridgeCollembola

    MAY R. BERENBAUMUniversity of IllinoisMovies, Insects in

    MARTIN B. BERGLoyola University ChicagoGrowth, Individual

    ELIZABETH A. BERNAYSUniversity of ArizonaHost Seeking, for PlantsPhytophagous Insects

    CHRISTER BJORKMANSwedish University of Agricultural Sciences Body Size

    SCOTT HOFFMAN BLACKThe Xerces Society, Portland, OregonEndangered Insects

    SETH S. BLAIRUniversity of Wisconsin, MadisonImaginal Discs

    WOLFGANG BLENAUUniversitt Potsdam, GermanyBiogenic Amines

    MURRAY S. BLUMUniversity of GeorgiaChemical Defense

    BRYONY C. BONNINGIowa State UniversityBiotechnology and Insects

    TIMOTHY J. BRADLEYUniversity of California, IrvineExcretion

    PAUL M. BRAKEFIELDLeiden University, The NetherlandsCrypsis

    JOHN E. BRITTAINUniversity of Oslo, NorwayEphemeroptera

    LINCOLN P. BROWERSweet Briar CollegeMonarchs

    ANDREAS BRUNEUniversitt Konstanz, GermanySymbionts Aiding Digestion

    WENDELL E. BURKHOLDERUniversity of Wisconsin, MadisonStored Products as Habitats Deceased.

    CONTRIBUTORS

  • GEORGE W. BYERSUniversity of KansasMecoptera

    RING T. CARDEUniversity of California, RiversideInsecta, OverviewOrientationPheromones

    REGINALD F. CHAPMANUniversity of ArizonaFeeding BehaviorLocustsMouthpartsRostrum

    LANNA CHENGScripps Institution of Oceanography, La JollaMarine Insects

    KENNETH A. CHRISTIANSENGrinnell College, IowaCollembola

    THOMAS M. CLARKIndiana University, South BendWater and Ion Balance, Hormonal Control of

    DONALD G. COCHRANVirginia Polytechnic Institute and State UniversityBlattodea

    EPHRAIM COHENThe Hebrew University of JerusalemChitinFat Body

    ANDREJ COKLNational Institute of Biology, Ljubljana, SloveniaVibrational Communication

    GREGORY W. COURTNEY Iowa State UniversityDiptera

    CHARLES V. COVELL, JR.University of LouisvilleCollection and PreservationStamps, Insects and

    CATHERINE L. CRAIGHarvard University/Tufts University Silk Production

    EVA CRANEInternational Bee Research AssociationApis SpeciesBeekeepingBee ProductsBeeswaxHoneyRoyal Jelly

    PETER S. CRANSTONUniversity of California, DavisPhylogeny of Insects

    GREGORY A. DAHLEMNorthern Kentucky UniversityHouse Fly

    DONALD L. DAHLSTENUniversity of California, BerkeleyGallmaking and Insects

    GENE R. DEFOLIARTUniversity of Wisconsin, MadisonFood, Insects as

    IAN DENHOLMRothamsted Research, United KingdomInsecticide and Acaricide Resistance

    DAVID L. DENLINGEROhio State UniversityCold/Heat ProtectionDiapause

    GREGOR J. DEVINERothamsted Research, United KingdomInsecticide and Acaricide Resistance

    MICHAEL DICKINSONCalifornia Institute of TechnologyFlight

    CHRISTOPHER H. DIETRICHIllinois Natural History SurveyAuchenorrhyncha

    HUGH DINGLEUniversity of California, DavisMigration

    ANGELA E. DOUGLASUniversity of York, United KingdomHoneydew

    ROBERT V. DOWELLCalifornia Department of Food and AgricultureRegulatory Entomology

    ROBERT DUDLEYUniversity of California, BerkeleyFlight

    JOHN D. EDMANUniversity of California, Davis Medical Entomology

    BRUCE F. ELDRIDGEUniversity of California, DavisMosquitoes

    JOSEPH S. ELKINTONUniversity of MassachusettsGypsy MothPopulation Ecology

    MICHAEL S. ENGELUniversity of KansasZoraptera

    JOACHIM ERBERTechnische Universitt Berlin, GermanyBiogenic Amines

    BRIAN A. FEDERICIUniversity of California, RiversidePathogens of Insects

    LEWIS J. FELDMANUniversity of California, Berkeley Insectivorous Plants

    CLELIA FERREIRAUniversity of So Paulo, BrazilDigestive System

    R. NELSON FOSTERU.S. Department of AgricultureBoll Weevil

    GORDON W. FRANKIEUniversity of California, BerkeleyPollination and Pollinators

    NIGEL R. FRANKSUniversity of Bristol, United KingdomAnts

    ANDREW S. FRENCHDalhousie University, CanadaMechanoreception

    DOUGLAS J. FUTUYMAUniversity of MichiganCoevolution

    ERIN C. GENTRYUniversity of FloridaBioluminescence

    HELEN GHIRADELLAState University of New York, AlbanyColoration

    ROSEMARY G. GILLESPIEUniversity of California, BerkeleyIsland Biogeography and EvolutionSpiders

    GONZALO GIRIBETHarvard UniversityDaddy-Long-Legs

    M. LEE GOFFChaminade University of Honolulu Forensic Entomology

    GORDON GORDHU.S. Department of AgricultureAnatomy: Head, Thorax, Abdomen, and Genitalia

    MIODRAG GRBICUniversity of Western Ontario, CanadaEmbryogenesis

    LES GREENBERGUniversity of California, RiversideFire Ants

    DAVID GRIMALDIAmerican Museum of Natural History, New York Fossil Record

    PENNY J. GULLANUniversity of California, DavisPhylogeny of InsectsSternorrhyncha

    DARRYL T. GWYNNEUniversity of Toronto, CanadaMating Behaviors

    xviii Contributors

  • JON F. HARRISONArizona State UniversityRespiratory SystemTracheal System

    MICHAEL W. HASTRITERBrigham Young UniversitySiphonaptera

    DAVID H. HEADRICKCalifornia Polytechnic State UniversityAnatomy: Head, Thorax, Abdomen, andGenitalia

    BERND HEINRICHUniversity of VermontThermoregulation

    DAVID W. HELDUniversity of KentuckyJapanese BeetleJune Beetles

    RONALD A. HELLENTHALUniversity of Notre DamePhthiraptera

    JORGE HENDRICHSFAO/IAE Division, Vienna, AustriaSterile Insect Technique

    ADAM D. HENKColorado State UniversitySex Determination

    NANCY C. HINKLEUniversity of GeorgiaCat FleasExtension Entomology

    M. S. HODDLEUniversity of California, RiversideBiological Control of Insect Pests

    JAMES N. HOGUECalifornia State University, NorthridgeCultural EntomologyFolk Beliefs and Superstitions

    MARILYN A. HOUCKTexas Tech UniversityPhoresy

    FRANCIS G. HOWARTHB. P. Bishop Museum, Honolulu, Hawaii Cave Insects

    RON HOYCornell UniversityHearing

    LAWRENCE E. HURDWashington and Lee UniversityMantodea

    MICHAEL E. IRWINUniversity of IllinoisCommercial Products from Insects

    RUDOLF JANDERUniversity of KansasMagnetic Sense

    ROBERT L. JEANNEUniversity of Wisconsin, MadisonNest Building

    MATHIEU JORONLeiden University, The NetherlandsAposematic ColorationMimicry

    ROBERT JOSEPHSONUniversity of California, IrvineMuscle System

    GAIL E. KAMPMEIERUniversity of IllinoisCommercial Products from Insects

    KENNETH Y. KANESHIROUniversity of HawaiiSexual Selection

    MICHAEL R. KANOSTKansas State UniversityHemolymph

    ALAN I. KAPLANEast Bay Regional Park District, Berkeley,CaliforniaEntomological Societies

    JOE B. KEIPERCleveland Museum of Natural HistoryDiptera

    GEORGE G. KENNEDYNorth Carolina State UniversityColorado Potato Beetle

    LAWRENCE R. KIRKENDALLUniversity of Bergen, NorwayParthenogenesis

    KLAUS-DIETER KLASSZoological Museum, Dresden, GermanyMantophasmatodea

    JOHN KLOTZUniversity of California, RiversideMagnetic Sense

    MARC J. KLOWDENUniversity of IdahoOviposition BehaviorReproduction, MaleReproduction, Male: Hormonal Control ofSpermathecaSpermatophore

    MARCOS KOGANOregon State UniversityAgricultural EntomologyIntegrated Pest Management

    VLADIMIR KOSTALInstitute of Entomology, Academy of Sciences,Czech RepublicTemperature, Effects on Developmentand Growth

    ANDREAS KRUESSUniversity of Gttingen, GermanyGrassland Habitats

    MICHAEL F. LANDUniversity of Sussex, United KingdomEyes and Vision

    ROBERT S. LANEUniversity of California, BerkeleyZoonoses, Arthropod-Borne

    STEPHEN G. A. LEAKInternational Trypanotolerance Centre, TheGambiaTsetse Fly

    RICHARD E. LEE, JR.Miami University, Oxford, Ohio DormancyHibernation

    M. J. LEHANEUniversity of Wales, BangorBlood Sucking

    NORMAN C. LEPPLAUniversity of FloridaRearing of Insects

    RICHARD J. LESKOSKYUniversity of IllinoisMovies, Insects in

    VERNARD R. LEWISUniversity of California, BerkeleyIsoptera

    JAMES K. LIEBHERRCornell UniversityColeoptera

    JAMES E. LLOYDUniversity of FloridaBioluminescence

    CATHERINE LOUDONUniversity of KansasAntennae

    DWIGHT E. LYNNU.S. Department of AgricultureCell Culture

    MICHAEL E. N. MAJERUSUniversity of Cambridge, United KingdomIndustrial MelanismLadybugsSymbionts, Bacterial

    JON H. MARTINThe Natural History Museum, LondonSternorrhyncha

    SINZO MASAKIHirosaki University, JapanAestivation

    LINDA J. MASON Purdue UniversityExtension Entomology

    FUMIO MATSUMURAUniversity of California, DavisDDTInsecticides

    Contributors xix

  • JOSEPH V. MCHUGHUniversity of GeorgiaColeoptera

    TERRI L. MEINKINGUniversity of Miami School of Medicine Lice, Human

    RICHARD W. MERRITTMichigan State UniversityAquatic HabitatsDipteraGrowth, Individual

    JOCELYN G. MILLAR University of California, RiversidePheromones

    THOMAS A. MILLERUniversity of California, Riverside Circulatory System

    NICK MILLSUniversity of California, BerkeleyParasitoids

    B. K. MITCHELLUniversity of Alberta, CanadaChemoreception

    THOMAS E. MITTLERUniversity of California, BerkeleyHoneydew

    EDWARD L. MOCKFORDIllinois State UniversityPsocoptera

    MARK W. MOFFETTUniversity of California, BerkeleyPhotography of Insects

    THOMAS P. MONATHAcambis Inc., Cambridge, Massachusetts Yellow Fever

    JOHN C. MORSEClemson UniversityTrichoptera

    MAX S. MOULDSAustralian Museum, SydneyCicadas

    LAURENCE A. MOUNDCSIRO, Canberra, AustraliaThysanoptera

    BRADLEY A. MULLENSUniversity of California, Riverside Veterinary Entomology

    WERNER NACHTIGALLUniversitt der Saarlandes, GermanySwimming, Lake Insects

    LISA NAGYUniversity of ArizonaEmbryogenesis

    OLDRICH NEDVEDInstitute of Entomology, Academy of Sciences,Czech RepublicTemperature, Effects on Developmentand Growth

    TIM R. NEWLa Trobe University, AustraliaConservation

    GORDON M. NISHIDAUniversity of California, BerkeleyMuseums and Display Collections

    BENJAMIN NORMARKUniversity of MassachusettsParthenogenesis

    DAVID A. OBROCHTAUniversity of Maryland Biotechnology InstituteGenetic Engineering

    BARRY M. OCONNORUniversity of MichiganMites

    SEAN ODONNELLUniversity of WashingtonCasteColonies

    PATRICK M. OGRADYAmerican Museum of Natural History, New York Drosophila melanogaster

    DANIEL OTTEPhiladelphia Academy of Natural SciencesCrickets

    TERRY L. PAGEVanderbilt UniversityCircadian Rhythms

    TIMOTHY D. PAINEUniversity of California, RiversideBorers

    JAMES O. PALMERAllegheny CollegeDermaptera

    BERNARD PANNETONAgriculture and Agri-Food Canada, QuebecPhysical Control of Insect Pests

    DANIEL R. PAPAJUniversity of ArizonaLearning

    NIPAM H. PATELUniversity of ChicagoSegmentation

    MATS W. PETTERSSONSwedish University of Agricultural Sciences Body Size

    JOHN D. PINTOUniversity of California, RiversideHypermetamorphosis

    RUDY PLARREFederal German Institute for Materials Researchand Testing, GermanyStored Products as Habitats

    EDWARD G. PLATZERUniversity of California, Riverside Dog Heartworm

    GEORGE POINAR JR.Oregon State UniversityAmber

    DANIEL A. POTTERUniversity of KentuckyJapanese Beetle June Beetles

    JERRY A. POWELLUniversity of California, BerkeleyLepidoptera

    ROGER D. PRICEUniversity of MinnesotaPhthiraptera

    RONALD PROKOPYUniversity of MassachusettsAgricultural EntomologyIntegrated Pest Management

    ALEXANDER H. PURCELLUniversity of California, BerkeleyPhytotoxemiaPlant Diseases and Insects

    DONALD L. J. QUICKEImperial College, University of London,United KingdomHymenoptera

    FRANK J. RADOVSKYOregon State UniversityNeosomy

    SUSAN M. RANKINAllegheny CollegeDermaptera

    WILLIAM K. REISENUniversity of California, DavisMalaria

    D. C. F. RENTZCalifornia Academy of Sciences, San FranciscoGrylloblattodeaOrthoptera

    VINCENT H. RESHUniversity of California, BerkeleyInsecta, OverviewPollution, Insect Response toRiver Blindness

    LYNN M. RIDDIFORDUniversity of WashingtonMolting

    JAMES RIDSDILL-SMITHCSIRO, Canberra, AustraliaDung Beetles

    xx Contributors

  • ROY E. RITZMANNCase Western Reserve UniversityWalking and Jumping

    ALAN ROBINSONIAEA Laboratories, Seibersdorf, Austria SterileInsect Technique

    GENE E. ROBINSONUniversity of Illinois, Urbana-ChampaignDivision of Labor in Insect Societies

    GEORGE K. RODERICKUniversity of California, BerkeleyGenetic VariationIsland Biogeography and Evolution

    DAVID M. ROSENBERGFreshwater Institute, Winnipeg, CanadaPollution, Insect Response to

    EDWARD S. ROSSCalifornia Academy of SciencesEmbiidina

    MICHAEL K. RUSTUniversity of California, RiversideCat FleasUrban Habitats

    MICHEL SARTORIMuseum of Zoology, Lausanne, SwitzerlandEphemeroptera

    LESLIE SAUL-GERSHENZCenter for Ecosystem Survival, San Francisco,CaliforniaInsect Zoos

    CARL W. SCHAEFERUniversity of ConnecticutProsorrhyncha

    KATHERINE N. SCHICKUniversity of California, BerkeleyGallmaking and Insects

    JUSTIN O. SCHMIDTSouthwestern Biological Institute, Tucson, ArizonaDefensive BehaviorVenomWasps

    MICHELLE PELLISSIER SCOTTUniversity of New HampshireParental Care

    THOMAS W. SCOTTUniversity of California, DavisDengue

    J. MARK SCRIBERMichigan State UniversityPlantInsect Interactions

    FRANTISEK SEHNALInstitute of Entomology, Academy of Sciences,Czech RepublicTemperature, Effects on Developmentand Growth

    IRWIN W. SHERMANUniversity of California, RiversideBubonic Plague

    RONALD A. SHERMANUniversity of California, IrvineMedicine, Insects in

    DANIEL SIMBERLOFFUniversity of TennesseeIntroduced Insects

    S. J. SIMPSONUniversity of Oxford, United KingdomNutrition

    SCOTT R. SMEDLEYTrinity College, ConnecticutPuddling Behavior

    EDWARD H. SMITHCornell University (Emeritus)Asheville, North CarolinaHistory of Entomology

    JANET R. SMITHAsheville, North CarolinaHistory of Entomology

    DANIEL E. SONENSHINEOld Dominion UniversityTicks

    JOHN T. SORENSONCalifornia Department of Food and AgricultureAphids

    JOSEPH C. SPAGNAUniversity of California, BerkeleySpiders

    BEVERLY SPARKSUniversity of GeorgiaExtension Entomology

    FELIX A. H. SPERLINGUniversity of Alberta, CanadaTeaching Resources

    BERNHARD STATZNERUniversit Lyon I, FranceSwimming, Stream Insects

    INGOLF STEFFANDEWENTERUniversity of Gttingen, GermanyGrassland Habitats

    FREDERICK W. STEHRMichigan State UniversityCaterpillarsChrysalisCocoonLarvaMetamorphosisOcelli and StemmataPupa and Puparium

    KENNETH W. STEWARTUniversity of North TexasPlecoptera

    PETER STILINGUniversity of South FloridaGreenhouse Gases, Global Warming, andInsects

    ANDREW J. STORERMichigan Technological University Forest Habitats

    NIGEL E. STORKJames Cook University, AustraliaBiodiversity

    RICHARD STOUTHAMERUniversity of California, RiversideWolbachia

    MICHAEL R. STRANDUniversity of GeorgiaPolyembryony

    NICHOLAS J. STRAUSFELDUniversity of ArizonaBrain and Optic Lobes

    HELMUT STURMUniversity Hildesheim, GermanyArchaeognathaZygentoma

    YOU NING SUAustralian National University, CanberraOrthoptera

    R. K. SUAREZUniversity of California, Santa BarbaraMetabolism

    DANIEL J. SULLIVANFordham UniversityHyperparasitism

    SATOSHI TAKEDANational Institute of Agrobiological Sciences,JapanBombyx moriSericulture

    CATHERINE A. TAUBERCornell UniversityNeuroptera

    MAURICE J. TAUBERCornell UniversityNeuroptera

    ORLEY R. TAYLORUniversity of KansasNeotropical African Bees

    WILLIAM H. TELFERUniversity of PennsylvaniaVitellogenesis

    K. J. TENNESSENTennessee Valley AuthorityOdonata

    WALTER R. TERRAUniversity of So Paulo, BrazilDigestionDigestive System

    CARSTEN THIESUniversity of Gttingen, GermanyGrassland Habitats

    Contributors xxi

  • xxii Contributors

    F. CHRISTIAN THOMPSONU.S. Department of AgricultureNomenclature and Classification, Principles of

    S. NELSON THOMPSONUniversity of California, Riverside MetabolismNutrition

    JAMES H. THORPUniversity of KansasArthropoda and Related Groups

    ROBBIN W. THORPUniversity of California, DavisPollination and Pollinators

    ERICH H. TILGNERUniversity of GeorgiaPhasmida

    PAIVI H. TORKKELIDalhousie University, CanadaMechanoreception

    JAMES F. A. TRANIELLOBoston UniversityRecruitment Communication

    TEJA TSCHARNTKEUniversity of Gttingen, GermanyGrassland Habitats

    KAREN M. VAILUniversity of Tennessee, Knoxville Extension Entomology

    R. G. VAN DRIESCHEUniversity of MassachusettsBiological Control of Insect Pests

    MACE VAUGHANThe Xerces Society, Portland, OregonEndangered Insects

    CHARLES VINCENTAgriculture and Agri-Food Canada, QuebecPhysical Control of Insect Pests

    META VIRANT-DOBERLETNational Institute of Biology, Ljubljana, SloveniaVibrational Communication

    P. KIRK VISSCHERUniversity of California, RiversideDance LanguageHomeostasis, Behavioral

    PATRICIA J. VITTUMUniversity of MassachusettsSoil Habitats

    GREGORY P. WALKERUniversity of California, RiversideSalivary Glands

    J. BRUCE WALLACEUniversity of GeorgiaAquatic Habitats

    GRAHAM C. WEBBThe University of Adelaide, AustraliaChromosomes

    STEPHEN C. WELTERUniversity of California, BerkeleyCodling Moth

    RONALD M. WESELOHConnecticut Agricultural Experiment StationHost Seeking, by ParasitoidsPredation/Predatory Insects

    DIANA E. WHEELERUniversity of Arizona Accessory GlandsEggsEgg CoveringsOvariolesReproduction, FemaleReproduction, Female: Hormonal Control of

    MICHAEL F. WHITINGBrigham Young UniversitySiphonapteraStrepsiptera

    KIPLING W. WILLUniversity of California, BerkeleyResearch Tools, Insects as

    STANLEY C. WILLIAMSSan Francisco State UniversityScorpions

    SHAUN L. WINTERTONNorth Carolina State UniversityScales and Setae

    DAVID L. WOODUniversity of California, BerkeleyForest Habitats

    ROBIN J. WOOTTONUniversity of Exeter, United KingdomWings

    JAYNE YACKCarleton University, Ottawa, CanadaHearing

    JAMES E. ZABLOTNYU.S. Department of AgricultureSociality

    SASHA N. ZILLMarshall UniversityWalking and Jumping

    PETER ZWICKMax-Planck-Institut fr Limnologie, GermanyBiogeographical Patterns

  • xxiii

    T he Encyclopedia of Insects is a complete source of informationon the subject of insects, contained within a single volume.Each article in the Encyclopedia provides an overview of theselected topic to inform a broad spectrum of readers, frominsect biologists and scientists conducting research in relatedareas, to students and the interested general public.

    In order that you, the reader, will derive the maximumbenefit from the Encyclopedia of Insects, we have provided thisGuide. It explains how the book is organized and how theinformation within its pages can be located.

    SUBJECT AREAS

    The Encyclopedia of Insects presents 271 separate articles onthe entire range of entomological study. Articles in the Ency-clopedia fall within twelve general subject areas, as follows:

    Anatomy Physiology Behavior Evolution Reproduction Development and Metamorphosis Major Groups and Notable Forms Interactions with Other Organisms Interactions with Humans Habitats Ecology History and Methodology

    ORGANIZATION

    The Encyclopedia of Insects is organized to provide the maximumease of use for its readers. All of the articles are arranged in asingle alphabetical sequence by title. An alphabetical Table of

    Contents for the articles can be found beginning on p. v ofthis introductory section.

    As a reader of the Encyclopedia, you can use this alphabeti-cal Table of Contents by itself to locate a topic. Or you can firstidentify the topic in the Contents by Subject Area (p. xiii) andthen go to the alphabetical Table to find the page location.

    So that they can be more easily identified, article titles beginwith the key word or phrase indicating the topic, with anydescriptive terms following this. For example, Temperature,Effects on Development and Growth is the title assigned tothis article, rather than Effects of Temperature on Develop-ment and Growth, because the specific term Temperature isthe key word.

    ARTICLE FORMAT

    Each article in the Encyclopedia begins with an introductoryparagraph that defines the topic being discussed and indicatesits significance. For example, the article Exoskeleton beginsas follows:

    The exoskeleton is noncellular material that is located on topof the epidermal cell layer and constitutes the outermost partof the integument. The local properties and appearance ofthe exoskeleton are highly variable, and nearly all visiblefeatures of an insect result from the exoskeleton. Theexoskeleton serves as a barrier between the interior of theinsect and the environment, preventing desiccation and thepenetration of microorganisms. Muscles governing themovements are attached to the exoskeleton.

    Major headings highlight important subtopics that are dis-cussed in the article. For example, the article Flight includesthese topics: Evolution of Flight; Aerodynamics; NeuralControl; Energetics; Ecology and Diversity.

    GUIDE TO THEENCYCLOPEDIA

  • CROSS-REFERENCES

    The Encyclopedia of Insects has an extensive system of cross-referencing. References to other articles may appear either asmarginal headings within the AZ topical sequence, or asindications of related topics at the end of a particular article.

    As an example of the first type of reference cited above, thefollowing marginal entry appears in the AZ article list betweenthe entries Beeswax and Biodiversity:

    Beetle see Coleoptera

    This reference indicates that the topic of Beetles is discussedelsewhere, under the article title Coleoptera, which is thename of the order including this group.

    An example of the second type, a cross-reference at the endof an article, can be found in the entry DDT. This articleconcludes with the statement:

    See Also the Following ArticlesInsecticides Integrated Pest Management Pollution

    This reference indicates that these three related articles allprovide some additional information about DDT.

    BIBLIOGRAPHY

    The Bibliography section appears as the last element of anarticle, under the heading Further Reading. This section listsrecent secondary sources that will aid the reader in locatingmore detailed or technical information on the topic at hand.Review articles and research papers that are important to amore detailed understanding of the topic are also listed here.

    The Bibliography entries in this Encyclopedia are for thebenefit of the reader, to provide references for further readingor additional research on the given topic. Thus they typicallyconsist of a limited number of entries. They are not intended

    to represent a complete listing of all the materials consulted bythe author or authors in preparing the article. The Biblio-graphy is in effect an extension of the article itself, and itrepresents the authors choice as to the best sources availablefor additional information.

    GLOSSARY

    The Encyclopedia of Insects presents an additional resource forthe reader, following the AZ text. A comprehensive glossaryprovides definitions for more than 750 specialized terms usedin the articles in this Encyclopedia. The terms were identifiedby the contributors as helpful to the understanding of theirentries, and they have been defined by these authorsaccording to their use in the actual articles.

    INDEX

    The Subject Index for the Encyclopedia of Insects contains morethan 7,000 entries. Within the entry for a given topic, refer-ences to general coverage of the topic appear first, such as acomplete article on the subject. References to more specificaspects of the topic then appear below this in an indented list.

    ENCYCLOPEDIA WEB SITE

    The Encyclopedia of Insects maintains its own editorial Web pageon the Internet at:

    http://www.apnet.com/insects/

    This site gives information about the Encyclopedia projectand features links to related sites that provide informationabout the articles of the Encyclopedia. The site will continueto evolve as more information becomes available.

    xxiv Giude to the Encyc lopedia

  • xxv

    I would say that creating an encyclopedia of insects was aherculean task, but I think that sells the enterprise short.After all, Hercules only had twelve labors assigned to him,and twelve years to complete themwith insects, there areover 900,000 different species and many, many more storiesto tell. Twelve years from now, there will likely be even more.Why, then, would anyone undertake the seemingly impos-sible task of compiling an encyclopedia of insects? To anentomologist, the answer is obvious. For one thing, theresthe numbers argumentover 70% of all known species areinsects, so if any group merits attention in encyclopedicform, surely its the one that happens to dominate the planet.Moreover, owing in large part to their staggering diversity,insects are in more different places in the world than virtuallyany other organism. There are insects in habitats rangingfrom the high Arctic to tropical rainforests to petroleumpools to glaciers to mines a mile below the surface to caves tosea lion nostrils and horse intestines. About the only placewhere insects are conspicuously absent is in the deep ocean(actually, in deep water in general), an anomaly that hasfrustrated more than a few entomologists who have grownaccustomed to world domination. Then theres the fact thatinsects have been around for longer than most other high-profile life-forms. The first proto-insects date back some 400million years; by contrast, mammals have been around onlyabout 230 million years and humans (depending on howtheyre defined) a measly one million years.

    Probably the best justification for an encyclopediadevoted to insects is that insects have a direct and especiallyeconomic impact on humans. In the United States alone,insects cause billions of dollars in losses to staple crops, fruitcrops, truck crops, greenhouse and nursery products, forestproducts, livestock, stored grain and packaged food,clothing, household goods and furniture, and just aboutanything else people try to grow or build for sale or for their

    own consumption. Beyond the balance sheet, they causeincalculable losses as vectors of human pathogens. Theyreinvolved in transmission of malaria, yellow fever, typhus,plague, dengue, various forms of encephalitis, relapsing fever,river blindness, filariasis, sleeping sickness, and innumerableother debilitating or even fatal diseases, not just abroad inexotic climes but here in the United States as well. All told,insects represent a drag on the economy unequaled by anyother single class of organisms, a seemingly compellingreason for keeping track of them in encyclopedic form.

    In the interests of fairness, however, it should bementioned that insects also amass economic benefits in amagnitude unequaled by most invertebrates (or even,arguably, by most vertebrates). Insect-pollinated crops in theUnited States exceed $9 billion in value annually, and insectproducts, including honey, wax, lacquer, silk, and so on,contribute millions more. Insect-based biological control ofboth insect and weed pests is worth additional millions inreclaimed land and crop production, and even insect disposalof dung and other waste materials, although decidedlyunglamorous, is economically significant in fields, rangelands,and forests throughout the country.

    So, for no reason other than economic self-interest, theresreason enough for creating an encyclopedia of insects. Butwhat can be learned from insects that cant be learned froman encyclopedia of any other abundant group of organisms?Basically, the biology of insects is the biology of small size.Small size, which has been in large part responsible for theoverwhelming success of the taxon, at the same time imposesmajor limits on the taxon. The range in size of livingorganisms, on earth at least, encompasses some 13 orders ofmagnitude (from a 100 metric ton blue whale to rotifersweighing less than 0.01 mg). Insects range over five ordersof magnitudefrom 30-g beetles to 0.03-g fairyfliessoeight orders of magnitude are missing in the class Insecta.

    FOREWORD

  • Problems at the upper limit involve support, transport, andovercoming inertia, issues clearly not critical for organisms,like insects, at the lower end of the range.

    We humans, in the grand scheme of things, are big creaturesand as a consequence we interact with the biological and phys-ical world entirely differently. Rules that constrain humanbiology often are suspended for insects, which operate by acompletely different set of rules. The constraints and benefitsof small size are reflected in every aspect of insect biology.They hear, smell, taste, and sense the world in every other waywith abilities that stagger the imagination. They are capableof physical feats that seem impossiblemost fly, some glowin the dark, and others control the sex of their offspring andeven occasionally engage in virgin birth, to cite a few examples.Their generation times are so short and reproductive rates sohigh that they can adapt and evolve at rates that continuallysurprise (and stymie) us. The environment is patchier tosmaller organisms, which can divide resources more finelythan can large, lumbering species. Thus, they can make aliving on resources so rare or so nutrient-poor that it defiesbelief, such as nectar, dead bodies, and even dung.

    So theyre profoundly different from humans and otherbig animals, and the study of insects can offer many insights

    into life on earth that simply couldnt be gained from a studyof big creatures. By the same token, though, they are cutfrom the same cloththe same basic building blocks of life,same genetic code, and the likeand their utility as researchorganisms has provided insights into all life on the planet.

    The Encyclopedia of Insects contains contributions fromsome of the greatest names in entomology today. Such a workhas to be a collective effort because nobody can be an expertin everything entomological. Even writing a foreword forsuch a wide-ranging volume is a daunting task. To be such anexpert would mean mastering every biological science frommolecular biology (in which the fruit fly Drosophila melanogasterserves as a premier model organism) to ecosystem ecology (inwhich insects play an important role in rates of nutrientturnover and energy flow). But, because insects, through theirubiquity and diversity, have had a greater influence on humanactivities than perhaps any other class of organisms, to be theultimate authority on insects also means mastering the minu-tiae of history, economics, art, literature, politics, and evenpopular culture. Nobody can master all of that informationand thats why this encyclopedia is such a welcome volume.

    May R. Berenbaum

    xxvi Foreword

  • xxvii

    I nsects are ever present in human lives. They are at once aweinspiring, fascinating, beautiful, and, at the same time, ascourge of humans because of food loss and disease. Yetdespite their negative effects, we depend on insects forpollination and for their products. As insects are the largestliving group on earth (75% of all animal species), anyunderstanding of ecological interactions at local or globalscales depends on our knowledge about them. Given thecurrent interest in biodiversity, and its loss, it must beremembered that insects represent the major part of existingbiodiversity. Aesthetically, insect images are often with us aswell: early images include Egyptian amulets of sacred scarabs;modern images include dragonfly jewelry, butterflystationery, and childrens puppets.

    The idea of an Encyclopedia of Insects is new, but theconcept of an encyclopedia is quite old. In 1745, Diderotand DAlembert asked the best minds of their eraincludingVoltaire and Montesquieuto prepare entries that wouldcompile existing human knowledge in one place: the worldsfirst encyclopedia. It took over 20 years to finish the firstedition, which became one of the worlds first best-sellingbooks and a triumph of the Enlightenment.

    What do we intend this encyclopedia to be? Our goal is toconvey the exciting, dynamic story of what entomology istoday. It is intended to be a concise, integrated summary ofcurrent knowledge and historical background on each of thenearly 300 entries presented. Our intention has been to makethe encyclopedia scientifically uncompromising; it is to becomprehensive but not exhaustive. Cross-references point thereader to related topics, and further reading lists at the end ofeach article allow readers to go into topics in more detail. Thepresence of a certain degree of overlap is intentional, becauseeach article is meant to be self-contained.

    The Encyclopedia of Insects also includes organisms that arerelated to insects and often included in the purview of

    entomology. Therefore, besides the members of the classInsectathe true insectsthe biology of spiders, mites, andrelated arthropods is included. The core of this encyclopediaconsists of the articles on the taxonomic groupsthe 30 orso generally accepted orders of insects, the processes thatinsects depend on for their survival and success, and therange of habitats they occupy. The fact that entomology is adynamic field is emphasized by the discovery of a new orderof insects, the Mantophasmatodea, just as this encyclopediawas being completed. This is the first order of insects to bedescribed in over 80 years, and we are pleased to be able toinclude it as an entry, further underscoring that there is muchleft to learn about insects. Some topics, especially the posterinsectsthose well-known taxa below the level of orders forwhich entries are presentedmay not cover all that aredesired by some readers. Given insect biodiversity, yourindulgence is requested.

    We have gathered over 260 experts worldwide to write onthe entries that we have selected for inclusion. Thesespecialists, of course, have depended on the contributions ofthousands of their entomological predecessors. Because themodern study of entomology is interdisciplinary, we enlistedexperts ranging from arachnologists to specialists in zoonoticdiseases. Given that the two of us have spent over 25 com-bined years as editors of the Annual Review of Entomology,many of our contributors were also writers for that peri-odical. We thank our contributors for putting up with ourcompulsive editing, requests for rewrites, and seeminglyendless questions.

    Our intended audience is not entomological specialistsbut entomological generalists, whether they be students,teachers, hobbyists, or interested nonscientists. Therefore, tocover the diverse interests of this readership, we haveincluded not just purely scientific aspects of the study ofinsects, but cultural (and pop-cultural) aspects as well.

    PREFACE

  • We thank the staff of Academic Press for their encour-agement and assistance on this project. Chuck Crumly hadthe original concept for this encyclopedia, convinced us of itsmerit, and helped us greatly in defining the format. ChrisMorris provided suggestions about its development. JocelynLofstrom and Joanna Dinsmore guided the book throughprinting. Gail Rice managed the flow of manuscripts andrevisions with skill and grace, and made many valuable sug-

    gestions. Julie Todd of Iowa State University provided a crucialfinal edit of the completed articles. All these professionalshave helped make this a rewarding and fascinating endeavor.

    We dedicate our efforts in editing the Encyclopedia ofInsects to our wives, Cheryl and Anja; their contributions toour entomological and personal lives have been indescribable.

    Vincent H. Resh and Ring T. Card

    xxviii Preface

  • Acarisee Mites; Ticks

    Accessory GlandsDiana E. WheelerUniversity of Arizona

    T he accessory glands of reproductive systems in both femaleand male insects produce secretions that function in spermmaintenance, transport, and fertilization. In addition, accessoryglands in females provide protective coatings for eggs. Accessoryglands can be organs distinct from the main reproductive tract,or they can be specialized regions of the gonadal ducts (ductsleading from the ovaries or testes). Typically, glandular tissue iscomposed of two cell types: one that is secretory and the otherthat forms a duct. The interplay between male and femalesecretions from accessory glands is a key element in the designof diverse mating systems.

    ACCESSORY GLANDS OF FEMALES

    Management of Sperm and Other Male Contributions

    Sperm management by females involves a wide range ofprocesses, including liberation of sperm from a sperma-tophore, digestion of male secretions and sperm, transport ofsperm to and from the spermatheca, maintenance of storedsperm, and fertilization.

    Accessory gland secretions can have digestive functionsimportant in sperm management. First, digestive breakdown

    of the spermatophore can free encapsulated sperm for fertil-ization and storage. Second, male contributions can providean important nutritional benefit to their mates. Female secre-tions can digest the secretory components of male seminal fluidto facilitate a nutritive role. In addition, females can digestunwanted sperm to transform it into nutrients. Third, femalesecretions in some species are required to digest sperm cover-ings that inhibit fertilization.

    Transfer of sperm to and from the spermatheca is generallyaccomplished by a combination of chemical signals and mus-cular contractions. Secretions of female accessory glands in somespecies increase sperm motility or appear to attract spermtoward the spermathecae. Transport of fluid out through thewall of the spermatheca may also create negative pressure thatdraws in sperm.

    Sperm can be stored for some length of time in sperma-thecae, with the record belonging to ant queens thatmaintain sperm viability for a decade or more. Secretions ofspermathecal glands are poorly characterized, and how spermis maintained for such extended periods is not known.Spermathecal tissue seems to create a chemical environmentthat maintains sperm viability, perhaps through reducedmetabolism. A nutritional function is also possible.

    Transport of sperm out of storage can be facilitated by thesecretions of the spermathecal gland, which presumably activatequiescent sperm to move toward the primary reproductivetract. One potential function of female accessory glands thathas been explored only slightly is the production of hormone-like substances that modulate reproduction functions.

    Production of Egg Coverings

    Female accessory glands that produce protective coverings foreggs are termed colleterial glands. Colleterial glands havebeen best characterized in cockroaches, which produce anoothecal case surrounding their eggs. Interestingly, the left

    1

    A

  • and right glands are anatomically different and have differentproducts. Separation of the chemicals permits reactions tobegin only at the time of mixing and ootheca formation. Otherprotective substances produced by glands include toxins andantibacterials.

    Nourishment for Embryos or Larvae

    Viviparous insects use accessory glands to provide nourish-ment directly to developing offspring. Tsetse flies and sheepkeds are dipterans that retain single larvae within their repro-ductive tracts and provide them with nourishment. They givebirth to mature larvae ready to pupate. The gland that producesthe nourishing secretion, rich in amino acids and lipids, isknown as the milk gland. The Pacific beetle roach, Diplopterapunctata, is also viviparous and provides its developing embryoswith nourishment secreted by the brood sac, an expandedportion of oviduct.

    ACCESSORY GLANDS OF MALES

    Accessory glands of the male reproductive tract have diversefunctions related to sperm delivery and to the design ofspecific mating systems.

    Sperm Delivery

    Males of many insects use spermatophores to transfer spermto females. A spermatophore is a bundle of sperm contained

    in a protective packet. Accessory glands secrete the structuralproteins necessary for the spermatophores construction.Males of the yellow mealworm, Tenebrio molitor, have twodistinct accessory glands, one bean-shaped and the othertubular (Fig. 1). Bean-shaped accessory glands contain cellsof at least seven types and produce a semisolid material thatforms the wall and core of the spermatophore. Tubularaccessory glands contain only one type of cell, and itproduces a mix of water-soluble proteins of unknownfunction. Spermatophores are not absolutely required forsperm transfer in all insects. In many insects, male secretionscreate a fluid medium for sperm transfer.

    Effects on Sperm Management and on the Female

    The effects of male accessory gland secretions in the femaleare best known for the fruit fly, Drosophila melanogaster, inwhich the function of several gene products has beenexplored at the molecular level. Since insects have a diversityof mating systems, the specific functions of accessory glandsecretions are likely to reflect this variation.

    In Drosophila, the accessory glands are simple sacsconsisting of a single layer of secretory cells around a centrallumen (Fig. 2). Genes for more than 80 accessory glandproteins have been identified so far. These genes code forhormonelike substances and enzymes, as well as many novel

    2 Acces sor y Glands

    FIGURE 1 Male reproductive system of T. molitor, showing testes (T),ejaculatory duct (EJD), tubular accessory gland (TAG), and bean-shapedaccessory gland (BAG). [From Dailey, P. D., Gadzama J. M., and Happ, G.M. (1980). Cytodifferentiation in the accessory glands of Tenebrio molitor. VI.A congruent map of cells and their secretions in the layered elastic product ofthe male bean-shaped accessory gland. J. Morphol. 166, 289322. Reprintedby permission of Wiley-Liss, Inc., a subsidiary of John Wiley & Sons, Inc.]

    FIGURE 2 Accessory gland of D. melanogaster. (A) The cells in this normalaccessory gland express b-galactosidase driven by a promoter of a gene for anaccessory gland protein. (B) A transgenic accessory gland, cells expressing thegene have been selectively killed after eclosion. These flies were used to explorethe function of accessory gland secretions. In transgenic males, accessory glandsare small and translationally inert. [From Kalb, J. M., DiBenedetto, A. J., andWolfner, M. F. (1993). Probing the function of Drosophila melanogasteraccessory glands by directed cell ablation. Proc. Natl. Acad. Sci. USA 90,80938097. Copyright 1993, National Academy of Sciences, U.S.A.]

  • proteins. The gene products or their derivatives have diversefunctions, including an increased egg-laying rate, a reducedinclination of females to mate again, increased effectivenessof sperm transfer to a females spermatheca, and various toxiceffects most likely involved in the competition of sperm fromdifferent males. A side effect of this toxicity is a shortened lifespan for females. Other portions of the reproductive tractcontribute secretions with diverse roles. For example, theejaculatory bulb secretes one protein that is a majorconstituent of the mating plug, and another that hasantibacterial activity.

    See Also the Following ArticlesEgg Coverings Spermatheca Spermatophore

    Further ReadingChen, P. S. (1984). The functional morphology and biochemistry of insect

    male accessory glands and their secretions. Annu. Rev. Entomol. 29,233255.

    Eberhard, W. G. (1996). Female Control: Sexual Selection by CrypticFemale Choice. Princeton University Press, Princeton, NJ.

    Gillott, C. (1988). ArthropodaInsecta. In Accessory Sex Glands, (Adiyodiand Adiyodi, eds.). Vol. 3 of Reproductive Biology of Invertebrates, pp. 319471. Wiley, New York.

    Happ, G. M. (1992). Maturation of the male reproductive system and itsendocrine regulation. Annu. Rev. Entomol. 37, 303320.

    Wolfner, M. F. (2001). The gifts that keep on giving: Physiological functionsand evolutionary dynamics of male seminal proteins in Drosophila.Heredity 88, 8593.

    Acoustic Behaviorsee Hearing

    Aedes Mosquitosee Mosquitoes

    AestivationSinzo MasakiHirosaki University

    A estivation is a dormant state for insects to pass the summerin either quiescence or diapause. Aestivating, quiescentinsects may be in cryptobiosis and highly tolerant to heat anddrought. Diapause for aestivation, or summer diapause, serves

    not only to enable the insect to tolerate the rigors of summerbut also to ensure that the active phase of the life cycle occursduring the favorable time of the year.

    QUIESCENCE

    Quiescence for aestivation may be found in arid regions. Forexample, the larvae of the African chironomid midge,Polypedilum vanderplanki, inhabit temporary pools inhollows of rocks and become quiescent when the waterevaporates. Dry larvae of this midge can revive whenimmersed in water, even after years of quiescence. Thequiescent larva is in a state of cryptobiosis and tolerates thereduction of water content in its body to only 4%, survivingeven brief exposure to temperatures ranging from +102C to270C. Moreover, quiescent eggs of the brown locust,Locustana pardalina, survive in the dry soil of South Africafor several years until their water content decreases to 40%.When there is adequate rain, they absorb water, synchronouslyresume development, and hatch, resulting in an outburst ofhopper populations. The above-mentioned examples aredramatic, but available data are so scanty that it is difficult tosurmise how many species of insects can aestivate in a state ofquiescence in arid tropical regions.

    SUMMER DIAPAUSE

    Syndrome

    The external conditions that insects must tolerate differsharply in summer and winter. Aestivating and hibernatinginsects may show similar diapause syndromes: cessation ofgrowth and development, reduction of metabolic rate,accumulation of nutrients, and increased protection by bodycoverings (hard integument, waxy material, cocoons, etc.),which permit them to endure the long period of dormancythat probably is being mediated by the neuroendocrine system.

    Migration to aestivation sites is another component ofdiapause syndrome found in some species of moths, butter-flies, beetles, and hemipterans. In southeastern Australia, theadults of the Bogong moth, Agrotis infusa, emerge in late springto migrate from the plains to the mountains, where they aes-tivate, forming huge aggregations in rock crevices and caves(Fig. 1).

    Seasonal Cues

    Summer diapause may be induced obligatorily or facultativelyby such seasonal cues as daylength (nightlength) and tem-perature. When it occurs facultatively, the response to thecues is analogous to that for winter diapause; that is, the cuesare received during the sensitive stage, which precedes theresponsive (diapause) stage. The response pattern is, however,almost a mirror image of that for winter diapause (Fig. 2).Aestivating insects themselves also may be sensitive to the

    Aest ivat ion 3

  • seasonal cues; a high temperature and a long daylength (shortnightlength) decelerate, and a short daylength (long night-length) and a low temperature accelerate the termination ofdiapause.

    The optimal range of temperature for physiogenesis duringsummer diapause broadly overlaps with that for morphogenesis,or extends even to a higher range of temperature. Aestivatingeggs of the brown locust, L. pardalina, can terminate diapauseat 35C and those of the earth mite, Halotydeus destructor, dothis even at 70C. The different thermal requirements for

    physiogenesis clearly distinguish summer diapause from winterdiapause, suggesting that despite the superficial similarity intheir dormancy syndromes, the two types of diapause involvebasically different physiological processes.

    See Also the Following ArticlesCold/Heat Protection Diapause Dormancy Migration

    Further ReadingCommon, I. F. B. (1954). A study of the biology of the adult Bogong moth,

    Agrotis infusa (Boisd.) (Lepidopera: Noctuidae), with special reference to itsbehaviour during migration and aestivation. Austral. J. Zool. 2, 223263.

    Furunishi, S., Masaki, S., Hashimoto, Y., and Suzuki, M. (1982). Diapauseresponse to photoperiod and night interruption in Mamestra brassicae(Lepidoptera: Noctuidae). Appl. Entomol. Zool. 17, 398409.

    Hinton, H. E. (1960). Cryptobiosis in the larva of Polypedilum vanderplankiHint. (Chironomidae). J. Insect Physiol. 5, 286300.

    Masaki, S. (1980). Summer diapause. Annu. Rev. Entomol. 25, 125.Matthe, J. J. (1951). The structure and physiology of the egg of Locustana

    pardalina (Walk.). Union S. Afr. Dept. Agric. Sci. Bull. 316, 183.Tauber, M. J., Tauber, C. A., and Masaki, S. (1986). Seasonal Adaptations

    of Insects. Oxford University Press, New York.

    Africanized Beessee Neotropical African Bees

    Agricultural EntomologyMarcos KoganOregon State University

    Ronald ProkopyUniversity of Massachusetts

    T he study of all economically important insects is theobject of the subdiscipline economic entomology.Agricultural entomology, a branch of economic entomology,is dedicated to the study of insects of interest to agriculturebecause they help increase crop production (e.g., pollinators);help produce a commodity (e.g., honey, silk, lacquer); causeinjury leading to economic losses to plants grown for food,feed, fiber, or landscaping; cause injury to farm animals; orare natural enemies of agricultural pests and, therefore,considered to be beneficial. Study of all fundamental aspectsof the ecology, life history, and behavior of insects associatedwith agricultural crops and farm animals falls within therealm of agricultural entomology. These studies provide thefoundation for the design and implementation of integratedpest management (IPM) programs (Fig. 1).

    4 Agricul tural Entomolog y

    FIGURE 2 Photoperiodic response in the noctuid M. brassicae controllingthe pupal diapause at 20C. Note the different ranges of photoperiod for theinduction of summer diapause (dashed line) and winter diapause (solid line).[From Furunishi et al., 1982, reproduced with permission.]

    FIGURE 1 Bogong moths, Agrotis infusa, aestivating in aggregation on theroof of a cave at Mt Gingera, A. C. T., Australia. [Photograph from Common,I. (1954). Aust. J. Zool. 2, 223263, courtesy of CSIRO Publishing.]

  • ECONOMIC ENTOMOLOGY

    Insects are regarded by some as the main competitors ofhumans for dominance on the earth. Humans depend oninsects for pollination of many crops, for production of honeyand silk, for the decomposition of organic matter and therecycling of carbon, and for many other vital ecological roles.But it is the negative impact of insect pests that has been ofgreatest concern to humans. There are no reliable estimates ofaggregate losses caused by insects as vectors of pathogens andparasites of humans and domestic animals, as agents causingdirect damage to dwellings and other human-made structures,and as pests of crop plants and farm animals, but the amountsrun to probably hundreds of billions of dollars annually. Lossescaused by insects and vertebrate pests worldwide in the pro-duction of only eight principal food and cash crops (barley,coffee, cotton, maize, potato, rice, soybean, and wheat) between1988 and 1990 have been estimated at $90.5 billion.

    In the late 1800s and early 1900s, entomology becameestablished in many academic and research institutions as adiscipline equal in rank with botany and zoology. Thediversity of insects and their economic importance was thejustification for ranking the study of a class of animals(Insecta) as being equivalent to the study of two kingdoms oforganisms (plants and animals other than insects). Throughthe first half of the twentieth century, there was a schismbetween basic and applied (or economic) entomology. Sincethen, common use of the expression economic entomologyhas declined, being replaced by designations of its principalbranches, such as agricultural entomology, forest entomology,

    urban entomology, and medical and veterinary entomology. Adetailed historical account is beyond the scope of this article,but Table I provides a chronology of some landmarks in thedevelopment of agricultural entomology through the ages.

    The realm of agricultural entomology includes all basicstudies of beneficial and pest insects associated with agriculturalcrops and farm animals. This article deals mainly with crops,but the general principles and concepts are equally applicableto farm animals. The starting point of such studies is a correctidentification of the insect species, in accordance with thescience known as biosystematics.

    BIOSYSTEMATICS

    Scientific nomenclature is a powerful tool for obtaining infor-mation about the basic biology of closely related species withina genus. When systematic studies have been extended beyondthe naming of species (taxonomy) and contain detailedinformation on geographic distribution, host records, andbiology of one or more species in a genus, it is often possibleto extrapolate the information to other closely related speciesof that genus. Although details of the biology must be ascer-tained for each individual species, biosystematics offers a blue-print to follow when dealing with a new pest. For example,the genus Cerotoma (Coleoptera: Chrysomelidae) contains 10to 12 species distributed from southern Brazil to the north-eastern United States. All seem to be associated with herbaceousplants in the family Fabaceae (bean family). The biology of twoof the species, C. trifurcata in North America and C. arcuatain South America (Fig. 2), has been studied extensively. Based

    Agricul tural Entomolog y 5

    FIGURE 1 A bridge metaphor: agricultural entomology is conceived as one of the main pillars, together with plant pathology and weed science, of supportingthe integrated pest management bridge. The bridge connects two-way traffic between crop production and crop protection. The other pillar is providedby the social sciences of economics and sociology. The main tension cables, which are system integration and information systems, hold the vertical lines thattogether 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.

    Bio

    logi

    cal c

    ontr

    ol

    Cul

    tura

    l con

    trol

    Gen

    etic

    con

    trol

    Pla

    nt r

    esis

    tanc

    e

    Che

    mic

    al c

    ontr

    ol

    Beh

    avio

    ral c

    ontr

    ol

    Weedscience

    Plantpathology

    Agriculturalentomology

    Biologicalsciences

    Ruralsociology

    Agriculturaleconomics

    Socialsciences

    Integrated Pest Management Bridge

    System integration and information systems

    The flowing river of societal needs and demands

    Cropproduction

    Cropprotection

  • on information for these two species, it is possible to inferthat the other species in the genus share at least some of thefollowing features: eggs are laid in the soil adjacent to growingleguminous plants; larvae feed on nitrogen-fixing root nodulesand pupate in soil inside pupal cases; first-generation adultsemerge when seedlings emerge, and second-generation adultsemerge when plants are in full vegetative growth, feeding firston foliage and, later on, switching to feeding on developingpods. The biosystematic information on the genus allowsstudents of agricultural entomology in South, Central, or

    North America to understand, at least in general terms, therole of any other species of Cerotoma within their particularagroecosystem.

    The flip side of this notion is recognition that closelyrelated and morphologically nearly undistinguishable(sibling) species may have many important biologicaldifferences. Examples of the critical need for reliablebiosystematics studies are found in the biological controlliterature. The present account is based on studies conductedby Paul DeBach, one of the leading biological controlspecialists of the twentieth century. The California red scale,Aonidella aurantii, is a serious pest of citrus in California andother citrus-producing areas of the world. Biological controlof the red scale in California had a long history of confusionand missed opportunities because of misidentification of itsparasitoids. The red scale parasitoid Aphytis chysomphali hadbeen known to occur in California and was not considered tobe a very effective control agent. When entomologists dis-covered parasitized scales during foreign exploration, theparasitoids were misidentified as A. chrysomphali and there-fore were not imported into California. It was laterdiscovered that the parasitoids were in fact two differentspecies, Aphytis lingnanensis and A. melinus, both moreefficient natural enemies of the California red scale than A.chrysomphali. Now A. lignanensis and A. melinus are theprincipal red scale parasitoids in California. Furtherbiosystematics studies have shown that what was oncethought to be single species, A. chrysomphali, parasitic on theCalifornia red scale in the Orient and elsewhere, and acci-dentally established in California, is in fact a complexincluding at least seven species having different biologicaladaptations but nearly indistinguishable morphologically.

    6 Agricul tural Entomolog y

    FIGURE 2 Morphological diversity and biological similarities in the genusCerotoma: four of the dozen known species are illustrated by male and femalespecimens. The species are clearly distinguishable by morphologicalcharacters, but they have similar life histories and behaviors. (Fromunpublished drawings by J. Sherrod, Illinois Natural History Survey.)

    TABLE I Some Landmarks in the Historical Development of Agricultural Entomologya

    Significant events Years ago from 2000 Date

    Beginnings of agriculture 10,000 8000 B.C.E.First records of insecticide use 4,500 2500 B.C.E.First descriptions of insect pests 3,500 1500 B.C.E.Soaps used to control insects in China 900 1100Beginning of scientific nomenclature10th edition of Linnaeus, Systema Naturae 242 1758Burgeoning descriptions of insects 100200 18th and 19th centuriesFirst record of plant resistance to an insect 169 1831Charles Darwin and Alfred Wallace jointly present paper on the theory of evolution 142 1858First successful case of biological control: the cottony cushion scale, on citrus, in California,

    by the vedalia beetle 112 1888First record of widespread damage of cotton in Texas by the cotton boll weevil 106 1894First record of an insect resistant to an insecticide 86 1914First edition of C. L. Metcalf and W. P. Flints Destructive and Useful Insects 72 1928Discovery of DDT and beginning of the insecticide era 61 1939First report of insect resistance to DDT 54 1946Term pheromone coined by P. Karlson and P. Butenandt, who identified first such substance in

    the silkworm moth 45 1959First edition of Rachel Carlsons Silent Spring 48 1962Expression integrated pest management first appears in the press 32 1968Rapid development of molecular biology 20 1980sRelease of Bt transgenic varieties of cotton, corn, and potato 5 1990s

    a Based in part on Norris et al. (2003).

  • Knowledge of the name of a species, however, is not anindication of its true potential economic impact or pest status.A next important phase in agricultural entomology is, there-fore, the assessment of benefits or losses caused by that species.

    PEST IMPACT ASSESSMENT

    The mere occurrence of an insect species in association witha crop or a farm animal does not necessarily mean that thespecies is a pest of that crop or animal. To be a pest it mustcause economic losses. The assessment of economic lossesfrom pests is the subject of studies conducted underconditions that match as closely as possible the conditionsunder which the crop is grown commercially or the animalsare raised. Much of the methodology used in crop lossassessment has been established under the sponsorship of theFood and Agriculture Organization (FAO) of the UnitedNations as a means of prioritizing budget allocations andresearch efforts. Key data for these studies relate to thedetermination of the yield potential of a crop. The geneticmakeup of a crop variety determines its maximum yield inthe absence of adverse environmental factors. This is knownas the attainable yield. To determine the attainable yield, thecrop is grown under nearly ideal conditions; the actual yieldis what occurs when the crop is grown under normal farmingconditions. The difference between attainable and actualyields is a measure of crop loss (Fig. 3).

    To assess crop losses and attribute the losses to a specificcause (e.g., the attack of a pest) requires setting up experimentsto isolate the effect of the pest from all other constraints.Methodologies vary with pest categorywhether the pestsare insects, vertebrates, plant pathogens, or weeds, forexample. The quantitative relationship between crop lossesand pest population levels is the basis for computing theeconomic injury level for the pest. The economic injury levelis a fundamental concept in IPM.

    LIFE HISTORY AND HABITS

    Once the identity and pest status of a species have been wellestablished, it becomes essential to extend the informationalbase on the life history and habits of the species to theconditions under which the crop is grown. Economically im-portant life history traits include information on develop-mental threshold temperatures and temperature-dependentdevelopmental rates. These data are used in modeling thephenology of the pest. Other essential studies include theorientation, feeding, host selection, and sexual behavior ofthe species. Many of these studies provide the foundation forstrategic planning in IPM and for the development of target-specific control tactics. For example, the study of sexualbehavior involves the definition of the role of pheromones inmating and the identification of those pheromones. These, inturn, may be used for monitoring pest incidence and abun-dance or in mating disruption, both valuable components ofIPM systems for many crops. The study of host selectionbehavior often leads to the identification of kairomones,equally important in IPM development.

    PHENOLOGY

    The life cycle of different insect species varies greatly, althoughall insects undergo the basic stages of development from eggto reproductive adult (or imago). Depending on the length ofthe life cycle, there is considerable variation in the number ofgenerations per year, a phenomenon called voltinism. A uni-voltine species has one generation per year; a multivoltinespecies may have many generations per year. The range ofvariation in the Insecta is evident when one considers that the17-year periodical cicada has one generation every 17 years,whereas whiteflies or mosquitoes may complete a generationin about 21 days. Under temperate climate conditions,generations often are discrete, but under warmer subtropicalconditions they frequently overlap. The definition of temporalperiodicity in an organisms develo


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Arthropoda 7 Ticks Mites

Amber Arthropod Key - Freeambre.jaune.free.fr/AMBRE_CLEF_identification_des_Arthropodes.pdf · Opiliones (Daddy long-legs) Acari (Mites and ticks) Scorpiones (Scorpions) Pseudoscorpionida

Amber Arthropod Key - Freeambre.jaune.free.fr/AMBRE_CLEF_identification_des_Arthropodes.pdf · Opiliones (Daddy long-legs) Acari (Mites and ticks) Scorpiones (Scorpions) Pseudoscorpionida

Oribatid mites (Acari, Oribatida) as intermediate hosts of tapeworms ...

Oribatid mites (Acari, Oribatida) as intermediate hosts of tapeworms ...

1 Ticks, Mites and Associated Pathogens Overview: Characteristics of the subclass Acari (ticks and mites) Metastigmatid mites = ticks Ticks and tick-borne.

1 Ticks, Mites and Associated Pathogens Overview: Characteristics of the subclass Acari (ticks and mites) Metastigmatid mites = ticks Ticks and tick-borne.

Prevention and control of ticks and mites

Prevention and control of ticks and mites

Ticks & Mites

Ticks & Mites

Ticks, Mites, Fleas, Lice as a Vector

Ticks, Mites, Fleas, Lice as a Vector

ACARI MITES & TICKS. Mites and ticks, collectively known as the Acari, constitute the second most diverse group of animals on the planet today and are.

ACARI MITES & TICKS. Mites and ticks, collectively known as the Acari, constitute the second most diverse group of animals on the planet today and are.

CHAPTER 9-1 ARTHROPODS: MITES (ACARI)

CHAPTER 9-1 ARTHROPODS: MITES (ACARI)

FOCUSED MULTIFACETED GROWTH · Acaricides are used to destroy or prevent the spread of pests such as ticks and mites, which belong to the Acari group. Brands Mitex, Colonel-S and

FOCUSED MULTIFACETED GROWTH · Acaricides are used to destroy or prevent the spread of pests such as ticks and mites, which belong to the Acari group. Brands Mitex, Colonel-S and

TICKS OF THE GENUS AMBLYOMMA (ACARI: IXODIDA: …

TICKS OF THE GENUS AMBLYOMMA (ACARI: IXODIDA: …

New records of mites Acari and harvestmen Opiliones from ... et al 2017... · New records of mites Acari and harvestmen Opiliones from Malta with a preliminar checklist of Maltese

New records of mites Acari and harvestmen Opiliones from ... et al 2017... · New records of mites Acari and harvestmen Opiliones from Malta with a preliminar checklist of Maltese

Natural protection against e p l us ticks, fleas & mites. t u rn a P · 2019. 1. 7. · Natural protection against ticks, fleas & mites. Ticks - Fleas - MiTes bogadual® spot-on bogacare

Natural protection against e p l us ticks, fleas & mites. t u rn a P · 2019. 1. 7. · Natural protection against ticks, fleas & mites. Ticks - Fleas - MiTes bogadual® spot-on bogacare