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ESSENTIALENTOMOLOGY

ESSENTIAL ENTOMOLOGY

SECONDEDITION

HonoraryResearchAssociateoftheOxfordUniversity MuseumofNaturalHistoryandHonoraryPrincipalResearchFellow, ImperialCollege

Leonidas-RomanosDavranoglou

LeverhulmeTrustEarlyCareerFellow, OxfordUniversityMuseumofNaturalHistory

GreatClarendonStreet,Oxford,OX26DP, UnitedKingdom

OxfordUniversityPressisadepartmentoftheUniversityofOxford. ItfurtherstheUniversity’sobjectiveofexcellenceinresearch,scholarship, andeducationbypublishingworldwide.Oxfordisaregisteredtrademarkof OxfordUniversityPressintheUKandincertainothercountries

Text©GeorgeC.McGavinandLeonidas-RomanosDavranoglou2022

Illustrations©RichardLewington2001exceptthoselistedonpagexi

Themoralrightsoftheauthorshavebeenasserted

FirstEditionpublishedin2001

SecondEditionpublishedin2022

Impression:1

Reprinted2003

Allrightsreserved.Nopartofthispublicationmaybereproduced,storedin aretrievalsystem,ortransmitted,inanyformorbyanymeans,withoutthe priorpermissioninwritingofOxfordUniversityPress,orasexpresslypermitted bylaw,bylicenceorundertermsagreedwiththeappropriatereprographics rightsorganization.Enquiriesconcerningreproductionoutsidethescopeofthe aboveshouldbesenttotheRightsDepartment,OxfordUniversityPress,atthe addressabove

Youmustnotcirculatethisworkinanyotherform andyoumustimposethissameconditiononanyacquirer

PublishedintheUnitedStatesofAmericabyOxfordUniversityPress 198MadisonAvenue,NewYork,NY10016,UnitedStatesofAmerica BritishLibraryCataloguinginPublicationData Dataavailable

LibraryofCongressControlNumber:2022921538

ISBN978–0–19–284311–1(hbk.) ISBN978–0–19–284312–8(pbk.)

DOI:10.1093/oso/9780192843111.001.0001

Printedandboundby

CPIGroup(UK)Ltd,Croydon,CR04YY

Linkstothird-partywebsitesareprovidedbyOxfordingoodfaithand forinformationonly.Oxforddisclaimsanyresponsibilityforthematerials containedinanythird-partywebsitereferencedinthiswork.

phyla pluralofphylum,themajor groupingsoftheanimal kingdom,composedof superclassesandclasses,that is,Mollusca,Arthropoda, Chordata,etc.

Preface

Trophiclevel thelevelsoffoodorenergy transferinfoodchains/webs. Foodproducersorautotrophs formthefirstlevel,herbivores formthesecondlevel,primary carnivoresthethirdlevel. Othercarnivoresformthe fourthandfifthlevels.

Lifebeganintheseas,andsaltwaterisstillhometomostoftheearth’s phyla. Onland,however,twogroupsdominate:theplantsandtheinsects.Theword insectcomesfromtheLatin, insectus,thepastparticipleof insecare,tocutor incise,andreferstothemajordivisionsofaninsect’sbody.

Insectshavebeenaroundforwellover400millionyears.Theywerethe firstanimalsonlandandthefirsttogetintotheair.TheyarethemostspeciesrichandabundantgroupofmulticellularanimalstheEarthhaseverseen. Morethanhalfofalldescribedspeciesandabout75%ofallanimalsareinsects.ButexactlyhowmanyspeciessharetheplanetEarthwithus?Thetruth is:wedon’thavethefoggiestidea.Someearlyguessesof30millionoreven 100millionhavebeenreplacedinthelastdecadewithmorerealisticestimatesofsomewherebetween5and10millionspecies.Despitethismassive uncertainty,onethingisindisputable:themajorityofEarth’sinhabitantsare insects.

Interrestrialecosystems,greenplantsaretheprimaryproducers,trappingthesun’senergyandusingittoconvertcarbondioxidetoorganic matter.Withoutphotosynthesistheplanetwouldbeaverydifferentplace. Autotrophslikeplantsformthebaseofterrestrialandfreshwaterfoodchains, butevery trophiclevel aboveisdominatedbyinsects.

Forexample,itmaycomeasasurprisetomanythatalltheheaving, snortingherdsofgrazingungulatesontheAfricansavannahsareentirely ‘out-munched’(perhapsbyafactoroftentoone)bymyriadsoftiny mandibles.Whataboutthemeateaters?Again,insectsconsumemanytimes moreanimalfleshthanallthevertebratecarnivoresputtogetherandants alonearethemajorcarnivorousspeciesinanyhabitatyoucouldmention. Ifthissoundsimplausible,considerthat,althoughinsectsareindividually small,thereareanawfullotofthem.

Insectspollinatethevastmajorityoftheworld’squarterofamillionorso speciesoffloweringplants.Pollinationisoneofthemostessentialsymbioses evertohaveevolved.Thisplant–insectversionof‘I’llscratchyourbackif youscratchmine’haslastedfor100millionyearsandithasgeneratedarich diversityofspecies.Twentythousandspeciesofbeeare,toalargeextent,responsibleforthecontinuedsurvivaloftheangiosperms,whichincludesa verylonglistofthethingsweeat:fruitandvegetablesfrompumpkins,plums, andpeastocherries,cucumbers,andcocoa.

Withouttheproteinprovidedbyinsectsmanygroupsofhigheranimals wouldsimplynotexist.Insectsfeatureexclusivelyorverylargelyinthediets ofawholebeastlybestiaryfromtheaardvarktothezorilla(anAfricanstriped polecat).Birds,thedescendantsofthedinosaurs,aremostlyinsectivorous. Abroodofninegreattitchickswillconsumearound120,000caterpillars whiletheyareinthenest.Asingleswallowchickmayconsumeupwards of20,000bugs,flies,andbeetlesbeforeitfledges,andevenspeciessuchas

Verticalbarchartshowing theapproximateproportions ofallspeciesonEarth.

Insects

Arachnids

Crustaceans

Other Arthropods 1.2% Molluscs

Nematodes 0.9%

Other Invertebrates

Vertebrates

Theimageshownhereisa compositeofphotographsof individualinsectsattracted toaseahollyfloweroverthe courseof15minutes.Itis obviousthatwithoutinsects, pollinationwouldbe impossibleformostplants, andtheworld’sterrestrial ecosystemswouldcollapse.

buntings,whichareseedfeedersasadults,reartheiryoungonanutritious dietofinsects.Theearliestprimateswereinsectivores,andmanyprimatesstill aretoday.Incountrieswhereinsectsarelargeorveryabundant,theyform animportantsourceofprotein,vitamins,andmineralsforhumans.Locusts andgrasshoppers,beetlegrubs,caterpillars,andtermitesfeatureregularlyin tropicaldietsandtheirnutritionalvalueexceedsthatofwestern-stylefast foods.

Atleastaquarterofallinsectspeciesareparasitesorpredatorsofother speciesofinsect.Insectsrecyclenutrients,enrichsoils,anddisposeofcarcassesanddung.Apartfromtheseessentialecosystemservices,insects providehumanswithusefulproductssuchassilk,honey,waxes,medicines, anddyes.Weusethemtocontrolpestorganismsandasmodelsystemstohelp usstudymanyaspectsofbiologyfromgeneticsandbehaviourtophysiology andecology.

Ofcourse,insectshaveadarkside.Herbivorousinsectsmayeatupto15–20%ofallcropsgrownforhumanconsumption,andlocally,thepercentage maybemuchhigher.Although,contrastthiswiththe30%ofallfoodgrown thatiswastedbyhumans.Inaddition,insectscarrytheviruses,bacteria, spirochaetes,rickettsiae,protozoans,roundworms,andfungithatareresponsibleforinnumerableplant,animal,andhumandiseases.Aboutoneinsix humanbeingsalivetodayisaffectedbyaninsect-borneillnessofsomekind. Besidesstingsandallergies,whichcanbefatalforsome,thelistofdiseases carriedbyinsectsincludesplague,typhus,sleepingsickness,riverblindness, Chagas’disease,yellowfever,Zikavirus,epidemictyphus,trenchfever,loiasis,filariasis,andleishmaniasis.Intheearlytwentiethcentury,SirRonald Rossshowedconclusivelythatmosquitoeswereresponsiblefortransmitting malaria.Althoughmuchmoreisknownaboutmalariatoday,itremainsa deadlydiseaseresponsibleforhundredsofthousandsofdeathseveryyear. Tocompletethedestructivesideoftheiractivities,insectscausegreatdamagetowoodenstructuresandawiderangeofnaturalmaterialsandfabrics. Thereareeven,muchtotheaggravationofmuseumcurators,beetleswhose larvaehaveavoraciousappetiteforcollectionsofdriedinsects.

Beingmuchsmaller,insectsmaynothavetheimmediateappealofthe ‘furriesorfeatheries’,butifyoulookcloser,youwillfindthattheyarealot morevariedandinteresting.Whatanimalsotherthanhumansbuildairconditionedcondominiums,useanunderwaterbreathingapparatus,constructacousticequipment,causeexplosions,makepaper,cultivategardens, andfarmotheranimals?Theworldofinsectsisendlesslyfascinating,and theyhaveextraordinaryrelationshipswithotherorganisms.Someoftheir lifecyclesbeggarbelief.Therearemothsthatsuckmammalianblood,flies thatbreedinpoolsofcrudepetroleum,flieswhoselarvaedeveloponlyin thetrachealpassagesofredkangaroos,andlicethatsolelyinhabitthethroat pouchesofcertaincormorantsandpelicans.

Despiteallthetalkoftheneedtoquantifybiodiversityinrecentyears, taxonomyandsystematicsarestilllessfashionabletodaythantheyoncewere andthereisalackoffundingforbasictaxonomicresearch.Howarewegoing tosavebiodiversity,ifwedonotknowwhatwearetryingtosaveinthefirst

place?Perhapswehavelostsightofthefactthat,withoutasystemofclassification,ourinvestigationscanonlybeaconfusionofunconnectedfacts.We surviveinacomplexworldbydefiningwhatthingsare,namingthem,and thusidentifyingthem.Imaginethecolossaltaskofhavingtoremembereach andeverynovelobjectindividuallybyitsuniquecharacteristics.Classificationisacentralprocessofthehumanbrain,andit,notmathematics,ashas sometimesbeensuggested,isthefoundationandcornerstoneofallscience. AsBritishbiologistandbeetletaxonomistRoyCrowsonpointedout,you needtoknowwhatyouaregoingtocountormeasurebeforeyournumbers aregoingtomakeanysense.

Sex,violence,andacastoftrillions—thestudyofinsectsisexcitingand intellectuallysatisfying,butwheredoyoustart?Withclassificationofcourse. Inanystudy,whetherecological,physiological,orbehavioural,thefirstthing youneedtoknowiswhatsortofinsectyou’redealingwith.Textbookssometimesemphasizesystemsbiologyattheexpenseofsystematicswiththeresult thatstudentsarefamiliarwiththebroaderpictureofinsectphysiologyor behaviourbutareconfusedwhenitcomestorecognizingaspecificinsect. Whetherthereare2or10millioninsectspecies,youdonotneedtoknow themallindividually.Atfirstglance,abeetlemayresembleanant,andsome fliesareexcellentmimicsofbees,butantsdonothavewingcasesandbees havetwopairsofwings.Withineachorderthereareconspicuousfeatures thatweusetoaidcorrectidentification.

Theamountofbiologicalinformationavailable,especiallyconcerninginsects,isincreasingrapidlyandnewcomerstothestudyofinsectscanbe putoffsimplybytheenormityofthesubject.Somespecieshavereceived ahugeamountofattentionforvariousreasons.TheAmericanCockroach, Periplanetaamericana,andlocustspecies,suchas Schistocercagregaria,are enduringlypopularasmodelsforresearch,andfruitflies(Drosophila spp.) areextraordinarilyusefulingenetics.Particularresearchershavehadtheir favourites.Wigglesworth’spioneeringworkoninsectcuticleusedaSouth American,cone-nosed,reduviidbug,Rhodniusprolixus,andDethier’schoice ofaparticularblowfly, Phormiaregina,forhisworkoninsectfeedingand sensorybiologyresultedfromalunchtimechanceobservationofafemaleof thisspecieslayinghereggsonaliverwurstsandwich.These,andahandfulof otherspecies,arerareexceptions.Theresthaveyettoenjoytheir15minutes offame.Themajorityoftheworld’slivinginsectsarecompletelyunknown andthebiologyofmanyofthoseweknowaboutareobscure.Thereisstill plentyofworktobedone.

Butthereisaproblem.Humanactivitieshavebroughtaboutabiodiversitycrisis.Theconversionofmoreandmoreoftheplanet’snaturalcapital intoproducinghumanbeingsandservingtheirneedsanddesireshascome atanenormouscost.Weseemtohavedevelopedaseriousdisconnectwith thenaturalworldandthisdisconnectmaybeourdownfall.Populationsof majorgroupsoforganisms,forexample,birds,reptiles,andmammals,have declinedbyanaverageof68%inthelast50years.Almosteveryrecent studyundertakenpointstoamarkedslumpininsectspecies’richnessand abundance.Thedecreasesseeninwell-studiedinsectgroupslikebeesand

butterfliesareoccurringinmanyothergroups.Thecausesofthesedeclines areveryclear:theacceleratinglossofnaturalhabitat,mostlyduetoagriculturalintensification,coupledwiththeprodigiousamountsofpesticidesused, aretakingaheavytoll.Ontopofthiswearefacingglobalclimatechange causedbyouruseoffossilfuels,whichaffectseveryspeciesandeveryhabitat onEarth.

Ithaslongbeenthoughtthattheworld’stropicalforestsarehometomore thanhalfofallextantspecies.Ifthesecomplexhabitatscontinuetobefelled fortimber,cashcrops,andranchingateventheslowestratesuggested,itwill stillonlybeamatterofafewhundredyearsbeforetheyarecompletelylost. Itisthereforeaninescapableconclusionthatourplanetcouldlosemorethan halfofallitslivingspeciesinthetimeittakesforanacorntobecomeaveteran oaktree.Weknowthatnospecieslastsforever.Ofallspeciesthathaveever lived,99%arenowextinctanditwasthenumerousmassextinctionevents thatpavedthewayfortheappearanceofthosecreaturesthatledtohumanbeings.Thedifferenceisthatnowweknowenoughtodosomethingthatmight prolongourownsurvival.

Therecanbelittledoubtthathumansarethemostintelligentandcapable speciesyettoevolveonEarth.Intheveryshorttimesinceourappearance, wecoveredtheentireglobeandestablishedcolonieswhereveritwaspossible tosurvive.Afewofushavewalkedonthesurfaceofthemoonandvisited thedeepestabyssesoftheoceans.Wespendvastsumsofmoneytoprobethe verymake-upofmatterandexaminetheuniverse.Wewanttounderstand thescienceofeverythingfromtheinfinitesimallysmalltotheastronomically largebutseemtobepayinglittleheedtothehealthoftheveryecosystems thatsupportus.

Theaimofthisbookistoprovideareadableintroductiontothemost abundantmulticellularlifeformsonEarth.Itisnotafieldguidebut,should youneedtoknowtheessentialfactsaboutaparticularorderofinsect(e.g. wheretheyoccur,howmanyspeciesareknown,whatmakesthemdifferent andinterestingfromallotherorders),thisbookwillprovidethemanddirect youtospecialisttextsandsourcesofinformationtoguideyoufurtheralong anyparticularpath.Ifyoureallywanttounderstandhowthenaturalworld works,youneedtoknowaboutinsects.

Thebookislaidoutinanuncomplicatedfashion.Thefirstsectiongives abriefintroductiontotheinsectsandcoverstopicssuchastheevolution ofthegroupandthefactorsthatmadethemsuchsuccessfulorganisms.It discussestheimportanceoftheirroleinterrestrialecosystemsandoutlines thefeaturesofstructure,function,andphysiologythattheyshare.Thesecondsectionprovidesasemi-pictorialkeytotheinsectorders.Unambiguous text,coupledwithcleardrawingsdesignedtohighlightkeyfeatures,allowthe readertoassignmostadultinsectstothecorrectorder.Thebulkofthebook isdevotedtotheessentialsofthe28livinginsectorders.Thefinalsection dealswithcollectingtechniques.

Acknowledgements

Wethankthefollowingpeopleforprovidingthephotographsthatdecorate theneweditionofourbook:NickyBay,PaulBrock,JohnGausas,Martin Gore,MattDoogue,PiotrNaskrecki,EdPhillips,MikePicker,HansPohl, AnneRiley,GillesSanMartin,JohnSmit,ZestinSoh,RupertSoskin,and ManfredUlitzka.

Figure Acknowledgments

Allillustrationsnotlistedbeloware©RichardLewington,2001and2023 PrefacepageviVerticalbarchart©OxfordUniversityPress2001.

Pages4-7Diagramsofthearthropod-likegroups©OxfordUniversityPress 2001.

Page13DiagramfromWilmer,P.,Stone,G.andJohnston,I.(2000) EnvironmentalPhysiologyofAnimals.BlackwellScience,Oxford.

Page15DiagramfromGullan,P.J.andCranston,P.S.(2000) Theinsects:an outlineofentomology,2ndedition,BlackwellScience,Oxford.

Page17DiagramsfromChapman,R.F.(1998) Theinsects:structureandfunction, 4thedition.CambridgeUniversityPress.©AfterHepburn,H.R.(1985) Structureoftheintegument,in Comprehensiveinsectphysiology,biochemistry andpharmacology, Vol.3(edsG.A.KerkutandL.I.Gilbert),PergamonPress, Oxford.

Page19Diagram©OxfordUniversityPress2001. Page20DiagramredrawnfromKukulova-Peck,J.(1991) TheinsectsofAustralia,2ndedition.MelbourneUniversityPress,Australia.©CSIROAustralia 1991.

Page21DiagramafterChapman,R.F.(1998) Theinsects:structureand function, 4thedition.CambridgeUniversityPress.

Page22Diagram©OxfordUniversityPress2001.

Page36Diagramofthephylogenyofhexapods.Adaptedfrom©HansPohl (2014).

Page286Diagram©GeorgeC.McGavin2001.

Biographies

GeorgeMcGavin studiedZoologyatEdinburghUniversity,followedbya PhDinentomologyatImperialCollegeandtheNaturalHistoryMuseum inLondon.After30yearsasanacademic,mostlyatOxfordUniversity,he becameanaward-winningtelevisionpresenter.GeorgeisanHonoraryResearchAssociateoftheOxfordUniversityMuseumofNaturalHistoryand anHonoraryPrincipalResearchFellowatImperialCollege.HeisaFellowof theLinneanSocietyanHonoraryFellowoftheRoyalSocietyofBiology,and anHonoraryLifeFellowoftheRoyalEntomologicalSociety.In2019George becamethePresidentoftheDorsetWildlifeTrust.

Leonidas-RomanosDavranoglou studiedZoologyatImperialCollege London.HethenundertookhisDPhilthesisinentomologyattheDepartmentofZoology,UniversityofOxford,fundedbytheprestigiousOxford NaturalMotionandtheOnassisFoundationscholarships.In2021,Leonidas wasawardedwithbothaLeverhulmeTrustEarlyCareerFellowshipandthe JohnFellOUPFundattheOxfordUniversityMuseumofNaturalHistoryto studytheevolutionofinsectcommunication.Leonidaswillberesearching

howinsectscommunicateusingsoundsbyexaminingthemorphologyand biomechanicsoflivingspecies,aswellastheirextinctrelativespreservedin 100-million-year-oldCretaceousamber.

Glossary

Aedeagus Theorganofcopulationinmaleinsects.

Ametabolous Developmentwithoutmetamorphosis,suchasinbristletails andsilverfish.

Apodous Withoutlegs.

Apodemes Internaltendon-likeextensionsofthecuticle,whichserveas muscleattachmentsites.

Apolysis Theseparationoftheoldandnewcuticles.

Aposematic Havingbrightcolourationtoserveasawarningtopredators. Usuallyred,yellow,andblack.

Apterous Lackingwings.

Apterygota (adj.apterygote)Primitivelywinglessinsects.Thesmaller subclassoftheInsecta.

Arrhenotoky Parthenogenesiswherehaploidmalesareproducedfrom unfertilizedeggs.

Asexual Ofreproductionwithoutseparatesexes.Reproductionoccurring byparthenogenesis.

Autotomy Thecastingorsheddingoffofapartofthebody,forexample, whenunderthreat.

Batesianmimicry Whereanedibleorpalatablespeciesmimicsthecolours andpatternsofaninediblespecies.Themimicsarerarerelativetothe models.

Biologicalcontrol Useofnaturalpredators,parasites,ordiseaseproducing organismstoeffectareductioninthepopulationsofpestspecies.

Biramous Havingtwobranches.

Bisexual Havingtwoseparatesexes.

Caste Anygroupofindividualsinacolonyofsocialinsectsthatis structurallyorbehaviourallydifferentfromindividualsinothergroups, thatis,soldiers,workers,andreproductivesasseenintermites,bees,and ants.

Cerci (sing.cercus)Apairofterminalabdominalappendages,which typicallyhaveasensoryfunction.

Chitin Ahighmolecularweight,unbranched,amino-sugarpolysaccharide madeupof β(1–4)linkedunitsofN-acetyl-D-glucosamine.Chitinisthe mainstructuralelementinarthropodexoskeletonsandisalsofoundin fungalcellwalls.

Chrysalis Thealternativenameforalepidopteranpupa.

Class Amajortaxonomicsubgroup,aboveOrderandbelowPhylum.

Coelom Themainbodycavitysurroundingthegutinanimalssuchas annelidworms,starfish,andvertebrates.Inarthropodsandmolluscs,the coelomismuchreduced,andthemainbodycavityisahaemocoel.

Colony Anaggregationofsocialinsectssharinganest.

Corbiculum (pl.corbicula)Thepollenbasketofhoneybees,whichisa concave,shinyareaonthehindtibiae,fringedwithstiffhairs.

Cosmopolitan Occurringthroughoutmostoftheworld.

Coxa (pl.coxae)Thefirstsegmentofaninsect’sleg,joiningtherestofthe legtothethorax.

Crop Anareaoftheforegutininsectswherefoodisstored.

Cuckoo Atermusedforaninsectthatusesthefoodstoredbyanotherto reartheirownyoung.

DDT Dichlorodiphenyltrichloroethane.

Dimorphic Occurringintwodistinctforms.Thesexesofsomeinsectsare differentlycolouredorshaped.

Ecdysis Thelaststageofmoulting,whichisthecastingorsloughingofthe oldcuticle.

Elytra (sing.elytron)Therigidfrontwingsofbeetles,modifiedascoversfor thehindwingsandnotusedinflight.

Endite Alobeofalegsegmentthatisdirectedinwardstowardsthemidline ofthebody.

Endopterygota (adj.endopterygote)Insectswherethewingsdevelop internally.Metamorphosisiscompleteandthereisapupalstage (holometabolous).

Entognathous Havingeversiblemouthpartscontainedinsidethehead withinasmallpocket.

Epidermis Ininsects,thesinglelayerofcellsthatsecretesthecuticle.

Exite Alobeofalegsegmentthatisdirectedoutwardsawayfromthe midlineofthebody.

Exopterygota (adj.exopterygote)Insectswherethewingsdevelopoutside thebody.Metamorphosisisincompleteandthereisnopupalstage (hemimetabolous).

Family AtaxonomicgroupbelowthatofOrderandcomprisedof subfamilies,genera,andspecies.

Fatbody Anaggregationofmetabolicandstoragecellsinthehaemocoel.

Femur (pl.femora)Theportionofaninsect’slegthatcorrespondstothe mammalianthigh.Theorderofsegmentsis coax–trochanter–femur–tibia–tarsus.

Fungivorous Eatingfungi.

GABA Gamma-aminobutyricacid.Typicallyaninhibitorytransmitterof neuromuscularjunctions.Gammaisoftenshownsymbolically: γ-aminobutyricacid.

Gall Anabnormalplantgrowthcausedbyavirus,bacterium,fungus,mite, orinsect.

Genitalia Thehardpartsofinsectreproductivesystemsthatengage betweenmalesandfemalesduringmating.

Genus (pl.genera)Alowtaxonomicgroupcomposedofspecies.

Haemocoel Thehaemolymph-filledinternalbodycavityofarthropods, suchasinsects,whichisessentiallyanenlargedpartofthecirculatory system.

Haemolymph Afluidininvertebrateanimalsthatistheequivalentof blood.

Haltere(s) ThegreatlymodifiedhindwingsofDiptera,whichserveas balancingorgans.

Hemimetabolous Developingbyincompleteorgradualmetamorphosis, forexample,intheOrthopteraandHemiptera.Immaturestagesare callednymphs(seeexopterygota).

Hemimetabola Thoseorderswithincompletemetamorphosis.

Holometabolous Developingbycompletemetamorphosis,forexample,in theDipteraandLepidoptera.Immaturestagesarecalledlarvae(see endopterygota).

Holometabola Thoseorderswithcompletemetamorphosis.

Homeoticgenes Genesinvolvedintheregulationofbodyplan.

Homology Characterstatesacquiredbydirectinheritancefroman ancestralform.

Homoplasy Characterstatesacquiredbyconvergentevolution.

Honeydew Thecarbohydrate-richexcrementofplantsapfeederssuchas aphids.

Hyperparasitoid Aparasitoidofanotherparasitoid.

Idiobiont Aparasitoidthatkillsorparalysesitshostwhenlayinganegg, andthuspreventingthehostfromdevelopinganyfurther.

Inquiline Aspeciesthatlivesinthenest,gall,orhomeofanother.

Insect ThewordinsectcomesfromtheLatin, insectus,pastparticipleof insecare,tocutinto,andreferstothemajordivisionsofaninsect’sbody.

Integument Thecuticularandcellularcoveringofinsectsandother arthropods(i.e.,theepidermisandcuticlecombined).

Koinobiont Aparasitoidthatdoesnotkillthehostwhenegglaying,thus allowingthehosttodevelopfurther.

Lobopods Protoarthropodswithantennae,annularbodysegmentation (notajointedexoskeleton),andsoft,fleshyappendagesbearingclaws.

Meconium Theaccumulatedsolidwastesofthelarvalstagesofsome insects,whichareexpelledwhentheadultstageisreached.

Metamorphosis Thetransformationbetweentheimmatureformandthe adultform,withtheformsbeingdifferent.

Monophagous Restrictedtoeatingasinglespeciesplantortypeof foodstuff.

Monophyletic Oforganismsthathavedescendedfromasinglecommon evolutionaryancestororancestralgroup.

Mullerianmimicry Whereseveralinedibleorunpalatablespeciesconverge incolourpattern.Asallthespeciesarenon-palatable,mimicryrings,as theyareknown,areenhancedbythefrequencyallringmembersand evendistantlyrelatedspeciescanbeinvolved.

Mycetocytes Specialcellsthatpermanentlyhouseintracellularbacterial symbionts.

Myrmecophilous Greekforant-loving.Anyspeciesthatliveswithand dependsonantsforfood,protection,andcare.

Neotenic Retainingfeaturesoftheimmaturestagesintheadult.

Ocelli Simplelight-receptiveorgansontheheadofmanyinsects.

Oligophagous Restrictedtoeatingasmallrangeofrelatedplantsor foodstuffs(seepolyphagous,monophagous).

Ommatidia Individuallightreceptiveunitsinthecompoundeyesof insects.Eachommatidiumcomprisesaclearlenssystemand light-sensitiveretinulacells.

Omnivorous Havingvariableeatingpreferences.

Ootheca Protectiveeggcase.Cancontainasingleegginsomeorders,and uptoseveralhundredeggsinotherorders.

Order Ataxonomicrankbelowaclassandaboveafamily.

Oviparous Producingyoungbymeansofeggsexpelledfromthebody beforetheyarehatched.

Ovoviviparous Theyoungareproducedbymeansofhatchingeggsinside thebody.

Palaeozoic(Paleozoic) ThefirstofthethreeerasofthePhanerozoic (havingvisiblelife).ThePalaeozoicincludestheCambrian,Ordovician, Silurian,Devonian,Carboniferous,andPermianperiods.ThePalaeozoic isfollowedbytheMesozoicandCenozoiceras.

Parasite (adj.parasitic)Aspecieslivingoffthebodyortissuesofanother, butnotcausingthedeathofthehost.

Paraphyletic Ofagroupofspeciesthatincludessome,butnotall,ofthe speciesthatshareasinglecommonancestor.

Parasitoid Aspeciesthatdevelopsbyconsumingthebodytissuesofa singlehost,eventuallycausingitsdeath(seeidiobiontandkoinobiont).

Parthenogenesis (adj.parthenogenetic)Reproductionwithouttheneedfor fertilization.

Perineurium Alayerofglialcellsinsidetheneurallamella,whichactsas theblood–brainbarrier.

Pheromone Avolatilesubstanceproducedbyinsectstocommunicatewith othersforthepurposesofreproduction,aggregation,defence,etc.

Phloem Themaintransportvesselsofplantstakingnutrientsfromthe leavestootherparts.

Phyla (sing.phylum)Themajorgroupingsoftheanimalkingdom, comprisedofsuperclassesandclasses,forexample,Mollusca, Arthropoda,Chordata,etc.

Physogastric Whentheabdomen(usuallyinfemales)becomesgreatly distendedandmembranous,typicallyafterameal,whenpregnant,or whenstoringfatorfood.

Polyembryony Asingleegggivingrisetomanyembryosbysubdivision.

Polymorphic Havingmorethantwoforms.

Polyphagous Eatingawiderangeofplantsorfoodstuffs.

Polyphyletic Ofagroupofspeciesthathavedifferentancestors.Suchtaxa are‘unnatural’orincorrect.

Predacious (n.predator)Eatingotheranimals.

Proleg Afleshyabdominallimbofinsectlarvae.

Pronotum Thedorsalcoverofthefirstsegmentofthethorax.

Pruinescent (colouration)Ofapowder,orpowdery.

Pterygota (adj.pterygote)Wingedinsects.Thelargersubclassofthe Insecta.

Rostrum Theprolongedpartoftheheadofweevilsandscorpionflies.Also thetubular,slender,suckingmouthpartsofinsectssuchastheHemiptera (alsoknownasthelabiuminthelatter).

Saprophagous Eatingdecayedorganicmaterial.

Sclerite Ahardenedchitinousbodyplatesurroundedbyflexible membranesorsutures(Gk. skleros—hard).

Sistergroups Groupsofthesametaxonomicrankresultingfromthe splittingofanancestrallineage.

Solitary Occurringsinglyorinpairs,notinsocialgroups.

Somite Asinglebodydivisionofasegmentedanimal.

Species Agroupoflivingorganismsconsistingofrelatedsimilar individualscapableofexchanginggenesorinterbreeding.Ataxonomic rankbelowagenus.

Spiracle(s) Thebreathingholesofinsectsleadingtothetracheaeoccurring atthesidesoftheabdomenandthorax.

Stridulation Theactofproducingsound,usuallybyrubbingtwopartsof thebodytogether.

Subfamily AtaxonomicsubdivisionofFamily.

Subimago Pre-adultwingedstageofEphemeroptera.

Suborder AtaxonomicsubdivisionofOrder.

Suture Agrooveontheoutsideofaninsectbodysegmentthatindicatesthe fusionoftwosclerites.

Symbiosis Differentspecieslivinginanassociationthatbringsmutual benefit.

Synanthropic Associatedwithhumansandtheirhabitations.

Synonymy Whereasinglespecieshasbeendescribedandnamedmore thanonce

Tagmata (sing.tagma)Agroupofbodysegmentsthattogetherforma majorfunctionalunitsuchasthehead,thorax,orabdomen.

Tarsus (pl.tarsi)Thefootofaninsect.Itisattachedtotheendofthetibia andmadeupofasmallnumberofsubsegmentsortarsomeres.

Thelytoky Parthenogenesiswhereonlyfemaleoffspringareproduced.

Tibia (pl.tibiae)Thelowerlegsegmentofinsects,correspondingtothe mammalianshin.

Tracheae (sing.trachea)Theinternalairwaysofinsects.

Trochanter Thesmallsegmentoftheinsectlegbetweenthecoxaandthe femur.

Trophallaxis Theexchangeoffoodbetweencolonymembersinsocial insects.

Trophiclevel Thelevelsoffoodorenergytransferinfoodchain.Food producersorautotrophsformthefirstlevel,herbivoresformthesecond level,andprimarycarnivoresthethirdlevel.Othercarnivoresformthe fourthandfifthlevels.

Vector Anintermediatehostthatcarriesadiseaseorganismincontactwith itstargetorganism.

SECTION1 Introductiontoinsect evolutionandbiology

radiodonts

Earlyarthropods.Thefront sectionoftheelongatebody carriedapairofappendages andaventralmouthwith distinctiveradiatingteeth.The trunkofthesegmentedbody hadlobe-orflap-like appendages.Somewere predatorswhileotherswere filterfeeders

lobopods earlyarthropodswith antennae,annularbody segmentation(notajointed exoskeleton)andsoft,fleshy appendagesbearingclaws. Earlierstudiessuggestedthat velvetworms(Onycophora) andwater-bears(Tardigrada) aredirectdescendantsof lobopods,althoughrecent workssuggestthattheir similaritiesarelikelyonly superficial

homeoticgenes genesinvolvedinthe regulationofbodyplan

Theevolutionofthearthropods

TheEarthformedfromgasanddebrisaround4,600millionyearsago(mya). Thefirsttracesoflifeappearasfossilsofblue-greenalgaearound3,600–3,400mya,andyetformorethan2billionyears,nothingmorecomplicated thanbacteria,algae,andminute,unicellularplanktonicorganismsexisted. Thenevolutionwentintooverdrive.Knownpopularlyasbiology’s‘bigbang’, thetimearoundthebeginningoftheCambrianperiod(543mya)sawthe ‘sudden’appearanceinthefossilrecordofanincredibleassemblageofmulticellularanimals,someofwhichweretheancestorsofallspeciesonEarth today.Chiefamongtheseanimalswereearlyarthropods,suchas radiodonts and lobopods.

AlthoughtheCambrianexplosionischaracterizedbysomeexceptionally well-preservedmaterial,suchasthosefoundintheBurgessShaledepositsof BritishColumbia,itisoftendifficulttosayexactlywhichmighthavebeen thedirectancestorsofmodernforms.Theprocessoffossilizationisunpredictable,andfinedetailisoftenrecordedonlyunderspecialconditions. Evolutionaryintermediatesmayneverbefound.Thesegmented,worm-like forebearsofthesecreatureshadcertainlyarisenbythelatePrecambrian(the Vendian),fortheearlyarthropodshadalreadyevolvedintoarangeofclearly recognizablegroupswithdistinctbodyplansbytheearlytomid-Cambrian.

Arthropodsdifferfromeachotherchieflyinthenumberanddesignof theirappendages.Thediscoveryof homeoticgenes andthefunctionofsome ofthesegeneshasprovidedsomeinsightintotheearlydiversificationofthe group.

Wenowknowhowcertainhomeoticgenesmayhavedeterminedtheevolutionofsegmentdiversity,aswellashowtheyledtotheevolutionofthe dazzlingrangeofbodyplansexhibitedbyarthropodsasawhole.

Justastheinventionofsimpleandthencompoundmicroscopesintheseventeenthcenturyadvancedthebiologicaltechniquesoftheday,andtherefore madeitinevitablethatoldclassificationshadtobeabandonedandnewones adopted,moleculartechniquesareproducingdatathatsometimescontradict viewsbasedontraditionalmorphology.Whilethephylogeny(evolutionary history)ofanygroupmaybeinferredfrommorphologicalormolecularstudies,bothapproachescansufferfrombeingunabletodistinguishwhether ornotparticularcharacterstatessharedbytwoorganismsareduetocommonevolutionarydescent(acquiredbydirectinheritancefromanancestral form—homology,orbyconvergentevolution—homoplasy).Verydifferent animalsmightevolvefeaturesthatmakethemlooksuperficiallysimilar(e.g. molesandmolecrickets;seefigureonnextpage),anditisnotunusualfor naturalselectionactingoveralongtimetomakecloselyrelatedspecieslook verydifferentfromeachother(e.g.elephantsandhyraxes). monophyletic (of)agroupofspeciesthat shareacommonancestor polyphyletic (of)agroupofspecieshaving differentancestors.Suchtaxa are‘unnatural’

Arthropodsarea monophyletic group,thatis,theyhaveasinglecommon ancestor.Thestaggeringmorphologicaldiversityfoundintoday’sarthropodsledsomeauthorsinthepasttosuggesta polyphyletic origin,where arthropodsevolvedfromfourdifferenttypesofwormsthatindependently gainedanexoskeleton.Thisviewisnowobsolete.Weknowthatarthropods

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