TrimSize:189mmx246mm Quicke ffirs.tex V1-October1,2014 5:59P.M. P.ii TrimSize:189mmx246mm Quicke ffirs.tex V1-October1,2014 5:59P.M. P.i THE BRACONID AND ICHNEUMONID PARASITOID WASPS TrimSize:189mmx246mm Quicke ffirs.tex V1-October1,2014 5:59P.M. P.ii TrimSize:189mmx246mm Quicke ffirs.tex V1-October1,2014 5:59P.M. P.iii THE BRACONID AND ICHNEUMONID PARASITOID WASPS BIOLOGY, SYSTEMATICS, EVOLUTION AND ECOLOGY Donald L. J. Quicke FacultyofScience,ChulalongkornUniversity,Bangkok,Thailand TrimSize:189mmx246mm Quicke ffirs.tex V1-October1,2014 5:59P.M. P.iv Thiseditionfirstpublished2015©2015byJohnWiley&Sons,Ltd Registeredoffice:JohnWiley&Sons,Ltd,TheAtrium,SouthernGate,Chichester,WestSussex,PO198SQ,UK Editorialoffices:9600GarsingtonRoad,Oxford,OX42DQ,UK TheAtrium,SouthernGate,Chichester,WestSussex,PO198SQ,UK 111RiverStreet,Hoboken,NJ07030-5774,USA Fordetailsofourglobaleditorialoffices,forcustomerservicesandforinformationabouthowtoapplyforpermissiontoreusethe copyrightmaterialinthisbookpleaseseeourwebsiteatwww.wiley.com/wiley-blackwell. TherightoftheauthortobeidentifiedastheauthorofthisworkhasbeenassertedinaccordancewiththeUKCopyright, DesignsandPatentsAct1988. Allrightsreserved.Nopartofthispublicationmaybereproduced,storedinaretrievalsystem,ortransmitted,inanyformorby anymeans,electronic,mechanical,photocopying,recordingorotherwise,exceptaspermittedbytheUKCopyright,Designsand PatentsAct1988,withoutthepriorpermissionofthepublisher. Designationsusedbycompaniestodistinguishtheirproductsareoftenclaimedastrademarks.Allbrandnamesandproduct namesusedinthisbookaretradenames,servicemarks,trademarksorregisteredtrademarksoftheirrespectiveowners.The publisherisnotassociatedwithanyproductorvendormentionedinthisbook. LimitofLiability/DisclaimerofWarranty:Whilethepublisherandauthor(s)haveusedtheirbesteffortsinpreparingthisbook, theymakenorepresentationsorwarrantieswithrespecttotheaccuracyorcompletenessofthecontentsofthisbookand specificallydisclaimanyimpliedwarrantiesofmerchantabilityorfitnessforaparticularpurpose.Itissoldontheunderstanding thatthepublisherisnotengagedinrenderingprofessionalservicesandneitherthepublishernortheauthorshallbeliablefor damagesarisingherefrom.Ifprofessionaladviceorotherexpertassistanceisrequired,theservicesofacompetentprofessional shouldbesought. LibraryofCongressCataloging-in-PublicationData Quicke,DonaldL.J. TheBraconidandIchneumonidParasitoidWasps:Biology,Systematics,EvolutionandEcology/byDonaldL.J.Quicke. pagescm Includesbibliographicalreferencesandindex. ISBN978-1-118-90705-4(cloth) 1.Parasitoids.2.Parasiticwasps.3.Braconidae.4.Ichneumonidae.I.Title. QL496.12.Q552015 595.7–dc23 2014015272 AcataloguerecordforthisbookisavailablefromtheBritishLibrary. Wileyalsopublishesitsbooksinavarietyofelectronicformats.Somecontentthatappearsinprintmaynotbeavailablein electronicbooks. Coverimage:©DonaldL.J.Quike Setin9/11ptPhotinabyLaserwordsPrivateLimited,Chennai,India 1 2015 TrimSize:189mmx246mm Quicke ffirs.tex V1-October1,2014 5:59P.M. P.v TothememoryofawonderfuldogcalledMii, wholivedatSaphanTaksinpier. TrimSize:189mmx246mm Quicke ffirs.tex V1-October1,2014 5:59P.M. P.vi TrimSize:189mmx246mm Quicke ftoc.tex V3-November4,2014 3:09P.M. P.vii CONTENTS Preface,xiii Number,positionandpossiblefunctionsofovipositor valvilli,50 Acknowledgements,xv Venomretentionanddelivery,52 Ovipositorsecretorypores,53 Ovipositorsensilla,54 1 INTRODUCTION,1 Ovipositorsheaths,55 Lifehistory,5 Systematics,6 4 INTERNALANDREPRODUCTIVE ANATOMY,57 Nervoussystem,58 PART1 MORPHOLOGYANDBIOLOGY,7 Digestivetract,58 Femaleinternalreproductivesystem,59 Ovaries,59 2 ADULTEXTERNALMORPHOLOGY,9 Timescaleofeggmaturation,60 Head,10 Spermatheca,61 Antennalsensilla,12 Commonoviductandvaginalgland,62 Antennalglandsandtyloids,14 Venomglandandreservoir,63 Palps,15 Dufour’sgland,64 Mesosoma,15 Cuticularhydrocarbons,66 Legs,17 Sexpheromones,67 Wings,wingvenationandwingcells,18 Maleinternalreproductivesystem,68 Confusingandsometimeserroneouslyappliedvein Spermultrastructure,69 names,26 Spermatogenyindex,70 Wingflexionlines,27 Metasoma,29 5 IMMATURESTAGES,71 Sexualdimorphism,30 Eggsandoögenesis,72 Maleexternalgenitalia,32 Hydropicandanhydropiceggs,72 Embryogenesis,73 3 THEOVIPOSITORANDOVIPOSITOR Embryonicmembranes,75 SHEATHS,35 Larva,76 Theactofoviposition,39 Larvalfeedingandnutrition,81 Functionalmorphologyofwood-drillers,41 Larvalfoodconsumptionanddietaryefficiency,82 Ovipositorstabilisationguidesandbucklingforce,43 Lipidmetabolism,82 Ovipositornotchesandendoparasitism,44 Respirationinendoparasitoids,83 Ovipositorsteeringmechanisms,44 Larvalsecretions,83 Proposedevolutionaryandrelatedovipositor Thepupalstage,84 transitions,48 Cocoons,84 vii TrimSize:189mmx246mm Quicke ftoc.tex V3-November4,2014 3:09P.M. P.viii viii Contents 6 IDIOBIONTS,KOINOBIONTSANDOTHER Passiveevasionofencapsulationbyparasitoideggs, LIFEHISTORYTRAITS,87 139 Avoidingencapsulationbyphysicalmeans,139 Parasitoidism,88 Effectofhostageandhaemocytenumber,141 Idiobiontandkoinobiontstrategies,88 Otherhostdefencemechanisms,141 Generalistsandspecialists,89 Venoms,141 Ecto-andendoparasitism,90 Neurophysiologicalvenomactions,143 Permanenthostparalysis,91 Venomeffectsonhostimmuneresponse,144 Gregariousdevelopment,91 Polydnaviruses,145 Superparasitism,92 Effectsofpolydnavirusesonhosts,152 Larvalcombatandphysiologicalsuppression,93 Otherreproductiveviruses,155 Adaptivesuperparasitism,95 Improvinghostquality,156 Multiparasitism,96 Hostcastrationandsimilareffects,156 Obligateandpreferentialmultiparasitism,99 Teratocytes,158 Hyperparasitismandpseudohyperparasitism,99 Intraspecificvariationinresistancetoparasitoids,159 Kleptoparasitism,100 Effectsonhostmoultingpattern,160 Evolutionoflifehistorystrategies,100 Parasitoid-inducedchangesinhostbehaviour,160 7 SEX,COURTSHIPANDMATING,107 Sexdetermination,108 10 CONVERGENTADAPTATIONS,163 Localmatecompetitionandavoidanceofinbreeding, Antennalhammersandvibrational 110 sounding,164 Sexallocation,110 Enlargedmandibles,167 Protandryandvirginity,112 Chisel-likemandibles,168 Thelytokyandcytoplasmicincompatibility,113 Concealednectarextractionapparatus,168 Matelocation,117 Reducednumberofpalpalsegments,169 Courtship,119 ‘Facial’protruberances,169 Swarmingandlekking,120 Frontaldepressions,170 Matingposition,121 Dorsalridgesonheadormesosoma,170 Multiplematingandspermcompetition,121 Brachypteryandaptery,170 Sex-relatedscentglands,123 Dorso-ventralflattening,171 Genomesizeandrecombination,125 Postpectalcarina,173 Cytogenetics,125 Propodealspines,173 ‘Fossorial’legs,173 8 HOSTLOCATION,ASSOCIATIVE Foretibialspines,174 LEARNINGANDHOSTASSESSMENT, Foretibialapicaltooth,174 127 Expandedhindbasitarsi,174 Toothedhindfemur,174 Tritrophicinteractions,129 Distitarsalscraper,175 Hostacceptance,130 Pectinateclawsandclawswithangularbasallobes, Associativelearning,130 175 Biosensors,134 Glabrouswingpatchesandwingmembrane Patchuse,134 scleromes,176 Carapacisation,177 9 OVERCOMINGHOSTIMMUNEREACTION Petiolatemetasomas,177 ANDPHYSIOLOGICALINTERACTIONS Modificationstotheposteriormetasomal WITHHOST,137 margin,178 Overcominghostimmunityinendoparasitoids,138 Spermathecalcolour,179