Donald L. J. Quicke is currently Visiting Professor at the Department of Biology, Faculty of Science, Chulalongkorn University, Thailand. He graduated from Oxford University with a degree in zoology and after doctoral and postdoctoral work on snail neurophysiology, sea anemone ecology and spider venoms, made parasitic wasps, and especially the ichneumonoid wasp family Braconidae, his main love and research interest. He held a lectureship at Sheffield University, moved to Imperial College London in 1993 and held a joint post between them and the Natural History Museum, London, until retiring in 2013 to live in Thailand. He was made Professor of Systematics in 2008. He has travelled widely collecting and studying parasitic wasps, especially in Africa. Over the past years he has described more than 560 new species and 76 new genera, including a number of fossil taxa, as well as making extensive studies of functional anatomy parasitic wasp ovipositors which are of enormous biological importance. A lot of his recent work has concerned global diversity estimation and patterns.

"Overall, this is a highly valuable compendium of known information, as well as currently unanswered questions, concerning ichneumonoid wasps.... Quicke is to be congratulated for producing a standard work that I, for one, will be consulting for a long time." (American Entomologist, 2016)"This is certainly a field with many pitfalls, but there is hardly a better guide through it than Professor Quicke." (International Journal of Environmental Studies, 9 March December 2015)"It sounds like a backhanded compliment to say that this is the best book of its kind, when I have already said that it is the only book of its kind. However, The Braconid and Ichneumonid Parasitoid Wasps goes beyond being the best of a limited field – it is a truly impressive assemblage of information on an intriguing and important group of insects. I hope that it inspires more people to work in the field." (Bulletin de la Société d'entomologie du Canada, 2015)

• Preface xiiiAcknowledgements xv1 Introduction 1Life history 5Systematics 6Part 1 Morphology and Biology 72 Adult External Morphology 9Head 10Antennal sensilla 12Antennal glands and tyloids 14Palps 15Mesosoma 15Legs 17Wings wing venation and wing cells 18Confusing and sometimes erroneously applied vein names 26Wing flexion lines 27Metasoma 29Sexual dimorphism 30Male external genitalia 323 The Ovipositor and Ovipositor Sheaths 35The act of oviposition 39Functional morphology of wood-drillers 41Ovipositor stabilisation guides and buckling force 43Ovipositor notches and endoparasitism 44Ovipositor steering mechanisms 44Proposed evolutionary and related ovipositor transitions 48Number position and possible functions of ovipositor valvilli 50Venom retention and delivery 52Ovipositor secretory pores 53Ovipositor sensilla 54Ovipositor sheaths 554 Internal and Reproductive Anatomy 57Nervous system 58Digestive tract 58Female internal reproductive system 59Ovaries 59Time scale of egg maturation 60Spermatheca 61Common oviduct and vaginal gland 62Venom gland and reservoir 63Dufour’s gland 64Cuticular hydrocarbons 66Sex pheromones 67Male internal reproductive system 68Sperm ultrastructure 69Spermatogeny index 705 Immature Stages 71Eggs and oögenesis 72Hydropic and anhydropic eggs 72Embryogenesis 73Embryonic membranes 75Larva 76Larval feeding and nutrition 81Larval food consumption and dietary efficiency 82Lipid metabolism 82Respiration in endoparasitoids 83Larval secretions 83The pupal stage 84Cocoons 846 Idiobionts Koinobionts and Other Life History Traits 87Parasitoidism 88Idiobiont and koinobiont strategies 88Generalists and specialists 89Ecto- and endoparasitism 90Permanent host paralysis 91Gregarious development 91Superparasitism 92Larval combat and physiological suppression 93Adaptive superparasitism 95Multiparasitism 96Obligate and preferential multiparasitism 99Hyperparasitism and pseudohyperparasitism 99Kleptoparasitism 100Evolution of life history strategies 1007 Sex Courtship and Mating 107Sex determination 108Local mate competition and avoidance of inbreeding 110Sex allocation 110Protandry and virginity 112Thelytoky and cytoplasmic incompatibility 113Mate location 117Courtship 119Swarming and lekking 120Mating position 121Multiple mating and sperm competition 121Sex-related scent glands 123Genome size and recombination 125Cytogenetics 1258 Host Location Associative Learning and Host Assessment 127Tritrophic interactions 129Host acceptance 130Associative learning 130Biosensors 134Patch use 1349 Overcoming Host Immune Reaction and Physiological Interactions With Host 137Overcoming host immunity in endoparasitoids 138Passive evasion of encapsulation by parasitoid eggs 139Avoiding encapsulation by physical means 139Effect of host age and haemocyte number 141Other host defence mechanisms 141Venoms 141Neurophysiological venom actions 143Venom effects on host immune response 144Polydnaviruses 145Effects of polydnaviruses on hosts 152Other reproductive viruses 155Improving host quality 156Host castration and similar effects 156Teratocytes 158Intraspecific variation in resistance to parasitoids 159Effects on host moulting pattern 160Parasitoid-induced changes in host behaviour 16010 Convergent Adaptations 163Antennal hammers and vibrational sounding 164Enlarged mandibles 167Chisel-like mandibles 168Concealed nectar extraction apparatus 168Reduced number of palpal segments 169‘Facial’ protruberances 169Frontal depressions 170Dorsal ridges on head or mesosoma 170Brachyptery and aptery 170Dorso-ventral flattening 171Postpectal carina 173Propodeal spines 173‘Fossorial’ legs 173Fore tibial spines 174Fore tibial apical tooth 174Expanded hind basitarsi 174Toothed hind femur 174Distitarsal scraper 175Pectinate claws and claws with angular basal lobes 175Glabrous wing patches and wing membrane scleromes 176Carapacisation 177Petiolate metasomas 177Modifications to the posterior metasomal margin 178Spermathecal colour 179Compression of apical part of metasoma 179The ‘ophionoid facies’ 179White antennal stripes and tips 180White ovipositor sheath stripes and tips 181Number of larval instars 182Egg-larval parasitism 182Disc-like larval antennae 182Reduction of larval hypostomal spur 183Wide and heavily sclerotised larval epistoma 184Suspended cocoons 184Polyembryony 184Phytophagy and cecidogenesis 184Part 2 Taxonomic and Systematic Treatment 18711 Overview of Ichneumonoidea: Relationships and Systematics 189Monophyly of Ichneumonoidea Ichneumonidae and Braconidae 190Relationship of Ichneumonoidea to other Hymenoptera 190Fossil history and family-level phylogeny 192Brief history of classification 194Ancestral biology of Ichneumonoidea 196Separating ichneumonids from braconids 197Identifying specimens 19812 Phylogeny and Systematics of The Braconidae 201Historical perspective 202Morphophylogenetic hypotheses 202Molecular phylogenetics 204Braconid classification 205Eoichneumoninae† 205Trachypetiformes 205Trachypetinae 205Cyclostomes incertae sedis 209Protorhyssalinae et al. 209Apozyginae 210The aphidioid clade or ‘Gondwanan’ complex 212Aphidiinae 212Maxfischeriinae 224Mesostoinae (including Canberreriini and Hydrangeocolini) 225The remaining cyclostomes 229Doryctinae (including Ypsistocerini) 231Pambolinae 236Rhysipolinae 237Rhyssalinae 238Rogadinae s.l. Hormiinae Lysiterminae 243Betylobraconinae 243Hormiinae 243Lysiterminae 245Rogadinae sensu stricto 246Alysioid subcomplex including Braconinae 250Alysiinae and Opiinae 250Alysiinae 251General Alysiinae biology 251Alysiini 253Dacnusini 255Opiinae 256Braconinae 260Exothecinae 269Gnamptodontinae (= Gnaptodontinae) 270Telengaiinae 271The non-cyclostomes 271Sigalphoid complex 271Agathidinae 272Sigalphinae 275Helconoid complex 278Helconinae 279Helconoid group incertae sedis 281Blacinae 282Acampsohelconinae 283Macrocentrine subcomplex 284Macrocentrinae 284Charmontiinae 287Amicrocentrinae 287Xiphozelinae 288Homolobinae 290Microtypinae 292Orgilinae 292Euphoroid complex 294Euphorinae 294Cenocoeliinae 310The microgastroids 311Cardiochilinae 312Cheloninae (including Adeliini) 315Dirrhopinae 319Ichneutinae 320Khoikhoiinae 322Mendesellinae 322Microgastrinae 322Miracinae 335Unplaced subfamilies 335Masoninae 335Meteorideinae 33713 Phylogeny and Systematics of The Ichneumonidae 341History of ichneumonid classification 342Henry Townes (1913–90) and his idiosyncratic nomenclature 344The extinct subfamilies 344Tanychorinae† 344Palaeoichneumoninae† 346Labenopimplinae† 348Pherombinae† 349Townesitinae† 349The xoridiformes 349Xoridinae 349The labeniformes 353Labeninae 353Groteini 355Labenini 355Poecilocryptini 356The pimpliformes 356Acaenitinae 356Collyriinae 359Cylloceriinae 360Diacritinae 360Diplazontinae 361Orthocentrinae (= Helictinae) 366Pimplinae 367Delomeristini 369Ephialtini (= Pimplini of Townes) 369Polysphincta group 371Pimplini 373Poemeniinae (= Neoxoridinae) 378Poemeniini 378Pseudorhyssini 378Rodrigamini 378Rhyssinae 379The ichneumoniformes 383Adelognathinae 383Agriotypinae 385Alomyinae 387Cryptinae 388Aptesini 391Cryptini 391Phygadeuontini 393Ichneumoninae 394The brachycyrtiformes 398Brachycyrtinae 398Claseinae (Clasinae) 398Pedunculinae 399The orthopelmatiformes 400Orthopelmatinae 400The ophioniformes 400Lower ophioniformes 402Banchinae 402Lycorininae 406Sisyrostolinae 407Stilbopinae 407Tryphoninae 411Middle ophioniformes 416Ctenopelmatinae 416Mesochorinae 421Metopiinae 422Oxytorinae 424Tatogastrinae 425Tersilochinae (including Neorhacodinae and Phrudinae s.s.) 426Higher ophioniformes 430Anomaloninae 430Campopleginae 432Cremastinae 438Hybrizontinae 439Nesomesochorinae 442Ophioninae 442Unplaced subfamilies 445Eucerotinae 445Microleptinae 447Part 3 Ecology and Diversity 45114 Ecology 453Adult diet 454Host-feeding 454Water sugar and pollen feeding 457Fecundity 460Voltinism and seasonality 462Daily activity patterns 462Diapause 463Cold hardiness hibernation and overwintering 465Coloration and thermoregulation 467Biological control 467Effect on host food consumption 471Artificial diets 474Artificial hosts 475Use of alternative hosts 475Hyperparasitism and kleptoparasitism 476Predation 477Pathogens 477Transmission of host pathogens 479Dispersal 480Coloration and mimetic rings 480Palatability and odours 481Competition 482Apparent competition 482Host ranges of parasitoids 483Parasitoid guilds and food webs 484Evolution of host ranges and speciation 48615 Local and Global Patterns In Diversity 489Field research in the tropics and anomalous diversity 490Estimation of global ichneumonoid species richness 492Distribution related to climate and latitude 496The nasty host hypothesis 497Biogeography 503Islands and their parasitoid faunas 505Species accumulation curves 506Altitudinal gradients 507Estimating local species diversity 508Ichneumonoidea as biodiversity indicators 510Conservation 510Effect of habitat degradation on ichneumonoid composition 511Significance of cryptic species 51116 Collecting and Rearing Ichneumonoidea 513Field collecting adults 516Pan traps 518Sweep netting 519Light trapping 521Canopy fogging 521Malaise traps 521Rearings from wild-collected hosts 523Rearing leaf rollers and tiers 524Substrate rearings 524Culturing 524Mating in captivity 525Mass rearing 525Mounting specimens for taxonomic study 526Preparing specimens from alcohol storage 526Direct pinning 527Side gluing 527Card rectangles and card points 527Secondary staging 528Labelling 528Preserving specimens for DNA analysis 528Packaging and posting specimens to other workers 53017 Epilogue 533Phylogenetic questions 534Host and parasitism questions 534Physiological questions 535Ecological questions 536Glossary 539References 547Author index 633General index 653Host index 659Ichneumonoid genus tribe and subfamily index 665Ichneumonoidea species index 677Color Plate Sections Are Inserted Between Pages Noted BelowFirst 13-page colour plate section (between pages 112 and 113)Second 13-page colour plate section (between pages 224 and 225)Third 13-page colour plate section (between pages 336 and 337)Fourth 13-page colour plate section (between pages 448 and 449)

Donald L. J. Quicke is currently Visiting Professor at the Department of Biology, Faculty of Science, Chulalongkorn University, Thailand. He graduated from Oxford University with a degree in zoology and after doctoral and postdoctoral work on snail neurophysiology, sea anemone ecology and spider venoms, made parasitic wasps, and especially the ichneumonoid wasp family Braconidae, his main love and research interest. He held a lectureship at Sheffield University, moved to Imperial College London in 1993 and held a joint post between them and the Natural History Museum, London, until retiring in 2013 to live in Thailand. He was made Professor of Systematics in 2008. He has travelled widely collecting and studying parasitic wasps, especially in Africa. Over the past years he has described more than 560 new species and 76 new genera, including a number of fossil taxa, as well as making extensive studies of functional anatomy parasitic wasp ovipositors which are of enormous biological importance. A lot of his recent work has concerned global diversity estimation and patterns.

"Overall, this is a highly valuable compendium of known information, as well as currently unanswered questions, concerning ichneumonoid wasps.... Quicke is to be congratulated for producing a standard work that I, for one, will be consulting for a long time." (American Entomologist, 2016)"This is certainly a field with many pitfalls, but there is hardly a better guide through it than Professor Quicke." (International Journal of Environmental Studies, 9 March December 2015)"It sounds like a backhanded compliment to say that this is the best book of its kind, when I have already said that it is the only book of its kind. However, The Braconid and Ichneumonid Parasitoid Wasps goes beyond being the best of a limited field – it is a truly impressive assemblage of information on an intriguing and important group of insects. I hope that it inspires more people to work in the field." (Bulletin de la Société d'entomologie du Canada, 2015)

• Preface xiiiAcknowledgements xv1 Introduction 1Life history 5Systematics 6Part 1 Morphology and Biology 72 Adult External Morphology 9Head 10Antennal sensilla 12Antennal glands and tyloids 14Palps 15Mesosoma 15Legs 17Wings wing venation and wing cells 18Confusing and sometimes erroneously applied vein names 26Wing flexion lines 27Metasoma 29Sexual dimorphism 30Male external genitalia 323 The Ovipositor and Ovipositor Sheaths 35The act of oviposition 39Functional morphology of wood-drillers 41Ovipositor stabilisation guides and buckling force 43Ovipositor notches and endoparasitism 44Ovipositor steering mechanisms 44Proposed evolutionary and related ovipositor transitions 48Number position and possible functions of ovipositor valvilli 50Venom retention and delivery 52Ovipositor secretory pores 53Ovipositor sensilla 54Ovipositor sheaths 554 Internal and Reproductive Anatomy 57Nervous system 58Digestive tract 58Female internal reproductive system 59Ovaries 59Time scale of egg maturation 60Spermatheca 61Common oviduct and vaginal gland 62Venom gland and reservoir 63Dufour’s gland 64Cuticular hydrocarbons 66Sex pheromones 67Male internal reproductive system 68Sperm ultrastructure 69Spermatogeny index 705 Immature Stages 71Eggs and oögenesis 72Hydropic and anhydropic eggs 72Embryogenesis 73Embryonic membranes 75Larva 76Larval feeding and nutrition 81Larval food consumption and dietary efficiency 82Lipid metabolism 82Respiration in endoparasitoids 83Larval secretions 83The pupal stage 84Cocoons 846 Idiobionts Koinobionts and Other Life History Traits 87Parasitoidism 88Idiobiont and koinobiont strategies 88Generalists and specialists 89Ecto- and endoparasitism 90Permanent host paralysis 91Gregarious development 91Superparasitism 92Larval combat and physiological suppression 93Adaptive superparasitism 95Multiparasitism 96Obligate and preferential multiparasitism 99Hyperparasitism and pseudohyperparasitism 99Kleptoparasitism 100Evolution of life history strategies 1007 Sex Courtship and Mating 107Sex determination 108Local mate competition and avoidance of inbreeding 110Sex allocation 110Protandry and virginity 112Thelytoky and cytoplasmic incompatibility 113Mate location 117Courtship 119Swarming and lekking 120Mating position 121Multiple mating and sperm competition 121Sex-related scent glands 123Genome size and recombination 125Cytogenetics 1258 Host Location Associative Learning and Host Assessment 127Tritrophic interactions 129Host acceptance 130Associative learning 130Biosensors 134Patch use 1349 Overcoming Host Immune Reaction and Physiological Interactions With Host 137Overcoming host immunity in endoparasitoids 138Passive evasion of encapsulation by parasitoid eggs 139Avoiding encapsulation by physical means 139Effect of host age and haemocyte number 141Other host defence mechanisms 141Venoms 141Neurophysiological venom actions 143Venom effects on host immune response 144Polydnaviruses 145Effects of polydnaviruses on hosts 152Other reproductive viruses 155Improving host quality 156Host castration and similar effects 156Teratocytes 158Intraspecific variation in resistance to parasitoids 159Effects on host moulting pattern 160Parasitoid-induced changes in host behaviour 16010 Convergent Adaptations 163Antennal hammers and vibrational sounding 164Enlarged mandibles 167Chisel-like mandibles 168Concealed nectar extraction apparatus 168Reduced number of palpal segments 169‘Facial’ protruberances 169Frontal depressions 170Dorsal ridges on head or mesosoma 170Brachyptery and aptery 170Dorso-ventral flattening 171Postpectal carina 173Propodeal spines 173‘Fossorial’ legs 173Fore tibial spines 174Fore tibial apical tooth 174Expanded hind basitarsi 174Toothed hind femur 174Distitarsal scraper 175Pectinate claws and claws with angular basal lobes 175Glabrous wing patches and wing membrane scleromes 176Carapacisation 177Petiolate metasomas 177Modifications to the posterior metasomal margin 178Spermathecal colour 179Compression of apical part of metasoma 179The ‘ophionoid facies’ 179White antennal stripes and tips 180White ovipositor sheath stripes and tips 181Number of larval instars 182Egg-larval parasitism 182Disc-like larval antennae 182Reduction of larval hypostomal spur 183Wide and heavily sclerotised larval epistoma 184Suspended cocoons 184Polyembryony 184Phytophagy and cecidogenesis 184Part 2 Taxonomic and Systematic Treatment 18711 Overview of Ichneumonoidea: Relationships and Systematics 189Monophyly of Ichneumonoidea Ichneumonidae and Braconidae 190Relationship of Ichneumonoidea to other Hymenoptera 190Fossil history and family-level phylogeny 192Brief history of classification 194Ancestral biology of Ichneumonoidea 196Separating ichneumonids from braconids 197Identifying specimens 19812 Phylogeny and Systematics of The Braconidae 201Historical perspective 202Morphophylogenetic hypotheses 202Molecular phylogenetics 204Braconid classification 205Eoichneumoninae† 205Trachypetiformes 205Trachypetinae 205Cyclostomes incertae sedis 209Protorhyssalinae et al. 209Apozyginae 210The aphidioid clade or ‘Gondwanan’ complex 212Aphidiinae 212Maxfischeriinae 224Mesostoinae (including Canberreriini and Hydrangeocolini) 225The remaining cyclostomes 229Doryctinae (including Ypsistocerini) 231Pambolinae 236Rhysipolinae 237Rhyssalinae 238Rogadinae s.l. Hormiinae Lysiterminae 243Betylobraconinae 243Hormiinae 243Lysiterminae 245Rogadinae sensu stricto 246Alysioid subcomplex including Braconinae 250Alysiinae and Opiinae 250Alysiinae 251General Alysiinae biology 251Alysiini 253Dacnusini 255Opiinae 256Braconinae 260Exothecinae 269Gnamptodontinae (= Gnaptodontinae) 270Telengaiinae 271The non-cyclostomes 271Sigalphoid complex 271Agathidinae 272Sigalphinae 275Helconoid complex 278Helconinae 279Helconoid group incertae sedis 281Blacinae 282Acampsohelconinae 283Macrocentrine subcomplex 284Macrocentrinae 284Charmontiinae 287Amicrocentrinae 287Xiphozelinae 288Homolobinae 290Microtypinae 292Orgilinae 292Euphoroid complex 294Euphorinae 294Cenocoeliinae 310The microgastroids 311Cardiochilinae 312Cheloninae (including Adeliini) 315Dirrhopinae 319Ichneutinae 320Khoikhoiinae 322Mendesellinae 322Microgastrinae 322Miracinae 335Unplaced subfamilies 335Masoninae 335Meteorideinae 33713 Phylogeny and Systematics of The Ichneumonidae 341History of ichneumonid classification 342Henry Townes (1913–90) and his idiosyncratic nomenclature 344The extinct subfamilies 344Tanychorinae† 344Palaeoichneumoninae† 346Labenopimplinae† 348Pherombinae† 349Townesitinae† 349The xoridiformes 349Xoridinae 349The labeniformes 353Labeninae 353Groteini 355Labenini 355Poecilocryptini 356The pimpliformes 356Acaenitinae 356Collyriinae 359Cylloceriinae 360Diacritinae 360Diplazontinae 361Orthocentrinae (= Helictinae) 366Pimplinae 367Delomeristini 369Ephialtini (= Pimplini of Townes) 369Polysphincta group 371Pimplini 373Poemeniinae (= Neoxoridinae) 378Poemeniini 378Pseudorhyssini 378Rodrigamini 378Rhyssinae 379The ichneumoniformes 383Adelognathinae 383Agriotypinae 385Alomyinae 387Cryptinae 388Aptesini 391Cryptini 391Phygadeuontini 393Ichneumoninae 394The brachycyrtiformes 398Brachycyrtinae 398Claseinae (Clasinae) 398Pedunculinae 399The orthopelmatiformes 400Orthopelmatinae 400The ophioniformes 400Lower ophioniformes 402Banchinae 402Lycorininae 406Sisyrostolinae 407Stilbopinae 407Tryphoninae 411Middle ophioniformes 416Ctenopelmatinae 416Mesochorinae 421Metopiinae 422Oxytorinae 424Tatogastrinae 425Tersilochinae (including Neorhacodinae and Phrudinae s.s.) 426Higher ophioniformes 430Anomaloninae 430Campopleginae 432Cremastinae 438Hybrizontinae 439Nesomesochorinae 442Ophioninae 442Unplaced subfamilies 445Eucerotinae 445Microleptinae 447Part 3 Ecology and Diversity 45114 Ecology 453Adult diet 454Host-feeding 454Water sugar and pollen feeding 457Fecundity 460Voltinism and seasonality 462Daily activity patterns 462Diapause 463Cold hardiness hibernation and overwintering 465Coloration and thermoregulation 467Biological control 467Effect on host food consumption 471Artificial diets 474Artificial hosts 475Use of alternative hosts 475Hyperparasitism and kleptoparasitism 476Predation 477Pathogens 477Transmission of host pathogens 479Dispersal 480Coloration and mimetic rings 480Palatability and odours 481Competition 482Apparent competition 482Host ranges of parasitoids 483Parasitoid guilds and food webs 484Evolution of host ranges and speciation 48615 Local and Global Patterns In Diversity 489Field research in the tropics and anomalous diversity 490Estimation of global ichneumonoid species richness 492Distribution related to climate and latitude 496The nasty host hypothesis 497Biogeography 503Islands and their parasitoid faunas 505Species accumulation curves 506Altitudinal gradients 507Estimating local species diversity 508Ichneumonoidea as biodiversity indicators 510Conservation 510Effect of habitat degradation on ichneumonoid composition 511Significance of cryptic species 51116 Collecting and Rearing Ichneumonoidea 513Field collecting adults 516Pan traps 518Sweep netting 519Light trapping 521Canopy fogging 521Malaise traps 521Rearings from wild-collected hosts 523Rearing leaf rollers and tiers 524Substrate rearings 524Culturing 524Mating in captivity 525Mass rearing 525Mounting specimens for taxonomic study 526Preparing specimens from alcohol storage 526Direct pinning 527Side gluing 527Card rectangles and card points 527Secondary staging 528Labelling 528Preserving specimens for DNA analysis 528Packaging and posting specimens to other workers 53017 Epilogue 533Phylogenetic questions 534Host and parasitism questions 534Physiological questions 535Ecological questions 536Glossary 539References 547Author index 633General index 653Host index 659Ichneumonoid genus tribe and subfamily index 665Ichneumonoidea species index 677Color Plate Sections Are Inserted Between Pages Noted BelowFirst 13-page colour plate section (between pages 112 and 113)Second 13-page colour plate section (between pages 224 and 225)Third 13-page colour plate section (between pages 336 and 337)Fourth 13-page colour plate section (between pages 448 and 449)

Donald L. J. Quicke is currently Visiting Professor at the Department of Biology, Faculty of Science, Chulalongkorn University, Thailand. He graduated from Oxford University with a degree in zoology and after doctoral and postdoctoral work on snail neurophysiology, sea anemone ecology and spider venoms, made parasitic wasps, and especially the ichneumonoid wasp family Braconidae, his main love and research interest. He held a lectureship at Sheffield University, moved to Imperial College London in 1993 and held a joint post between them and the Natural History Museum, London, until retiring in 2013 to live in Thailand. He was made Professor of Systematics in 2008. He has travelled widely collecting and studying parasitic wasps, especially in Africa. Over the past years he has described more than 560 new species and 76 new genera, including a number of fossil taxa, as well as making extensive studies of functional anatomy parasitic wasp ovipositors which are of enormous biological importance. A lot of his recent work has concerned global diversity estimation and patterns.

"Overall, this is a highly valuable compendium of known information, as well as currently unanswered questions, concerning ichneumonoid wasps.... Quicke is to be congratulated for producing a standard work that I, for one, will be consulting for a long time." (American Entomologist, 2016)"This is certainly a field with many pitfalls, but there is hardly a better guide through it than Professor Quicke." (International Journal of Environmental Studies, 9 March December 2015)"It sounds like a backhanded compliment to say that this is the best book of its kind, when I have already said that it is the only book of its kind. However, The Braconid and Ichneumonid Parasitoid Wasps goes beyond being the best of a limited field – it is a truly impressive assemblage of information on an intriguing and important group of insects. I hope that it inspires more people to work in the field." (Bulletin de la Société d'entomologie du Canada, 2015)

• Preface xiiiAcknowledgements xv1 Introduction 1Life history 5Systematics 6Part 1 Morphology and Biology 72 Adult External Morphology 9Head 10Antennal sensilla 12Antennal glands and tyloids 14Palps 15Mesosoma 15Legs 17Wings wing venation and wing cells 18Confusing and sometimes erroneously applied vein names 26Wing flexion lines 27Metasoma 29Sexual dimorphism 30Male external genitalia 323 The Ovipositor and Ovipositor Sheaths 35The act of oviposition 39Functional morphology of wood-drillers 41Ovipositor stabilisation guides and buckling force 43Ovipositor notches and endoparasitism 44Ovipositor steering mechanisms 44Proposed evolutionary and related ovipositor transitions 48Number position and possible functions of ovipositor valvilli 50Venom retention and delivery 52Ovipositor secretory pores 53Ovipositor sensilla 54Ovipositor sheaths 554 Internal and Reproductive Anatomy 57Nervous system 58Digestive tract 58Female internal reproductive system 59Ovaries 59Time scale of egg maturation 60Spermatheca 61Common oviduct and vaginal gland 62Venom gland and reservoir 63Dufour’s gland 64Cuticular hydrocarbons 66Sex pheromones 67Male internal reproductive system 68Sperm ultrastructure 69Spermatogeny index 705 Immature Stages 71Eggs and oögenesis 72Hydropic and anhydropic eggs 72Embryogenesis 73Embryonic membranes 75Larva 76Larval feeding and nutrition 81Larval food consumption and dietary efficiency 82Lipid metabolism 82Respiration in endoparasitoids 83Larval secretions 83The pupal stage 84Cocoons 846 Idiobionts Koinobionts and Other Life History Traits 87Parasitoidism 88Idiobiont and koinobiont strategies 88Generalists and specialists 89Ecto- and endoparasitism 90Permanent host paralysis 91Gregarious development 91Superparasitism 92Larval combat and physiological suppression 93Adaptive superparasitism 95Multiparasitism 96Obligate and preferential multiparasitism 99Hyperparasitism and pseudohyperparasitism 99Kleptoparasitism 100Evolution of life history strategies 1007 Sex Courtship and Mating 107Sex determination 108Local mate competition and avoidance of inbreeding 110Sex allocation 110Protandry and virginity 112Thelytoky and cytoplasmic incompatibility 113Mate location 117Courtship 119Swarming and lekking 120Mating position 121Multiple mating and sperm competition 121Sex-related scent glands 123Genome size and recombination 125Cytogenetics 1258 Host Location Associative Learning and Host Assessment 127Tritrophic interactions 129Host acceptance 130Associative learning 130Biosensors 134Patch use 1349 Overcoming Host Immune Reaction and Physiological Interactions With Host 137Overcoming host immunity in endoparasitoids 138Passive evasion of encapsulation by parasitoid eggs 139Avoiding encapsulation by physical means 139Effect of host age and haemocyte number 141Other host defence mechanisms 141Venoms 141Neurophysiological venom actions 143Venom effects on host immune response 144Polydnaviruses 145Effects of polydnaviruses on hosts 152Other reproductive viruses 155Improving host quality 156Host castration and similar effects 156Teratocytes 158Intraspecific variation in resistance to parasitoids 159Effects on host moulting pattern 160Parasitoid-induced changes in host behaviour 16010 Convergent Adaptations 163Antennal hammers and vibrational sounding 164Enlarged mandibles 167Chisel-like mandibles 168Concealed nectar extraction apparatus 168Reduced number of palpal segments 169‘Facial’ protruberances 169Frontal depressions 170Dorsal ridges on head or mesosoma 170Brachyptery and aptery 170Dorso-ventral flattening 171Postpectal carina 173Propodeal spines 173‘Fossorial’ legs 173Fore tibial spines 174Fore tibial apical tooth 174Expanded hind basitarsi 174Toothed hind femur 174Distitarsal scraper 175Pectinate claws and claws with angular basal lobes 175Glabrous wing patches and wing membrane scleromes 176Carapacisation 177Petiolate metasomas 177Modifications to the posterior metasomal margin 178Spermathecal colour 179Compression of apical part of metasoma 179The ‘ophionoid facies’ 179White antennal stripes and tips 180White ovipositor sheath stripes and tips 181Number of larval instars 182Egg-larval parasitism 182Disc-like larval antennae 182Reduction of larval hypostomal spur 183Wide and heavily sclerotised larval epistoma 184Suspended cocoons 184Polyembryony 184Phytophagy and cecidogenesis 184Part 2 Taxonomic and Systematic Treatment 18711 Overview of Ichneumonoidea: Relationships and Systematics 189Monophyly of Ichneumonoidea Ichneumonidae and Braconidae 190Relationship of Ichneumonoidea to other Hymenoptera 190Fossil history and family-level phylogeny 192Brief history of classification 194Ancestral biology of Ichneumonoidea 196Separating ichneumonids from braconids 197Identifying specimens 19812 Phylogeny and Systematics of The Braconidae 201Historical perspective 202Morphophylogenetic hypotheses 202Molecular phylogenetics 204Braconid classification 205Eoichneumoninae† 205Trachypetiformes 205Trachypetinae 205Cyclostomes incertae sedis 209Protorhyssalinae et al. 209Apozyginae 210The aphidioid clade or ‘Gondwanan’ complex 212Aphidiinae 212Maxfischeriinae 224Mesostoinae (including Canberreriini and Hydrangeocolini) 225The remaining cyclostomes 229Doryctinae (including Ypsistocerini) 231Pambolinae 236Rhysipolinae 237Rhyssalinae 238Rogadinae s.l. Hormiinae Lysiterminae 243Betylobraconinae 243Hormiinae 243Lysiterminae 245Rogadinae sensu stricto 246Alysioid subcomplex including Braconinae 250Alysiinae and Opiinae 250Alysiinae 251General Alysiinae biology 251Alysiini 253Dacnusini 255Opiinae 256Braconinae 260Exothecinae 269Gnamptodontinae (= Gnaptodontinae) 270Telengaiinae 271The non-cyclostomes 271Sigalphoid complex 271Agathidinae 272Sigalphinae 275Helconoid complex 278Helconinae 279Helconoid group incertae sedis 281Blacinae 282Acampsohelconinae 283Macrocentrine subcomplex 284Macrocentrinae 284Charmontiinae 287Amicrocentrinae 287Xiphozelinae 288Homolobinae 290Microtypinae 292Orgilinae 292Euphoroid complex 294Euphorinae 294Cenocoeliinae 310The microgastroids 311Cardiochilinae 312Cheloninae (including Adeliini) 315Dirrhopinae 319Ichneutinae 320Khoikhoiinae 322Mendesellinae 322Microgastrinae 322Miracinae 335Unplaced subfamilies 335Masoninae 335Meteorideinae 33713 Phylogeny and Systematics of The Ichneumonidae 341History of ichneumonid classification 342Henry Townes (1913–90) and his idiosyncratic nomenclature 344The extinct subfamilies 344Tanychorinae† 344Palaeoichneumoninae† 346Labenopimplinae† 348Pherombinae† 349Townesitinae† 349The xoridiformes 349Xoridinae 349The labeniformes 353Labeninae 353Groteini 355Labenini 355Poecilocryptini 356The pimpliformes 356Acaenitinae 356Collyriinae 359Cylloceriinae 360Diacritinae 360Diplazontinae 361Orthocentrinae (= Helictinae) 366Pimplinae 367Delomeristini 369Ephialtini (= Pimplini of Townes) 369Polysphincta group 371Pimplini 373Poemeniinae (= Neoxoridinae) 378Poemeniini 378Pseudorhyssini 378Rodrigamini 378Rhyssinae 379The ichneumoniformes 383Adelognathinae 383Agriotypinae 385Alomyinae 387Cryptinae 388Aptesini 391Cryptini 391Phygadeuontini 393Ichneumoninae 394The brachycyrtiformes 398Brachycyrtinae 398Claseinae (Clasinae) 398Pedunculinae 399The orthopelmatiformes 400Orthopelmatinae 400The ophioniformes 400Lower ophioniformes 402Banchinae 402Lycorininae 406Sisyrostolinae 407Stilbopinae 407Tryphoninae 411Middle ophioniformes 416Ctenopelmatinae 416Mesochorinae 421Metopiinae 422Oxytorinae 424Tatogastrinae 425Tersilochinae (including Neorhacodinae and Phrudinae s.s.) 426Higher ophioniformes 430Anomaloninae 430Campopleginae 432Cremastinae 438Hybrizontinae 439Nesomesochorinae 442Ophioninae 442Unplaced subfamilies 445Eucerotinae 445Microleptinae 447Part 3 Ecology and Diversity 45114 Ecology 453Adult diet 454Host-feeding 454Water sugar and pollen feeding 457Fecundity 460Voltinism and seasonality 462Daily activity patterns 462Diapause 463Cold hardiness hibernation and overwintering 465Coloration and thermoregulation 467Biological control 467Effect on host food consumption 471Artificial diets 474Artificial hosts 475Use of alternative hosts 475Hyperparasitism and kleptoparasitism 476Predation 477Pathogens 477Transmission of host pathogens 479Dispersal 480Coloration and mimetic rings 480Palatability and odours 481Competition 482Apparent competition 482Host ranges of parasitoids 483Parasitoid guilds and food webs 484Evolution of host ranges and speciation 48615 Local and Global Patterns In Diversity 489Field research in the tropics and anomalous diversity 490Estimation of global ichneumonoid species richness 492Distribution related to climate and latitude 496The nasty host hypothesis 497Biogeography 503Islands and their parasitoid faunas 505Species accumulation curves 506Altitudinal gradients 507Estimating local species diversity 508Ichneumonoidea as biodiversity indicators 510Conservation 510Effect of habitat degradation on ichneumonoid composition 511Significance of cryptic species 51116 Collecting and Rearing Ichneumonoidea 513Field collecting adults 516Pan traps 518Sweep netting 519Light trapping 521Canopy fogging 521Malaise traps 521Rearings from wild-collected hosts 523Rearing leaf rollers and tiers 524Substrate rearings 524Culturing 524Mating in captivity 525Mass rearing 525Mounting specimens for taxonomic study 526Preparing specimens from alcohol storage 526Direct pinning 527Side gluing 527Card rectangles and card points 527Secondary staging 528Labelling 528Preserving specimens for DNA analysis 528Packaging and posting specimens to other workers 53017 Epilogue 533Phylogenetic questions 534Host and parasitism questions 534Physiological questions 535Ecological questions 536Glossary 539References 547Author index 633General index 653Host index 659Ichneumonoid genus tribe and subfamily index 665Ichneumonoidea species index 677Color Plate Sections Are Inserted Between Pages Noted BelowFirst 13-page colour plate section (between pages 112 and 113)Second 13-page colour plate section (between pages 224 and 225)Third 13-page colour plate section (between pages 336 and 337)Fourth 13-page colour plate section (between pages 448 and 449)