The Ichneumonoidea is a vast and important superfamily of parasitic wasps, with some 60,000 described species and estimated numbers far higher, especially for small-bodied tropical taxa. The superfamily comprises two cosmopolitan families - Braconidae and Ichneumonidae - that have largely attracted separate groups of researchers, and this, to a considerable extent, has meant that understanding of their adaptive features has often been considered in isolation. This book considers both families, highlighting similarities and differences in their adaptations. The classification of the whole of the Ichneumonoidea, along with most other insect orders, has been plagued by typology whereby undue importance has been attributed to particular characters in defining groups. Typology is a common disease of traditional taxonomy such that, until recently, quite a lot of taxa have been associated with the wrong higher clades. The sheer size of the group, and until the last 30 or so years, lack of accessible identification materials, has been a further impediment to research on all but a handful of ‘lab rat’ species usually cultured initially because of their potential in biological control. New evidence, largely in the form of molecular data, have shown that many morphological, behavioural, physiological and anatomical characters associated with basic life history features, specifically whether wasps are ecto- or endoparasitic, or idiobiont or koinobiont, can be grossly misleading in terms of the phylogeny they suggest. This book shows how, with better supported phylogenetic hypotheses entomologists can understand far more about the ways natural selection is acting upon them. This new book also focuses on this superfamily with which the author has great familiarity and provides a detailed coverage of each subfamily, emphasising  anatomy, taxonomy and systematics, biology, as well as pointing out the importance and research potential of each group. Fossil taxa are included and it also has sections on biogeography, global species richness, culturing and rearing and preparing specimens for taxonomic study. The book highlights areas where research might be particularly rewarding and suggests systems/groups that need investigation. The author provides a large compendium of references to original research on each group. This book is an essential workmate for all postgraduates and researchers working on ichneumonoid or other parasitic wasps worldwide. It will stand as a reference book for a good number of years, and while rapid advances in various fields such as genomics and host physiological interactions will lead to new information, as an overall synthesis of the current state it will stay relevant for a long time.

Table of Contents:
Preface xiii Acknowledgements xv 1 Introduction 1 Life history 5 Systematics 6 Part 1 Morphology and Biology 7 2 Adult External Morphology 9 Head 10 Antennal sensilla 12 Antennal glands and tyloids 14 Palps 15 Mesosoma 15 Legs 17 Wings wing venation and wing cells 18 Confusing and sometimes erroneously applied vein names 26 Wing flexion lines 27 Metasoma 29 Sexual dimorphism 30 Male external genitalia 32 3 The Ovipositor and Ovipositor Sheaths 35 The act of oviposition 39 Functional morphology of wood-drillers 41 Ovipositor stabilisation guides and buckling force 43 Ovipositor notches and endoparasitism 44 Ovipositor steering mechanisms 44 Proposed evolutionary and related ovipositor transitions 48 Number position and possible functions of ovipositor valvilli 50 Venom retention and delivery 52 Ovipositor secretory pores 53 Ovipositor sensilla 54 Ovipositor sheaths 55 4 Internal and Reproductive Anatomy 57 Nervous system 58 Digestive tract 58 Female internal reproductive system 59 Ovaries 59 Time scale of egg maturation 60 Spermatheca 61 Common oviduct and vaginal gland 62 Venom gland and reservoir 63 Dufour’s gland 64 Cuticular hydrocarbons 66 Sex pheromones 67 Male internal reproductive system 68 Sperm ultrastructure 69 Spermatogeny index 70 5 Immature Stages 71 Eggs and oögenesis 72 Hydropic and anhydropic eggs 72 Embryogenesis 73 Embryonic membranes 75 Larva 76 Larval feeding and nutrition 81 Larval food consumption and dietary efficiency 82 Lipid metabolism 82 Respiration in endoparasitoids 83 Larval secretions 83 The pupal stage 84 Cocoons 84 6 Idiobionts Koinobionts and Other Life History Traits 87 Parasitoidism 88 Idiobiont and koinobiont strategies 88 Generalists and specialists 89 Ecto- and endoparasitism 90 Permanent host paralysis 91 Gregarious development 91 Superparasitism 92 Larval combat and physiological suppression 93 Adaptive superparasitism 95 Multiparasitism 96 Obligate and preferential multiparasitism 99 Hyperparasitism and pseudohyperparasitism 99 Kleptoparasitism 100 Evolution of life history strategies 100 7 Sex Courtship and Mating 107 Sex determination 108 Local mate competition and avoidance of inbreeding 110 Sex allocation 110 Protandry and virginity 112 Thelytoky and cytoplasmic incompatibility 113 Mate location 117 Courtship 119 Swarming and lekking 120 Mating position 121 Multiple mating and sperm competition 121 Sex-related scent glands 123 Genome size and recombination 125 Cytogenetics 125 8 Host Location Associative Learning and Host Assessment 127 Tritrophic interactions 129 Host acceptance 130 Associative learning 130 Biosensors 134 Patch use 134 9 Overcoming Host Immune Reaction and Physiological Interactions With Host 137 Overcoming host immunity in endoparasitoids 138 Passive evasion of encapsulation by parasitoid eggs 139 Avoiding encapsulation by physical means 139 Effect of host age and haemocyte number 141 Other host defence mechanisms 141 Venoms 141 Neurophysiological venom actions 143 Venom effects on host immune response 144 Polydnaviruses 145 Effects of polydnaviruses on hosts 152 Other reproductive viruses 155 Improving host quality 156 Host castration and similar effects 156 Teratocytes 158 Intraspecific variation in resistance to parasitoids 159 Effects on host moulting pattern 160 Parasitoid-induced changes in host behaviour 160 10 Convergent Adaptations 163 Antennal hammers and vibrational sounding 164 Enlarged mandibles 167 Chisel-like mandibles 168 Concealed nectar extraction apparatus 168 Reduced number of palpal segments 169 ‘Facial’ protruberances 169 Frontal depressions 170 Dorsal ridges on head or mesosoma 170 Brachyptery and aptery 170 Dorso-ventral flattening 171 Postpectal carina 173 Propodeal spines 173 ‘Fossorial’ legs 173 Fore tibial spines 174 Fore tibial apical tooth 174 Expanded hind basitarsi 174 Toothed hind femur 174 Distitarsal scraper 175 Pectinate claws and claws with angular basal lobes 175 Glabrous wing patches and wing membrane scleromes 176 Carapacisation 177 Petiolate metasomas 177 Modifications to the posterior metasomal margin 178 Spermathecal colour 179 Compression of apical part of metasoma 179 The ‘ophionoid facies’ 179 White antennal stripes and tips 180 White ovipositor sheath stripes and tips 181 Number of larval instars 182 Egg-larval parasitism 182 Disc-like larval antennae 182 Reduction of larval hypostomal spur 183 Wide and heavily sclerotised larval epistoma 184 Suspended cocoons 184 Polyembryony 184 Phytophagy and cecidogenesis 184 Part 2 Taxonomic and Systematic Treatment 187 11 Overview of Ichneumonoidea: Relationships and Systematics 189 Monophyly of Ichneumonoidea Ichneumonidae and Braconidae 190 Relationship of Ichneumonoidea to other Hymenoptera 190 Fossil history and family-level phylogeny 192 Brief history of classification 194 Ancestral biology of Ichneumonoidea 196 Separating ichneumonids from braconids 197 Identifying specimens 198 12 Phylogeny and Systematics of The Braconidae 201 Historical perspective 202 Morphophylogenetic hypotheses 202 Molecular phylogenetics 204 Braconid classification 205 Eoichneumoninae† 205 Trachypetiformes 205 Trachypetinae 205 Cyclostomes incertae sedis 209 Protorhyssalinae et al. 209 Apozyginae 210 The aphidioid clade or ‘Gondwanan’ complex 212 Aphidiinae 212 Maxfischeriinae 224 Mesostoinae (including Canberreriini and Hydrangeocolini) 225 The remaining cyclostomes 229 Doryctinae (including Ypsistocerini) 231 Pambolinae 236 Rhysipolinae 237 Rhyssalinae 238 Rogadinae s.l. Hormiinae Lysiterminae 243 Betylobraconinae 243 Hormiinae 243 Lysiterminae 245 Rogadinae sensu stricto 246 Alysioid subcomplex including Braconinae 250 Alysiinae and Opiinae 250 Alysiinae 251 General Alysiinae biology 251 Alysiini 253 Dacnusini 255 Opiinae 256 Braconinae 260 Exothecinae 269 Gnamptodontinae (= Gnaptodontinae) 270 Telengaiinae 271 The non-cyclostomes 271 Sigalphoid complex 271 Agathidinae 272 Sigalphinae 275 Helconoid complex 278 Helconinae 279 Helconoid group incertae sedis 281 Blacinae 282 Acampsohelconinae 283 Macrocentrine subcomplex 284 Macrocentrinae 284 Charmontiinae 287 Amicrocentrinae 287 Xiphozelinae 288 Homolobinae 290 Microtypinae 292 Orgilinae 292 Euphoroid complex 294 Euphorinae 294 Cenocoeliinae 310 The microgastroids 311 Cardiochilinae 312 Cheloninae (including Adeliini) 315 Dirrhopinae 319 Ichneutinae 320 Khoikhoiinae 322 Mendesellinae 322 Microgastrinae 322 Miracinae 335 Unplaced subfamilies 335 Masoninae 335 Meteorideinae 337 13 Phylogeny and Systematics of The Ichneumonidae 341 History of ichneumonid classification 342 Henry Townes (1913–90) and his idiosyncratic nomenclature 344 The extinct subfamilies 344 Tanychorinae† 344 Palaeoichneumoninae† 346 Labenopimplinae† 348 Pherombinae† 349 Townesitinae† 349 The xoridiformes 349 Xoridinae 349 The labeniformes 353 Labeninae 353 Groteini 355 Labenini 355 Poecilocryptini 356 The pimpliformes 356 Acaenitinae 356 Collyriinae 359 Cylloceriinae 360 Diacritinae 360 Diplazontinae 361 Orthocentrinae (= Helictinae) 366 Pimplinae 367 Delomeristini 369 Ephialtini (= Pimplini of Townes) 369 Polysphincta group 371 Pimplini 373 Poemeniinae (= Neoxoridinae) 378 Poemeniini 378 Pseudorhyssini 378 Rodrigamini 378 Rhyssinae 379 The ichneumoniformes 383 Adelognathinae 383 Agriotypinae 385 Alomyinae 387 Cryptinae 388 Aptesini 391 Cryptini 391 Phygadeuontini 393 Ichneumoninae 394 The brachycyrtiformes 398 Brachycyrtinae 398 Claseinae (Clasinae) 398 Pedunculinae 399 The orthopelmatiformes 400 Orthopelmatinae 400 The ophioniformes 400 Lower ophioniformes 402 Banchinae 402 Lycorininae 406 Sisyrostolinae 407 Stilbopinae 407 Tryphoninae 411 Middle ophioniformes 416 Ctenopelmatinae 416 Mesochorinae 421 Metopiinae 422 Oxytorinae 424 Tatogastrinae 425 Tersilochinae (including Neorhacodinae and Phrudinae s.s.) 426 Higher ophioniformes 430 Anomaloninae 430 Campopleginae 432 Cremastinae 438 Hybrizontinae 439 Nesomesochorinae 442 Ophioninae 442 Unplaced subfamilies 445 Eucerotinae 445 Microleptinae 447 Part 3 Ecology and Diversity 451 14 Ecology 453 Adult diet 454 Host-feeding 454 Water sugar and pollen feeding 457 Fecundity 460 Voltinism and seasonality 462 Daily activity patterns 462 Diapause 463 Cold hardiness hibernation and overwintering 465 Coloration and thermoregulation 467 Biological control 467 Effect on host food consumption 471 Artificial diets 474 Artificial hosts 475 Use of alternative hosts 475 Hyperparasitism and kleptoparasitism 476 Predation 477 Pathogens 477 Transmission of host pathogens 479 Dispersal 480 Coloration and mimetic rings 480 Palatability and odours 481 Competition 482 Apparent competition 482 Host ranges of parasitoids 483 Parasitoid guilds and food webs 484 Evolution of host ranges and speciation 486 15 Local and Global Patterns In Diversity 489 Field research in the tropics and anomalous diversity 490 Estimation of global ichneumonoid species richness 492 Distribution related to climate and latitude 496 The nasty host hypothesis 497 Biogeography 503 Islands and their parasitoid faunas 505 Species accumulation curves 506 Altitudinal gradients 507 Estimating local species diversity 508 Ichneumonoidea as biodiversity indicators 510 Conservation 510 Effect of habitat degradation on ichneumonoid composition 511 Significance of cryptic species 511 16 Collecting and Rearing Ichneumonoidea 513 Field collecting adults 516 Pan traps 518 Sweep netting 519 Light trapping 521 Canopy fogging 521 Malaise traps 521 Rearings from wild-collected hosts 523 Rearing leaf rollers and tiers 524 Substrate rearings 524 Culturing 524 Mating in captivity 525 Mass rearing 525 Mounting specimens for taxonomic study 526 Preparing specimens from alcohol storage 526 Direct pinning 527 Side gluing 527 Card rectangles and card points 527 Secondary staging 528 Labelling 528 Preserving specimens for DNA analysis 528 Packaging and posting specimens to other workers 530 17 Epilogue 533 Phylogenetic questions 534 Host and parasitism questions 534 Physiological questions 535 Ecological questions 536 Glossary 539 References 547 Author index 633 General index 653 Host index 659 Ichneumonoid genus tribe and subfamily index 665 Ichneumonoidea species index 677 Color Plate Sections Are Inserted Between Pages Noted Below First 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)