Braconid and Ichneumonid Parasitoid Wasps
-15%
portes grátis
Braconid and Ichneumonid Parasitoid Wasps
Biology, Systematics, Evolution and Ecology
Quicke, Donald L. J.
John Wiley and Sons Ltd
01/2015
704
Dura
Inglês
9781118907054
15 a 20 dias
1604
Descrição não disponível.
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 ooegenesis 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)
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 ooegenesis 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)
Este título pertence ao(s) assunto(s) indicados(s). Para ver outros títulos clique no assunto desejado.
Ichneumonidae; Braconidae; Classification; Systematics; Morphology; Evolution; Biological control; Development; Ecology; Biodiversity
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 ooegenesis 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)
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 ooegenesis 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)
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