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Tables of Contents for Parasitic Wasps
Chapter/Section Title
Page #
Page Count
Preface
xv
2
Acknowledgements
xvii
1 Introduction to the parasitic Hymenoptera
1
18
1.1 Introduction
1
2
1.2 Parasitoidism among other insects
3
4
1.3 A brief evolutionary history and phylogeny of the Hymenoptera
7
1
1.4 Origin of parasitism in the Hymenoptera
8
2
1.5 Oviposition: a crucial behaviour
10
1
1.6 Transition to secondary phytophagy
11
1
1.7 Transition to cecidogenesis
12
1
1.8 Evolution of inquilinism
13
1
1.9 Fig wasps
14
1
1.10 Evolution of parasitism of adults
15
1
1.11 Evolution of predatory relationships
16
1
1.12 Transition to aculeate strategy and to sociality
17
1
1.13 Secondary evolution of parasitism
18
1
2 Sex and genetics
19
48
2.1 Haplodiploidy and sex determination
19
1
2.2 Complementary sex determination (CSD)
19
2
2.3 Non-CSD systems
21
2
2.4 Diploid males
23
2
2.5 Polyploidy
25
1
2.6 Gynandromorphs and mosaics
26
1
2.7 Cytogenetics and karyotypic variation
26
3
2.8 Recombination rates
29
1
2.9 Genome size
30
1
2.10 Genomic molecular organization
30
1
2.11 Retroelements
31
1
2.12 Dosage compensation
32
1
2.13 Genetic diversity
33
3
2.14 Relationship between hosts and heterozygosity
36
1
2.15 Heritability of behaviours and life history parameters
37
1
2.16 Mating strategy and virginity
38
2
2.17 Effect of mating status on oviposition behaviour
40
1
2.18 Protandry
40
1
2.19 Sperm competition
40
2
2.20 Sex ratio
42
1
2.21 Sex allocation depending on host type and size
43
1
2.22 Sib-sib, mother-son mating and local mate competition
43
2
2.23 Determination of primary sex ratio
45
1
2.24 Signs of fertilization
46
1
2.25 Sex ratio in gregarious broods
47
2
2.26 Heteronmy and heterotrophy
49
1
2.27 Cytoplasmic incompatibility
50
1
2.28 Parthenogenesis - thelytoky and deuterotoky
51
1
2.29 Cytogenetics of thelytoky
52
2
2.30 Gene flow between thelytokous and arrhenotokous strains
54
1
2.31 Thelytoky resulting from crossing strains
55
1
2.32 Deuterotoky
55
1
2.33 Sex-ratio distorters - B-chromosomes and other factors
56
1
2.34 Sex-ratio distorters - Wolbachia, Arsenophonus and others
57
3
2.35 Long-term effects of Wolbachia-mediated thelytoky
60
1
2.36 Horizontal transmission of Wolbachia
61
1
2.37 Thelytoky and biocontrol
61
1
2.38 Sex determination and biocontrol
61
1
2.39 Genetic markers and mutations
62
1
2.40 Mutation rate
63
1
2.41 Genetic transformation
63
1
2.42 Resistance to pesticides and use in integrated pest management (IPM)
64
2
2.43 Population size and biocontrol success
66
1
3 Life history strategy: idiobionts and koinobionts
67
12
3.1 Introduction
67
1
3.2 Ectoparasitism vs. endoparasitism
68
1
3.3 Concealed vs. exposed hosts
69
1
3.4 Generalist vs. specialist
69
2
3.5 Synovigeny vs. pro-ovigeny
71
1
3.6 Oosorption
71
1
3.7 Adult longevity
71
1
3.8 Larval developmental period
72
1
3.9 Relative sizes of parasitoid and host
72
1
3.10 Host paralysis
73
1
3.11 Host-feeding
73
1
3.12 Diurnal vs. nocturnal
74
1
3.13 Egg and pupal parasitism
74
1
3.14 Egg-larval and egg-pupal parasitism
75
1
3.15 Offspring sex determination
76
3
4 Preimaginal development: from gametogenesis to syngamy
79
23
4.1 Spermatogenesis
79
1
4.2 Structure and ultrastructure of mature sperm
80
2
4.3 Sperm polymorphism
82
2
4.4 Spermatophores and sperm bundles
84
1
4.5 Insemination potential
85
1
4.6 Oogenesis
86
3
4.7 Meiosis
89
1
4.8 Accessory nuclei
89
1
4.9 Egg maturation
90
2
4.10 Chorion
92
2
4.11 Different egg types
94
2
4.12 Hydropic and anhydropic eggs
96
1
4.13 Sticky eggs
97
1
4.14 Anchored eggs
98
1
4.15 Respiratory adaptations of eggs
99
1
4.16 The micropyle and fertilization
99
1
4.17 Polyspermy
100
1
4.18 Oosorption
101
1
4.19 Egg secretions
101
1
5 Preimaginal development: from embryo initiation to pupa
102
46
5.1 Initiation of embryogenesis
102
2
5.2 Embryology
104
1
5.3 Polyembryony
105
1
5.4 Embryonic membranes
106
2
5.5 Teratocytes
108
2
5.6 Hatching
110
1
5.7 General larval morphology
111
3
5.8 Planidial larvae
114
1
5.9 Precocious or killer larvae
115
1
5.10 Number of larval instars
116
2
5.11 Larval feeding and digestive system
118
4
5.12 Larval dietary physiology and metabolism
122
3
5.13 Sequestering unusual compounds and metals
125
2
5.14 Larval excretory systems and products
127
1
5.15 In vitro rearing of parasitic wasp larvae
128
3
5.16 Larval respiration
131
2
5.17 Larval endocrinology and endocrine system
133
2
5.18 Diapause
135
1
5.19 Ectoparasitic vs. endoparasitic larval features
136
2
5.20 Cold tolerance
138
1
5.21 Larval movements
138
1
5.22 Protective behaviours of larvae
138
1
5.23 Larval competition
139
1
5.24 Silk and salivary glands
139
1
5.25 Cocoons
140
1
5.26 Silk cocoons and silk
140
2
5.27 Non-silk cocoons
142
1
5.28 Host mummification
143
2
5.29 Imaginal bud development
145
1
5.30 Timing of metamorphosis
145
1
5.31 Developmental abnormalities
146
1
5.32 Pupae
146
1
5.33 Pupation site
146
2
6 Adult morphology and adaptations
148
73
6.1 Reproductive systems
148
1
6.2 Male internal reproductive system
149
1
6.3 Seminal vesicle
149
1
6.4 Accessory glands
150
1
6.5 Male external genitalia
151
3
6.6 Female genital system
154
1
6.7 Ovaries and oviducts
155
3
6.8 Calyx gland
158
1
6.9 Colleterial, accessory or uterus glands
158
1
6.10 Spermathecal structure
159
2
6.11 Spermathecal function
161
2
6.12 Venom apparatus
163
2
6.13 Dufour's gland
165
1
6.14 Ovipositor structure
166
6
6.15 Mechanics of oviposition
172
3
6.16 Long ovipositors
175
1
6.17 Ovipositor guides and ovipositor buckling
176
1
6.18 Ovipositor steering mechanisms
177
3
6.19 Telescopic and concealed ovipositor mechanisms
180
2
6.20 Valvilli
182
1
6.21 Miscellaneous ovipositor specializations
183
1
6.22 Sensory systems
184
1
6.23 Antennae
184
2
6.24 Antennal sensilla
186
1
6.25 Placoid or multiporous plate sensilla
187
1
6.26 Gustatory sensilla
188
1
6.27 Ovipositor sense organs
189
1
6.28 Ovipositor sheath sensilla
190
1
6.29 Cerci
190
1
6.30 Eyes and ocelli
191
1
6.31 Other sense organs
191
2
6.32 Alimentary canal
193
1
6.33 Destination of dietary heavy metal contaminants
194
1
6.34 Pigments and coloration
195
1
6.35 Circulatory systems
196
1
6.36 Skeletomuscular system
196
1
6.37 Ferritin
197
1
6.38 Wings
198
3
6.39 Flight
201
1
6.40 Brachyptery and aptery
201
2
6.41 Grooming structures and behaviour
203
2
6.42 Exocrine scent glands
205
2
6.43 Antennal glands
207
1
6.44 Defensive odours
207
2
6.45 Cuticular hydrocarbons and chemical mimicry
209
1
6.46 Adult defensive structures
210
2
6.47 Modifications of the mouthparts
212
1
6.48 Adaptations for grasping and manipulating hosts
212
2
6.49 Adaptations associated with emergence
214
1
6.50 Adaptations to parasitism of hosts within wood
215
1
6.51 Adaptations for parasitoidism of adult insects
216
1
6.52 Adaptations to nocturnal activity
217
1
6.53 Aquatic parasitic wasps
218
1
6.54 Adaptations of fig wasps
219
2
7 Physiological interactions of parasitic wasps and their hosts
221
35
7.1 Introduction
221
1
7.2 Venoms
221
2
7.3 Specificity of venom action
223
1
7.4 Paralysing venoms
224
1
7.5 Neurobiological actions and chemistry of paralysing venoms
225
2
7.6 Effects of non-paralysing venoms on host development and physiology
227
1
7.7 Chemistry of non-paralysing venoms
228
1
7.8 Host castration and pseudoparasitism
229
3
7.9 Endocrine interactions
232
1
7.10 Symbiotic viruses, virus-like particles and fungi
233
1
7.11 Polydnaviruses
234
4
7.12 Physiological effects of polydnaviruses
238
2
7.13 Other viruses
240
1
7.14 Virus-like particles
241
1
7.15 Symbiotic yeasts
242
1
7.16 Stimulation of koinobiont development
243
1
7.17 Teratocyte function
243
1
7.18 Physiologically active larval secretions
244
2
7.19 Encapsulation and other host defences
246
3
7.20 Non-encapsulating host defences
249
1
7.21 Use of immunologically favourable sites or host stages
250
1
7.22 Physiological suppression of supernumeraries
251
1
7.23 Effects on host behaviour
252
1
7.24 Effects on host food consumption
253
2
7.25 Host relationships of parasitic wasps at family level
255
1
8 Adult behaviour
256
38
8.1 Host location
256
1
8.2 Host frass
257
1
8.3 Honeydew and other aphid secretions
258
1
8.4 Trail following
258
1
8.5 Host pheromones
259
2
8.6 The role of silk as a source of kairomones
261
1
8.7 Host-created vibrations
261
1
8.8 Temperature
262
1
8.9 Visual cues
262
1
8.10 Host plants and tritrophic effects
263
2
8.11 Oviposition stimuli
265
1
8.12 Kleptoparasitism
265
1
8.13 Host acceptance
266
1
8.14 Discrimination and superparasitism
266
3
8.15 Host marking
269
1
8.16 Multiparasitism
270
2
8.17 Learning
272
1
8.18 Adult diet
273
1
8.19 Host-feeding
274
3
8.20 Feeding tubes
277
1
8.21 Other types of food
277
1
8.22 Oviposition site
278
1
8.23 Phoresy as adults
279
1
8.24 Aggregations and hibernation of parasitic wasps
279
1
8.25 Adult diapause
280
1
8.26 Daily activity patterns
281
1
8.27 Host guarding, manipulation and maternal care
281
2
8.28 Sex pheromones
283
4
8.29 Courtship and mating
287
1
8.30 Mate choice
288
1
8.31 Male mating swarms
289
1
8.32 Post-copulation courtship
290
1
8.33 Role of courtship in species recognition
290
1
8.34 Polyandry
291
2
8.35 Male aggression
293
1
9 Non-physiological host defence strategies
294
8
9.1 Physical barriers
294
2
9.2 Behavioural defence
296
2
9.3 Host escape responses
298
1
9.4 Chemical defence
299
1
9.5 Dangerous hosts
299
1
9.6 Association with ants
300
1
9.7 Ecological factors
301
1
10 Ecology and diversity of parasitic wasps
302
27
10.1 Introduction
302
1
10.2 Parasitoid guilds
302
2
10.3 Host stage attacked
304
1
10.4 Life expectancy
304
1
10.5 Fecundity
305
2
10.6 Effects of host plants on parasitism success
307
1
10.7 Parasitoid species richness and diversity
308
3
10.8 Host ranges and parasitoid loads
311
1
10.9 Competition
312
1
10.10 Factors affecting parasitoid diversity
313
1
10.11 Effects of host niche on parasitoid diversity
314
1
10.12 `Nasty host' hypothesis - effects of secondary plant substances
315
2
10.13 Food webs and trophic interactions
317
1
10.14 Predators
317
2
10.15 Pathogens
319
1
10.16 Viruses
320
1
10.17 Microsporidians
320
1
10.18 Neogregarines
321
1
10.19 Nematodes
321
1
10.20 Fungi
322
1
10.21 Epizootiology
322
1
10.22 Coloration and mimicry
323
2
10.23 Factors affecting the success of biocontrol introductions
325
1
10.24 Succession and colonization
326
1
10.25 Dispersal and labelling of parasitoids
327
2
11 Parasitic wasp phylogeny and taxonomy
329
26
11.1 Introduction
329
1
11.2 Phylogenetic hypotheses
330
7
11.3 Relationships at family level
337
2
11.4 Morphological characters
339
1
11.5 The fossil record and extinct families of parasitic Hymenoptera
340
2
11.6 Molecular phylogenetic studies
342
2
11.7 Speciation
344
1
11.8 Effect of host and environment on wasp morphology
345
1
11.9 Evidence for genetic isolation
346
1
11.10 Crossing experiments
347
1
11.11 Sibling and cryptic species
348
1
11.12 Taxonomic implications of thelytoky
348
1
11.13 Molecular tools at the species level
349
1
11.14 Species discrimination and identification
350
2
11.15 Sex associations
352
1
11.16 Importance of correct identification in biocontrol
352
1
11.17 Specimen preparation and vouchers
353
2
Appendix A Parasitic Hymenoptera Frequently studied in culture
355
9
Appendix B Utilization of host groups by different families of parasitic Hymenoptera
364
3
Glossary
367
13
References
380
81
Index
461