The Insects/ P.J. Gullan and P.S. Cranston

List of colour plates ix List of boxes xiii Preface to the fifth edition xv Preface to the fourth edition xvii Preface to the third edition xix Preface to the second edition xxi Preface and acknowledgments for first edition xxiii About the companion website xxv 1 THE IMPORTANCE DIVERSITY AND CONSERVATION OF INSECTS 1 1.1 What is entomology? 2 1.2 The importance of insects 2 1.3 Insect biodiversity 6 1.4 Naming and classification of insects 10 1.5 Insects in popular culture and commerce 11 1.6 Culturing insects 13 1.7 Insect conservation 14 1.8 Insects as food 20 Further reading 25 2 EXTERNAL ANATOMY 26 2.1 The cuticle 27 2.2 Segmentation and tagmosis 33 2.3 The head 35 2.4 The thorax 45 2.5 The abdomen 52 Further reading 55 3 INTERNAL ANATOMY AND PHYSIOLOGY 56 3.1 Muscles and locomotion 57 3.2 The nervous system and co-ordination 63 3.3 The endocrine system and the function of hormones 66 3.4 The circulatory system 69 3.5 The tracheal system and gas exchange 73 3.6 The gut digestion and nutrition 77 3.7 The excretory system and waste disposal 86 3.8 Reproductive organs 90 Further reading 93 4 SENSORY SYSTEMS AND BEHAVIOUR 95 4.1 Mechanical stimuli 96 4.2 Thermal stimuli 105 4.3 Chemical stimuli 107 4.4 Insect vision 117 4.5 Insect behaviour 122 Further reading 124 5 REPRODUCTION 125 5.1 Bringing the sexes together 126 5.2 Courtship 128 5.3 Sexual selection 128 5.4 Copulation 131 5.5 Diversity in genitalic morphology 136 5.6 Sperm storage fertilization and sex determination 139 5.7 Sperm competition 140 5.8 Oviparity (egg-laying) 144 5.9 Ovoviviparity and viviparity 150 5.10 Other modes of reproduction 150 5.11 Physiological control of reproduction 153 Further reading 154 6 INSECT DEVELOPMENT AND LIFE HISTORIES 156 6.1 Growth 157 6.2 Life-history patterns and phases 158 6.3 Process and control of moulting 169 6.4 Voltinism 172 6.5 Diapause 173 6.6 Dealing with environmental extremes 174 6.7 Migration 178 6.8 Polymorphism and polyphenism 180 6.9 Age-grading 181 6.10 Environmental effects on development 183 Further reading 188 7 INSECT SYSTEMATICS: PHYLOGENY AND CLASSIFICATION 190 7.1 Systematics 191 7.2 The extant Hexapoda 201 7.3 Informal group Entognatha: Collembola (springtails) Diplura (diplurans) and Protura (proturans) 202 7.4 Class Insecta (true insects) 203 Further reading 224 8 INSECT EVOLUTION AND BIOGEOGRAPHY 227 8.1 Relationships of the Hexapoda to other Arthropoda 228 8.2 The antiquity of insects 229 8.3 Were the first insects aquatic or terrestrial? 236 8.4 Evolution of wings 238 8.5 Evolution of metamorphosis 241 8.6 Insect diversification 242 8.7 Insect biogeography 244 8.8 Insect evolution in the Pacific 245 Further reading 247 9 GROUND-DWELLING INSECTS 249 9.1 Insects of litter and soil 250 9.2 Insects and dead trees or decaying wood 260 9.3 Insects and dung 261 9.4 Insect carrion interactions 264 9.5 Insect fungal interactions 265 9.6 Cavernicolous insects 268 9.7 Environmental monitoring using ground-dwelling hexapods 268 Further reading 270 10 AQUATIC INSECTS 271 10.1 Taxonomic distribution and terminology 272 10.2 The evolution of aquatic lifestyles 275 10.3 Aquatic insects and their oxygen supplies 277 10.4 The aquatic environment 282 10.5 Environmental monitoring using aquatic insects 284 10.6 Functional feeding groups 285 10.7 Insects of temporary waterbodies 286 10.8 Insects of the marine intertidal and littoral zones 287 Further reading 288 11 INSECTS AND PLANTS 289 11.1 Coevolutionary interactions between insects and plants 291 11.2 Phytophagy (or herbivory) 293 11.3 Insects and plant reproductive biology 313 11.4 Insects that live mutualistically in specialized plant structures 318 Further reading 320 12 INSECT SOCIETIES 322 12.1 Subsociality in insects 323 12.2 Eusociality in insects 327 12.3 Inquilines and parasites of social insects 345 12.4 Evolution and maintenance of eusociality 348 12.5 Success of social insects 351 Further reading 353 13 INSECT PREDATION AND PARASITISM 354 13.1 Prey/host location 355 13.2 Prey/host acceptance and manipulation 361 13.3 Prey/host selection and specificity 364 13.4 Population biology predator/parasitoid and prey/host abundance 372 13.5 The evolutionary success of insect predation and parasitism 375 Further reading 376 14 INSECT DEFENCE 377 14.1 Defence by hiding 379 14.2 Secondary lines of defence 380 14.3 Mechanical defences 382 14.4 Chemical defences 384 14.5 Defence by mimicry 388 14.6 Collective defences in gregarious and social insects 392 Further reading 396 15 MEDICAL AND VETERINARY ENTOMOLOGY 397 15.1 Insects as causes and vectors of disease 398 15.2 Generalized disease cycles 399 15.3 Pathogens 399 15.4 Forensic entomology 413 15.5 Insect nuisance and phobia 414 15.6 Venoms and allergens 416 Further reading 417 16 PEST MANAGEMENT 418 16.1 Insects as pests 419 16.2 The effects of insecticides 425 16.3 Integrated pest management 428 16.4 Chemical control 429 16.5 Biological control 435 16.6 Host-plant resistance to insects 447 16.7 Physical control 451 16.8 Cultural control 451 16.9 Pheromones and other insect attractants 452 16.10 Genetic manipulation of insect pests 454 Further reading 455 17 INSECTS IN A CHANGING WORLD 457 17.1 Models of change 458 17.2 Economically significant insects under climate change 463 17.3 Implications of climate change for insect biodiversity and conservation 467 17.4 Global trade and insects 468 Further reading 473 18 METHODS IN ENTOMOLOGY:COLLECTING PRESERVATIONCURATION AND IDENTIFICATION 474 18.1 Collection 475 18.2 Preservation and curation 478 18.3 Identification 488 Further reading 491 TAXOBOXES 493 1 Entognatha: non-insect hexapods(Collembola Diplura and Protura) 493 2 Archaeognatha (Microcoryphia; archaeognathans or bristletails) 495 3 Zygentoma (silverfish) 496 4 Ephemeroptera (mayflies) 497 5 Odonata (damselflies anddragonflies) 498 6 Plecoptera (stoneflies) 500 7 Dermaptera (earwigs) 500 8 Zoraptera (zorapterans or angelinsects) 501 9 Orthoptera (grasshoppers locustskatydids and crickets) 502 10 Embioptera (Embiidina Emboidea;embiopterans or webspinners) 503 11 Phasmatodea (phasmids stick-insects or walking sticks) 503 12 Grylloblattodea (Grylloblattaria or Notoptera; grylloblattids ice crawlers or rock crawlers) 504 13 Mantophasmatodea (heelwalkers) 505 14 Mantodea (mantids mantises or praying mantids) 506 15 Blattodea: roach families (cockroaches or roaches) 507 16 Blattodea: epifamily Termitoidae (former order Isoptera; termites white ants ) 508 17 Psocodea: Psocoptera (bark lice and book lice) 509 18 Psocodea: Phthiraptera (chewing lice and sucking lice) 510 19 Thysanoptera (thrips) 511 20 Hemiptera (bugs moss bugs cicadas leafhoppers planthoppers spittle bugs treehoppers aphids jumping plant lice scale insects and whiteflies) 512 21 Neuropterida: Neuroptera (lacewings owlflies and antlions) Megaloptera (alderflies dobsonflies and fishflies) and Raphidioptera (snakeflies) 514 22 Coleoptera (beetles) 516 23 Strepsiptera (strepsipterans) 517 24 Diptera (true flies) 519 25 Mecoptera (hangingflies scorpionflies and snowfleas) 520 26 Siphonaptera (fleas) 521 27 Trichoptera (caddisflies) 522 28 Lepidoptera (butterflies and moths) 523 29 Hymenoptera (ants bees wasps sawflies and wood wasps) 524 Glossary 526 References 555 Index 563 Appendix: A reference guide to orders 589