The Insects An Outline of Entomology TIA01 5/20/04 4:37 PM Page i TIA01 5/20/04 4:37 PM Page ii Third Edition The Insects An Outline of Entomology P.J. Gullan and P.S. Cranston Department of Entomology, University of California, Davis, USA With illustrations by K. Hansen McInnes TIA01 6/3/04 2:10 PM Page iii © 2005 by Blackwell Publishing Ltd Previous editions © P.J. Gullan and P.S. Cranston 350 Main Street, Malden, MA 02148-5020, USA 108 Cowley Road, Oxford OX4 1JF, UK 550 Swanston Street, Carlton, Victoria 3053, Australia The right of P.J. Gullan and P.S. Cranston to be identified as the Authors of this Work has been asserted in accordance with the UK Copyright, Designs, and Patents Act 1988. All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, mechanical, photocopying, recording or otherwise, except as permitted by the UK Copyright, Designs, and Patents Act 1988, without the prior permission of the publisher. First published 1994 by Chapman & Hall Second edition published 2000 by Blackwell Publishing Ltd Third edition published 2005 Library of Congress Cataloging-in-Publication Data Gullan, P.J. The insects: an outline of entomology/P.J. Gullan & P.S. Cranston; with illustrations by K. Hansen McInnes. – 3rd ed. p. cm. Includes bibliographical references and index. ISBN 1-4051-1113-5 (hardback: alk. paper) 1. Insects. I. Cranston, P.S. II. Title. QL463.G85 2004 595.7–dc22 2004000124 A catalogue record for this title is available from the British Library. Set in 9/11pt Photina by Graphicraft Limited, Hong Kong Printed and bound in the United Kingdom by The Bath Press Cover and text illustrations © Karina Hansen McInnes For further information on Blackwell Publishing, visit our website: http://www.blackwellpublishing.com TIA01 5/20/04 4:37 PM Page iv List of color plates, viii List of boxes, x Preface to the third edition, xii Preface to the second edition, xiv Preface and acknowledgments for first edition, xvi 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, 4 1.4 Naming and classification of insects, 8 1.5 Insects in popular culture and commerce, 9 1.6 Insects as food, 10 1.7 Insect conservation, 13 Further reading, 20 2 EXTERNAL ANATOMY, 21 2.1 The cuticle, 22 2.2 Segmentation and tagmosis, 28 2.3 The head, 30 2.4 The thorax, 38 2.5 The abdomen, 45 Further reading, 48 3 INTERNAL ANATOMY AND PHYSIOLOGY, 49 3.1 Muscles and locomotion, 50 3.2 The nervous system and co-ordination, 56 3.3 The endocrine system and the function of hormones, 59 3.4 The circulatory system, 61 3.5 The tracheal system and gas exchange, 65 3.6 The gut, digestion, and nutrition, 68 3.7 The excretory system and waste disposal, 77 3.8 Reproductive organs, 81 Further reading, 84 4 SENSORY SYSTEMS AND BEHAVIOR, 85 4.1 Mechanical stimuli, 86 4.2 Thermal stimuli, 94 4.3 Chemical stimuli, 96 4.4 Insect vision, 105 4.5 Insect behavior, 109 Further reading, 111 5 REPRODUCTION, 113 5.1 Bringing the sexes together, 114 5.2 Courtship, 117 5.3 Sexual selection, 117 5.4 Copulation, 118 5.5 Diversity in genitalic morphology, 123 5.6 Sperm storage, fertilization, and sex determination, 128 5.7 Sperm competition, 128 5.8 Oviparity (egg-laying), 129 5.9 Ovoviviparity and viviparity, 135 5.10 Atypical modes of reproduction, 135 5.11 Physiological control of reproduction, 138 Further reading, 139 6 INSECT DEVELOPMENT AND LIFE HISTORIES, 141 6.1 Growth, 142 6.2 Life-history patterns and phases, 143 6.3 Process and control of molting, 153 6.4 Voltinism, 156 6.5 Diapause, 157 6.6 Dealing with environmental extremes, 158 6.7 Migration, 161 6.8 Polymorphism and polyphenism, 163 CONTENTS TIA01 5/20/04 4:37 PM Page v vi Contents 6.9 Age-grading, 164 6.10 Environmental effects on development, 166 6.11 Climate and insect distributions, 171 Further reading, 175 7 INSECT SYSTEMATICS: PHYLOGENY AND CLASSIFICATION, 177 7.1 Phylogenetics, 178 7.2 The extant Hexapoda, 180 7.3 Protura (proturans), Collembola (springtails), and Diplura (diplurans), 183 7.4 Class Insecta (true insects), 184 Further reading, 199 8 INSECT BIOGEOGRAPHY AND EVOLUTION, 201 8.1 Insect biogeography, 202 8.2 The antiquity of insects, 203 8.3 Were the first insects aquatic or terrestrial? 208 8.4 Evolution of wings, 208 8.5 Evolution of metamorphosis, 211 8.6 Insect diversification, 213 8.7 Insect evolution in the Pacific, 214 Further reading, 216 9 GROUND-DWELLING INSECTS, 217 9.1 Insects of litter and soil, 218 9.2 Insects and dead trees or decaying wood, 221 9.3 Insects and dung, 223 9.4 Insect–carrion interactions, 224 9.5 Insect–fungal interactions, 226 9.6 Cavernicolous insects, 229 9.7 Environmental monitoring using ground- dwelling hexapods, 229 Further reading, 237 10 AQUATIC INSECTS, 239 10.1 Taxonomic distribution and terminology, 240 10.2 The evolution of aquatic lifestyles, 240 10.3 Aquatic insects and their oxygen supplies, 241 10.4 The aquatic environment, 245 10.5 Environmental monitoring using aquatic insects, 248 10.6 Functional feeding groups, 249 10.7 Insects of temporary waterbodies, 250 10.8 Insects of the marine, intertidal, and littoral zones, 251 Further reading, 261 11 INSECTS AND PLANTS, 263 11.1 Coevolutionary interactions between insects and plants, 265 11.2 Phytophagy (or herbivory), 265 11.3 Insects and plant reproductive biology, 281 11.4 Insects that live mutualistically in specialized plant structures, 286 Further reading, 297 12 INSECT SOCIETIES, 299 12.1 Subsociality in insects, 300 12.2 Eusociality in insects, 304 12.3 Inquilines and parasites of social insects, 318 12.4 Evolution and maintenance of eusociality, 320 12.5 Success of eusocial insects, 324 Further reading, 324 13 INSECT PREDATION AND PARASITISM, 327 13.1 Prey/host location, 328 13.2 Prey/host acceptance and manipulation, 334 13.3 Prey/host selection and specificity, 338 13.4 Population biology – predator/parasitoid and prey/host abundance, 345 13.5 The evolutionary success of insect predation and parasitism, 347 Further reading, 353 14 INSECT DEFENSE, 355 14.1 Defense by hiding, 356 14.2 Secondary lines of defense, 359 14.3 Mechanical defenses, 360 14.4 Chemical defenses, 360 14.5 Defense by mimicry, 365 14.6 Collective defenses in gregarious and social insects, 369 Further reading, 373 15 MEDICAL AND VETERINARY ENTOMOLOGY, 375 15.1 Insect nuisance and phobia, 376 15.2 Venoms and allergens, 376 15.3 Insects as causes and vectors of disease, 377 15.4 Generalized disease cycles, 378 15.5 Pathogens, 379 15.6 Forensic entomology, 388 Further reading, 393 16 PEST MANAGEMENT, 395 16.1 Insects as pests, 396 TIA01 5/20/04 4:37 PM Page vi 16.2 The effects of insecticides, 400 16.3 Integrated pest management, 403 16.4 Chemical control, 404 16.5 Biological control, 407 16.6 Host-plant resistance to insects, 417 16.7 Physical control, 420 16.8 Cultural control, 420 16.9 Pheromones and other insect attractants, 421 16.10 Genetic manipulation of insect pests, 422 Further reading, 423 17 METHODS IN ENTOMOLOGY: COLLECTING, PRESERVATION, CURATION, AND IDENTIFICATION, 427 17.1 Collection, 428 17.2 Preservation and curation, 431 17.3 Identification, 440 Further reading, 443 Glossary, 445 References, 469 Index, 477 Appendix: A reference guide to orders, 499 Color plates fall between pp. 14 and 15 Contents vii TIA01 5/20/04 4:37 PM Page vii PLATE 1 1.1 An atlas moth, Attacus atlas (Lepidoptera: Saturniidae), which occurs in southern India and south-east Asia, is one of the largest of all lepidopterans, with a wingspan of about 24 cm and a larger wing area than any other moth (P.J. Gullan). 1.2 A violin beetle, Mormolyce phyllodes (Coleoptera: Carabidae), from rainforest in Brunei, Borneo (P.J. Gullan). 1.3 The moon moth, Argema maenas (Lepidoptera: Saturniidae), is found in south-east Asia and India; this female, from rainforest in Borneo, has a wingspan of about 15 cm (P.J. Gullan). 1.4 The mopane emperor moth, Imbrasia belina (Lepidoptera: Saturniidae), from the Transvaal in South Africa (R. Oberprieler). 1.5 A “worm” or “phane” – the caterpillar of Imbrasia belina – feeding on the foliage of Schotia brachypetala, from the Transvaal in South Africa (R. Oberprieler). 1.6 A dish of edible water bugs, Lethocerus indicus (Hemiptera: Belostomatidae), on sale at a market in Lampang Province, Thailand (R.W. Sites). PLATE 2 2.1 Food insects at a market stall in Lampang Province, Thailand, displaying silk moth pupae (Bombyx mori), beetle pupae, adult hydrophiloid beetles, and water bugs, Lethocerus indicus (R.W. Sites). 2.2 Adult Richmond birdwing (Troides richmondia) butterfly and cast exuvial skin on native pipevine (Pararistolochia sp.) host (see p. 15) (D.P.A. Sands). 2.3 A bush coconut or bloodwood apple gall of Cystococcus pomiformis (Hemiptera: Eriococcidae), cut open to show the cream-colored adult female and her numerous, tiny nymphal male offspring covering the gall wall (P.J. Gullan). 2.4 Close-up of the second-instar male nymphs of Cystococcus pomiformis feeding from the nutritive tissue lining the cavity of the maternal gall (see p. 12) (P.J. Gullan). 2.5 Adult male scale insect of Melaleucococcus phacelopilus (Hemiptera: Margarodidae), showing the setiferous antennae and the single pair of wings (P.J. Gullan). 2.6 A tropical butterfly, Graphium antiphates itamputi (Lepidoptera: Papilionidae), from Borneo, obtaining salts by imbibing sweat from a training shoe (refer to Box 5.2) (P.J. Gullan). PLATE 3 3.1 A female katydid of an undescribed species of Austrosalomona (Orthoptera: Tettigoniidae), from northern Australia, with a large spermatophore attached to her genital opening (refer to Box 5.2) (D.C.F. Rentz). 3.2 Pupa of a Christmas beetle, Anoplognathus sp. (Coleoptera: Scarabaeidae), removed from its pupation site in the soil in Canberra, Australia (P.J. Gullan). 3.3 Egg mass of Tenodera australasiae (Mantodea: Mantidae) with young mantid nymphs emerging, from Queensland, Australia (refer to Box 13.2) (D.C.F. Rentz). 3.4 Eclosing (molting) adult katydid of an Elephantodeta species (Orthoptera: Tettigoniidae), from the Northern Territory, Australia (D.C.F. Rentz). 3.5 Overwintering monarch butterflies, Danaus plexippus (Lepidoptera: Nymphalidae), from Mill Valley in California, USA (D.C.F. Rentz). 3.6 A fossilized worker ant of Pseudomyrmex oryctus (Hymenoptera: Formicidae) in Dominican amber from the Oligocene or Miocene (P.S. Ward). 3.7 A diversity of flies (Diptera), including calliphorids, are attracted to the odor of this Australian phalloid fungus, Anthurus archeri, which produces a foul-smelling slime containing spores that are LIST OF COLOR PLATES TIA01 5/20/04 4:37 PM Page viii consumed by the flies and distributed after passing through the insects’ guts (P.J. Gullan). PLATE 4 4.1 A tree trunk and under-branch covered in silk galleries of the webspinner Antipaluria urichi (Embiidina: Clothodidae), from Trinidad (refer to Box 9.5) (J.S. Edgerly-Rooks). 4.2 A female webspinner of Antipaluria urichi defending the entrance of her gallery from an approaching male, from Trinidad (J.S. Edgerly-Rooks). 4.3 An adult stonefly, Neoperla edmundsi (Plecoptera: Perlidae), from Brunei, Borneo (P.J. Gullan). 4.4 A female thynnine wasp of Zaspilothynnus trilobatus (Hymenoptera: Tiphiidae) (on the right) compared with the flower of the sexually deceptive orchid Drakaea glyptodon, which attracts pollinating male wasps by mimicking the female wasp (see p. 282) (R. Peakall). 4.5 A male thynnine wasp of Neozeloboria cryptoides (Hymenoptera: Tiphiidae) attempting to copulate with the sexually deceptive orchid Chiloglottis trapeziformis (R. Peakall). 4.6 Pollination of mango flowers by a flesh fly, Australopierretia australis (Diptera: Sarcophagidae), in northern Australia (D.L. Anderson). 4.7 The wingless adult female of the whitemarked tussock moth, Orgyia leucostigma (Lepidoptera: Lymantriidae), from New Jersey, USA (D.C.F. Rentz). PLATE 5 5.1 Mealybugs of an undescribed Planococcus species (Hemiptera: Pseudococcidae) on an Acacia stem attended by ants of a Polyrhachis species (Hymenoptera: Formicidae), coastal Western Australia (P.J. Gullan). 5.2 A camouflaged late-instar caterpillar of Plesanemma fucata (Lepidoptera: Geometridae) on a eucalypt leaf in eastern Australia (P.J. Gullan). 5.3 A female of the scorpionfly Panorpa communis (Mecoptera: Panorpidae) from the UK (P.H. Ward). 5.4 The huge queen termite (approximately 7.5 cm long) of Odontotermes transvaalensis (Isoptera: Termitidae: Macrotermitinae) surrounded by her king (mid front), soldiers, and workers, from the Transvaal in South Africa ( J.A.L. Watson). 5.5 A parasitic Varroa mite (see p. 320) on a pupa of the bee Apis cerana (Hymenoptera: Apidae) in a hive from Irian Jaya, New Guinea (D.L. Anderson). 5.6 An adult moth of Utetheisa ornatrix (Lepidoptera: Arctiidae) emitting defensive froth containing pyrrolizidine alkaloids that it sequesters as a larva from its food plants, legumes of the genus Crotalaria (T. Eisner). 5.7 A snake-mimicking caterpillar of the spicebush swallowtail, Papilio troilus (Lepidoptera: Papilionidae), from New Jersey, USA (D.C.F. Rentz). PLATE 6 6.1 The cryptic adult moths of four species of Acronicta (Lepidoptera: Noctuidae): A. alni, the alder moth (top left); A. leporina, the miller (top right); A. aceris, the sycamore (bottom left); and A. psi, the grey dagger (bottom right) (D. Carter and R.I. Vane-Wright). 6.2 Aposematic or mechanically protected caterpillars of the same four species of Acronicta: A. alni (top left); A. leporina (top right); A. aceris (bottom left); and A. psi (bottom right); showing the divergent appearance of the larvae compared with their drab adults (D. Carter and R.I. Vane-Wright). 6.3 A blister beetle, Lytta polita (Coleoptera: Meloidae), reflex-bleeding from the knee joints; the hemolymph contains the toxin cantharidin (sections 14.4.3 & 15.2.2) (T. Eisner). 6.4 One of Bates’ mimicry complexes from the Amazon Basin involving species from three different lepidopteran families – Methona confusa confusa (Nymphalidae: Ithomiinae) (top), Lycorea ilione ilione (Nymphalidae: Danainae) (second from top), Patia orise orise (Pieridae) (second from bottom), and a day-flying moth of Gazera heliconioides (Castniidae) (R.I. Vane- Wright). 6.5 An aposematic beetle of the genus Lycus (Coleoptera: Lycidae) on the flower spike of Cussonia (Araliaceae) from South Africa (P.J. Gullan). 6.6 A mature cottony-cushion scale, Icerya purchasi (Hemiptera: Margarodidae), with a fully formed ovisac, on the stem of a native host plant from Australia (P.J. Gullan). 6.7 Adult male gypsy moth, Lymantria dispar (Lepidoptera: Lymantriidae), from New Jersey, USA (D.C.F. Rentz). List of color plates ix TIA01 5/20/04 4:37 PM Page ix Box 1.1 Collected to extinction? 16 Box 1.2 Tramp ants and biodiversity, 17 Box 1.3 Sustainable use of mopane worms, 19 Box 3.1 Molecular genetic techniques and their application to neuropeptide research, 60 Box 3.2 Tracheal hypertrophy in mealworms at low oxygen concentrations, 69 Box 3.3 The filter chamber of Hemiptera, 71 Box 3.4 Cryptonephric systems, 79 Box 4.1 Aural location of host by a parasitoid fly, 91 Box 4.2 The electroantennogram, 97 Box 4.3 Reception of communication molecules, 99 Box 4.4 Biological clocks, 106 Box 5.1 Courtship and mating in Mecoptera, 116 Box 5.2 Nuptial feeding and other “gifts”, 121 Box 5.3 Sperm precedence, 126 Box 5.4 Control of mating and oviposition in a blow fly, 130 Box 5.5 Egg-tending fathers – the giant water bugs, 132 Box 6.1 Molecular insights into insect development, 148 Box 6.2 Calculation of day-degrees, 168 Box 6.3 Climatic modeling for fruit flies, 174 Box 7.1 Relationships of the Hexapoda to other Arthropoda, 181 Box 9.1 Ground pearls, 222 Box 9.2 Non-insect hexapods (Collembola, Protura, and Diplura), 230 Box 9.3 Archaeognatha (bristletails) and Zygentoma (Thysanura; silverfish), 232 Box 9.4 Grylloblattodea (Grylloblattaria, Notoptera; grylloblattids, ice or rock crawlers), 233 Box 9.5 Embiidina or Embioptera (embiids, webspinners), 234 Box 9.6 Zoraptera, 234 Box 9.7 Dermaptera (earwigs), 235 Box 9.8 Blattodea (Blattaria; cockroaches, roaches), 236 Box 10.1 Ephemeroptera (mayflies), 252 Box 10.2 Odonata (damselflies and dragonflies), 253 Box 10.3 Plecoptera (stoneflies), 255 Box 10.4 Trichoptera (caddisflies), 255 Box 10.5 Diptera (true flies), 257 Box 10.6 Other aquatic orders, 258 Box 11.1 Induced defenses, 268 Box 11.2 The grape phylloxera, 276 Box 11.3 Salvinia and phytophagous weevils, 280 Box 11.4 Figs and fig wasps, 284 Box 11.5 Orthoptera (grasshoppers, locusts, katydids, and crickets), 289 Box 11.6 Phasmatodea (phasmatids, phasmids, stick-insects or walking sticks), 290 Box 11.7 Thysanoptera (thrips), 291 Box 11.8 Hemiptera (bugs, cicadas, leafhoppers, spittle bugs, planthoppers, aphids, jumping plant lice, scale insects, whiteflies), 292 Box 11.9 Psocoptera (booklice, barklice, or psocids), 294 Box 11.10 Coleoptera (beetles), 295 Box 11.11 Lepidoptera (butterflies and moths), 296 Box 12.1 The dance language of bees, 310 Box 12.2 Hymenoptera (bees, ants, wasps, sawflies, and wood wasps), 325 Box 12.3 Isoptera (termites), 326 Box 13.1 Viruses, wasp parasitoids, and host immunity, 337 Box 13.2 Mantodea (mantids), 348 Box 13.3 Mantophasmatodea (heel walkers), 349 Box 13.4 Neuropterida, or neuropteroid orders, 350 Box 13.5 Mecoptera (scorpionflies, hangingflies), 351 Box 13.6 Strepsiptera, 352 LIST OF BOXES TIA01 5/20/04 4:37 PM Page x [...]... that the species richness of insects is so great that, to a near approximation, all organisms can be considered to be insects Although dominant on land and in freshwater, few insects are found beyond the tidal limit of oceans In this opening chapter, we outline the significance of insects and discuss their diversity and classification and their roles in our economic and wider lives First, we outline the. .. the field of entomology and the role of entomologists, and then introduce the ecological functions of insects Next, we explore insect diversity, and then discuss how we name and classify this immense diversity Sections follow in which we consider past and some continuing cultural and economic aspects of insects, their aesthetic and tourism appeal, and their importance as foods for humans and animals... while feeding and, with numerous alate queens in the nest, can be mistaken for a queen and attacked and eaten by nurse ants Now that we understand the intricacies of the relationship, we can see that the well-meaning creation of reserves that lacked rabbits and excluded other grazers created vegetational and microhabitat changes that altered the dominance of ant species, to the detriment of the butterfly’s... OF INSECTS Charles Darwin inspecting beetles collected during the voyage of the Beagle (After various sources, especially Huxley & Kettlewell 1965 and Futuyma 1986.) TIC01 5/20/04 4:49 PM Page 2 2 The importance, diversity, and conservation of insects Curiosity alone concerning the identities and lifestyles of the fellow inhabitants of our planet justifies the study of insects Some of us have used insects. .. plant chemical barrier has been overcome Waves of specialization followed by breakthrough and radiation must have been a major factor in promoting the high species richness of phytophagous insects Another explanation for the high species numbers of insects is the role of sexual selection in the diversification of many insects The propensity of insects to become isolated in small populations (because of. .. item in central Angola, Africa (Larva after Santos Oliveira et al 1976.) caterpillars is high, with their protein content ranging from 45 to 80%, and they are a rich source of iron For instance, caterpillars are the most important source of animal protein in some areas of the Northern Province TIC01 5/20/04 4:49 PM Page 12 12 The importance, diversity, and conservation of insects of Zambia The edible caterpillars... (buckwheat), and control of alien plants plus limitation on human disturbance Southern Californian coastal dune systems are seriously endangered habitats, and management of this reserve for the El Segundo blue conserves other threatened species Land conservation for butterflies is not an indulgence of affluent southern Californians: the world’s largest butterfly, the Queen Alexandra’s birdwing (Ornithoptera alexandrae),... reintroduction of the large blue to England from continental Europe The larva of the large blue butterfly in England and on the European continent is an obligate predator in colonies of red ants belonging to species of Myrmica Larval large blues must enter a Myrmica nest, in which they feed on larval ants Similar predatory behavior, and/or tricking ants into feeding them as if they were the ants’ own brood,... expose them to other potential diversifying influences that enhance their species richness Interactions between certain groups of insects and other organisms, such as plants in the case of herbivorous insects, or hosts for parasitic insects, may promote the genetic diversification of eater and eaten These interactions are often called coevolutionary and are discussed in more detail in Chapters 11 and 13 The. .. with their surveillance and control The big-headed ant (Pheidole megaceph- ala), the long legged or yellow crazy ant (Anoplolepis longipes), the Argentine ant (Linepithema humile), the “electric” or little fire ant (Wasmannia auropunctata), and tropical fire ants (Solenopsis species) are considered the most serious of these ant pests Invasive ant behavior threatens biodiversity, especially on islands . The Insects An Outline of Entomology TIA01 5/20/04 4:37 PM Page i TIA01 5/20/04 4:37 PM Page ii Third Edition The Insects An Outline of Entomology P.J. Gullan and P.S. Cranston Department of. First, we outline the field of entomology and the role of ento- mologists, and then introduce the ecological functions of insects. Next, we explore insect diversity, and then discuss how we name and. past and some continuing cultural and economic aspects of insects, their aesthetic and tourism appeal, and their import- ance as foods for humans and animals. We conclude with a review of the