PJ Gullan and PS Cranston
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Trim size: 189mm x 246mm Gullan f01.tex V2 - 05/20/2014 1:46 P.M. Page ii Trim size: 189mm x 246mm Gullan f01.tex V2 - 05/20/2014 1:46 P.M. Page i THE INSECTS Trim size: 189mm x 246mm Gullan f01.tex V2 - 05/20/2014 1:46 P.M. Page ii Trim size: 189mm x 246mm Gullan f01.tex V2 - 05/20/2014 1:46 P.M. Page iii Fifth Edition THE INSECTS AN OUTLINE OF ENTOMOLOGY P.J. Gullan and P.S. Cranston Research School of Biology, The Australian National University, Canberra, Australia & Department of Entomology and Nematology, University of California, Davis, USA With illustrations by Karina H. McInnes Trim size: 189mm x 246mm Gullan f01.tex V2 - 05/20/2014 1:46 P.M. Page iv This edition first published 2014 © 2014 by John Wiley & Sons, Ltd Fourth edition published 2010 © 2010 by P.J. Gullan and P.S. Cranston Third edition published 2005 by Blackwell Publishing Ltd Second edition published 2000 by Blackwell Science First edition published 1994 by Chapman & Hall Registered office: John Wiley & Sons, Ltd, The Atrium, Southern Gate, Chichester, West Sussex, PO19 8SQ, UK Editorial offices: 9600 Garsington Road, Oxford, OX4 2DQ, UK The Atrium, Southern Gate, Chichester, West Sussex, PO19 8SQ, UK 111 River Street, Hoboken, NJ 07030-5774, USA For details of our global editorial offices, for customer services and for information about how to apply for permission to reusethe copyright material in this book please see our website at www.wiley.com/wiley-blackwell. The right of the author to be identified as the author of this work has been asserted in accordance with the UKCopyright, 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. Designations used by companies to distinguish their products are often claimed as trademarks. All brand names and product names used in this book are trade names, service marks, trademarks or registered trademarks of their respective owners. The publisher is not associated with any product or vendor mentioned in this book. Limit of Liability/Disclaimer of Warranty: While the publisher and author(s) have used their best efforts in preparing this book, they make no representations or warranties with respect to the accuracy or completeness of the contents of this book and specifically disclaim any implied warranties of merchantability or fitness for a particular purpose. It is sold on the understanding that the publisher is not engaged in rendering professional services and neither the publisher nor the author shall be liable for damages arising herefrom. If professional advice or other expert assistance is required, the services of a competent professional should be sought. Library of Congress Cataloging-in-Publication Data Gullan, P.J. The insects : an outline of entomology / P.J. Gullan and P.S. Cranston ; with illustrations by Karina H. McInnes. – Fifth edition. pages cm Includes bibliographical references and index. ISBN 978-1-118-84615-5 (cloth) 1. Insects. 2. Entomology. I. Cranston, P.S. II. Title. QL463.G85 2014 595.7– dc23 2014013797 A catalogue record for this book is available from the British Library. Wiley also publishes its books in a variety of electronic formats. Some content that appears in print may not be available in electronic books. Cover image: A cartoon depicting some accidental passenger insects – a consequence of increased global commerce. Cover design by Peter Cranston and Karina McInness. Set in 9/11pt, PhotinaMTStd by Laserwords Private Limited, Chennai, India 1 2014 Trim size: 189mm x 246mm Gullan ftoc.tex V3 - 07/16/2014 7:28 A.M. Page v CONTENTS List of colour plates, ix 3 INTERNAL ANATOMY AND PHYSIOLOGY, 56 List of boxes, xiii 3.1 Muscles and locomotion, 57 3.2 The nervous system and Preface to the fifth edition, xv co-ordination, 63 Preface to the fourth edition, xvii 3.3 The endocrine system and the function of hormones, 66 Preface to the third edition, xix 3.4 The circulatory system, 69 3.5 The tracheal system and gas Preface to the second edition, xxi exchange, 73 3.6 The gut, digestion and nutrition, 77 Preface and acknowledgments for first edition, xxiii 3.7 The excretory system and waste About the companion website, xxv disposal, 86 3.8 Reproductive organs, 90 1 THE IMPORTANCE, DIVERSITY Further reading, 93 AND CONSERVATION OF INSECTS, 1 4 SENSORY SYSTEMS 1.1 What is entomology?, 2 AND BEHAVIOUR, 95 1.2 The importance of insects, 2 4.1 Mechanical stimuli, 96 1.3 Insect biodiversity, 6 4.2 Thermal stimuli, 105 1.4 Naming and classification of insects, 10 4.3 Chemical stimuli, 107 1.5 Insects in popular culture and 4.4 Insect vision, 117 commerce, 11 4.5 Insect behaviour, 122 1.6 Culturing insects, 13 Further reading, 124 1.7 Insect conservation, 14 1.8 Insects as food, 20 5 REPRODUCTION, 125 Further reading, 25 5.1 Bringing the sexes together, 126 5.2 Courtship, 128 2 EXTERNAL ANATOMY, 26 5.3 Sexual selection, 128 2.1 The cuticle, 27 5.4 Copulation, 131 2.2 Segmentation and tagmosis, 33 5.5 Diversity in genitalic morphology, 136 2.3 The head, 35 5.6 Sperm storage, fertilization and sex 2.4 The thorax, 45 determination, 139 2.5 The abdomen, 52 5.7 Sperm competition, 140 Further reading, 55 5.8 Oviparity (egg-laying), 144 Trim size: 189mm x 246mm Gullan ftoc.tex V3 - 07/16/2014 7:28 A.M. Page vi vi Contents 5.9 Ovoviviparity and viviparity, 150 9 GROUND-DWELLING 5.10 Other modes of reproduction, 150 INSECTS, 249 5.11 Physiological control of 9.1 Insects of litter and soil, 250 reproduction, 153 9.2 Insects and dead trees or decaying Further reading, 154 wood, 260 9.3 Insects and dung, 261 6 INSECT DEVELOPMENT 9.4 Insect–carrion interactions, 264 AND LIFE HISTORIES, 156 9.5 Insect–fungal interactions, 265 6.1 Growth, 157 9.6 Cavernicolous insects, 268 6.2 Life-history patterns and 9.7 Environmental monitoring using phases, 158 ground-dwelling hexapods, 268 6.3 Process and control of moulting, 169 Further reading, 270 6.4 Voltinism, 172 6.5 Diapause, 173 10 AQUATIC INSECTS, 271 6.6 Dealing with environmental 10.1 Taxonomic distribution and extremes, 174 terminology, 272 6.7 Migration, 178 10.2 The evolution of aquatic lifestyles, 275 6.8 Polymorphism and polyphenism, 180 10.3 Aquatic insects and their oxygen 6.9 Age-grading, 181 supplies, 277 6.10 Environmental effects on 10.4 The aquatic environment, 282 development, 183 10.5 Environmental monitoring using aquatic Further reading, 188 insects, 284 10.6 Functional feeding groups, 285 7 INSECT SYSTEMATICS: 10.7 Insects of temporary waterbodies, 286 PHYLOGENY AND 10.8 Insects of the marine, intertidal and CLASSIFICATION, 190 littoral zones, 287 7.1 Systematics, 191 Further reading, 288 7.2 The extant Hexapoda, 201 7.3 Informal group Entognatha: Collembola 11 INSECTS AND PLANTS, 289 (springtails), Diplura (diplurans) and 11.1 Coevolutionary interactions between Protura (proturans), 202 insects and plants, 291 7.4 Class Insecta (true insects), 203 11.2 Phytophagy (or herbivory), 293 Further reading, 224 11.3 Insects and plant reproductive biology, 313 8 INSECT EVOLUTION 11.4 Insects that live mutualistically in AND BIOGEOGRAPHY, 227 specialized plant structures, 318 8.1 Relationships of the Hexapoda to other Further reading, 320 Arthropoda, 228 8.2 The antiquity of insects, 229 12 INSECT SOCIETIES, 322 8.3 Were the first insects aquatic or 12.1 Subsociality in insects, 323 terrestrial?, 236 12.2 Eusociality in insects, 327 8.4 Evolution of wings, 238 12.3 Inquilines and parasites of social 8.5 Evolution of metamorphosis, 241 insects, 345 8.6 Insect diversification, 242 12.4 Evolution and maintenance of 8.7 Insect biogeography, 244 eusociality, 348 8.8 Insect evolution in the Pacific, 245 12.5 Success of social insects, 351 Further reading, 247 Further reading, 353 Trim size: 189mm x 246mm Gullan ftoc.tex V3 - 07/16/2014 7:28 A.M. Page vii Contents vii 13 INSECT PREDATION 17 INSECTS IN A CHANGING AND PARASITISM, 354 WORLD, 457 13.1 Prey/host location, 355 17.1 Models of change, 458 13.2 Prey/host acceptance and 17.2 Economically significant insects under manipulation, 361 climate change, 463 13.3 Prey/host selection and specificity, 364 17.3 Implications of climate change for insect 13.4 Population biology – predator/parasitoid biodiversity and conservation, 467 and prey/host abundance, 372 17.4 Global trade and insects, 468 13.5 The evolutionary success of insect Further reading, 473 predation and parasitism, 375 Further reading, 376 18 METHODS IN ENTOMOLOGY: COLLECTING, PRESERVATION, 14 INSECT DEFENCE, 377 CURATION AND 14.1 Defence by hiding, 379 IDENTIFICATION, 474 14.2 Secondary lines of defence, 380 18.1 Collection, 475 14.3 Mechanical defences, 382 18.2 Preservation and curation, 478 14.4 Chemical defences, 384 18.3 Identification, 488 14.5 Defence by mimicry, 388 Further reading, 491 14.6 Collective defences in gregarious and social insects, 392 TAXOBOXES, 493 Further reading, 396 1 Entognatha: non-insect hexapods (Collembola, Diplura and Protura), 493 15 MEDICAL AND VETERINARY 2 Archaeognatha (Microcoryphia; ENTOMOLOGY, 397 archaeognathans or bristletails), 495 15.1 Insects as causes and vectors of 3 Zygentoma (silverfish), 496 disease, 398 4 Ephemeroptera (mayflies), 497 15.2 Generalized disease cycles, 399 5 Odonata (damselflies and 15.3 Pathogens, 399 dragonflies), 498 15.4 Forensic entomology, 413 6 Plecoptera (stoneflies), 500 15.5