OUTLINES OF ENTOMOLOGY

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1 OUTLINES OF ENTOMOLOGY

2 OUTLINES OF ENTOMOLOGY R.G. Davies Emeritus Professor of Entomology Imperial College, University of London SEVENTH EDITION SPRINGER-SCIENCE+BUSINESS MEDIA, B.V.

3 1988 R.G. Davies Originally published by Chapman & Hall in 1988 Softcover reprint of the hardcover 1 st edition 1988 Typeset in 10.5/12 Bembo by Photoprint, Torquay ISBN ISBN (ebook) DOI / This title is available in both hardbound and paperback editions. The paperback edition is sold subject to the condition that it shall not, by way of trade or otherwise, be lent, resold, hired out, or otherwise circulated without the publisher's prior consent in any form of binding or cover other than that in which it is published and without a similar condition including this condition being imposed on the subsequent purchaser. All rights reserved. No part of this book may be reprinted, or reproduced or utilized in any form or by any electronic, mechanical or other means, now known or hereafter invented, including photocopying and recording, or in any information storage and retrieval system, without permission in writing from the publisher. British Library Cataloguing in Publication Data Davies, R.G. (Richard Gareth), Outlines of entomology. - 7th cd. 1. Insects I. Title II. Imms, Augustus Daniel. Imms' outlines of entomology ISBN Library of Congress Cataloging in Publication Data Davies, R. G. (Richard Gareth) Outlines of entomology. - 7th cd. / R.G. Davies. p. cm. En!. cd. of: Outlines of entomology / A.D. Imms. Bibliography: p. Includes index. ISBN Insects. I. Imms, A. D. (Augustus Daniel) Outlines of entomology. II. Title. QL dcI CIP

4 Contents Preface viii 1 Introduction 1 2 Insect structure and function 7 The integument and its derivatives; coloration 7 Segmentation and body regions 12 The head and its appendages 13 The thorax, legs and wings 17 The abdomen and genitalia 30 The muscular system 35 The nervous system 43 The sense organs and perception 48 Alimentary canal, digestion and nutrition 61 The respiratory system 67 The circulatory system and associated tissues 77 The excretory organs, fat-body and other haemocoelic structures 82 The glands or organs of secretion 86 The reproductive system and reproduction 88 3 Development and metamorphosis 97 Embryonic development 97 Post-embryonic development Classification and biology 115 Order 1. Diplura 118 Order 2. Protura 120 Order 3. Collembola 121 Order 4. Archaeognatha 122 Order 5. Zygentoma 124 Order 6. Ephemeroptera 126 Order 7. Odonata 128

5 CONTENTS Order 8. Plecoptera 130 Order 9. Grylloblattodea 131 Order 10. Orthoptera 132 Order 11. Phasmida 134 Order 12. Dermaptera 135 Order 13. Embioptera 135 Order 14. Dictyoptera 137 Order 15. Isoptera 138 Order 16. Zoraptera 139 Order 17. Psocoptera 140 Order 18. Phthiraptera 141 Order 19. Hemiptera 142 Order 20. Thysanoptera 148 Order 21. Neuroptera 149 Order 22. Coleoptera 151 Order 23. Strepsiptera 156 Order 24. Mecoptera 157 Order 25. Siphonaptera 158 Order 26. Diptera 159 Order 27. Lepidoptera 168 Order 28. Trichoptera 173 Order 29. Hymenoptera Evolutionary relationships of insects 182 Insects and other Arthropods 183 The ancestry of insects 186 Apterygote relationships 188 Phylogeny of the Pterygota Some important modes of life in insects 195 Relationships between insects and plants 196 Leaf-mining insects 201 Insects and plant galls 203 Insects and the pollination of flowers 207 Insectivorous plants 214 Insects and micro-organisms 215 Predacious insects 218 Insect parasitoids 222 Ectoparasitic insects 227 Blood-sucking insects 230 Aquatic insects 232 Protective coloration and mimicry 237 Social insects 241 VI

6 CONTENTS 7 The biology of insect populations 248 Estimation of insect population size 249 Models of insect population growth 251 Population fluctuations 257 Life tables 258 Key factor analysis 260 Regulation of population size; density-dependent processes 262 Competition 266 Competitive exclusion; resource partitioning; the ecological niche 271 Host-parasitoid and predator-prey relationships 272 Food webs and the trophic structure of communities 276 Diversity, stability and succession in insect communities 281 Genetic variation in insect populations Biology and control of injurious insects 290 Insect pests of cultivated plants 291 Insect pests of forestry, timber and stored products 305 Insects of medical and veterinary importance 313 Insecticidal control 326 Biological control methods Select classified bibliography 343 Index 384 VB

7 Preface to the seventh edition The present edition may be regarded as a descendant, much changed and greatly enlarged, of the late Dr A.D. Imms' Outlines of Entomology, first published in This went through three further editions without much change, but after the death of the original author a fifth, revised edition by Professor o. W. Richards and myself appeared in 1959 and a sixth in The book now appears in a considerably extended version in which I have tried to provide a more balanced introduction to the whole field of modern entomology by dealing with several aspects of the subject not discussed at all in previous editions. Thus, in addition to innumerable lesser changes in the sections on insect structure, function, development, classification and phylogeny, I have completely recast the earlier chapter on some important modes of life in insects. This now includes a far wider range of biological topic;s well exemplified by the insects and should, I hope, appeal not only to, those already dedicated to entomology but also to others with more general biological interests. A completely new chapter on the biology of insect populations has also been added and may serve to indicate the debt which modern ecological theory owes to work on insect populations. It should hardly be necessary to apologize for introducing a certain amount of elementary mathematics into this account of a subject which is now among the most highly quantitative of biological disciplines. A third new section is that devoted to the biology of a selection of injurious insects and the principles that underlie modern integrated methods of pest management. The examples here are drawn from many parts of the world and should help to demonstrate the extent to which practical control of insect pests depends on an understanding of their biology. The book is now more fully illustrated, with most of the figures newly redrawn and relettered so as to make them more immediately intelligible. Lastly, a much larger classified bibliography aims at introducing more advanced students to the ever-increasing range of specialized publications on entomology. R.G.D. London, November 1987

IMMS' GENERAL TEXTBOOK OF ENTOMOLOGY. Volume I

IMMS' GENERAL TEXTBOOK OF ENTOMOLOGY Volume I IMMS' GENERAL TEXTBOOK OF ENTOMOLOGY TENTH EDITION Volume I: Structure, Physiology and Development O. W. RICHARDS M.A., D.Sc., F.R.S. Emeritus Professor of