The Insects

529 Pages·2005·26.32 MB·English

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The Insects
An Outline of Entomology
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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
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© 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
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CONTENTS
List of color plates, viii 3.7 The excretory system and waste disposal, 77
3.8 Reproductive organs, 81
List of boxes, x Further reading, 84
Preface to the third edition, xii 4 SENSORY SYSTEMS AND
BEHAVIOR,85
Preface to the second edition, xiv 4.1 Mechanical stimuli, 86
4.2 Thermal stimuli, 94
Preface and acknowledgments for first edition, xvi 4.3 Chemical stimuli, 96
4.4 Insect vision, 105
1 THE IMPORTANCE, DIVERSITY, AND 4.5 Insect behavior, 109
CONSERVATION OF INSECTS, 1 Further reading, 111
1.1 What is entomology? 2
1.2 The importance of insects, 2 5 REPRODUCTION,113
1.3 Insect biodiversity, 4 5.1 Bringing the sexes together, 114
1.4 Naming and classification of insects, 8 5.2 Courtship, 117
1.5 Insects in popular culture and commerce, 9 5.3 Sexual selection, 117
1.6 Insects as food, 10 5.4 Copulation, 118
1.7 Insect conservation, 13 5.5 Diversity in genitalic morphology, 123
Further reading, 20 5.6 Sperm storage, fertilization, and sex
determination, 128
2 EXTERNAL ANATOMY,21 5.7 Sperm competition, 128
2.1 The cuticle, 22 5.8 Oviparity (egg-laying), 129
2.2 Segmentation and tagmosis, 28 5.9 Ovoviviparity and viviparity, 135
2.3 The head, 30 5.10 Atypical modes of reproduction, 135
2.4 The thorax, 38 5.11 Physiological control of reproduction, 138
2.5 The abdomen, 45 Further reading, 139
Further reading, 48
6 INSECT DEVELOPMENT AND
3 INTERNAL ANATOMY AND LIFE HISTORIES,141
PHYSIOLOGY, 49 6.1 Growth, 142
3.1 Muscles and locomotion, 50 6.2 Life-history patterns and phases, 143
3.2 The nervous system and co-ordination, 56 6.3 Process and control of molting, 153
3.3 The endocrine system and the function of 6.4 Voltinism, 156
hormones, 59 6.5 Diapause, 157
3.4 The circulatory system, 61 6.6 Dealing with environmental extremes, 158
3.5 The tracheal system and gas exchange, 65 6.7 Migration, 161
3.6 The gut, digestion, and nutrition, 68 6.8 Polymorphism and polyphenism, 163
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vi Contents
6.9 Age-grading, 164 11 INSECTS AND PLANTS,263
6.10 Environmental effects on development, 166 11.1 Coevolutionary interactions between insects
6.11 Climate and insect distributions, 171 and plants, 265
Further reading, 175 11.2 Phytophagy (or herbivory), 265
11.3 Insects and plant reproductive biology, 281
7 INSECT SYSTEMATICS: PHYLOGENY 11.4 Insects that live mutualistically in specialized
AND CLASSIFICATION,177 plant structures, 286
7.1 Phylogenetics, 178 Further reading, 297
7.2 The extant Hexapoda, 180
7.3 Protura (proturans), Collembola (springtails), 12 INSECT SOCIETIES,299
and Diplura (diplurans), 183 12.1 Subsociality in insects, 300
7.4 Class Insecta (true insects), 184 12.2 Eusociality in insects, 304
Further reading, 199 12.3 Inquilines and parasites of social insects, 318
12.4 Evolution and maintenance of eusociality, 320
8 INSECT BIOGEOGRAPHY AND 12.5 Success of eusocial insects, 324
EVOLUTION,201 Further reading, 324
8.1 Insect biogeography, 202
8.2 The antiquity of insects, 203 13 INSECT PREDATION AND
8.3 Were the first insects aquatic or terrestrial? 208 PARASITISM,327
8.4 Evolution of wings, 208 13.1 Prey/host location, 328
8.5 Evolution of metamorphosis, 211 13.2 Prey/host acceptance and manipulation, 334
8.6 Insect diversification, 213 13.3 Prey/host selection and specificity, 338
8.7 Insect evolution in the Pacific, 214 13.4 Population biology – predator/parasitoid and
Further reading, 216 prey/host abundance, 345
13.5 The evolutionary success of insect predation
9 GROUND-DWELLING INSECTS,217 and parasitism, 347
9.1 Insects of litter and soil, 218 Further reading, 353
9.2 Insects and dead trees or decaying wood, 221
9.3 Insects and dung, 223 14 INSECT DEFENSE,355
9.4 Insect–carrion interactions, 224 14.1 Defense by hiding, 356
9.5 Insect–fungal interactions, 226 14.2 Secondary lines of defense, 359
9.6 Cavernicolous insects, 229 14.3 Mechanical defenses, 360
9.7 Environmental monitoring using ground- 14.4 Chemical defenses, 360
dwelling hexapods, 229 14.5 Defense by mimicry, 365
Further reading, 237 14.6 Collective defenses in gregarious and social
insects, 369
10 AQUATIC INSECTS,239 Further reading, 373
10.1 Taxonomic distribution and terminology,
240 15 MEDICAL AND VETERINARY
10.2 The evolution of aquatic lifestyles, 240 ENTOMOLOGY,375
10.3 Aquatic insects and their oxygen supplies, 15.1 Insect nuisance and phobia, 376
241 15.2 Venoms and allergens, 376
10.4 The aquatic environment, 245 15.3 Insects as causes and vectors of disease, 377
10.5 Environmental monitoring using aquatic 15.4 Generalized disease cycles, 378
insects, 248 15.5 Pathogens, 379
10.6 Functional feeding groups, 249 15.6 Forensic entomology, 388
10.7 Insects of temporary waterbodies, 250 Further reading, 393
10.8 Insects of the marine, intertidal, and littoral
zones, 251 16 PEST MANAGEMENT,395
Further reading, 261 16.1 Insects as pests, 396
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Contents vii
16.2 The effects of insecticides, 400 17.2 Preservation and curation, 431
16.3 Integrated pest management, 403 17.3 Identification, 440
16.4 Chemical control, 404 Further reading, 443
16.5 Biological control, 407
16.6 Host-plant resistance to insects, 417 Glossary, 445
16.7 Physical control, 420
16.8 Cultural control, 420 References, 469
16.9 Pheromones and other insect attractants, 421
16.10 Genetic manipulation of insect pests, 422 Index, 477
Further reading, 423
Appendix: A reference guide to orders, 499
17 METHODS IN ENTOMOLOGY:
COLLECTING, PRESERVATION, Color plates fall between pp. 14and 15
CURATION, AND IDENTIFICATION,427
17.1 Collection, 428
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LIST OF
COLOR PLATES
PLATE 1 lining the cavity of the maternal gall (see p. 12)
1.1 An atlas moth, Attacus atlas(Lepidoptera: (P.J. Gullan).
Saturniidae), which occurs in southern India 2.5 Adult male scale insect of Melaleucococcus
and south-east Asia, is one of the largest of all phacelopilus (Hemiptera: Margarodidae), showing
lepidopterans, with a wingspan of about 24cm and a the setiferous antennae and the single pair of wings
larger wing area than any other moth (P.J. Gullan). (P.J. Gullan).
1.2 A violin beetle, Mormolyce phyllodes (Coleoptera: 2.6 A tropical butterfly, Graphium antiphates itamputi
Carabidae), from rainforest in Brunei, Borneo (Lepidoptera: Papilionidae), from Borneo, obtaining
(P.J. Gullan). salts by imbibing sweat from a training shoe (refer to
1.3 The moon moth, Argema maenas (Lepidoptera: Box 5.2) (P.J. Gullan).
Saturniidae), is found in south-east Asia and India; this
female, from rainforest in Borneo, has a wingspan of PLATE 3
about 15cm (P.J. Gullan). 3.1 A female katydid of an undescribed species of
1.4 The mopane emperor moth, Imbrasia belina Austrosalomona (Orthoptera: Tettigoniidae), from
(Lepidoptera: Saturniidae), from the Transvaal in northern Australia, with a large spermatophore
South Africa (R. Oberprieler). attached to her genital opening (refer to Box 5.2)
1.5 A “worm” or “phane” – the caterpillar of Imbrasia (D.C.F. Rentz).
belina– feeding on the foliage of Schotia brachypetala, 3.2 Pupa of a Christmas beetle, Anoplognathus sp.
from the Transvaal in South Africa (R. Oberprieler). (Coleoptera: Scarabaeidae), removed from its
1.6 A dish of edible water bugs, Lethocerus indicus pupation site in the soil in Canberra, Australia
(Hemiptera: Belostomatidae), on sale at a market in (P.J. Gullan).
Lampang Province, Thailand (R.W. Sites). 3.3 Egg mass of Tenodera australasiae (Mantodea:
Mantidae) with young mantid nymphs emerging,
PLATE 2 from Queensland, Australia (refer to Box 13.2)
2.1 Food insects at a market stall in Lampang (D.C.F. Rentz).
Province, Thailand, displaying silk moth pupae 3.4 Eclosing (molting) adult katydid of an
(Bombyx mori), beetle pupae, adult hydrophiloid Elephantodeta species (Orthoptera: Tettigoniidae),
beetles, and water bugs, Lethocerus indicus(R.W. Sites). from the Northern Territory, Australia (D.C.F. Rentz).
2.2 Adult Richmond birdwing (Troides richmondia) 3.5 Overwintering monarch butterflies, Danaus
butterfly and cast exuvial skin on native pipevine plexippus (Lepidoptera: Nymphalidae), from Mill Valley
(Pararistolochiasp.) host (see p. 15) (D.P.A. Sands). in California, USA (D.C.F. Rentz).
2.3 A bush coconut or bloodwood apple gall of 3.6 A fossilized worker ant of Pseudomyrmex oryctus
Cystococcus pomiformis(Hemiptera: Eriococcidae), cut (Hymenoptera: Formicidae) in Dominican amber from
open to show the cream-colored adult female and her the Oligocene or Miocene (P.S. Ward).
numerous, tiny nymphal male offspring covering the 3.7 A diversity of flies (Diptera), including
gall wall (P.J. Gullan). calliphorids, are attracted to the odor of this Australian
2.4 Close-up of the second-instar male nymphs of phalloid fungus, Anthurus archeri, which produces
Cystococcus pomiformisfeeding from the nutritive tissue a foul-smelling slime containing spores that are
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List of color plates ix
consumed by the flies and distributed after passing 5.5 A parasitic Varroa mite (see p. 320) on a pupa of
through the insects’ guts (P.J. Gullan). the bee Apis cerana (Hymenoptera: Apidae) in a hive
from Irian Jaya, New Guinea (D.L. Anderson).
PLATE 4 5.6 An adult moth of Utetheisa ornatrix (Lepidoptera:
4.1 A tree trunk and under-branch covered in Arctiidae) emitting defensive froth containing
silk galleries of the webspinner Antipaluria urichi pyrrolizidine alkaloids that it sequesters as a larva
(Embiidina: Clothodidae), from Trinidad (refer to from its food plants, legumes of the genus Crotalaria
Box 9.5) (J.S. Edgerly-Rooks). (T. Eisner).
4.2 A female webspinner of Antipaluria urichi 5.7 A snake-mimicking caterpillar of the spicebush
defending the entrance of her gallery from an swallowtail, Papilio troilus (Lepidoptera: Papilionidae),
approaching male, from Trinidad (J.S. Edgerly-Rooks). from New Jersey, USA (D.C.F. Rentz).
4.3 An adult stonefly, Neoperla edmundsi (Plecoptera:
Perlidae), from Brunei, Borneo (P.J. Gullan). PLATE 6
4.4 A female thynnine wasp of Zaspilothynnus 6.1 The cryptic adult moths of four species of Acronicta
trilobatus (Hymenoptera: Tiphiidae) (on the right) (Lepidoptera: Noctuidae): A. alni, the alder moth (top
compared with the flower of the sexually deceptive left); A. leporina, the miller (top right); A. aceris, the
orchid Drakaea glyptodon,which attracts pollinating sycamore (bottom left); and A. psi, the grey dagger
male wasps by mimicking the female wasp (see p. 282) (bottom right) (D. Carter and R.I. Vane-Wright).
(R. Peakall). 6.2 Aposematic or mechanically protected
4.5 A male thynnine wasp of Neozeloboria cryptoides caterpillars of the same four species of Acronicta:A. alni
(Hymenoptera: Tiphiidae) attempting to copulate with (top left); A. leporina (top right); A. aceris (bottom left);
the sexually deceptive orchid Chiloglottis trapeziformis and A. psi (bottom right); showing the divergent
(R. Peakall). appearance of the larvae compared with their drab
4.6 Pollination of mango flowers by a flesh fly, adults (D. Carter and R.I. Vane-Wright).
Australopierretia australis (Diptera: Sarcophagidae), 6.3 A blister beetle, Lytta polita (Coleoptera:
in northern Australia (D.L. Anderson). Meloidae), reflex-bleeding from the knee joints;
4.7 The wingless adult female of the whitemarked the hemolymph contains the toxin cantharidin
tussock moth, Orgyia leucostigma (Lepidoptera: (sections 14.4.3 & 15.2.2) (T. Eisner).
Lymantriidae), from New Jersey, USA (D.C.F. Rentz). 6.4 One of Bates’ mimicry complexes from the
Amazon Basin involving species from three different
PLATE 5 lepidopteran families – Methona confusa confusa
5.1 Mealybugs of an undescribed Planococcus (Nymphalidae: Ithomiinae) (top), Lycorea ilione ilione
species (Hemiptera: Pseudococcidae) on an Acacia (Nymphalidae: Danainae) (second from top), Patia orise
stem attended by ants of a Polyrhachis species orise (Pieridae) (second from bottom), and a day-flying
(Hymenoptera: Formicidae), coastal Western moth of Gazera heliconioides (Castniidae) (R.I. Vane-
Australia (P.J. Gullan). Wright).
5.2 A camouflaged late-instar caterpillar of 6.5 An aposematic beetle of the genus Lycus
Plesanemma fucata(Lepidoptera: Geometridae) on a (Coleoptera: Lycidae) on the flower spike of Cussonia
eucalypt leaf in eastern Australia (P.J. Gullan). (Araliaceae) from South Africa (P.J. Gullan).
5.3 A female of the scorpionfly Panorpa communis 6.6 A mature cottony-cushion scale, Icerya purchasi
(Mecoptera: Panorpidae) from the UK (P.H. Ward). (Hemiptera: Margarodidae), with a fully formed
5.4 The huge queen termite (approximately 7.5cm ovisac, on the stem of a native host plant from
long) of Odontotermes transvaalensis (Isoptera: Australia (P.J. Gullan).
Termitidae: Macrotermitinae) surrounded by her king 6.7 Adult male gypsy moth, Lymantria dispar
(mid front), soldiers, and workers, from the Transvaal (Lepidoptera: Lymantriidae), from New Jersey, USA
in South Africa (J.A.L. Watson). (D.C.F. Rentz).
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LIST OF BOXES
Box 1.1 Collected to extinction? 16 Box 9.8 Blattodea (Blattaria; cockroaches, roaches),
Box 1.2 Tramp ants and biodiversity, 17 236
Box 1.3 Sustainable use of mopane worms, 19 Box 10.1 Ephemeroptera (mayflies), 252
Box 3.1 Molecular genetic techniques and their Box 10.2 Odonata (damselflies and dragonflies), 253
application to neuropeptide research, 60 Box 10.3 Plecoptera (stoneflies), 255
Box 3.2 Tracheal hypertrophy in mealworms at low Box 10.4 Trichoptera (caddisflies), 255
oxygen concentrations, 69 Box 10.5 Diptera (true flies), 257
Box 3.3 The filter chamber of Hemiptera, 71 Box 10.6 Other aquatic orders, 258
Box 3.4 Cryptonephric systems, 79 Box 11.1 Induced defenses, 268
Box 4.1 Aural location of host by a parasitoid fly, 91 Box 11.2 The grape phylloxera, 276
Box 4.2 The electroantennogram, 97 Box 11.3 Salvinia and phytophagous weevils, 280
Box 4.3 Reception of communication molecules, 99 Box 11.4 Figs and fig wasps, 284
Box 4.4 Biological clocks, 106 Box 11.5 Orthoptera (grasshoppers, locusts,
Box 5.1 Courtship and mating in Mecoptera, 116 katydids, and crickets), 289
Box 5.2 Nuptial feeding and other “gifts”, 121 Box 11.6 Phasmatodea (phasmatids, phasmids,
Box 5.3 Sperm precedence, 126 stick-insects or walking sticks), 290
Box 5.4 Control of mating and oviposition in a Box 11.7 Thysanoptera (thrips), 291
blow fly, 130 Box 11.8 Hemiptera (bugs, cicadas, leafhoppers,
Box 5.5 Egg-tending fathers – the giant water bugs, spittle bugs, planthoppers, aphids, jumping plant
132 lice, scale insects, whiteflies), 292
Box 6.1 Molecular insights into insect development, Box 11.9 Psocoptera (booklice, barklice, or psocids),
148 294
Box 6.2 Calculation of day-degrees, 168 Box 11.10 Coleoptera (beetles), 295
Box 6.3 Climatic modeling for fruit flies, 174 Box 11.11 Lepidoptera (butterflies and moths), 296
Box 7.1 Relationships of the Hexapoda to other Box 12.1 The dance language of bees, 310
Arthropoda, 181 Box 12.2 Hymenoptera (bees, ants, wasps, sawflies,
Box 9.1 Ground pearls, 222 and wood wasps), 325
Box 9.2 Non-insect hexapods (Collembola, Protura, Box 12.3 Isoptera (termites), 326
and Diplura), 230 Box 13.1 Viruses, wasp parasitoids, and host
Box 9.3 Archaeognatha (bristletails) and Zygentoma immunity, 337
(Thysanura; silverfish), 232 Box 13.2 Mantodea (mantids), 348
Box 9.4 Grylloblattodea (Grylloblattaria, Notoptera; Box 13.3 Mantophasmatodea (heel walkers), 349
grylloblattids, ice or rock crawlers), 233 Box 13.4 Neuropterida, or neuropteroid orders,
Box 9.5 Embiidina or Embioptera (embiids, 350
webspinners), 234 Box 13.5 Mecoptera (scorpionflies, hangingflies),
Box 9.6 Zoraptera, 234 351
Box 9.7 Dermaptera (earwigs), 235 Box 13.6 Strepsiptera, 352