Table Of ContentHandbook of Agricultural
Entomology
Handbook of Agricultural
Entomology
H. F. van Emden
Emeritus Professor of Horticulture
School of Agriculture, Policy and Development
University of Reading
Reading
UK
A John Wiley & Sons, Ltd., Publication
This edition first published 2013 © 2013 by John Wiley & Sons, Ltd.
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Library of Congress Cataloging-in-Publication Data
Van Emden, Helmut Fritz.
Handbook of agricultural entomology/H.F. Van Emden.
p. cm.
Includes bibliographical references and index.
ISBN 978-0-470-65913-7 (hardback: alk. paper)
1. Insect pests-identification. I. Title.
SB931.V25 2013
632'.7-dc23
21012025032
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 credits: Middle left: Soft scales (Coccidae) on citrus: (a) Coccus hesperidum and (b)
olive scale (Saissetia oleae) (from Bayer 1968, with permission). Bottom left: Rhyssa persuasoria
(from Mandahl-Barth 1974, with permission).
Top right: Pea moth (Cydia nigricana) and pod opened to show damage with caterpillars (from
Bayer 1968, with permission). Middle right: Side view of head of (a) a short-nosed and (b) a long-
nosed weevil (from Reitter 1908–1916). Bottom right: A spider beetle (Ptinidae) (from Reitter
1908–1916).
Cover design by Design Deluxe (e: [email protected])
Set in 9.5/12 pt Berkeley by Toppan Best-set Premedia Limited
1 2013
Contents
Companion Website details xv
Preface xvii
Acknowledgements xxi
1 The world of insects 1
1.1 The diversity of insects 1
1.2 The impact of insects on us 1
1.3 The impact we have on insects 5
1.3.1 World distribution 5
1.3.2 Climate change 5
1.3.3 Land management practices 5
1.4 Exploitation of insects 6
1.5 Other uses humans make of insects 6
1.6 Insect classification 7
2 External features of insects – structure and function 9
2.1 Introduction 9
2.2 The exoskeleton 10
2.3 The basic body plan of the insect 11
2.4 The head 11
2.4.1 Sense organs 12
2.4.1.1 Antennae 12
2.4.1.2 Ocelli 14
2.4.1.3 Compound eyes 14
2.4.2 Basic structure of the mouthparts 15
2.4.2.1 Mandibles 16
2.4.2.2 Maxillae 16
2.4.2.3 Labium 16
2.5 The thorax 16
2.5.1 Thoracic sclerites 16
2.5.2 Legs 18
2.5.3 Wings 19
2.5.4 Spiracles 22
2.6 The abdomen 22
2.6.1 Abdominal sclerites 22
2.6.2 Appendages 22
2.6.3 Genitalia 22
2.6.4 Spiracles 23
v
vi CONTENTS
3 The major divisions of the Insecta 25
3.1 Introduction 25
3.2 Class Insecta, Subclass Apterygota or Phylum Arthropoda, Class
Entognatha 25
3.3 Subclass Pterygota 26
3.3.1 Division Exopterygota 26
3.3.1.1 Palaeopteran Orders (example: dragonflies) 28
3.3.1.2 Orthopteroid Orders (example:
grasshoppers) 28
3.3.1.3 Hemipteroid Orders (example: plant bugs) 28
3.3.2 Division Endopterygota 28
4 Subclass Apterygota 31
4.1 Introduction 31
4.2 Order Diplura (two-pronged bristle-tails) 31
4.3 Order Protura 31
4.4 Order Thysanura (silverfish) 32
4.5 Order Collembola (springtails) 33
4.5.1 Suborder Arthropleona 34
4.5.2 Suborder Symphypleona 34
5 Subclass Pterygota, Division Exopterygota, Palaeopteran Orders 36
5.1 Introduction 36
5.2 Order Ephemeroptera (mayflies) 36
5.3 Order Odonata (dragonflies) 38
5.3.1 Suborder Zygoptera (damselflies) 39
5.3.2 Suborder Anisoptera (dragonflies) 40
6 Subclass Pterygota, Division Exopterygota, Orthopteroid Orders 42
6.1 Introduction 42
6.2 Order Plecoptera (stoneflies) 43
6.3 Order Grylloblattodea 43
6.4 Order Mantophasmatodea (gladiators or heelwalkers) 44
6.5 Order Zoraptera (angel insects) 44
6.6 Order Orthoptera (grasshoppers and crickets) 44
6.6.1 Suborder Ensifera 45
6.6.1.1 Superfamily Tettigonioidea (long-horned
grasshoppers) – the main Family is the
Tettigoniidae 45
6.6.1.2 Superfamily Grylloidea (crickets) – the main
Family is the Gryllidae 46
6.6.2 Suborder Caelifera 48
6.6.2.1 Superfamily Acridoidea (short-horned
grasshoppers) – the main Family is the
Acrididae 48
6.7 Order Phasmida (stick and leaf insects) 51
6.8 Order Dermaptera (earwigs) 52
CONTENTS vii
6.9 Order Embioptera (web spinners) 53
6.10 Order Dictyoptera (cockroaches and mantids) 54
6.10.1 Suborder Blattaria (cockroaches) 54
6.10.2 Suborder Mantodea (mantids) 56
6.11 Order Isoptera (termites) 57
6.11.1 Family Kalotermitidae (dry-wood termites) 60
6.11.2 Family Hodotermitidae 60
6.11.3 Family Rhinotermitidae (wet-wood termites) 61
6.11.4 Family Termitidae (mound-building termites) 61
7 Subclass Pterygota, Division Exopterygota, Hemipteroid Orders 63
7.1 Introduction 63
7.2 Order Psocoptera (booklice) 63
7.3 Order Mallophaga (biting lice) 64
7.4 Order Anoplura (= Siphunculata) (sucking lice) 64
7.5 Order Hemiptera (true bugs) 65
7.5.1 Suborder Heteroptera (land and water bugs) 67
7.5.1.1 Series Cryptocerata (water bugs) 68
Water boatmen (Families Notonectidae and
Corixidae) 68
Water scorpions and allies (Family Nepidae) 69
7.5.1.2 Series Gymnocerata 69
Pond skaters (Families Gerridae and
Veliidae) 70
Family Miridae (mirids or capsids) 70
Family Lygaeidae (chinch bugs) 72
Superfamily Pentatomoidea (shield bugs) 74
Family Pentatomidae (stink bugs) 74
Family Acanthosomatidae 76
Family Scutelleridae 76
Family Coreidae 77
Family Pyrrhocoridae 77
Family Tingidae (lace bugs) 78
Family Anthocoridae (flower bugs) 79
Family Cimicidae (bed bugs) 79
Family Nabidae (damsel bugs) 80
Family Reduviidae (assassin bugs) 80
7.5.2 Suborder Auchenorryncha (hoppers) 81
7.5.2.1 Superfamily Fulgoroidea 81
Family Fulgoridae (lantern flies) 81
Family Delphacidae (planthoppers) 82
7.5.2.2 Superfamily Cicadelloidea 83
Family Membracidae (treehoppers) 83
Family Cicadellidae (leafhoppers) 83
7.5.2.3 Superfamily Cicadoidea (Family Cicadidae –
cicadas) 85
7.5.2.4 Superfamily Cercopoidea (Family Cercopidae
– froghoppers) 86
viii CONTENTS
7.5.3 Suborder Sternorryncha (plant lice) 87
7.5.3.1 Superfamily Psylloidea (Family Psyllidae
– suckers or jumping plant lice) 87
7.5.3.2 Superfamily Aphidoidea (aphids) 88
Family Lachnidae 89
Family Aphididae (aphids, greenfly,
blackfly) 89
Family Phylloxeridae 99
7.5.3.3 Superfamily Aleyrodoidea (whiteflies)
– Family Aleyrodidae 99
7.5.3.4 Superfamily Coccoidea (scale insects and
mealybugs) 101
Family Pseudococcidae (mealybugs) 102
Family Margarodidae (cushion scales) 102
Family Kerridae 103
Family Asterolecaniidae 103
Family Coccidae (soft scales) 104
Family Diaspididae (armoured scales) 105
7.6 Order Thysanoptera (thrips or thunderflies) 106
7.6.1 Suborder Terebrantia 107
7.6.1.1 Family Thripidae 107
7.6.1.2 Family Aeolothripidae 110
7.6.2 Suborder Tubulifera 110
7.6.2.1 Family Phlaeothripidae 110
8 Subclass Pterygota, Division Endopterygota, Lesser Orders 112
8.1 Introduction 112
8.2 Order Mecoptera (scorpion flies) 112
8.3 Order Siphonaptera (fleas) 113
8.4 Order Neuroptera 114
8.4.1 Suborder Megaloptera 115
8.4.1.1 Family Corydalidae 115
8.4.1.2 Family Sialidae (alder flies) 115
8.4.1.3 Family Raphidiidae (snake flies) 116
8.4.2 Suborder Planipennia 116
8.4.2.1 Superfamily Coniopterygoidea 116
Family Coniopterygidae 116
8.4.2.2 Superfamily Osmyloidea 116
Family Osmylidae 116
8.4.2.3 Superfamily Mantispoidea 116
Family Sisyridae 116
Family Mantispidae (mantis flies) 117
8.4.2.4 Superfamily Hemeroboidea (lacewings) 117
Family Hemerobiidae (brown lacewings) 117
Family Chrysopidae (green lacewings) 117
8.4.2.5 Superfamily Myrmeleontoidea (ant lions) 118
8.5 Order Trichoptera (caddis flies) 119
8.6 Order Strepsiptera (stylops) 120
CONTENTS ix
9 Subclass Pterygota, Division Endopterygota, Order Lepidoptera
(butterflies and moths) 124
9.1 Introduction 124
9.2 Suborder Glossata 128
9.2.1 Infraorder Exoporia 128
9.2.1.1 Superfamily Hepialoidea 128
Family Hepialidae (swift moths) 128
9.2.2 Infraorder Heteroneura – Division Monotrysia 129
9.2.2.1 Superfamily Nepticuloidea 129
Family Nepticulidae 129
9.2.2.2 Superfamily Incurvarioidea 129
Family Incurvariidae 129
9.2.3 Infraorder Heteroneura – Division Ditrysia 129
9.2.3.1 Superfamily Tineoidea 129
Family Gracillariidae 130
9.2.3.2 Superfamily Gelechioidea 131
Family Oecophoridae 131
Family Gelechiidae 131
Family Coleophoridae (casebearer
moths) 133
9.2.3.3 Superfamily Yponomeutoidea 133
Family Plutellidae 133
Family Yponomeutidae 134
9.2.3.4 Superfamily Pyraloidea 134
Family Pyralidae 135
9.2.3.5 Superfamily Sesioidea 139
Family Sesiidae (clearwings) 139
9.2.3.6 Superfamily Cossoidea 139
Family Cossidae 139
9.2.3.7 Superfamily Tortricoidea 140
Family Tortricidae 140
9.2.3.8 Superfamily Geometroidea 143
Family Geometridae 143
9.2.3.9 Superfamily Papilionoidea (butterflies) 145
Family Hesperiidae (skippers) 145
Family Papilionidae (swallowtails) 145
Family Pieridae (whites) 146
Family Lycaenidae (blues) 147
Family Nymphalidae 147
9.2.3.10 Superfamily Bombycoidea 148
Family Bombycidae 148
Family Lasiocampidae 148
9.2.3.11 Superfamily Sphingoidea 149
Family Sphingidae (hawk moths) 149
9.2.3.12 Superfamily Noctuoidea 150
Family Lymantriidae 150
Family Arctiidae 151
Family Noctuidae (owlet moths) 152
x CONTENTS
10 Subclass Pterygota, Division Endopterygota, Order Diptera (true flies) 159
10.1 Introduction 159
10.2 Suborder Nematocera 161
10.2.1 Family Tipulidae (crane flies) 161
10.2.2 Family Psychodidae 163
10.2.3 Family Cecidomyiidae (gall midges) 163
10.2.4 Family Culicidae (mosquitoes) 165
10.2.5 Family Chironomidae (non-biting midges) 168
10.2.6 Family Ceratopogonidae (biting midges) 169
10.2.7 Family Simuliidae (black flies) 171
10.2.8 Family Sciaridae (mushroom flies, dark-winged fungus
gnats) 172
10.2.9 Family Mycetophilidae (fungus gnats) 172
10.2.10 Family Bibionidae 172
10.3 Suborder Brachycera 173
10.3.1 Family Tabanidae (horse flies) 173
10.3.2 Family Stratiomyidae (soldier flies) 173
10.3.3 Family Asilidae (robber flies) 174
10.3.4 Family Empididae (assassin flies) 174
10.4 Suborder Cyclorrhapha 174
10.4.1 Series Aschiza 175
10.4.1.1 Family Syrphidae (hover flies) 175
10.4.2 Series Schizophora 177
10.4.2.1 Division Acalyptrata 178
Family Psilidae 178
Family Sepsidae (semaphore
flies) 179
Family Ephydridae (shore flies) 179
Family Chloropidae 180
Family Opomyzidae 181
Family Agromyzidae 181
Family Diopsidae (stalk-eyed flies) 182
Family Tephritidae (or Trypetidae)
(fruit flies) 183
10.4.2.2 Division Calyptrata 185
Family Scathophagidae (or
Cordiluridae) 185
Family Anthomyiidae 185
Family Muscidae 187
Family Calliphoridae (bluebottles and
greenbottles) 189
Family Sarcophagidae (flesh flies) 189
Family Tachinidae 190
Family Glossinidae (tsetse flies) 190
Family Oestridae 191
10.4.2.3 Division Pupipara (louse flies, keds) 192
Family Hippoboscidae 192
CONTENTS xi
11 Subclass Pterygota, Division Endopterygota, Order Hymenoptera
(sawflies, ants, bees and wasps) 193
11.1 Introduction 193
11.2 Suborder Symphyta (sawflies) 194
11.2.1 Family Siricidae (wood wasps) 195
11.2.2 Family Cephidae 196
11.2.3 Family Tenthredinidae 196
11.2.4 Family Diprionidae 198
11.3 Suborder Apocrita (ants, bees, wasps and parasitic
wasps) 198
11.3.1 The social Hymenoptera and their allies 198
11.3.1.1 Superfamily Apoidea 199
Family Andrenidae 199
Family Megachilidae (leaf-cutter and mason
bees) 199
Family Apidae 200
Family Sphecidae (digger wasps) 202
11.3.1.2 Superfamily Vespoidea (wasps) 203
Family Vespidae 203
Family Mutillidae (velvet ants) 205
Family Pompilidae (spider-hunting
wasps) 205
Family Scoliidae 206
Family Tiphiidae 206
Family Formicidae (ants) 206
11.3.1.3 Superfamily Chrysidoidea 210
11.3.2 Other Superfamilies 210
11.3.2.1 Superfamily Ichneumonoidea 212
Family Ichneumonidae 212
Family Braconidae 213
11.3.2.2 Superfamily Chalcidoidea 214
Family Agaonidae (fig wasps) 215
Family Aphelinidae 215
Family Chalcidae 216
Family Pteromalidae 216
Family Torymidae 216
Family Encyrtidae 216
Family Trichogrammatidae 217
Family Mymaridae (fairy flies) 217
Family Tanaostigmatidae 218
11.3.2.3 Superfamily Proctotrupoidea 218
11.3.2.4 Superfamily Evanioidea 218
Family Evaniidae 218
Family Gasteruptiidae 218
11.3.2.5 Superfamily Cynipoidea 219
Family Charipidae 219
Family Cynipidae (gall wasps) 219
