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
3 The major divisions ol 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
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
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
9 Subclass Pterygota, Division Endopterygola, 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
10 Subclass Pterygota, Division Endopterygola, Order
Diplera (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
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
12 Subclass Pterygota, Division Endopterygota, Order
Coleoptera (beetles) ..................................... 221
12.1 Introduction ............................................. 221
12.1.1 Adult morphology .................................. 221
12.1.2 Larvae and pupae .................................. 224
12.2 Suborder Adephaga ........................................ 225
12.2.1 Family Gyrinidae (whirligig beetles) .............. 227
12.2.2 Families Haliplidae and Hygrobiidae ............... 227
12.2.3 Family Dytiscidae ................................. 227
12.2.4 Family Cicindelidae (tiger beetles) ............... 228
12.2.5 Family Carabidae (ground beetles) ................. 228
12.3 Suborder Polyphaga ....................................... 231
12.3.1 Superfamily Staphylinoidea ........................ 231
12.3.1.1 Family Staphylinidae (rove beetles) ........... 231
12.3.1.2 Family Silphidae (burying, sexton or carrion
beetles) ............................................. 232
12.3.1.3 Family Pselaphidae ............................ 232
12.3.1.4 Family Ptiliidae ............................. 233
12.3.2 Superfamily Hydrophiloidea - Family
Hydrophilidae .......................................... 233
12.3.3 Superfamily Histeroidea - Family Histeridae ....... 234
12.3.4 Superfamily Scarabaeoidea ......................... 234
12.3.4.1 Family Lucanidae (stag beetles) ............... 234
12.3.4.2 Family Geotrupidae (dung beetles) ............. 235
12.3.4.3 Family Scarabaeidae ........................... 235
12.3.5 Superfamily Buprestoidea - Family Buprestidae ..... 238
12.3.6 Superfamily Elateroidea - Family Elateridae ....... 238
12.3.7 Superfamily Cantharoidea .......................... 239
12.3.7.1 Family Cantharidae (soldier beetles) .......... 240
12.3.7.2 Family Lampyridae (glow-worms) ................ 240
12.3.8 Superfamily Dermestoidea .......................... 240
12.3.9 Superfamily Bostrychoidea ......................... 241
12.3.9.1 Family Lyctidae (powder-post beetles) ......... 241
12.3.9.2 Family Bostrychidae ........................... 242
12.3.9.3 Family Anobiidae .............................. 242
12.3.9.4 Family Ptinidae (spider beetles) .............. 243
12.3.10 Superfamily Cleroidea ............................ 243
12.3.10.1 Family Cleridae .............................. 243
12.3.10.2 Family Trogossitidae ......................... 243
12.3.10.3 Family Melyridae ............................. 243
12.3.11 Superfamily Cucujoidea ........................... 244
12.3.11.1 Series Heteromera ........................... 244
Family Anthicidae .................................... 244
Family Meloidae (oil beetles, blister beetles) ....... 244
Family Ripiphoridae (previously Rhipiphoridae) ....... 245
Family Alleculidae ................................... 246
Family Oedemeridae ................................... 246
Family Pyrochroidae .................................. 246
Family Tenebrionidae (darkling beetles) .............. 246
12.3.11.2 Series Clavicornia ........................... 247
Family Byturidae ..................................... 248
Family Coccinellidae (ladybirds or ladybeetles
(USA)) ............................................. 248
Family Cryptophagidae (fungus beetles) ............... 250
Family Cucujidae ..................................... 251
Family Lathridiidae .................................. 251
Family Mycetophagidae ................................ 251
Family Nitidulidae ................................... 252
12.3.12 Superfamily Chrysomeloidea ....................... 252
12.3.12.1 Family Cerambycidae (longhorn beetles) ....... 253
12.3.12.2 Family Chrysomelidae (leaf beetles) .......... 255
Subfamily Bruchinae (seed beetles) ................... 255
Subfamily Alticinae (previously Halticinae) .......... 257
Subfamily Chrysomelinae .............................. 258
Subfamily Criocerinae ................................ 259
Subfamily Galerucinae ................................ 260
Subfamily Hispinae ................................... 260
Subfamily Cassidinae ................................. 261
Subfamily Clytrinae .................................. 261
12.3.13 Superfamily Curculionoidea ....................... 262
12.3.13.1 Family Curculionidae ......................... 262
Subfamily Scolytinae (bark beetles) .................. 262
Subfamily Platypodinae ............................... 264
Subfamily Apioninae (seed weevils) ................... 265
Other Subfamilies (true weevils) ..................... 265
Subfamily Entiminae .................................. 266
Subfamily Curculioninae .............................. 268
Subfamily Rhynchophorinae ............................ 269
Subfamily Ceutorhynchinae ............................ 270
Subfamily Hyperinae .................................. 271
13 Class Arachnida .......................................... 272
13.1 Introduction ............................................. 272
13.2 Subclass Araneida (spiders) .............................. 272
13.2.1 Family Araneidae .................................. 273
13.2.2 Family Lycosidae .................................. 273
13.3 Subclass Acarina (mites and ticks) ....................... 274
13.3.1 Order Mesostigmata ................................ 274
13.3.1.1 Family Phytoseiidae ........................... 274
13.3.1.2 Family Varroidae .............................. 275
13.3.2 Order Ixodida (ticks) ............................. 275
13.3.2.1 Family Ixodidae (hard ticks) .................. 275
13.3.2.2 Family Argasidae (soft ticks) ................. 276
13.3.3 Order Prostigmata ................................. 276
13.3.3.1 Family Tetranychidae (spider mites) ........... 276
13.3.3.2 Family Tenuipalpidae (false spider mites) ..... 280
13.3.3.3 Family Tarsonemidae ........................... 280
13.3.3.4 Family Eriophyidae ('gall mites', but there
are also free-living species) ........................ 281
13.3.4 Order Astigmata ................................... 282
13.3.4.1 Family Acaridae ............................... 282
13.3.4.2 Family Sarcoptidae ............................ 283
13.3.5 Order Cryptostigmata (beetle mites) ............... 283
Bibliography .................................................. 284
Index ......................................................... 287
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