Part I General Aspects
1 Systematics .................................................. 3
2 Biology of Mosquitoes ........................................ 9
2.1 Oviposition ............................................ 10
2.2 Embryonic Development .................................. 13
2.3 Hatching ............................................... 13
2.4 Larvae ................................................. 16
2.5 Pupae .................................................. 19
2.6 Adults ................................................. 19
2.6.1 Emergence ....................................... 19
2.6.2 Mating .......................................... 20
2.6.3 Dispersal and Host-Seeking Behaviour ............ 21
2.6.4 Feeding ......................................... 22
2.7 Survival During Dry Seasons and Hibernation ............ 24
2.7.1 Egg Stage ....................................... 24
2.7.2 Larval Stage .................................... 24
2.7.3 Adult Stage ..................................... 24
3 Medical Importance of Mosquitoes ............................ 25
3.1 Malaria ................................................ 25
3.2 Arboviruses ............................................ 30
3.2.1 Togaviridae (Alphavirus) ........................ 31
3.2.2 Flaviviridae (Flavivirus) ....................... 34
3.2.3 Bunyaviridae (Bunyavirus) ....................... 37
3.3 Filariasis ............................................. 39
3.4 Future Perspectives and Conclusions .................... 40
4 Mosquito Research Techniques ................................ 43
4.1 Sampling Mosquito Eggs ................................. 43
4.1.1 Anopheles Eggs .................................. 43
4.1.2 Egg Rafts ....................................... 43
4.1.3 Aedes/Ochlerotatus Eggs ......................... 44
4.1.4 Eggs in Artificial Oviposition Sites ............ 45
4.2 Sampling Mosquito Larvae and Pupae ..................... 46
4.3 Sampling Adult Mosquitoes in the Field ................. 47
4.3.1 Sampling Flying Mosquitoes ...................... 47
4.3.2 Adult Mosquito-Outdoor Resting Catches .......... 47
4.3.3 Adult Mosquito-Indoor Catches ................... 47
4.3.4 Bait Catches .................................... 48
4.3.5 Adult Mosquito Traps ............................ 49
4.3.6 Mark-Release-Recapture Techniques ............... 51
4.4 Laboratory Based Research Techniques ................... 51
4.4.1 Rearing Mosquitoes .............................. 51
4.4.2 Preserving Mosquitoes ........................... 53
4.4.3 Mosquito Blood Meal Identification .............. 55
4.4.4 Methods for Measuring the Physiological Stage ... 55
4.4.5 Morphological and Taxonomic Techniques .......... 56
4.5 Assessing the Activity of Insecticides and Repellents
on Mosquitoes .......................................... 58
4.5.1 Insecticide Susceptibility Testing .............. 58
4.5.2 Assays of Insecticide Deposits on Surfaces
(e.g. Walls or Nets) ............................ 60
4.5.3 Assays of Efficacy of ULV Insecticide
Treatments in the Field ......................... 60
4.5.4 Assays of Efficacy of Mosquito Repellents ....... 60
4.6 Conclusion ............................................. 61
5 Morphology of Mosquitoes .................................... 63
5.1 Adults ................................................. 63
5.1.1 Head ............................................ 63
5.1.2 Thorax .......................................... 67
5.1.3 Abdomen ......................................... 72
5.2 Larvae ................................................. 76
5.2.1 Head ............................................ 78
5.2.2 Thorax .......................................... 80
5.2.3 Abdomen ......................................... 80
5.3 Pupae .................................................. 84
Part II Identification Keys; Morphology; Ecology and
Distribution of European Species
6 Key to Female Mosquitoes .................................... 91
Genera ...................................................... 91
6.1 Genus Anopheles ........................................ 92
6.2 Genera Aedes and Ochlerotatus .......................... 95
6.3 Genus Culex ........................................... 106
6.4 Genus Culiseta ........................................ 108
6.5 Genus Coquillettidia .................................. 111
7 Key to Male Mosquitoes ..................................... 113
Genera ..................................................... 113
7.1 Genus Anopheles ....................................... 115
7.2 Genera Aedes and Ochlerotatus ......................... 118
7.3 Genus Culex ........................................... 128
7.4 Genus Culiseta ........................................ 132
7.5 Genus Coquillettidia .................................. 134
8 Key to Mosquito Fourth Instar Larvae ....................... 135
Genera ..................................................... 135
8.1 Genus Anopheles ....................................... 137
8.2 Genera Aedes and Ochlerotatus ......................... 141
8.3 Genus Culex ........................................... 154
8.4 Genus Culiseta ........................................ 159
8.5 Genus Coquillettidia .................................. 162
9 Subfamily Anophelinae ...................................... 163
9.1 Genus Anopheles (Meigen) .............................. 164
9.1.1 Subgenus Anopheles (Meigen) .................... 165
Anopheles algeriensis (Theobald 1903) .......... 165
Anopheles Claviger Complex ..................... 166
Anopheles claviger s.s. (Meigen 1804) ....... 166
Anopheles petragnani (Del Vecchio 1939) ..... 168
Anopheles hyrcanus (Pallas 1771) ............... 169
Anopheles Maculipennis Complex ................. 170
Anopheles atroparvus (Van Thiel 1927) ....... 173
Anopheles beklemishevi (Stegnii and
Kabanova 1976) .............................. 173
Anopheles labranchiae (Falleroni 1926) ...... 174
Anopheles maculipennis s.s. (Meigen 1818) ... 174
Anopheles melanoon (Hackett 1934) ........... 175
Anopheles messeae (Falleroni 1926) .......... 175
Anopheles sacharovi (Favre 1903) ............ 175
Anopheles subalpinus (Hackett and Lewis
1935) ....................................... 176
Anopheles marten (Senevet and Prunelle 1927) ... 177
Anopheles plumbeus (Stephens 1828) ............. 178
9.1.2 Subgenus Cellia (Theobald 1902) ................ 180
Anopheles cinereus (Theobald 1901) ............. 180
Anopheles cinereus hispaniola (Theobald
1903) .......................................... 181
Anopheles multicolor (Cambouliu 1902) .......... 182
Anopheles sergentii (Theobald 1907) ............ 183
Anopheles superpictus (Grassi 1899) ............ 185
10 Subfamily Culicinae ........................................ 187
10.1 Genus Aedes (Meigen) .................................. 187
10.1.1 Subgenus Aedes (Meigen) ........................ 188
Aedes cinereus (Meigen 1818) ................... 189
Aedes geminus (Peus 1970) ...................... 191
Aedes rossicus (Dolbeskin, Gorickaja and
Mitrofanova 1930) .............................. 192
10.1.2 Subgenus Aedimorphus (Theobald) ................ 193
Aedes vexans (Meigen 1830) ..................... 194
10.1.3 Subgenus Fredwardsius (Reinert) ................ 196
Aedes vittatus (Bigot 1861) .................... 196
10.1.4 Subgenus Stegomyia (Theobald) .................. 198
Aedes aegypti (Linnaeus 1762) .................. 198
Aedes albopictus (Skuse 1895) .................. 201
Aedes cretinus (Edwards 1921) .................. 203
10.2 Genus Ochlerotatus (Lynch Arribalzaga) ................ 204
10.2.1 Subgenus Finlaya (Theobald) .................... 205
Ochlerotatus echinus (Edwards 1920) ............ 205
Ochlerotatus geniculatus (Olivier 1791) ........ 206
10.2.2 Subgenus Ochlerotatus Lynch Arribalzaga ........ 208
Ochlerotatus annulipes (Meigen 1830) ........... 209
Ochlerotatus behningi (Martini 1926) ........... 211
Ochlerotatus berlandi (Seguy 1921) ............. 212
Ochlerotatus cantans (Meigen 1818) ............. 214
Ochlerotatus Caspius Complex ................... 216
Ochlerotatus caspius (Pallas 1771) .......... 216
Ochlerotatus cataphylla (Dyar 1916) ............ 218
Ochlerotatus Communis Complex .................. 219
Ochlerotatus communis (De Geer 1776) ........ 220
Ochlerotatus cyprius (Ludlow 1920) ............. 221
Ochlerotatus Detritus Complex .................. 222
Ochlerotatus detritus (Haliday 1833) ........ 223
Ochlerotatus diantaeus (Howard, Dyar and Knab
1913) .......................................... 224
Ochlerotatus dorsalis (Meigen 1830) ............ 226
Ochlerotatus Excrucians Complex ................ 228
Ochlerotatus euedes (Howard, Dyar and Knab
1913) ....................................... 228
Ochlerotatus excrucians (Walker 1856) ....... 229
Ochlerotatus surcoufi (Theobald 1912) ....... 231
Ochlerotatus flavescens (Muller 1764) .......... 231
Ochlerotatus hexodontus (Dyar 1916) ............ 233
Ochlerotatus hungaricus (Mihalyi 1955) ......... 234
Ochlerotatus impiger (Walker 1848) ............. 236
Ochlerotatus intrudens (Dyar 1919) ............. 237
Ochlerotatus leucomelas (Meigen 1804) .......... 238
Ochlerotatus Mariae Complex .................... 240
Ochlerotatus mariae (Sergent and Sergent
1903) ....................................... 240
Ochlerotatus zammitii (Theobald 1903) ....... 242
Ochlerotatus mercurator (Dyar 1920) ............ 242
Ochlerotatus nigrinus (Eckstein 1918) .......... 243
Ochlerotatus nigripes (Zetterstedt 1838) ....... 245
Ochlerotatus pionips (Dyar 1919) ............... 246
Ochlerotatus puleritarsis (Rondani 1872) ....... 248
Ochlerotatus pullatus (Coquillett 1904) ........ 249
Ochlerotatus punctodes (Dyar 1922) ............. 250
Ochlerotatus punctor (Kirby 1837) .............. 251
Ochlerotatus riparius (Dyar and Knab 1907) ..... 253
Ochlerotatus sticticus (Meigen 1838) ........... 255
10.2.3 Subgenus Rusticoidus (Shevchenko and
Prudkina) ...................................... 256
Ochlerotatus lepidonotus (Edwards 1920) ........ 257
Ochlerotatus quasirusticus (Torres Canamares
1951) .......................................... 258
Ochlerotatus refiki (Medschid 1928) ............ 259
Ochlerotatus rusticus (Rossi 1790) ............. 261
Ochlerotatus subdiversus (Martini 1926) ........ 263
10.3 Genus Culex Linnaeus .................................. 264
10.3.1 Subgenus Barraudius (Edwards) .................. 265
Culex modestus (Ficalbi 1889) .................. 265
Culex pusillus (Macquart 1850) ................. 267
10.3.2 Subgenus Culex (Linnaeus) ...................... 268
Culex brumpti (GaWiard 1931) ................... 269
Culex laticinctus (Edwards 1913) ............... 270
Culex mimetlcus (Noe 1899) ..................... 271
Culex perexiguus (Theobald 1903) ............... 273
Culex Pipiens Complex .......................... 275
Culex pipiens pipiens (Linnaeus 1758) ....... 275
Culex pipiens pipiens biotype molestus
(Forskal 1775) .............................. 277
Culex pipiens quinquefasciatus (Say 1832) ... 278
Culex torrentium (Martini 1925) ............. 279
Culex theileri (Theobald 1903) ................. 280
10.3.3 Subgenus Maillotia (Theobald) .................. 282
Culex hortensis (Ficalbi 1889) ................. 282
10.3.4 Subgenus Neoculex (Dyar) ....................... 283
Culex impudicus (Ficalbi 1890) ................. 284
Culex martinii (Medschid 1930) ................. 285
Culex territans (Walker 1856) .................. 286
10.4 Genus Culiseta Felt ................................... 288
10.4.1 Subgenus Allotheobaldia (Broelemann) ........... 289
Culiseta longiareolata (Marcquart 1838) ........ 289
10.4.2 Subgenus Culicella (Felt) ...................... 290
Culiseta fumipennis (Stephens 1825) ............ 291
Culiseta litorea (Shute 1928) .................. 292
Culiseta morsitans (Theobald 1901) ............. 294
Culiseta ochroptera (Peus 1935) ................ 296
10.4.3 Subgenus Culiseta (Felt) ....................... 297
Culiseta alaskaensis (Ludlow 1906) ............. 298
Culiseta annulata (Schrank 1776) ............... 299
Culiseta bergrothi (Edwards 1921) .............. 301
Culiseta glaphyroptera (Schiner 1864) .......... 303
Culiseta subochrea (Edwards 1921) .............. 305
10.5 Genus Coquillettidia (Dyar) ........................... 306
10.5.1 Subgenus Coquillettidia (Dyar) ................. 307
Coquillettidia buxtoni (Edwards 1923) .......... 307
Coquillettidia richiardii (Ficalbi 1889) ....... 308
10.6 Genus Orthopodomyia (Theobald) ........................ 310
Orthopodomyia pulcripalpis (Rondani 1872) ...... 310
10.7 Genus Uranotaenia (Lynch Arribalzaga) ................. 312
10.7.1 Subgenus Pseudoficalbia (Theobald) ............. 312
Uranotaenia unguiculata (Edwards 1913) ......... 312
Part III Identification Keys, Morphology, Ecology, and
Distribution of Important Vector and Nuisance
Species - World-Wide
11 Africa ..................................................... 317
11.1 Key to African Female Mosquitoes ...................... 318
11.2 Species Description ................................... 322
Anopheles (Cellia) funestus (Giles 1900) .............. 322
Anopheles Gambiae Complex ............................. 323
Anopheles (Cellia) gambiae s.s. (Giles 1902) ....... 324
Anopheles (Cellia) arabiensis (Patton 1905) ........ 325
Anopheles (Cellia) quadriannulatus s.l. (Theobald
1911) .............................................. 325
Anopheles (Cellia) bwambae (White 1985) ............ 326
Anopheles (Cellia) melas (Theobald 1903) ........... 326
Anopheles (Cellia) merus (Donitz 1902) ............. 326
Anopheles (Cellia) pharoensis (Theobald 1901) ......... 326
Culex (Culex) antennatus (Becker 1903) ................ 328
Culex (Culex) univittatus (Theobald 1901) ............. 329
Mansonia (Mansonioides) africana (Theobald 1901) ...... 330
Mansonia (Mansonioides) uniformis (Theobald 1901) ..... 330
12 Asia ....................................................... 333
12.1 Key to Asian Female Mosquitoes ........................ 334
12.2 Species Description ................................... 339
Anopheles (Anopheles) lesteri (Baisas and Hu 1936) .... 339
Anopheles (Anopheles) sinensis (Wiedemann 1828) ....... 340
Anopheles (Cellia) culicifacies (Giles 1901) .......... 341
Anopheles (Cellia) dims (Peyton and Harrison 1979) .... 342
Anopheles (Cellia) flavirostris (Ludlow 1914) ......... 343
Anopheles (Cellia) fluviatilis (James 1902) ........... 344
Anopheles (Cellia) maculatus (Theobald 1901) .......... 345
Anopheles (Cellia) minimus (Theobald 1901) ............ 346
Anopheles (Cellia) stephensi (Liston 1901) ............ 347
Anopheles (Cellia) sundaicus (Rodenwaldt 1925) ........ 348
Culex (Culex) tritaeniorhynchus (Giles 1901) .......... 349
13 Australia .................................................. 351
13.1 Key to Australian Female Mosquitoes ................... 351
13.2 Species Description ................................... 356
Anopheles (Cellia) farauti s.l. (Laveran 1902) ........ 356
Ochlerotatus (Mucidus) alternans (Westwood 1835) ...... 357
Ochlerotatus (Finlaya) notoscriptus (Skuse 1889) ...... 358
Ochlerotatus (Ochlerotatus) camptorhynchus (Thompson
1868) ................................................. 360
Ochlerotatus (Ochlerotatus) normanensis (Taylor
1915) ................................................. 360
Ochlerotatus (Ochlerotatus) sagax (Skuse 1889) ........ 362
Ochlerotatus (Ochlerotatus) theobaldi (Taylor 1914) ... 363
Ochlerotatus (Ochlerotatus) vigilax (Skuse 1889) ...... 364
Culex (Culex) annulirostris (Skuse 1889) .............. 365
Culex (Culex) sitiens (Wiedemann 1828) ................ 366
Coquillettidia (Coquillettidia) xanthogaster
(Edwards 1924) ........................................ 367
14 Central and South America .................................. 369
14.1 Key to Central and South American Female Mosquitoes ... 369
14.2 Species Description ................................... 374
Anopheles (Anopheles) calderoni (Wilkerson 1991) ...... 374
Anopheles (Anopheles) pseudopunctipennis (Theobald
1901) ................................................. 374
Anopheles (Anopheles) punctimacula (Dyar and Knab
1906) ................................................. 375
Anopheles (Nyssorhynchus) albimanus (Wiedemann
1820) ................................................. 376
Anopheles (Nyssorhynchus) aquasalis (Curry 1932) ...... 377
Anopheles (Nyssorhynchus) darlingi (Root 1926) ........ 378
Anopheles (Nyssorhynchus) nuneztovari (Gabaldon
1940) ................................................. 380
Haemagogus (Haemagogus) janthinomys (Dyar 1921) ....... 381
Ochlerotatus (Ochlerotatus) albifasciatus (Macquart
1838) ................................................. 382
Ochlerotatus (Ochlerotatus) scapularis (Rondani
1848) ................................................. 383
Culex (Culex) nigripalpus (Theobald 1901) ............. 384
Mansonia (Mansonia) titillans (Walker 1848) ........... 385
15 North America .............................................. 387
15.1 Key to North American Female Mosquitoes ............... 387
15.2 Species Description ................................... 391
Ochlerotatus (Ochlerotatus) sollicitans (Walker
1856) ................................................. 391
Ochlerotatus (Ochlerotatus) taeniorhynchus
(Wiedemann 1821) ...................................... 393
Psorophora (Grabhamia) columbiae (Dyar and Knab
1906) ................................................. 394
Psorophora (Janthinosoma) ferox (von Humboldt 1819) ... 395
Culex (Culex) restuans (Theobald 1901) ................ 396
Culex (Culex) salinarius (Coquillett 1904) ............ 397
Culex (Culex) tarsalis (Coquillett 1896) .............. 398
Coquillettidia (Coquillettidia) perturbans (Walker
1856) ................................................. 400
Part IV Control of Mosquitoes
16 Biological Control ......................................... 405
16.1 Introduction .......................................... 405
16.2 Predators ............................................. 406
16.2.1 Vertebrate Predators ........................... 406
16.2.2 Invertebrate Predators ......................... 410
16.3 Parasites ............................................. 415
16.3.1 Nematodes ...................................... 415
16.4 Pathogens ............................................. 416
16.4.1 Fungi .......................................... 416
16.4.2 Protozoa ....................................... 418
16.4.3 Bacteria ....................................... 419
16.4.4 Viruses ........................................ 431
17 Environmental Management of Mosquitoes ..................... 433
17.1 Introduction .......................................... 433
17.2 Environmental Management of Mosquitoes in Urban
Areas ................................................. 434
17.2.1 Denying Mosquito Access to Urban Areas,
and into Homes ................................. 434
17.2.2 Construction Sites ............................. 434
17.2.3 Water Storage Containers ....................... 434
17.2.4 Drainage Systems ............................... 435
17.2.5 Sewage and Waste-Water Processing .............. 435
17.2.6 Cemeteries ..................................... 436
17.2.7 Urban Sanitation ............................... 436
17.3 Environmental Management of Mosquitoes in Rural
Areas ................................................. 436
17.3.1 Agriculture .................................... 436
17.3.2 Natural Wetlands ............................... 437
17.4 Environmental Management of Mosquitoes and Human
Issues ................................................ 438
17.4.1 Community Participation ........................ 438
17.4.2 Regulations for Environmental Management
of Mosquitoes .................................. 439
18 Chemical Control ........................................... 441
18.1 History ............................................... 441
18.2 Insecticide ........................................... 442
18.2.1 Classification of Insecticides ................. 443
18.2.2 Insecticide Formulations ....................... 446
18.2.3 Insecticide Application Techniques ............. 448
18.3 Chemical Groups of Insecticides ....................... 449
18.3.1 Chlorinated Hydrocarbons ....................... 449
18.3.2 Organophosphates ............................... 452
18.3.3 Carbamates ..................................... 458
18.3.4 Pyrethroids .................................... 459
18.3.5 Insect Growth Regulators (IGRs) ................ 466
18.3.6 Novel Insecticide Classes ...................... 470
18.4 Management and Monitoring of Insecticide Resistance ... 471
18.4.1 Resistance Mechanisms .......................... 472
18.4.2 Resistance Surveillance ........................ 472
18.4.3 Resistance Management .......................... 474
19 Physical Control ........................................... 477
19.1 Introduction .......................................... 477
19.2 Physical Control of Immature Mosquitoes ............... 477
19.2.1 Oil ............................................ 477
19.2.2 Surface Films and Polystyrene Beads ............ 478
19.3 Physical Control of Adult Mosquitoes .................. 481
19.3.1 Other Technology: A Novel Technology or New
Tool ........................................... 481
19.4 Conclusions ........................................... 481
20 Genetic Control of Mosquitoes .............................. 483
20.1 Introduction .......................................... 483
20.2 Population Elimination Via the Sterile Insect
Technique (SIT) ....................................... 484
20.2.1 Introduction ................................... 484
20.2.2 Rearing and Sexing ............................. 484
20.2.3 Male Sterilization ............................. 485
20.2.4 SIT in Practice ................................ 485
20.3 Population Replacement ................................ 486
20.3.1 The principle of Population Replacement ........ 486
20.3.2 Refractoriness to Pathogens .................... 486
20.3.3 Genetic Drivers ................................ 487
20.4 Ethical, Legal and Social Implications of the Use
of Genetic Techniques for Mosquito Control ............ 488
20.4.1 Absence of Community Participation ............. 489
20.4.2 International Committee for Genetic Control
Work ........................................... 489
21 Personal Protection ........................................ 491
21.1 Introduction .......................................... 491
21.2 Impregnated Bednets ................................... 491
21.3 Repellents Against Mosquitoes ......................... 493
21.3.1 Repellents on Skin or Clothing ................. 493
21.3.2 Mosquito Coils ................................. 494
21.3.3 Vapourizing Mats ............................... 496
21.3.4 Liquid Vapourizers ............................. 496
22 Implementation and Integration of Mosquito Control
Measures ................................................... 499
22.1 Introduction .......................................... 499
22.2 Prerequisites for the Successful Implementation
of the IVM programme .................................. 501
22.2.1 Entomological Research (Monitoring) ............ 501
22.2.2 Mapping of the Breeding Sites .................. 503
22.2.3 Selection of Appropriate Tools ................. 504
22.2.4 Effective Dosage Assessment .................... 505
22.2.5 Design of the Control Strategy ................. 505
22.2.6 Training of Field Staff ........................ 506
22.2.7 Governmental Application Requirements .......... 506
22.2.8 Community Participation ........................ 506
22.2.9 Registration of Insecticides ................... 507
22.2.10 Routine Treatments ............................ 508
22.2.11 Public Information Systems .................... 509
References .................................................... 511
Subject Index ................................................. 553
Taxonomic Index ............................................... 565
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