Family Culicidae Meigen, 1818

Type genus: 

Culex Linnaeus, 1758. [Culiciformes Meigen, 1818 is the earliest family-group name that established the date of priority for Culicidae, not Billberg, 1820 (as Culicides).]

Classification: 

Mosquitoes, family Culicidae, belong to the order Diptera, the two-winged flies. The family is a large and abundant group which occurs throughout temperate and tropical regions of the world, and well beyond the Arctic Circle. The family includes 3,726 extant species (see Valid Species List) classified in two subfamilies and 113 genera. The subfamily Anophelinae has three genera and Culicinae has 110 genera segregated into 11 tribes. Many species traditionally included in genera Aedes and Ochlerotatus are regarded as incertae sedis in 'Aedes' and 'Ochlerotatus' sensu auctorum. NOTE: Regarding the traditional classification of tribe Aedini, the family and subfamily Culicinae include 41 and 38 genera, respectively.

Characteristics: 

Mosquitoes are slender, long-legged insects that are easily recognised by their long proboscis and the presence of scales on most parts of the body. Larvae are distinguished from other aquatic insects by the absence of legs, the presence of a distinct head bearing mouth brushes and antennae, a bulbous thorax that is wider than the head and abdomen, posterior anal papillae and either a pair of respiratory openings (subfamily Anophelinae) or an elongate siphon (subfamily Culicinae) borne near the end of the abdomen. Mosquitoes are usually, and most reliably, identified as final (fourth) instar larvae and adults. See Anophelinae and Culicinae.

Phylogenetic relationships: 

Family Culicidae is monophyletic but deeper relationships are largely unresolved. Subfamily Anophelinae is a monophyletic lineage basal to all other Culicidae, and genus Chagasia is a monophyletic lineage basal to other Anophelinae. Subfamily Culicinae is not demonstrably monophyletic in relation to genus Toxorhynchites. Tribes Aedini, Culicini and Sabethini are monophyletic, but the monophyly of the other tribes has not been tested and their phyletic relationships are uncertain. Although based on few species, the analysis of COX1, COX2, NADH4, NADH5 and CYOB gene sequences of mtDNA in the study de Oliveira Aragão et al. (2018) support the monophyly of the Anophelinae, Aedini, Culicini and Sabethini.See Harbach & Kitching (1998) and Harbach (2007).

Bionomics and disease relations: 

Mosquito larvae occupy a spectrum of aquatic environments. The majority of larvae feed on suspended particulate matter and microorganisms which they extract from the water with filamentous mouth brushes. Other species are obligatory or facultative predators which capture and feed largely on the immature stages of other mosquitoes by means of modified mouth brushes or grasping mandibles or maxillae. Some larvae resort to scavenging or cannibalism when food is scarce. The larvae of most mosquitoes obtain oxygen from the atmosphere by coming to the water surface. All species of Mansonia and Coquillettidia and some species of Mimomyia obtain oxygen from the air vessels of aquatic plants, which they pierce with a specialised siphon. Aedeomyia species apparently use their enlarged antennae for respiration. Some species have greatly enlarged anal papillae that are well supplied with tracheae, and these species seldom come to the surface and probably obtain dissolved oxygen from the water.

Since mosquitoes are delicate insects, they are always found where the air is relatively cool and the humidity is high. Many species live within a few meters of the ground, whereas many forest species occur primarily in the forest canopy. Vertical distribution is largely dependent on feeding preferences. All males and the females of many species feed exclusively on plant liquids, including nectar, honeydew, fruit juices and exudates. Females of a great number of species feed on the blood of living animals, but some that are normally blood feeders may produce eggs without a blood meal. Warm-blooded vertebrates are a common source of blood for most species, but many species also attack cold-blooded animals such as snakes, turtles, toads, frogs, and other insects, including nymphal cicadas, lepidopterous larvae and mantids. A species of Uranotaenia, Ur. sapphirina, is known to feed on earthworms and leeches. The time of flight and feeding activity is usually quite specific for most species. Some species are active at night (nocturnal) or twilight (crepuscular) while others are active during the daylight hours (diurnal).

Mosquitoes are important because the females of many species are bloodsucking, they annoy humans and other animals, and they may transmit the pathogens that cause human and animal diseases. The pathogens transmitted by mosquitoes include viruses (arboviruses), filarial worms (helminths) and protozoa. Mosquitoes are the indirect cause of more morbidity and mortality among humans that any other group of organisms.

Distribution: 

Culicidae is a large and abundant group that occurs throughout temperate and tropical regions of the world, and well beyond the Arctic Circle. Mosquitoes are most diverse in tropical forest environments.

Principal references: 

Barraud, 1934 (southern Asia); Bates, 1949 (biology); Lane, 1953 (Neotropical Region); Mattingly & Knight, 1956 (Arabia); Belkin, 1962 (classification, South Pacific); DuBose & Curtin, 1965 (keys, Mediterranean area); Cova-Garcia et al., 1966 (Venezuela); Belkin, 1968a (New Zealand); Belkin et al., 1970 (Jamaica); Gillett, 1971 (biology); Mattingly, 1971 (generic keys, world); Horsfall, 1972 (bionomics); Gutsevich et al., 1974 (former USSR); Knight & Stone, 1977 (catalogue); Knight, 1978 (catalogue supplement); Tanaka et al., 1979 (Japan); Wood et al., 1979 (Canada); Degallier & Claustre, 1980 (French Guiana,species list, taxonomic notes); Lee et al., 1980 (Australasian Region); Darsie & Ward, 1981, 2005 (keys, North America); Lu & Li, 1982 (China); Clark-Gil & Darsie, 1983 (Guatemala); Ward, 1984, 1992 (catalogue supplements); Danilov, 1985a, 1985b (keys, Afghanistan); Darsie, 1985 (keys, Argentina); Gaffigan & Ward, 1985 (catalogue supplement); Lounibos et al., 1985 (bionomics); Mitchell & Darsie, 1985 (Argentina); Cranston et al., 1987 (Britain); Laird, 1988 (larval bionomics); Debenham et al., 1989 (Australasian Region); Darsie & Pradhan, 1990 (Nepal); Clements, 1992, 1999, 2012 (biology); Service, 1990 (Afrotropical Region); Service, 1993a, 1993b (bionomics, taxonomy); Rattanarithikul & Panthusiri, 1994 (keys, medically important species, Thailand); Harbach & Sandlant, 1997 (genera, keys); Harbach & Kitching, 1998 (phylogeny); Hutchings et al., 2002 (Querari, Amazonas, Brazil, species list); Becker et al., 2003, 2010 (Europe); Rattanarithikul et al., 2005 (genera, Thailand); Harbach, 2007 (systematics); Azari-Hamidian & Harbach, 2009 (keys, Iran); Reidenbach et al., 2009 (phylogeny); Severini et al., 2009 (Italy, A keys, morphology, biology & distribution of species); Borkent, 2012 (pupal morphology); Jeffery et al., 2012 (Peninsula Malaysia, keys to adults, references, checklist, taxonomic notes); Tuten et al., 2012 (pyloric armature, phylogeny); Tantely et al., 2016 (Madagascar, checklist); Webb et al., 2016 (Australia, biology, ecology, taxonomy); Lemine et al., 2017 (Mauritania); Coetzee, 2017 (Afrotropical Region, morphology, bionomics, A,L keys to genera); Namgay et al., 2018 (Bhutan, checklist); Ruzzante et al., 2019 (evolutionary genomics); Tsuda, 2019 (Japan, illustrated genus and species keys for adults, distributions, bionomics); Yeo et al., 2019 (Singapore, adult females, key to genera); da Silva et al., 2020 (Neotropics, family-groups, phylogeny); Talaga et al., 2020 (French Guiana, species described from the country, taxonomic notes); Talaga et al., 2020 (French Guiana, species originally described from the country, type data); van der Beek et al., 2020 (Dutch Leeward Islands, key for adults and larvae, ecology, distribution); Wilkerson et al., 2021 (World, biology, taxonomy, systematics, morphology, medical importance, catalogue); Nugroho & Mujiyono, 2021 (Indonesia, keys to genera); Bromley-Schnur, 2021 (Israel and surrounding areas, A,L keys, distributions, bionomics, history); Merabti et al., 2021 (Algeria, checklist, taxonomy); Ndiaye et al., 2021 (Burkina Faso, Chad, Gambia, Mauritania, Niger, Senegal, species distributions); Robert et al., 2022 (Burkina Faso, Cape Verde, Gambia, Mali, Mauritania, Niger, Senegal and Chad, keys for adults and larvae); Harbach & Wilkerson, 2023 (exclusion of subspecies from culicid classification);Chen  et al., 2023 (Eastern Asia, family-groups, phylogeny); Russell & Burkot, 2023 (Pacific Island countries and territories, species lists, distributions, bionomics);Soghigian et al., 2023 (World, family-groups, phylogeny); Dawah et al., 2023 (Saudi Arabia, distributions, notes, medical importance, keys for adult females).

Subfamilies: 
Scratchpads developed and conceived by (alphabetical): Ed Baker, Katherine Bouton Alice Heaton Dimitris Koureas, Laurence Livermore, Dave Roberts, Simon Rycroft, Ben Scott, Vince Smith