Subfamily Culicinae Meigen, 1818
Culex Linnaeus, 1758. [Meigen (1818) introduced the first family-group name (Culiciformes) that established the date of priority for Culicidae and its coordinate subfamily name Culicinae (International Commission on Zoological Nomenclature, 1999: Article 36.1).]
Culicinae is the largest subfamily of mosquitoes, containing 3,117 species in 110 genera and two groups of incertae sedis, i.e. 'Aedes' sensu auctorum, 'Ochlerotatus' sensu auctorum. The genera are grouped into 11 tribes as shown in the list below. Species belonging to this subfamily are referred to as 'culicines', however species of Aedini are sometimes referred to as 'aedines' and species of the tribe Sabethini are known as 'sabethines'.
Aedeomyiini: Aedeomyia (7 species).
Aedini: Abraedes (1), Acartomyia (3), Aedes (12), 'Aedes' sensu auctorum (1), Aedimorphus (67), Alanstonea (2), Albuginosus (9), Armigeres (58), Ayurakitia (2), Aztecaedes (1), Belkinius (1), Bifidistylus (2), Borichinda (1), Bothaella (6), Bruceharrisonius (9), Cancraedes (10), Catageiomyia (28), Catatassomyia (1), Christophersiomyia (5), Collessius (9), Cornetius (1), Dahliana (3), Danielsia (3), Dendroskusea (5), Diceromyia (8), Dobrotworskyius (7), Downsiomyia (31), Edwardsaedes (3), Elpeytonius (2), Eretmapodites (48), Finlaya (36), Fredwardsius (1), Georgecraigius (3), Geoskusea (10), Gilesius (2), Gymnometopa (1), Haemagogus (28), Halaedes (3), Heizmannia (40), Himalaius (2), Hopkinsius (7), Howardina (34), Huaedes (3), Hulecoeteomyia (18), Indusius (1), Isoaedes (1), Jarnellius (5), Jihlienius (3), Kenknightia (12), Kompia (1), Leptosomatomyia (1), Levua (1), Lewnielsenius (1), Lorrainea (5), Luius (1), Macleaya (11), Molpemyia (3), Mucidus (14), Neomelaniconion (28), Nyctomyia (2); Ochlerotatus (199), 'Ochlerotatus' sensu auctorum (70), Opifex (2), Paraedes (9), Patmarksia (13), Paulianius (8), Petermattinglyius (5), Phagomyia (16), Polyleptiomyia (2), Pseudarmigeres (5), Psorophora (49), Rampamyia (3), Rhinoskusea (4), Sallumia (2), Scutomyia (9), Skusea (4), Stegomyia (128), Tanakaius (2), Tewarius (4), Udaya (3), Vansomerenis (3), Verrallina (95), Zavortinkius (11), Zeugnomyia (4). NOTE: Only the 10 genera indicated in boldface are recognised in the traditional classification of Aedini. The others are recognised as subgenera within a very large polyphyletic genus Aedes.
Culicini: Culex (789), Deinocerites (18), Galindomyia (1), Lutzia (9).
Culisetini: Culiseta (39).
Ficalbiini: Ficalbia (8), Mimomyia (45).
Hodgesiini: Hodgesia (11).
Mansoniini: Coquillettidia (58), Mansonia (25).
Orthopodomyiini: Orthopodomyia (36).
Sabethini: Isostomyia (4), Johnbelkinia (3), Kimia (5), Limatus (9), Malaya (12), Maorigoeldia (1), Onirion (7), Runchomyia (8), Sabethes (42), Shannoniana (3), Topomyia (68), Trichoprosopon (14), Tripteroides (123), Wyeomyia (140).
Toxorhynchitini: Toxorhynchites (92).
Uranotaeniini: Uranotaenia (272).
In a catalogue of the mosquitoes of Japan, Tanaka (2014, 2018) ranked the tribes Aedini, Culicini, Culisetini, Ficalbiini, Mansoniini and Orthopodomyiini as subtribes of a more encompassing tribe Culicini. In addition to subtribes Aedina, Culicina, Culisetina, Ficalbiina, Mansoniina and Orthopodomyiina, he created subtribe Heizmanniina for the genus Heizmannia. His subtribe Aedina included the genera Aedes, Aedimorphus, Armigeres, Bruceharrisonius, Collessius, Downsiomyia, Edwardsaedes, Empihals, Geoskusea, Hopkinsius, Hulecoeteomyia, Neomelaniconion, Ochlerotatus, Phagomyia, Stegomyia, Tanakaius and Verrallina for aedine species that occur in Japa. Newly recognised subgeneric taxa included Aedimorphus subgenera Ecculex and Lepidotomyia (currently synonyms of Aedimorphus), Bruceharrisonius subgenera Bruceharrisonius and koreicoides-group (informal groups were treated as subgenera without formal names); Phagomyia subgenera Phagomyia and oreophila-group; Ochlerotatus subgenera communis-group, dorsalis-group, excrucians-group, impiger-group, punctor-group, sticticus-group and Woodius; Stegomyia subgenus Quasistegomyia (currently a synonym of Stegomyia). The subtribe Culicina included the genera Culex and Lutzia, and Culex included the subgenera Barraudius, Culex, Culiciomyia, Lophoceraomyia, Mochthogenes (currently a synonym of Eumelanomyia), Neoculex, Oculeomyia, Protomelanoconion (also currently a synonym of Eumelanomyia) and Sirivanakarnius for species that occur in Japan As expected, the subtribes Culisetini, Ficalbiina, Mansoniina and Orthopodomyiina included the genera that are currently included in the coordinate tribal groups Culisetini, Ficalbiini, Mansoniini and Orthopodomyiini, respectively. However, the validity (availability) of the subtribal names is doubtful. The availability of two of the names, Aedina Lahille, 1904 and Culisetina Maslov, 1964, was not mentioned, and no action was taken to reinstate and define them. The other subtribal names were introduced without explanation or justification, and are herewith treated as nomina nuda because family-group names published after 1930 must be accompanied by a verbal description or definition of characters that purport to distinguish them (International Commission on Zoological Nomenclature, 1999: Articles 13.1 and 13.2).
Culicine females have maxillary palpi that are usually much shorter than the proboscis. Males have long palpi with numerous long setae, but they are not swollen apically like those of anophelines. The scutellum has three lobes with setae confined to each lobe. The wing veins are usually entirely dark-scaled, but speckles or patches of white or yellow scales are present in some species. Some species of the genera Aedeomyia, Culex (subgenus Culex, Mimeticus Group), Finlaya, Lutzia (subgenus Lutzia), Orthopodomyia and Uranotaenia have spots of pale scales similar to those of many Anopheles. An emargination on the posterior margin of the wing opposite vein CuA (cubitus anterior) is found only in species of tribe Toxorhynchitini. The abdominal terga and sterna are densely covered with scales, which distinguishes them immediately from almost all anophelines. Adult culicines stand with the body parallel to the surface on which they are resting. All culicine larvae have a respiratory siphon, the dorsal and ventral abdominal setae arise separately and usually without basal sclerites, and the mouth brushes are composed of numerous, usually slender filaments (except genera Lutzia, Mucidus and Toxorhynchites). See Culicidae.
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. See Harbach & Kitching (1998) and Harbach (2007).
Culicine larvae occupy a wide variety of aquatic habitats ranging from small, temporary to large, permanent bodies of water. The majority of species occur in collections of ground water, but many species inhabit a variety of plant cavities (phytotelmata) and artificial containers of different sizes. Most larvae use their siphons to suspend themselves from the water surface, but larvae of Mansonia, Coquillettidia and some Mimomyia insert the modified tips of their siphons into plants to obtain air. Most species feed by extracting suspended particulate matter from the water with filamentous mouth brushes, but grazing on submerged surfaces commonly occurs and the larvae of some species are facultative or obligate predators of other mosquito larvae and other kinds of insects. Culicine adults are active during the day or night, but the majority of species are probably active during the daylight hours, including crepuscular periods.
The genera Culex, Mansonia and Stegomyia are the most important from a medical point of view because they contain the majority of culicine mosquitoes that transmit helminths and arboviruses. Species of many genera are annoying pests of humans and domestic animals.
Members of the subfamily are found in all zoogeographic regions of the world, but the majority of species occur in tropical regions.
Barraud, 1934 (southern Asia); Edwards, 1941 (adults, Afrotropical Region); Hopkins, 1952 (larvae, Afrotropical Region); Lane, 1953 (Neotropical Region); Thurman, 1959 (Thailand); Belkin, 1962 (taxonomy, South Pacific); Cova Garcia et al., 1966 (Venezuela); Delfinado, 1966 (Philippines); Tanaka et al., 1979 (taxonomy, Japan); Ibrahim et al., 1983 (larvae, keys Iraq); Lee et al., 1984, 1987, 1988a, 1988b, 1988c, 1989a, 1989b, 1989c (Australasian Region); Hudson & Abul-Hab, 1987 (adults, keys, Iraq); Service, 1990 (Afrotropical Region); Lu Baolin et al., 1997 (China); Harbach & Kitching, 1998 (phylogeny); Becker et al., 2003, 2010 (Europe); Azari-Hamidian & Harbach, 2009 (keys, Iran); Reidenbach et al., 2009 (phylogeny).