A tremendous amount of research has been done on Culex mosquitoes, but much of the taxonomic work has been directed primarily toward discriminating species and not on organising them into natural groups. The result is that the classification of Culex is based on the intuitive interpretation of morphological similarity and few attempts have been made to resolve phylogenetic relationships using modern techniques.
The genus has a cosmopolitan distribution and includes 768 species divided among 26 subgenera. The current system of subgeneric classification is based primarily on external adult characters, especially features of the male genitalia. The species of the larger subgenera are arranged in informal classifications that variously include Sections, Series, Groups, Subgroups and Complexes. The infrasubgeneric categories are often based on superficial similarities that may not reflect natural relationships. In general, the larger the group, the less likely it is to be a monophyletic assemblage of species.
Subgenus Acalleomyia Leicester
Acalleomyia was originally proposed as a distinct genus by Leicester (1908), with obscurus Leicester, 1908 as its type and the only included species. Edwards (1913c) transferred the species to genus Micraedes where it remained until Edwards (1922a) recognised Acalleomyia as a subgenus of Culex.
Subgenus Acallyntrum Stone & Penn
Acallyntrum was originally proposed as a subgenus of Culex by Stone & Penn (1948), with the new species perkinsi Stone & Penn, 1948 as the type species. Acallyntrum currently includes eight species that are considered to comprise a homogeneous group without subdivision.
Subgenus Aedinus Lutz
Aedinus was originally proposed as a distinct genus in 1904 but its authorship was not resolved until Belkin (1968b). It was classified as a subgenus of Culex by Edwards (1930a). Stone et al. (1959) included subgenus Aedinus Lutz along with subgenera Anoedioporpa, Micraedes and Tinolestes (as synonyms) in subgenus Aedinus Bourroul, 1904 based on the short maxillary palpus of males. Belkin (1968b) recognised that this character had evolved independently in several unrelated groups, and elevated Aedinus Lutz, as well Anoedioporpa, Micraedes and Tinolestes, to subgeneric rank in Culex.
Subgenus Afroculex Danilov
Afroculex was originally proposed as a subgenus of Culex by Danilov (1989), with Pseudohowardina lineata Theobald, 1912 as its type and the only included species. Edwards (1914b) transferred Pseudohowardina lineata to genus Culex and gave it the replacement name of pulchrithorax because lineatus was preoccupied by Cx. lineatus von Humboldt, 1819. The species was eventually placed in subgenus Neoculex, but its taxonomic position was open to question (Edwards, 1941). Its provisional placement in Neoculex continued until Sirivanakarn (1971a) transferred it to subgenus Maillotia. Danilov (1989) realised that Cx. lineatus von Humboldt is actually a species of genus Psorophora, as recorded by Knight & Stone (1977), reinstated lineatus as the valid name of the species and transferred it to the new subgenus Afroculex based on unique features of the adults and male genitalia. The immature stages remain unknown.
Subgenus Allimanta Casal & García
Allimanta was originally proposed as a subgenus of Culex by Casal & Garcia (1968a), with tramazayguesi Duret, 1954 as its type and the only included species. Culex tramazayguesi was originally described as a species of subgenus Culex.
Subgenus Anoedioporpa Dyar
Anoedioporpa was originally proposed as a subgenus of Culex by Dyar (1923e), with Culex conservator Dyar & Knab, 1906 as its type species. Most of the species currently included in the subgenus were previously assigned to other groups that have been recognised as subgenera of Culex, including Isostomyia (Dyar, 1918b; Edwards, 1932a), Melanoconion (Dyar, 1925f, 1928) andTinolestes (Lane, 1953). Stone et al. (1959) included Anoedioporpa, along with Tinolestes and Micraedes, in subgenus Aedinus, which at the time was attributed to Bourroul (1904). Belkin (1968b) recognised Anoedioporpa as a distinct taxon and restored it to subgeneric rank in genus Culex. Berlin & Belkin (1980) divided the subgenus into two groups, the Conservator Group, which currently includes 11 species, and the monobasic Restrictor Group.
Subgenus Barraudius Edwards
Barraudius was originally proposed as a subgenus of Culex by Edwards (1921d), with Culex pusillus Macquart, 1850 as its type species. Barraudius currently includes four species that are considered to comprise a homogeneous group without subdivision.
Subgenus Belkinomyia Adames & Galindo
Belkinomyia was originally proposed as a subgenus of Culex by Adames & Galindo (1973), with the new species eldridgei Adames & Galindo, 1973 as its type and the only included species.
Subgenus Carrollia Lutz
Carrollia was originally proposed as a distinct genus by Lutz (1905), with iridescens Lutz, 1905 as its type and the only originally included species. Dyar (1918b) appears to have been the first person to regard Carrollia as a subgenus of Culex, followed by Aguilar (1931), Edwards (1932a) and all later authors. The current internal division of the subgenus into two species groups, the Bihaicola and Iridescens Groups, the latter with two subgroups, the Urichii and Iridescens Subgroups, is attributable to Valencia (1973).
Subgenus Culex Linnaeus
The internal classification of subgenus Culex is in a chaotic condition. The subgenus has only been examined on a worldwide basis by Edwards (1932a), who divided it into two groups: the Sitiens Group (Old World) and the Pipiens Group (cosmopolitan). Both groups are highly complex assemblages and include species that do not readily fit into either group. Four additional species groups have been recognised subsequently: the Guiarti Group (Edwards, 1941) for several Afrotropical species, the Atriceps Group (Belkin, 1962) for three South Pacific species, the Coronator Group (Forattini, 1965a; Bram, 1967b) for a number of apparently related Neotropical species and the Duttoni Group (Harbach, 1988) for the unusual Afrotropical Cx. duttoni. Heinemann & Belkin (1977a, and later publications) recognised two groups in the Neotropical Region, the Declarator and Inflictus Groups, but did not indicate which species they include. Strickman (1990) made reference to the Declarator Group, but he also did not mention which species comprise the group. The internal classification of the subgenus presented here is based principally on information extracted and integrated from the works of Edwards (1932a, 1941), Belkin (1962), Forattini (1965a), Bram (1967a,b), Sirivanakarn (1976), Tanaka et al. (1979) and Harbach (1988), but the inclusion of many species in groups and subgroups, especially New World species, is problematic.
Subgenus Culiciomyia Theobald
Culiciomyia was originally proposed as a distinct genus by Theobald (1907). Designation of the type species, Culiciomyia inornata Theobald, 1907 (subjective synonym of Culex fragilis Ludlow, 1903), is attributed to Edwards (1912c). Culiciomyia was reduced to subgeneric status in Culex by Edwards (1921d). Edwards (1932a) recognised two species groups in the subgenus: group A, the Fragilis Group, with species in the Oriental, Indomalayan and Australasian Regions, and group B, the Nebulosus Group, with species restricted to the Afrotropical Region. Three additional groups have since been recognised for species in the Oriental Region: the Dispectus Group (Bram, 1969), Tricuspis Group (Harrison, 1987) and the Shebbearei Group, which is the name given here, based on nomenclatural priority, for the unnamed "group or complex" of Sirivanakarn (1977c).
Subgenus Eumelanomyia Theobald
Eumelanomyia was originally proposed as a distinct genus by Theobald (1909a), with inconspicuosa Theobald, 1908 as its type and the only included species. Eumelanomyia was described as being similar to Culiciomyia but the true identity of the type species was not known until Edwards (1922d) re-examined the specimens and identified them as Culex. The name inconspicuosa was thus found to be preoccupied, and Culex albiventris was proposed for the species. Theobald (1910c) recognised Protomelanoconion Theobald, 1909 as a distinct genus based on a misidentified species of Culex which he named Protomelanoconion fusca. Edwards (1922d) also found this name to be preoccupied and proposed the replacement name Culex horridus for the species. In addition to these corrections, Edwards suggested that both Eumelanomyia and Protomelanoconion should be treated as subgenera of Culex. Eight years later, Edwards (1930a) established Mochthogenes as a subgenus of Culex with Aedes malayi Leicester, 1908 as the designed type species. No further changes were made to the taxonomy of these groups until Edwards (1932a) examined the classification of Culex on a worldwide basis. As a result, Edwards He retained Mochthogenes as a subgenus and included Eumelanomyia, Protomelanoconion and a number of other species within subgenus Neoculex Dyar, 1905, which he divided into three groups: Group A (Neoculex or apicalis-group), Group B (Eumelanomyia oralbiventris-group) and Group C (Protomelanoconion or uniformis-group). In his later treatise on the Afrotropical Culicinae, Edwards (1941) recognised two additional groups, the pulchrithorax and rima groups, for species previously included in hisapicalis group. No further changes were made to Edwards's classification until Sirivanakarn (1971a) revised the classification of Neoculex to include nearly all species previously placed in subgenus Mochthogenes and a number of species previously included in subgenus Neoculex. Sirivanakarn synonymised Protomelanoconion and Mochthogenes with Eumelanomyia and recognised subgenus Maillotia in addition to subgenera Eumelanomyia and Neoculex. The current internal classification of subgenus Eumelanomyia was developed by Sirivanakarn (1971a, 1972).
Subgenus Kitzmilleria Danilov
Kitzmilleria was originally proposed as a subgenus of Culex by Danilov (1989), with moucheti Evans, 1923 as its type and the only included species. Culex moucheti was originally placed in the Pipiens Group of subgenus Culex, and was considered a member of the Decens Series (Edward, 1932a) until Danilov (1989) proposed subgenus Kitzmilleria based on its distinct adult, larval and pupal morphology.
Subgenus Lasiosiphon Kirkpatrick
Lasiosiphon was originally proposed as a subgenus of Culex by Kirkpatrick (1925a), with adairi Kirkpatrick, 1926, a replacement name for pluvialis Kirkpatrick, 1925, as its type and the only included species.
Subgenus Lophoceraomyia Theobald
Lophoceraomyia was originally proposed as a distinct genus by Theobald (1905h), with uniformis Theobald, 1905 as its type and the only included species. It was reduced to a subgenus of Culex by Edwards (1917). Edwards (1932a) divided the subgenus (as subgenus Lophoceratomyia) into three groups: Group A (minutissimus-group), Group B (Lophoceratomyia or fraudatrix-group) and Group C (Cyathomyia or mammilifer-group). Edwards later (1934, in Barraud, 1934), amalgamated Groups A and B and subdivided Group C. Colless (1965), however, preferred to recognise only two major groups, with the second divided into two subgroups. Sirivanakarn (1977a) modified the classifications of Edwards (1932a, 1934) and Colless (1965) to include three groups, the Fraudatrix, Mammilifer and Wilfredi Groups, based principally on structures of the antennae and genitalia of males. The division of these groups into subgroups and complexes by Sirivanakarn (1968, 1977a) forms the backbone of the current classification of the subgenus.
Subgenus Maillotia Theobald
Maillotia was originally proposed as a distinct genus by Theobald (1907), with pilifera Theobald, 1907 (subjective synonym ofCulex hortensis Ficalbi, 1889) as its type and the only included species. It was implicitly synonymised with Culex by Edwards (1911b) and placed in synonymy with Neoculex Dyar, 1905 by Edwards (1932a), which was treated as a subgenus of Culex. Sirivanakarn (1971a) removed Maillotia from synonymy to accommodate eight species divided between three groups, the Hortensis Group with three species, the monobasic Pulchrithorax Group for pulchrithorax Edwards, 1914, and the Seyrigi Group with four species. The subgenus currently includes two species group and an unplaced species. The Pulchrithorax Group was eliminated when Danilov removed pulchrithorax from Maillotia and proposed subgenus Afroculex to accommodate it.
Subgenus Melanoconion Theobald
Melanoconion was originally proposed as a distinct genus by Theobald (1903a). The type species, Culex atratus Theobald, 1901, was subsequently designated by Dyar (1905a). Dyar & Knab (1906d) synonymised Melanoconion with Culex and proposed Mochlostyrax as a distinct genus with caudelli Dyar & Knab, 1906 as its type species. Howard et al. (1915) considered both Melanoconion and Mochlostyrax as synonyms of Culex, and three years later Dyar (1918b) recognised them as separate subgenera of Culex. In the same paper, Dyar also proposed Choeroporpa as a subgenus of Culex, with anips Dyar, 1916 as its type species. Choeroporpa included most of the species that Dyar has previously placed in Culex or Mochlostyrax. In a second paper published in the same year, Dyar (1918b) proposed Helcoporpa as another subgenus of Culex, with menytes Dyar, 1918 as its type species. Five years later, Dyar (1923e) instated Gnophdeomyia Theobald, 1905 as a subgenus (previously synonymised with Culex by ?Brunetti, 1914) and proposed Anoedioporpa as a replacement name for subgenus Isostomyia Coquillett, 1906. Dyar (1928) made significant changes to the classification of New World Culex. He recognised Melanoconion and Mochlostyraxas subgenera and reduced the other nominal generic-level groups to informal sections: Choeroporpa, Helcoporpa and the newly proposed Dinoporpa became sections of Mochlostyrax, and Tinolestes Coquillett, Gnophodeomyia and Anoedioporpa became sections of Melanoconion, which also included americanus (Neveu-Lamaaire) and antillummagnorum Dyar to subgenus Micraedes Coquillett, 1906. Edwards (1932a), in his treatment of world Culicidae, reinterpreted the taxonomy of Melanoconion and Mochlostyrax. He considered Melanoconion as a subgenus with Gnophodeomyia, Asebeomyia Aiken, 1911, Tinolestes, Choeroporpa, Helcoporpa and Dinoporpa as its synonyms; restricted subgenus Mochlostyrax to include species included in the Mochlostyrax section of Dyar (1928); synonymised Anoedioporpa with subgenus Isostomyia (currently a valid genus in tribe Sabethini); and transferred americanus and antillummagnorum to subgenus Micraedes. During the same year Komp & Curry (1932) proposed Upsiloporpa as a new subgenus of Culex, with the new species haynei Komp & Curry, 1932 as its type and only included species. Komp (1935) found haynei to be conspecific with menytes, thus Upsiloporpa became another synonym of Melanoconion. Except for the transfer of ocellatus Theobald from subgenus Microculex Theobald to subgenus Melanoconion by Lane & Whitman (1943), Edwards's classification remained unchanged until Rozeboom & Komp (1950a) treated Melanoconion and Mochlostyrax as a single subgenus. Lane (1953) followed Rozeboom & Komp's classification but resurrected Tinolestes from synonymy with Melanoconion as a separated subgenus. A year later, Foote (1954) determined that Mochlostyrax was distinct based on larval morphology and considered it to be a subgenus separate from Melanoconion. Foote's separation of Mochlostyrax and Melanoconion prevailed until Belkin (1968b), Belkin et al. (1970) and Sirivanakarn (1983) considered Melanoconion and Mochlostyrax to form a single subgenus.
Dyar (1928) recognised four sections in subgenus Mochlostyrax, the Dinoporpa, Helcoporpa, Mochlostyrax and Choeroporpa sections, and four sections in subgenus Melanoconion, the Tinolestes, Gnophodeomyia, Melanoconion and Anoedioporpa sections. Edwards (1932a) recognised subgenus Mochlostyrax, without sections, and divided subgenus Melanoconion into three groups (Groups A, B and C) based on external features of adults. Rozeboom & Komp (1950a) disagreed with Edwards's classification and largely adopted Dyar's (1928) scheme based chiefly on features of the male genitalia for their concept of subgenus Melanoconion, which included Mochlostyrax and excluded Anoedioporpa. Hence, Rozeboom & Komp divided the subgenus into seven sections, namely the Choeroporpa, Dinoporpa, Gnophodeomyia, Helcoporpa, Melanoconion, Mochlostyrax and Tinolestes sections. Nearly two decades later, Galindo (1969) established the Spissipes Group based on male genitalia and larval characters and Duret (1969d) recognised the Ocellatus Group based on distinctive features of adults and male genitalia. Both groups were retained and redefined in the revised scheme of classification proposed by Sirivanakarn (1983).
Sirivanakarn (1983) distinguished three sections within the subgenus, the Melanoconion, Ocellatus and Spissipes Sections, and divided the Melanoconion and Spissipes Sections into Groups and Subgroups based principally on structural differences of the male genitalia, characteristics of the scaling on the head and scutum of adults and features of the larvae. Pecor et al. (1992) removed the Ocellatus Section from the subgenus, and it remains without subgeneric placement within genus Culex. More recently, Sallum & Forattini (1996) refined the Spissipes Section to include eight Groups and three Subgroups.
Subgenus Micraedes Coquillett
Micraedes was proposed as a distinct genus by Coquillett (1906c), with the new species bisulcatus Coquillett, 1906 as the type and only included species. Howard et al. (1915) synonymised Micraedes with Culex where it remained until Dyar (1918b) elevated it to the subgeneric rank. Dyar (1928) synonymised bisculatus with Culex (Melanoconion) americanus (Neveu-Lemaire, 1902), thus Micraedes became a synonym of Melanoconion., where it remained until Edwards (1932a) restored it to subgeneric rank. Lane (1953) synonymised it with subgenus Tinolestes and Stone et al. (1959) placed it in subgenus Aedinus Bourroul, 1904 along with Anoedioporpa and Tinolestes. Berlin (1969c), following Foote (1954) and Belkin (1968b), once again treated Micraedesas a distinct subgenus of Culex.
Subgenus Microculex Theobald
Microculex was proposed as a distinct genus by Theobald (1907), with argenteoumbrosus Theobald, 1907 [subjective synonym of Culex imitator Theobald, 1903] as the type and only included species. Brunetti (1914) regarded Microculex to be a synonym ofCulex, but it seems the synonymy was never recognised. It has been treated as a subgenus of Culex since Dyar (1918b). Lane & Whitman (1951) recognised four groups (series) of species known to occur in Brazil, but no attempt has been made to develop a classification for all species of the subgenus.
Subgenus Nicaromyia González Broche & Rodríguez R.
Nicaromyia was originally proposed as a subgenus of Culex by González Broche & Rodríguez R. (2001), with nicaroensis Duret, 1967 as its type and the only included species. Culex nicaroensis was originally described as a species of subgenus Melanoconion. Sallum & Forattini (1996) exculded it from Melanoconion and it remained without subgeneric placement until Nicaromyia was proposed to accommodate it.
Subgenus Neoculex Dyar
Neoculex was originally proposed as a distinct genus by Dyar (1905a), with Culex territans Walker, 1856 as its type species. It was regarded as a synonym of Culex by Brunetti (1914) and treated as a subgenus of Culex by Dyar (1918b). In his comprehensive treatment of Culex, Edwards (1932a) included Maillotia Theobald, 1907, Eumelanomyia Theobald, 1909 and Protomelanoconion Theobald, 1909 as synonyms of Neoculex and divided the subgenus into three groups: Group A (Neoculex or apicalis-group), Group B (Eumelanomyia or albiventris-group) and Group C (Protomelanoconion or uniformis-group). In his later work on the Afrotropical Culicinae, Edwards (1941) retained the albiventris and uniformis groups and split Group A into three groups, the apicalis, pulchrithorax and rima groups. King & Hoogstraal (1947b) followed this scheme and recognised a sixth group, Group F, for pedicellus King & Hoogstraal, 1947 and crassistylus Brug, 1934 from New Guinea. As indicated by Mattingly & Marks (1955) and Belkin (1962), the groups recognised by Edwards (1932a, 1941) and King & Hoogstraal (1947) give little idea of natural relationships because they are based on superficial characters that greatly overlap with characters exhibited by members of other subgenera of Culex. This is obvious from his treatment of Mochthogenes as a subgenus separated from the Protomelanoconion (i.e. uniformis group) of Neoculex based on the relative length of the male palpi. As pointed out by Bram (1969), these groups are so similar in the larval stage that they should be included in the same subgenus. With this as background, Sirivanakarn (1971a) proposed a reclassification of Neoculex based principally on structural differences observed in the genitalia of males. Sirivanakarn removed Eumelanomyia and Maillotia from synonymy with Neoculex, established them as separate subgenera of Culex, and synonymised Mochthogenes with Eumelanomyia. The restricted concept of Neoculex that resulted from these actions, including the recognition of three subordinate species groups, still stands today.
Subgenus Oculeomyia Theobald
Oculeomyia was proposed as a distinct genus by Theobald (1907), with sarawaki Theobald, 1907 (subjective synonym of Culex infula Theobald, 1901) as the type and only included species. Brunetti (1914) considered Oculeomyia to be a genus of "uncertain validity". Edwards (1911) synonymised sarawaki with Culex agar Giles, 1901, and subsequently (Edwards, 1913c) with Culex bitaeniorhynchus Giles, 1901, thus relegating Oculeomyia to synonymy with Culex. Oculeomyia remained in synonymy with Culex, specifically subgenus Culex as bitaeniorhynchus was classified as a member of the Bitaeniorhynchus Series/Subgroup of the Sitiens Group (Edwards, 1932a, 1941; Belkin, 1962; Bram, 1967a; Sirivanakarn, 1976), until Tanaka (2004) resurrected it from synonymy and validated it as a subgenus to include bitaeniohynchus and other species previously included in the Bitaeniorhynchus Subgroup.
Subgenus Phenacomyia Harbach & Peyton
Phenacomyia was originally proposed as a subgenus of Culex by Harbach & Peyton (1992b), with corniger Theobald, 1903 as its type species. Prior to the recognition of Phenacomyia, Cx. corniger and its two related species, Cx. lactator and Cx. airozai, were included in subgenus Culex.
Subgenus Phytotelmatomyia Rossi & Harbach
Phytotelmatomyia was originally proposed as a subgenus of Culex by Rossi & Harbach (2008), with renatoi Lane & Ramalho, 1960 as its type species. Prior to the recognition of Phytotelmatomyia, Cx. renatoi and its related species were included in subgenus Culex.
Subgenus Sirivanakarnius Tanaka
Sirivanakarnius was originally proposed as a subgenus of Culex by Tanaka (2004), with boninensis Bohart, 1957 as its type and the only included species. Culex boninensis was regarded as a member of the Sitiens Group of subgenus Culex until Tanaka (2004) established subgenus Sirivanakarnius based on distinct characters of the adults and male genitalia.
Subgenus Tinolestes Coquillett
Tinolestes was proposed as a distinct genus by Coquillett (1906c), with the new species latisquama Coquillett, 1906 as its type and only included species. Howard et al. (1915) synonymised Tinolestes with Culex, and Dyar (1918b) resurrected it to subgeneric rank. Dyar (1928) placed latisquama in subgenus Melanoconion, and as a consequence Tinolestes became a synonym of Melanoconion. Lane (1953) restored Tinolestes to subgeneric rank and synonymised subgenera Micraedes Coquillett, Isostomyia Coquillett and Anoedioporpa Dyar with it. Stone et al. (1959) included Tinolestes, along with Micraedes and Anoedioporpa, in subgenus Aedinus Bourroul, 1904 based on the short palpus in males, but Belkin (1968b) noted that this character occurs independently in several obviously unrelated groups and reinstated Tinolestes as a monobasic subgenus of Culex. Two species, breviculus Senevet & Abonnenc, 1939 and cauchensis Floch & Abonnenc, 1945 were transferred from subgenus Melanoconionto subgenus Tinolestes by Sirivanakarn (1983).
Subgenus uncertain
Five species of the Ocellatus Group of Sirivanakarn (1983), i.e flochi Duret, 1969, inornata (Theobald, 1905), nigrimacula Lane & Whitman, 1943, ocellatus Theobald, 1903 and punctiscapularis Floch & Abonnenc, 1946, were removed from subgenus Melanoconion by Pecor et al. (1992), and are retained in genus Culex without subgeneric placement.
Mattingly & Marks (1955) noted that Pseudoskusea cairnsensis Taylor, 1905 was a species of Culex, probably of subgenus Lophoceraomyia, but its subgeneric placement must await a revision of the Australian species of that subgenus.
According to Belkin (1970b), the identity of Gnophodeomyia inornata Theobald, 1905 "may never be determined with certainty as the type series consists of females only".
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