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 (1913) transferred the species to genus Micraedes where it remained until
Edwards (1922) 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
(1968). It was classified as a subgenus of Culex
by Edwards (1930). 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 (1968) recognised that this character had evolved independently
in several unrelated groups, and elevated Aedinus
Lutz, as well as 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 (1914) 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
(1971) 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 & García (1968), 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 (1923), 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 Culex,
including Isostomyia (Dyar, 1918;
Edwards, 1932), Melanoconion (Dyar,
1925, 1928) and Tinolestes (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
(1968) 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 (1921), 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 (1918) appears to have been the first person
to regard Carrollia as a subgenus of Culex, followed by Aguilar (1931),
Edwards (1932) 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 (1932), 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, 1965; Bram, 1967) for a number of apparently related
Neotropical spcies 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 (1989) 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 (1932, 1941),
Belkin (1962), Forattini (1965), 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 (1912). Culiciomyia was reduced to subgeneric status in Culex by Edwards (1921). Edwards (1932)
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 (1977).
Subgenus Eumelanomyia
Theobald
Eumelanomyia was originally proposed as a distinct genus by Theobald (1909), 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 (1922) 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 (1910) recognised Protomelanoconion Theobald, 1909 as a distinct genus based on a
misidentified species of Culex which
he named Protomelanoconion fusca.
Edwards (1922) 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 (1930)
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 (1932) 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 or albiventris-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 his apicalis
group. No further changes were made to Edwards's classification until Sirivanakarn
(1971) 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 recognised, in addition to
subgenus Maillotia in addition to
subgenera Eumelanomyia and Neoculex. The current internal
classification of subgenus Eumelanomyia
was developed by Sirivanakarn (1971, 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, 1932) 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 (1925), 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 (1905), with uniformis Theobald, 1905 as its type and the
only included species. It was reduced to a subgenus of Culex by Edwards (1917). Edwards (1932) 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 (1977) modified the classifications of Edwards (1932,
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 (1977, 1978) 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 of Culex
hortensis Ficalbi, 1889) as its type and the only included
species. It was implicitly synonymised with Culex
by Edwards (1911) and placed in synonymy with Neoculex Dyar, 1905 by Edwards (1932), which was treated as a
subgenus of Culex. Sirivanakarn (1971)
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 (1903). The type
species, Culex atratus
Theobald, 1901, was subsequently designated by Dyar (1905). Dyar &
Knab (1906) 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 (1918a)
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 (1923) instated Gnophodeomyia 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 Mochlostyrax as 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-Lemaire) and antillummagnorum
Dyar of subgenus Micraedes Coquillett,
1906. Edwards (1932), 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
(1950) 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
(1968), 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 (1932)
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 (1950) 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 (1969) 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) 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 (1906), 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
(1918) elevated it to the subgeneric rank. Dyar (1928) synonymised bisulcatus with Culex (Melanoconion) americanus (Neveu-Lemaire, 1902), thus Micraedes became a synonym of Melanoconion., where it remained until
Edwards (1932) 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
(1969), following Foote (1954) and Belkin (1968), once again treated Micraedes as 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 of Culex, but it
seems the synonymy was never recognised. It has been treated as a subgenus of Culex since Dyar (1918). 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) excluded 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 (1905), 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 (1918). In
his comprehensive treatment of Culex,
Edwards (1932) 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 (1947) 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 (1932, 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 (1971) 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, 1913) 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, 1932, 1941; Belkin, 1962; Bram, 1967; Sirivanakarn,
1976), until Tanaka (2004) resurrected it from synonymy and validated it as a
subgenus to include bitaeniorhynchus
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 (1992), 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 (1906), with the new species latisquama Coquillett, 1906 as its type
and only included species. Howard et al.
(1915) synonymised Tinolestes with Culex, and Dyar (1918) 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 (1968) 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 Melanoconion
to 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 (1970), 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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