Subgenus Culex Linnaeus, 1758

Type species: 

Culex pipiens Linnaeus, 1758.

Classification: 

Subfamily Culicinae, genus Culex. Subgenus Culex, as presently defined, includes 215 extant species and one extinct (fossil) species. Subgenus abbreviation – Cux.

Characteristics: 

ADULTS – Generally medium-sized mosquitoes; decumbent scales of vertex predominantly narrow; eyes narrowly separated or contiguous above antennae; antenna slightly longer than proboscis, antenna of males without tufts of specialised setae or scales; maxillary palpus of females with 3 palpomeres, palpus of males with 5 palpomeres, usually distinctly longer than proboscis, palpomere 3 with row of ventral setae and variable number of longer ventrolateral setae, palpomeres 4 and 5 upturned and usually with numerous setae; proboscis usually as long or slightly longer than forefemur, with or without median pale ring or pale scaling on ventral surface, proboscis of males with or without median ventral cluster of setae but usually with false joint before mid-length; scutal scaling varied; acrostichal setae present in more or less double row, sometimes short; dorsocentral setae, scutal fossal setae, antealar setae and supraalar setae well developed; thoracic pleura usually with extensive scaling, usually present at least on antepronotum, postpronotum, upper proepisternum, mesokatepisternum and mesepimeron, present or absent on lower proepisternum, anteprocoxal membrane, postspiracular area and prealar area; pleural setae varied, postspiracular setae rarely present, lower mesepimeral setae present or absent; wing most often entirely dark-scaled, sometimes with scattered pale scales or distinct pale spots; legs varied, tarsi with or without pale bands; ungues small, equal and simple in females, outer unguis of fore- and midlegs toothed and larger than inner unguis in males; pulvilli well developed; abdominal terga usually with basal pale bands and basolateral pale spots, sometimes with apical pale bands, apicolateral pale patches or entirely dark. MALE GENITALIA – Ninth tergal lobes usually short and broad, with 1 or 2 rows of setae; gonocoxite with or without scales; subapical lobe not distinctly divided, setae a-c rod-like, setae d-f and seta h varied, 1 or all of setae d-f sometimes absent, seta g foliform or simple; gonostylus generally simple, curved and tapered; gonostylar claw weak, subapical, usually shoehorn-shaped; phallosome complex, varied, lateral plate usually with distinct inner and outer divisions bearing variously developed teeth and processes; paraproct with crown of numerous spicules of varying size and character, basal sternal process varied, sometimes absent; cercal setae present. LARVAE – Antenna shorter or as long as head; seta 1-A usually strongly developed; setae 2,3-A single, usually subapical, sometimes apical; median labral plate usually distinctly delimited from dorsal apotome; seta 1-C slender or stout; setae 1–3-P subequal and long, usually single, seta 3-P rarely distinctly shorter than setae 1,2-P; abdominal chaetotaxy varied; setae 6-I,II and 7-I strongly developed; seta 6-IIIVI moderately to strongly developed; seta 7-II large like seta 7-I or small like seta 7-IIIV; seta 1-III-VII varied; comb varied from numerous fringed scales to a few strong spines; siphon varied, short and stout to long and slender; pecten varied from a few to many spines, denticulation varied; seta 1-S in 3–7 posterolateral pairs, 1 or more distal elements sometimes out of line; seta 2-S single, sometimes hook-like and large; saddle usually complete, rarely incomplete; seta 1-X varied, not very long; setae 2,3-X strongly developed, seta 2-X branched, seta 3-X single; ventral brush (seta 4-X) comprised of 6 or 7 pairs of setae, all inserted on grid. PUPAE – Trumpet varied in length, generally cylindrical or distally flared with large pinna; meatus usually without slit; tracheoid area variable; cephalothoracic setae varied in development, all shorter than trumpet; abdominal chaetotaxy varied, seta 1-III–VII and 5-IV–VI varied in development; seta 2-III–VI single, very small, always inserted mesad of seta 1, seta 2-VII usually inserted laterad of seta 1, sometimes variable in position; seta 6-I,II usually distinctly longer than seta 7-I,II; paddle generally broad and oval, apex rounded, slightly produced or slightly emarginate, outer margin minutely spiculate proximally; seta 1-Pa and seta 2-Pa small and usually single, seta 2-Pa sometimes absent. See genus Culex

Phylogenetic relationships: 

Culex is almost certainly polyphyletic and will remain a "taxon of convenience" until the monophyly, phylogeny and classification of the included taxa and their relationships with other genera are resolved. Subgenera Phenacomyia, Oculeomyia, Phytotelmatomyia and Sirivanakarnius have been placed among species of subgenus Culex in both morphological and molecular studies that included species of these subgenera (Deus, 2009; Vesgueiro et al., 2011; Demari-Silva et al., 2011; Harbach et al., 2012; Laurito & Almirón, 2013). Analyses of the mitochondrial genomes of four species of the Coronator Group suggest that the ATP6, ATP8 and NADH5 genes may be useful for resolving phylogenetic relationships within the subgenus (Demari-Silva et al., 2015).

Harbach et al. (2017) conducted a phylogenetic analysis of the subgenus using implied and equally weighted parsimony methods to analyse a data set comprised of 286 characters of the larval, pupal and adult stages of 150 species of the subgenus and an outgroup of 17 species. The analyses largely failed to resolve relationships among the species and the informal groups in which they are currently placed; thus, the internal classification of the subgenus based on the intuitive assessment of morphological similarities and differences does not seem, in large part, to reflect evolutionary relationships. It is notable that the phylogenetic analysis of Sun et al. (2019) based on the nucleotide sequences of 13 protein-coding genes obtained from the complete mitochondrial genomes of 10 species of Culex strongly supported the suppositions of Edwards (1932), Sirivanakarn (1976) and Harbach et al. (2017) that the Sitiens Group possibly should be recognized as a genus-group taxon. It is also interesting to note that their study did not support the monophyly of the Pipiens Group, which was recovered as a polyphyletic group in the studies of Laurito & Almirón (2013) and Harbach et al. (2017), but did support the monophyly of the Coronator Group.

Bionomics and disease relations: 

The immature stages typically occur in ground-water habitats, but several species are frequently found in large artificial containers. A number of species inhabit rock holes, crab holes, tree holes, bamboo and the leaf axils of plants. Many species are notorious pests of humans and domesticated animals. Females are primarily active during crepuscular or nocturnal periods. The hosts of most species are unknown but probably principally include birds and mammals.

The subgenus includes most of the medically important and pest species of Culicini. Many species vector pathogens that cause human diseases, including Japanese encephalitis in the Oriental Region, several encephalitic diseases in North America, Murray Valley encephalitis and Ross River fever in Australia, West Nile fever in Africa, Europe and North America, and Rift Valley fever in Africa. A few species are important vectors of Wuchereria bancrofti, and some species may be involved in the transmission of Brugia malayi.

Distribution: 

Cosmopolitan.

Principal references: 

Belkin, 1962 (South Pacific, keys, taxonomy, bionomics, distributions); Bram, 1967 (Thailand, keys, taxonomy, bionomics, distributions); Bram, 1967 (Neotropical Region, keys, taxonomy, distributions); Sirivanakarn, 1976 (Oriental Region, keys, taxonomy, bionomics, distributions); Harbach, 1988 (southwestern Asia and Egypt, taxonomy, bionomics, distributions); Lee et al., 1989 (Australasian Region, keys, literature, distributions, bionomics); Laurito & Almirón, 2013 (morphology, phylogenetic relationships of species of the subgenus in Argentina); Harbach et al., 2017 (World, morphology, phylogeny): Sun et al. 2019 (phylogenetic relationships based on mitochondrial genomes); González et al., 2023 (Neotropical subgenera of Culex, comparative morphology, molecular associations).

Species: 
abnormalis Lane, 1936
abyssinicus Edwards, 1941
acharistus Root, 1927
aenescens Edwards, 1941
alani Forattini, 1965
alienus Colless, 1957
alis Theobald, 1903
ameliae Casal, 1967
andersoni Edwards, 1914
annulirostris Skuse, 1889
annuliventris (Blanchard, 1852)
antennatus (Becker, 1903)
aquarius Strickman, 1990
archegus Dyar, 1929
argenteopunctatus (Ventrillon, 1905)
articularis Philippi, 1865
asteliae Belkin, 1968
astridianus de Meillon, 1942
atriceps Edwards, 1926
australicus Dobrotworsky & Drummond, 1953
bahamensis Dyar & Knab, 1906
banksensis Maffi & Tenorio, 1977
barraudi Edwards, 1922
beta Séguy, 1924
bhutanensis Somboon, Namgay & Harbach, 2021 (in Somboon et al., 2021)
bickleyi Forattini, 1965
bidens Dyar, 1922
bihamatus Edwards, 1926
bilineatus Theobald, 1903
bonneae Dyar & Knab, 1919
brami Forattini, Rabello & Souza Lopes, 1967
brethesi Dyar, 1919
brevispinosus Bonne-Wepster & Bonne, 1920
brumpti Galliard, 1931
bukavuensis Wolfs, 1947
bwambanus Edwards, 1941
calurus Edwards, 1935
camposi Dyar, 1925
carcinoxenus de Oliveira Castro, 1932
carleti Brunhes & Ravaonjanahary, 1971
castelli Hamon, 1957
chidesteri Dyar, 1921
chitae Duret, 1967
chorleyi Edwards, 1941
comorensis Brunhes, 1977
coronator Dyar & Knab, 1906
covagarciai Forattini, 1965
crinicauda Edwards, 1921
curvibrachius Angulo, 1993 (in Angulo & Olivares, 1993)
cuyanus Duret, 1968
decens Theobald, 1901
declarator Dyar & Knab, 1906
delys Howard, Dyar & Knab, 1915
demeilloni Doucet, 1950
diengensis Brug, 1931
diplophyllum Dyar, 1929
dohenyi Hogue, 1975
dolosus (Lynch Arribálzaga, 1891)
draconis Ingram & de Meillon, 1927
duplicator Dyar & Knab, 1909
duttoni Theobald, 1901
eduardoi Casal & García, 1968
edwardsi Barraud, 1923
elgonicus Edwards, 1941
erikae Szadziewski & Szadziewska, 1985 (fossil species, 33.9–40.4 Mya, Cenozoic, Late Eocene)
erythrothorax Dyar, 1907
eschirasi Galliard, 1931
farakoensis Hamon, 1955
fasyi Baisas, 1938
fernandezi Casal, García & Cavalieri, 1966
foliaceus Lane, 1945
fuscocephala Theobald, 1907
gameti Bailly-Choumara, 1966
garciai Broche, 2000
gediensis Edwards, 1941
gelidus Theobald, 1901
globocoxitus Dobrotworsky, 1953
grahamii Theobald, 1910
guayasi Leví-Castillo, 1953
guiarti Blanchard, 1905
guizhouensis Chen & Zhao, 1985
habilitator Dyar & Knab, 1906
hancocki Edwards, 1930
hopkinsi Edwards, 1932
huangae Meng, 1958
hutchinsoni Barraud, 1924
incognitus Baisas, 1938
inflictus Theobald, 1901
ingrami Edwards, 1916
interfor Dyar, 1928
interrogator Dyar & Knab, 1906
invidiosus Theobald, 1901
iyengari Mattingly & Rageau, 1958
jacksoni Edwards, 1934 (in Barraud, 1934)
janitor Theobald, 1903
kartalae Brunhes, 1977
kesseli Belkin, 1962]
kingii (Theobald, 1913)
lahillei Bachmann & Casal, 1962
laticinctus Edwards, 1913
laticlasper Galindo & Blanton, 1954
levicastilloi Lane, 1945
litoralis Bohart, 1946
litwakae Harbach, 1985
longitubus Somboon, Namgay & Harbach, 2021 (in Somboon et al., 2021)
lygrus Root, 1927
maracayensis Evans, 1923
macrophyllus Edwards & Gibbins, 1939
marianae Bohart & Ingram, 1946
marquesensis Stone & Rosen, 1953
mattinglyi Knight, 1953
mauesensis Lane, 1945
maxi Dyar, 1928
mimeticus Noè, 1899
mimuloides Barraud, 1924
mimulus Edwards, 1915
miraculosus Bonne-Wepster, 1937
mirificus Edwards, 1913
mollis Dyar & Knab, 1906
murrelli Lien, 1968
musarum Edwards, 1932
nakuruensis Mattingly, 1951
neavei Theobald, 1906
nigripalpus Theobald, 1901 
nilgiricus Edwards, 1916
ninagongoensis Edwards, 1928
omani Belkin, 1962
orientalis Edwards, 1921
ornatothoracis Theobald, 1909
ousqua Dyar, 1918
pacificus Edwards, 1916
pajoti da Cunha Ramos & Ribeiro, 1982
pallens Coquillett, 1898
palpalis Taylor, 1912
paramaxi Duret, 1968
perexiguus Theobald, 1903
perfidiosus Edwards, 1914
perfuscus Edwards, 1914
perplexus Leicester, 1908
pervigilans Bergroth, 1889
philipi Edwards, 1929
philippinensis Sirivanakarn, 1976
pinarocampa Dyar & Knab, 1908
pipiens Linnaeus, 1758
plicatus Olivares, 1993 (in Angulo & Olivares, 1993)
propinquus Colless, 1955
prosecutor Séguy, 1927
pruina Theobald, 1901
pseudopruina van Someren, 1951
pseudostigmatosoma Strickman, 1990
pseudovishnui Colless, 1957
quasiguiarti Theobald, 1910
quinquefasciatus Say, 1823
quitensis Levi-Castillo, 1953
restuans Theobald, 1901
riojanus Duret, 1968
roseni Belkin, 1962
rotoruae Belkin, 1968
salinarius Coquillett, 1904
saltanensis Dyar, 1928
scheuberi Carpintero & Leguizamón, 2004
schwetzi Edwards, 1929
scimitar Branch & Seabrook, 1959
scottii Theobald, 1912
sechani Brunhes & Boussès, 2009
secutor Theobald, 1901
seldeslachtsi Wolfs, 1947
shoae Hamon & Ovazza, 1954
sinaiticus Kirkpatrick, 1925
sitiens Wiedemann, 1828
solitarius Bonne-Wepster, 1938
sphinx Howard, Dyar & Knab, 1913
spinosus Lutz, 1905
stenolepis Dyar & Knab, 1908
stigmatosoma Dyar, 1907
striatipes Edwards, 1941
summorosus Dyar, 1920
surinamensis Dyar, 1918
tamsi Edwards, 1934
tarsalis Coquillett, 1896
tatoi Casal & García, 1971
telesilla de Meillon & Lavoipierre, 1945 (in De Meillon et al., 1945)
tenagius van Someren, 1954
terzii Edwards, 1941
thalassius Theobald, 1903
theileri Theobald, 1903
thriambus Dyar, 1921
tianpingensis Chen, 1981
toroensis Edwards & Gibbins, 1939
torrentium Martini, 1925
toviiensis Klein, Rivière & Séchan, 1984
trifilatus Edwards, 1914
trifoliatus Edwards, 1914
tritaeniorhynchus Giles, 1901
tsengi Lien, 1968
ugandae van Someren, 1967
umbripes Edwards, 1941
univittatus Theobald, 1901
usquatissimus Dyar, 1922
usquatus Dyar, 1918
vagans Wiedemann, 1828
vansomereni Edwards, 1926
ventrilloni Edwards, 1920
verutus Harbach, 1987
vexillatus Edwards, 1941
vicinalis de Meillon & Lavoipierre, 1944
vicinus (Taylor, 1916)
vishnui Theobald, 1901
watti Edwards, 1920
weschei Edwards, 1935
whitei Barraud, 1923
whitmorei (Giles, 1904)
whittingtoni Belkin, 1962
yojoae Strickman, 1990
zombaensis Theobald, 1901

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