Culex Linnaeus, 1758

GLOBAL DISTRIBUTION

Family
Subfamily
Tribe

Generic abbreviation: Cx.

Type species: Culex pipiens Linneaus (Latreille 1810: 442)

Etymology: gnat, midge (L)

The genus Culex has 768 species, subdivided into 26 formally recognized subgenera including: Acalleomyia (1 species, Indo-Malaya), Acallyntrum (8 species, Australasia), Aedinus (4 species, Neotropics), Afroculex (1 species, South Africa), Allimanta (1 species, Argentina), Anoedioporpa (12 species, Neotropics), Barraudius (4 species, Palaearctic & Afrotropics), Belkinomyia (1 species, Colombia), Carrollia (18 species, Neotropics), Culex (198 species, global), Culiciomyia (55 species, Afrotropics, Orient & Australasia), Eumelanomyia (77 species, Afrotropics, Orient & Australasia), Kitzmilleria (1 species, Africa), Lasiosiphon (1 species, Africa, Southwest Asia), Lophoceraomyia (126 species, Orient & Australasia), Maillotia (9 species, Africa, Southwest Asia), Melanoconion (160 species, Nearctic, Neotropics), Micraedes (8 species, Caribbean), Microculex (33 species, Neotropics), Neoculex (26 species, Nearctic and Old World), Nicaromyia (1 species, Cuba), Oculeomyia (19 species, Afrotropics, Palearctic, Orient, Australasia), Phenacomyia (3 species, Neotropics), Phytotelmatomyia (4 species, Neotropics), Sirivanakarnius (1 species, Bonin Islands), Tinolestes (3 species, New World) and 7 species remain unplaced. Many Culex species are of biomedical importance.

 

DIAGNOSTIC CHARACTERS   (Click photos to view; mouse over and click large photo to zoom in.)

ADULT (illustrated): Thorax: Prespiracular and postspiracular setae absent. Wings: Alula with narrow fringe of scales. Legs: Well-developed pulvillus present

LARVA (not illustrated): Head: Seta 2-C usually absent; seta 3-C located dorsallyAbdomen: Seta 12-I always present. Terminal segments: Pecten present on siphon, seta 1-S with three or more pairs of prominent setae; saddle usually complete, sometimes incomplete and greatly reduced, but never divided into dorsal and ventral sclerites; ventral brush (seta 4-X) with at least three pairs of setae.

 

TAXONOMIC KEYS

Barraud 1934 (south Asia, British India)

Clark-Gil & Darsie 1983 (Guatemala)

Darsie & Pradhan 1991 (Nepal)

Darsie & Ward 1981 (North America)

Darsie & Ward 2005 (North America)

Darsie 1985 (Argentina)

Dubose & Curtin 1965 (Mediterranean)

Edwards 1941 (Afrotropics)

Hopkins 1952 (Afrotropics)

Lane 1953 (revision; Neotropics)

Rattanarithikul & Harrison et al. 2005 (Thailand)

Rattanarithikul & Panthusiri 1994 (Thailand)

Senevet & Andarelli 1954a: 36 (North Africa)

Senevet 1947b: 212 (North Africa)

Sicart 1954: 27 (Tunisia)

Wood et al. 1979 (Canada)

WRBU LUCID KEYS

 

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WRBU – Genera – Global – Adult

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WRBU – Genera – Global – Larva

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WRBU – Genera – Afrotropical – Adult

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WRBU – Genera – Afrotropical – Larva

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WRBU – Culex – Subgenera – Afrotropical – Adult

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WRBU – Culex – Subgenera – Afrotropical – Larva

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WRBU – Culex (Cux.) – East Africa – Adult

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WRBU – Culex (Cux.) – West Africa – Adult

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WRBU – Culex (Ocu.) – Afrotropical – Adult

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WRBU – Genera – Australasian – Adult

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WRBU – Genera – Australasian – Larva

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WRBU – Culex – Subgenera – Australasia – Adult

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WRBU – Culex – Subgenera – Australasia – Larva

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WRBU – Culex (Cux.) – Subgenera – Australasia – Adult

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WRBU – Culex (Cux.) – Subgenera – Australasia – Larva

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WRBU – Culex (Ocu.) – Subgenera – Australasia – Adult

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WRBU – Culex (Ocu.) – Subgenera – Australasia – Larva

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WRBU – Genera – IndoMalayan – Adult

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WRBU – Genera – IndoMalayan – Larva

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WRBU – Culex – Subgenera – IndoMalayan – Adult

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WRBU – Culex – Subgenera – IndoMalayan – Larva

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WRBU – Culex (Cux.) – Subgenera – IndoMalayan – Adult

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WRBU – Culex (Cux.) – Subgenera – IndoMalayan – Larva

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WRBU – Culex (Ocu.) – Subgenera – IndoMalayan – Adult

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WRBU – Culex (Ocu.) – Subgenera – IndoMalayan – Larva

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WRBU – Genera – Oriental – Adult

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WRBU – Genera – Oriental – Larva

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WRBU – Culex – Subgenera – Oriental – Adult

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WRBU – Culex – Subgenera – Oriental – Larva

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WRBU – Culex (Cux.) – Subgenera – Oriental – Adult

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WRBU – Culex (Cux.) – Subgenera – Oriental – Larva

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WRBU – Culex (Ocu.) – Subgenera – Oriental – Adult

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WRBU – Culex (Ocu.) – Subgenera – Oriental – Larva

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WRBU – Culex Subgenera & Culex (Cux.) species – Southwest Asia – Adult

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WRBU – Culex Subgenera & Culex (Cux.) species – Southwest Asia – Larva

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WRBU – Genera – Eastern Palearctic – Adult

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WRBU – Genera – Eastern Palearctic – Larva

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WRBU – Culex – Subgenera – Eastern Palearctic – Adult

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WRBU – Culex – Subgenera – Eastern Palearctic – Larva

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WRBU – Culex (Cux.) – Subgenera – Eastern Palearctic – Adult

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WRBU – Culex (Cux.) – Subgenera – Eastern Palearctic – Larva

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WRBU – Culex (Ocu.) – Subgenera – Eastern Palearctic – Adult

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WRBU – Culex (Ocu.) – Subgenera – Eastern Palearctic – Larva

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WRBU – Genera – Western Palearctic – Adult

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WRBU – Genera – Western Palearctic – Larva

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WRBU – Culex – Nearctic – Adult

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WRBU – Culex – Nearctic – Larva

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WRBU – Genera – Neotropical – Adult

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WRBU – Genera – Neotropical – Larva

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WRBU – Culex – Subgenera – Neotropical – Adult

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WRBU – Culex – Subgenera – Neotropical – Larva

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WRBU – Culex (Cux.) – Neotropical – Adult

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WRBU – Culex (Cux.) – Neotropical – Larva

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WRBU – Culex (Mel.) – Neotropical – Adult

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WRBU – Culex (Mel.) – Neotropical – Larva

Exemplar DNA sequences

Cx. (Barraudius) modestus COI: KJ012102

Cx. (Culex) laticinctus COI: KJ012096–101

Cx. (Cux.) mimeticus COI: KJ012096–101

Cx. (Cux.) pipiens COI: KJ012110–148

Culex (Cux.) quinquefasciatus  COI: KJ012162–173; whole genome GCF_000209185

Cx. (Cux.) theileri COI: KJ012174–234)

Cx. (Cux.) torrentium COI: KJ012235–242

Cx. (Cux.) tritaeniorhynchus COI: KJ012243–250

Cx. (Maillotia) hortensis COI: KJ012068–080

All Culex DNA sequences

 

BIONOMICS

Immatures

Typically, immature Culex typically occupy fresh, brackish, clean or heavily polluted groundwater habitats, where eggs are laid in rafts. Some—e.g., Cx. pipiens molestus—utilize underground water sources while others occupy natural containers, including phylotelmata (e.g., tree holes, leaf axils, flower bracts, bamboo, pitcher plants, fruit husks, and fallen spathes or leaves), and rock holes. Immature Belkinomyia, Nicaromyia and Tinolestes species rear in crabholes. Australasian Cx. (Ocu.) starckeae immatures are found in fast-flowing streams where they anchor onto algal mats using their modified siphonal setae. Some Microculex lay gel-covered individual eggs in small plant container habitats that are likely to dry out. In colony settings, Cx. pipiens can be cannibalistic when food resources are low.

Adults

Bionomic traits of Culex are extremely diverse. Adults typically feed on a wide variety of birds, reptiles, amphibians and mammals including man. They usually comprise the largest density of mosquitoes collected in light traps.

Associated Pathogens

*Associated pathogens: This list reports bacteria, viruses, and parasites recovered from, or experimentally passed through this species, and does not imply field vector status.

 

IMPORTANT REFERENCES (full citations below)

Linnaeus 1758: 602 (as genus)

Coquillett 1906b (taxonomy)

Edwards 1911b (Culex, taxonomy, Africa)
Barraud 1934 (M*, F*, L*; keys, tax.; south Asia (British India)

Edwards 1941 (M*, F*, P*; keys; Afrotropics)

Michener 1944: 263 (cibarial armature)

Senevet 1947b: 212 (L key; North Africa)

Hopkins 1952 (L*; key; Afrotropics)

Lane 1953 (revision; Neotropics)

Sicart 1954: 27 (P key; Tunisia)

Senevet & Andarelli 1954a: 36 (key; North Africa)

Mattingly & Knight 1956 (taxonomy; Arabia)

Belkin 1962 (South Pacific)

Dubose & Curtin 1965 (F, L; keys; Mediterranean)

Forattini 1965 (Neotropics)

 Cova-García et al. 1966 (Venezuela)

Delfinado 1966a (Philippines)

Bram 1967b (Thailand)

Bram 1967a (taxonomy, subgenus Culex; New World)

Belkin 1968a (revision; New Zealand)

Sirivanakarn 1968 (subgenus Lophoceraomyia)

Belkin et al. 1970 (Jamaica)

Berlin 1969a (revision; subgenus Howardina)

Berlin 1969c (revision; subgenus Micraedes)

Sirivanakarn 1971b (revision, subgenus Neoculex; SE Asia)

Sirivanakarn 1972 (revision, subgenus Eumelanomyia; SE Asia)

Adames & Galindo 1973 (taxonomy, subgenus Belkinomyia)

Valencia 1973 (revision, subgenus Carrollia)

Gutsevich et al. 1974 (revision; former USSR)

Sirivanakarn 1976 (revision, subgenus Culex; Oriental Region)

Sirivanakarn 1977a (revision, various subgenera; Oriental Region)

Tanaka et al. 1979 (revision; Japan, Korea)

Wood et al. 1979 (taxonomic keys; Canada)

Berlin & Belkin 1980 (revision, subg. Aedinus, Tinolestes, Anoedioporpa; Neotropics)

Ribeiro & da Cunha Ramos 1980 (taxonomy; Angola)

Darsie & Ward 1981 (F, L; keys; North America)

Lu & Li 1982a (taxonomy; People’s Republic of China)

Clark-Gil & Darsie 1983 (F, L; keys; Guatemala)

Sirivanakarn 1983 (revision; subgenus Melanoconion)

Darsie 1985 (F, L; keys, Argentina)

Lu & Su 1987 (tax.; People’s Republic of China)

Lee et al. 1989a (F; review; Australasia)

Lee et al. 1989b (F; review; Australasia)

Harbach 1988 (revision; subgenus Culex; SW Asia & Egypt)

Harbach & Peyton 1992b (taxonomy,  subgenus Phenacomyia)

Pecor et al. 1992 (tax. catalog; subgenus Melanoconion)

Rattanarithikul & Panthusiri 1994 (keys, medically important species; Thailand)

Reuben et al. 1994 (keys, JE vectors; Southeast Asia)

Sallum & Forattini 1996 (revision, Spissipes Section, subgenus Melanoconion)

Miller et al. 1996 (molecular phylogeny)

Lu et al. 1997 (taxonomy, People’s Republic of China)

Navarro & Liria 2000 (L mouthparts; taxonomy, phylogeny)

González Broche & Rodríguez R. 2001 (subgenus Nicaromyia)

Tanaka 2004b (subgenus Oculeomyia, Sirivanakarnius)

Darsie & Ward 2005 (F, L; keys, distribution; North America)

Rattanarithikul & Harrison et al. 2005 (F, L; keys, distribution; Thailand)

Rattanarithikul & Harrison et al. 2005 (taxonomic background; Thailand)

St John 2007 (morphological phylogeny)

Rossi & Harbach 2008 (subg. Phytotelmatomyia)

Harbach 2011 (classification, taxonomy, subgenus Culex)

Vesgueiro et al. 2011 (molecular phylogeny)

Demari-Silva et al. 2011 (molecular phylogeny)

Harbach et al. 2012 (morphological phylogeny)

Laurito et al. 2013 (morphological phylogeny)

Harbach & Culverwell 2016 (phylogeny)

 

VALID SUBGENERA

Acalleomyia Leicester [Aca.]

Acallyntrum Stone & Penn [Acl.]

Aedinus Lutz [Ads.]

Afroculex Danilov [Afc.]

Allimanta Casal & Garcia [Alm.]

Anoedioporpa Dyar [And.]

Barraudius Edwards [Bar.]

Belkinomyia Adames & Galindo [Bel.]

Carrollia Lutz [Car.]

Culex Linnaeus [Cux.]

Culiciomyia Theobald [Cui.]

Eumelanomyia Theobald [Eum.]

Kitzmilleria Danilov [Kit.]

Lasiosiphon Kirkpatrick [Las.]

Lophoceraomyia Theobald [Lop.]

Maillotia Theobald [Mai.]

Melanoconion Theobald [Mel.]

Micraedes Coquillett [Mca.]

Microculex Theobald [Mcx.]

Neoculex Dyar [Ncx.]

Nicaromyia Broche & Rodríguez Rodríguez [Nic.]

Oculeomyia Theobald [Ocu.]

Phenacomyia Harbach & Peyton [Phc.]

Phytotelmatomyia Rossi & Harbach [Phy.]

Sirivanakarnius Tanaka [Sir.]

Tinolestes Coquillett [Tin.]

 

CURRENT GENERIC SYNONYMS

syn. Heteronycha Lynch Arribálzaga 1891a: 373 1891b: 155 (as genus). Type species: Heteronycha dolosa Lynch Arribálzaga.

syn. Lasioconops Theobald 1903c: viii (as genus). Type species: Lasioconops poicilipes Theobald. Edwards 1911b: 256 (synonymy).

syn. Heptaphlebomyia Lynch Arribálzaga 1903a: 336 (as genus). Type species: Heptaphlebomyia simplex Theobald. Edwards 1911b: 262 (synonymy of simplex with univittatus Theobald).

syn. Pseudoheptaphlebomyia Ventrillon 1905: 427 (as genus). Type species: Pseudoheptaphlebomyia madagascariensis Ventrillon. Edwards 1932a: 200 (synonymy).

syn. Trichopronomyia Theobald 1905f: 98 (as genus). Type species: Trichopronomyia annulata Theobald. Edwards 1932a: 200 (synonymy).

syn. Aporoculex Theobald 1907: 316. Type species: Aporoculex punctipes Theobald.

syn. Leucomyia Theobald 1907: 372 (as genus) (not Brauer & Bergenstam 1891). Type species: Culex gelidus Theobald. Edwards 1932a: 200 (synonymy).

syn. Theobaldiomyia Brunetti 1912: 462 (as genus). Type species: Culex gelidus Theobald. Theobald 1901c: 20 (fixed type of replacement name). Edwards 1932a: 200 (synonymy)

syn. Phalangomyia Dyar & Knab 1914: 58 (as genus). Type species: Phalangomyia debilis Dyar & Knab. Edwards 1932a: 200 (synonymy).

syn. Laiomyia Izquierdo 1916: 65 (as genus). Type species: Culex stigmatosoma Dyar. Edwards 1932a: 200 (synonymy).

syn. Cacoculex Dyar 1918b (as subgenus of genus Culex): 100. Type species: Culex habilitator Dyar & Knab. Edwards 1932a: 200 (synonymy).

syn. Transculicia Dyar 1918 (1917)e (as subgenus of genus Culex): 184. Type species: Culex eleuthera Dyar. Edwards 1932a: 205 (synonymy).

 

CITED REFERENCES

Adames, A.J., & Galindo, P. (1973). Mosquito studies (Diptera, Culicidae). XXX. A new subgenus and species of Culex from Colombia. Contributions of the American Entomological Institute, 9(3), 55–61.

Belkin, J.N. (1962). The mosquitoes of the South Pacific (Diptera, Culicidae) (Vols. 1 &2). Berkeley, California: University of California Press.

Belkin, J.N. (1968a). Mosquito studies (Diptera, Culicidae) VII. The Culicidae of New Zealand. Contributions of the American Entomological Institute, 3(1), 1–182.

Belkin, J.N., Heinemann, S.J., & Page, W.A. (1970). The Culicidae of Jamaica (Mosquito studies. XXI) Contributions of the American Entomological Institute, 6(1), 458.

Berlin, O.G.W. (1969a). Mosquito studies (Diptera, Culicidae). XII. A revision of the Neotropical subgenus Howardina of Aedes. Contributions of the American Entomological Institute, 4(2), 1–190.

Berlin, O.G.W. (1969c). Mosquito studies (Diptera, Culicidae). XVIII. The subgenus Micraedes of Culex. Contributions of the American Entomological Institute, 5(1), 21–63.

Berlin, O.G.W., & Belkin, J.N. (1980). Mosquito studies (Diptera, Culicidae). 36. Subgenera Aedinus, Tinolestes and Anoedioporpa of Culex. Contributions of the American Entomological Institute, 17(2), 1–104.

Bram, R.A. (1967a). Classification of Culex subgenus Culex in the New World (Diptera: Culicidae). Proceedings of the United States National Museum, 120(3557), 1–122.

Bram, R.A. (1967b). Contributions to the mosquito fauna of Southeast Asia. II. The genus Culex in Thailand (Diptera: Culicidae). Contributions of the American Entomological Institute, 2(1), 1–296.

Brunetti, E. (1912). Annotated catalogue of Oriental Culicidae - Supplement. Rec. Indian Mus, 4, 403–517.

Clark-Gil, S., & Darsie, Jnr., R.F. (1983). The mosquitoes of Guatemala: their identification, distribution and bionomics, with keys to adult females and larvae, in English and Spanish. Mosquito Systematics, 15(3), 151–284.

Coquillett, D.W. (1906b). On the breaking-up of the old genus Culex. Science (NS), 23, 312–314.

Darsie, R.F., Jr. (1985). Mosquitoes of Argentina. Part I. Keys for identification of adult females and fourth stage larvae in English and Spanish (Diptera, Culicidae). Mosquito Systematics, 17(3), 153–253.

Darsie, R.F., Jr., & Ward, R.A. (1981). Identification and geographical distribution of the mosquitoes of North America, north of Mexico. Mosquito Systematics Supplement, Supplement 1, 1–313.

Darsie, R.F., Jr., & Ward, R.A. (2000). Summary of new distribution records for mosquito species in the United States and Canada for the period 1981–99. Journal of the American Mosquito Control Association, 16(1), 1–4.

Darsie, R.F., Jr., & Ward, R.A. (2005). Identification and geographical distribution of the mosquitoes of North America, north of Mexico. Gainesville: University Press of Florida.

Darsie, R.F., Jr., Pradhan, S.P., & Vaidya, R G. (1991). Notes on the mosquitoes of Nepal. 1. New country records and revised Aedes keys (Diptera, Culicidae). Mosquito Systematics, 23(1), 39–45.

Darsie, R.F., Pradhan, S.P., & Vaidya, R.G. (1992). Notes on the mosquitoes of Nepal. II. New species records from 1991 collections. Mosquito Systematics, 24, 23–28.

Delfinado, M.D. (1966a). The culicine mosquitoes of the Philippines, tribe Culicini (Diptera, Culicidae). Memoirs of the American Entomological Institute, 7, 1–252.

Demari-Silva, B., Vesgueiro, F.T., Sallum, M. A. M., & Marrelli, M.T. (2011). Taxonomic and phylogenetic relationships between species of the genus Culex (Diptera: Culicidae) from Brazil inferred from the cytochrome c oxidase I mitochondrial gene. Journal of Medical Entomology, 48(2), 272–279.

Dubose, W.P., & Curtin, T.J. (1965). Identification keys to the adult and larval mosquitoes of the Mediterranean area. Journal of Medical Entomology, 1(4), 349–355.

Dyar, H.G. (1918b). A revision of the American species of Culex on the male genitalia. Insecutor Inscitiae Menstruus, 86–111.

Dyar, H.G. (1918e). A second note on the species of Culex of the Bahamas (Diptera, Culicidae). Insecutor Inscitiae Menstruus, 5 (1917), 183–187.

Dyar, H.G., & Knab, F. (1914). New mosquitoes from Peru (Diptera, Culicidae). Insecutor Inscitiae Menstruus, 2, 58–62.

Edwards, F.W. (1911b). The African species of Culex and allied genera. Bulletin of Entomological Research, 2, 241–268.

Edwards, F.W. (1932a). Diptera family Culicidae. In P. Wytsman (Ed.), Genera insectorum (194th Fascicule) (pp. 1–258). Brussels, Belgium: Louis Desmet-Verteneuil.

Edwards, F.W. (1941). Mosquitoes of the Ethiopian Region. III. Culicine adults and pupae. Bulletin of the British Museum (Natural History) Entomology.

Forattini, O. P. (1965). Entomologia Médica. Vol. 2. Culicini: Culex, Aedes e Psorophora. São Paulo: Universidade de São Paulo.

González Broche, R., & Rodriguez Rodriguez, J. (2001). Nuevo subgenero de Culex, descripción de la pupa y de la larva y redescripçion de la hembra de Culex nicaroensis Duret (Diptera: Culicidae). Avicennia, 14, 65–74.

Gutsevich, A.V., Monchadskii, A.S., & Shtakel’berg, A.A. (1974). Fauna of the USSR. New series No. 100 Diptera. Vol. III, No. 4. Mosquitoes. Family Culicidae. Jerusalem, Israel: Keter Publishing House Jerusalem Ltd. (Original work published 1971).

Harbach, R.E. (1988). Mosquitoes of the subgenus Culex in southwestern Asia and Egypt (Diptera: Culicidae). Contributions of the American Entomological Institute, 24(1), 1–240.

Harbach, R.E. (2011). Classification within the cosmopolitan genus Culex (Diptera: Culicidae): The foundation for molecular systematics and phylogenetic research. Acta Tropica, 120(1–2), 1–14.

Harbach, R.E., & Peyton, E.L. (1992b). A new subgenus of Culex in the Neotropical Region (Diptera: Culicidae). Mosquito Systematics, 24(3), 242–252.

Harbach, R.E., Culverwell, C.L., & Kitching, I.J. (2016). Phylogeny of the nominotypical subgenus of Culex (Diptera: Culicidae): insights from analyses of anatomical data into interspecific relationships and species groups in an unresolved tree. Systematics and biodiversity, 15(4), 1–11.

Harbach, R.E., Kitching, I.J., Culverwell, C.L., Dubois, J., & Linton, Y.-M. (2012). Phylogeny of mosquitoes of tribe Culicini (Diptera: Culicidae) based on morphological diversity. Zoologica Scripta, 41(5), 499–514.

Hopkins, G.H.E. (1952). Mosquitoes of the Ethiopian region. I. Larval bionomics of mosquitoes and taxonomy of culicine larvae (2nd ed.). London, UK: British Museum (Natural History).

Lane, J. (1953). Neotropical Culicidae (Vols. I, II). São Paulo: University of São Paulo.

Laurito, M., de Oliveira, T.M.P., Almiron, W.R., & Sallum, M.A.M. (2013). COI barcode versus morphological identification of Culex (Culex) (Diptera: Culicidae) species: a case study using samples from Argentina and Brazil. Memórias do Instituto Oswaldo Cruz, 108 (Suppl. 1), 110–122.

Lee, D.J., Hicks, M.M., Debenham, M.L., Griffiths, M., Marks, E.N., Bryan, J.H., & Russell, R.C. (1989a). The Culicidae of the Australasian region. Volume 7. Commonwealth Department of Health, School of Public Health and Tropical Medicine Monograph Series, 2.

Lee, D.J., Hicks, M.M., Debenham, M.L., Griffiths, M., Marks, E.N., & Bryan, J.H. (1989b). The Culicidae of the Australasian region. Volume 8. Commonwealth Department of Health, School of Public Health and Tropical Medicine Monograph Series 2.

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Linnaeus, C. (1762). Zweyter Theil, enthalt Beschreibungen verschiedener wichtiger Naturalien. In Reise nach Palestina in den Jahren von 1749 bis 1752. Rostock, Germany: J.C. Koppe.

Linnaeus, C. (1767). Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis (Twelfth ed. Vol. 1, Part 2). Holmiae.

Linnaeus, C. (1792). Flora Lapponica . . . Editio altera, aucta et emendata sudio & cura Jacobis Edwardi Smith. London.

Lu, B.-L. (1997). Fauna Sinica, Insecta Vol. 8, Diptera: Culicidae I. Beijing: Science Press.

Lu, B.-L., & Li, B. S. (1982a). Identification of Chinese mosquitoes. In B. S. Lu (Ed.), Identification handbook for medically important animals in China [in Chinese] (pp. 1–159). Beijing: People’s Health Publication Company.

Lu, B.-L., & Su, L. (1987). A handbook for the identification of Chinese aedine mosquitoes. Science, 1–16.

Lu, B.-L.L. (1997). Fauna Sinica, Insecta Vol. 9, Diptera: Culicidae II. Beijing: Science Press.

Lu, B., Li, B., Ji, S., Chen, H., Meng, Q., Su, L., . . . eds. (1997). Diptera: Culicidae 1. Fauna Sinica, Insecta [in Chinese] (Vol. 8). Beijing, China: Science Press.

Lynch Arribálzaga, F. (1891a). Dipterologia argentine. Revista del Museo de La Plata, 1, 345–377.

Mattingly, P.F., & Knight, K.L. (1956). The mosquitoes of Arabia. I Bull Brit Mus, 89–141.

Michener, C.D. (1944). Differentiation of females of certain species of Culex by the cibarial armature. Journal of the New York Entomological Society, 52, 263–266.

Miller, B. R., Crabtree, M. B., & Savage, H. M. (1996). Phylogeny of fourteen Culex mosquito species, including the Culex pipiens complex, inferred from the internal transcribed spacers of ribosomal DNA. Insect Molecular Biology, l5(2), 93–107.

Navarro, J.C., & Liria, J. (2000). Phylogenetic relationships among eighteen neotropical Culicini species. Journal of the American Mosquito Control Association, 16(2), 75–85.

Pecor, J.E., Mallampalli, V.L., Harbach, R.E., & Peyton, E.L. (1992). Catalog and illustrated review of the subgenus Melanoconion of Culex (Diptera: Culicidae). Contributions of the American Entomological Institute, 27, 1–228.

Rattanarithikul, R., & Panthusiri, P. (1994). Illustrated keys to the medically important mosquitoes of Thailand. Southeast Asian Journal of Tropical Medicine and Public Health, 66(Supplement 1), 1.

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Walter Reed Biosystematics Unit (Year). Culex genus page. Walter Reed Biosystematics Unit Website, http://wrbu.si.edu/vectorspecies/genera/culex, accessed on [date (e.g. 03 February 2020) when you last viewed the site].