PALEARCTIC REGION
Etymology: club-bearing (Latin); ref. to shape of male palpus
Anopheles claviger is a large, drab species with entirely dark wings and legs. The Claviger Complex comprises two species: Anopheles claviger and An. petragnani del Vecchio. These taxa are purportedly distinguishable by subtle larval characters, but are most reliably identified using molecular techniques. During collections made in the United Kingdom, one of the authors determined another novel cryptic taxon, collected and link-reared from a woodland pool in sympatry with An. claviger s.s.. The new species was recovered on the basis of mtDNA COI and nuclear ITS2 sequence differences and is currently pending formal description. Seven synonyms exist for An. claviger s.s. and it remains unclear if the new taxon conforms to any of these available names. An. petragnani is largely restricted to the Mediterranean basin, where it has sometimes been reported in sympatry with An. claviger.
Type locality: [Germany], Europe
Type depository: Type non-existent (NE)
DIAGNOSTIC CHARACTERS (Click photos to view; mouse over and click large photo to zoom in.)
ADULT (illustrated): Head: Vertex with distinct tuft of broad white scales, lateral and posterior scales dark brown; labellum and proboscis dark, concolorous; palpomere MPlp5 short, not longer than half length of MPlp4. Thorax: Scutum with narrow pale scales on broad median area. Lower proepisternal setae present. Wing: Scales uniformly distributed, without concentrations of dark scale. Legs: Ta-I1 long, about equal in length to Ta-I2–5 combined.
LARVA (not illustrated): Head: Antenna (A) spiculate; seta 1-A basally branched; seta 2-C inserted close together; setae 2–3-C single or with 2–3 apical branches; seta 4-C shorter than 3-C, 2–5 branched; setae 5–7-C branched, long and plumose. Abdomen: Seta 1-I not palmate; seta 1-II palmate with 10–15 leaflets; setae 2-IV,V 3–5 branched; setae 6-IV–VI branched but not plumose; tergal plates narrow, height ≤3.0 x width.
TAXONOMIC KEYS
WRBU - Anopheles - Western Palearctic - Adult
WRBU - Genera - Global - Adult
WRBU - Genera - Global - Larva
Exemplar DNA sequences
Anopheles claviger COI: KM258237–44; KP942734–36; JX255719–JX255720.
BIONOMICS
Immatures
In the Mediterranean, An. claviger s.l. larvae are commonly collected in cisterns, yet typical habitats for immature An. claviger s.s. include cool, clean groundwaters, such as shaded woodland pools, springs and wells. Notably lower densities of An. claviger s.s. were correlated to the disappearance of spring-water sources in Tajikistan in the 1960s. In colder northerly climates, An. claviger s.s. overwinter as larvae, and are able to develop and pupate in winter. By contrast, An. petragnani tolerates hotter larval habitat waters.
Adults
Anopheles claviger s.s. is a competent malaria vector, preferentially biting man and large mammals outdoors after dusk, whereas An. petragnani appears solely zoophilic and has never been implicated in human disease transmission. Blooded female An. claviger commonly rest in animal shelters. The species is multivoltine, with northern populations peaking in May and September. Females are relatively long-lived, averaging 4–5 egg cycles per generation, although egg batches are notably smaller in the late summer populations. Unlike most Anopheles that oviposit directly on the water surface, Anopheles claviger s.s. lays its eggs in the damp substrate slightly above the waterline. In the warmer Mediterranean Region and southern parts of their distribution, the species is active year-round.
DISTRIBUTION NOTES
Afghanistan, Albania, Algeria, Armenia, Austria, Azerbaijan, Belarus, Belgium, Bosnia & Herzegovina, Bulgaria, Crimean Peninsula, Croatia, Cyprus, Czech Republic, Denmark, Estonia, Finland, France (includes Corsica), FYRO Macedonia, Georgia, Germany, Greece (includes Crete), Hungary, India, Iran, Iraq, Ireland, Israel (and Gaza Strip & West Bank), Italy (includes Sicily), Jordan, Kazakhstan, Kyrgyzstan, Latvia, Lebanon, Libya, Liechtenstein, Lithuania, Luxembourg, Moldova, Montenegro, Morocco, Netherlands, Norway, Pakistan, Palestine, People's Republic of China, Poland, Portugal, Romania, Russia (Central, Northwestern, Southern Districts, Volga), Serbia, Slovakia, Slovenia, Spain, Sweden, Switzerland, Syria, Tajikistan, Tunisia, Turkey, Turkmenistan, Ukraine, United Kingdom, Uzbekistan.
WRBU VECTOR HAZARD REPORTS
None; View other WRBU Vector Hazard Reports
Available GIS Models
IMPORTANT REFERENCES (full citations below)
Meigen 1804: 4 (M, F; Culex)
Marshall 1938: 126 (M*, F*, P*, L*, E*; bionomics)
Ross & Roberts 1943b: 3 (M*, F*, L*; taxonomy, distribution, bionomics)
Torres Cañamares, F. 1945 (M*, F, P*, L*; taxonomy, bionomics)
Bates et al. 1949: 422 (A, L: taxonomy)
Aitken 1954: 451 (A, P, L, E)
Senevet & Andarelli 1955b (taxonomy)
Service 1973 (bionomics)
Gutsevich et al. 1974: 86 (M*, F*, L*)
Utrio 1975: 63 (distribution)
Utrio 1976 (L*)
Glick 1992 (F key, distribution)
Ramsdale & Snow 2000: 3 (distribution)
Hadjivassilis 2000 7: 38 (distribution; Cyprus)
Samanidou-Voyadjoglou & Harbach, 2001 (keys; Greece)
Beck et al. 2003: 24 (distribution; Luxembourg)
Trari et al. 2002: 330 (distribution; Morocco)
Qu & Zhu 2008 (distribution; People's Republic of China)
Becker et al. 2010: 166 (M*, F*, P, L*; keys, taxonomy, distribution, bionomics)
Habirov et al. 2012 (molecular taxonomy, bionomics, distribution; Tajikistan)
Robert et al. 2019 (distribution; western Palaearctic)
CURRENT SYNONYMS
syn. villosus Robineau-Desvoidy
1827: 411 (M, F). Type locality: Paris, France (NE).
syn. grisescens Robineau-Desvoidy
1827 : 503 (two syntypes). Type locality: Ripley, Surrey, England (NHMUK). References: Edwards 1932a (synonomy); Townsend 1990: 80 (type information).
syn. turkestani Shingarev
1926: 47 (M, F; algeriensis var.). Type locality: Turkestan (LU). References: Edwards 1932a: 38 (synonomy suggested).
syn. amaurus Martini
1929: 135 (M*, F). Type locality: [Ben Anen, Taschkent], Turkestan (NHMUK). References: Senevet & Andarelli 1955a: 128 (taxonomy); Stone et al. 1959: 16 (synonomy).
syn. habibi Mulligan & Puri
1936: 68 (F*). Type locality: Hudda village on banks of Habib Nullah, Quetta, Baluchistan [Pakistan] (NHMUK). References: Aslamkhan 1971b (distribution; Pakistan); Glick 1992: 126 (synonomy).
syn. missiroli Del Vecchio
1939: 34 (E*; claviger var.). Type locality: Latium, Italy (LU). References: Bates et al. 1949: 422 (syn.).
syn. pollutus Cañamares
1945: 232 (L*; claviger var.). Type locality: Nr. Cuenca, Spain (LU). References: M. Bates in Boyd 1949 (syn.); Senevet & Andarelli 1955a: 128 (P, L; taxonomy); Stone et al. 1959: 17 (synonomy).
CITED REFERENCES
Aitken, T.H.G. (1954). The Culicidae of Sardinia and Corsica (Diptera). Bulletin of Entomological Research, 45(3), 436–494.
Aslamkhan, M. (1971b). The mosquitoes of Pakistan I. A checklist. Mosquito Systematics, 3(4), 147–159.
Bates, M., Beklemishev, W. N., & La Face, L. (1949). Anophelines of the Palearctic region. In M. F. Boyd (Ed.), Malariology: a comprehensive survey of all aspects of this group of diseases from a global standpoint, by 65 contributors (pp. 419–442). Philadelphia: W.B. Saunders Co.
Beck, M., Galm, M., Weitzel, T., Fohlmeister, V., Kasiser, A., Arnold, A., & Becker, N. (2003). Preliminary studies on the mosquito fauna of Luxembourg. European Mosquito Bulletin, 14, 21–24.
Becker, N. (2008). Influence of climate change on mosquito development and mosquito-borne diseases in Europe. Parasitology Research, 103 (Suppl. 1), 519–528.
Boyd, M.F. (Ed.) (1949). Malariology: A comprehensive survey of all aspects of this group of diseases from a global standpoint. By 65 contributors (Vol. I.). Philadelphia, PA: Saunders.
Del Vecchio, G. (1939). Sulle varieta de A. claviger (bifurcatus) nota I. Rivista di Malariologia, 3, 27–37.
Edwards, F.W. (1932a). Diptera family Culicidae. In P. Wytsman (Ed.), Genera insectorum (194th Fascicule) (pp. 1–258). Brussels, Belgium: Louis Desmet-Verteneuil.
Glick, J.I. (1992). Illustrated key to the female Anopheles of southwestern Asia and Egypt (Diptera: Culicidae). Mosquito Systematics, 24(2), 125–153.
Gutsevich, A.V. (1974). On the identification of mosquito females (Culicidae) by microscopic preparations of the head. IV. Key to species of the genus Aedes. Parazitologiya (St. Petersburg), 8(4), 329–335.
Habirov, Z., Kadamov, D., Iskandarov, F., Komilova, S., Cook, S., McAlister, E., & Harbach, R.E. (2012). Malaria and the Anopheles mosquitoes of Tajikistan. Journal of Vector Ecology, 37(2), 419–427.
Hadjivassilis, A. (2000). Mosquitoes (Diptera: Culicidae) of Cyprus. European Mosquito Bulletin, 7, 38.
Marshall, J.F. (1938). The British Mosquitoes. London: The British Museum (Natural History).
Martini, E. (1929). 11. u. 12. Culicidae, pp. 1–144. In: Lindner, E. (ed.), Die Fliegen der palaerktischen Region. Band III. E. Schweizerbart’sche Verlagsbuchhandlung (Erwin Nagele), Stuttgard.
Mulligan, H.W., & Puri, I.M. (1936). Description of Anopheles (Anopheles) habibi n. sp. from Ouetta, Baluchistan. Records of the Malaria Survey of India, 6, 67–71.
Qu, F., & Zhu, H. (2008). On a checklist of the Anopheline mosquitoes in China with rectification for some specific names. Chinese Journal of Parasitology and Parasitic Diseases, 26(3), 210–216.
Ramsdale, C.D., & Snow, K. (2000). Distribution of the genus Anopheles in Europe. European Mosquito Bulletin, 7, 1–26.
Robert, V., Günay, F., Le Goff, G., Boussès, P., Sulesco, T., Khalin, A., Medlock, J.M., Kampen, H., Petrić, D. & F. Schaffner. (2019). Distribution chart for Euro-Mediterranean mosquitoes (western Palaearctic region). Journal of the European Mosquito Control Association, 37, 1–28.
Robineau-Desvoidy, A.J.B. (1827). Essai sur la tribu des culicides. Société d’histoire naturelle, 3, 390–413.
Ross, E.S., & Roberts, H.R. (1943b). Mosquito atlas. Part II. Eighteen old world anophelines important to malaria. Contributions of the American Entomological Institute.
Samanidou-Voyadjoglou, A., & Harbach, R. E. (2001). Keys to the adult female mosquitoes (Culicidae) of Greece. European Mosquito Bulletin, 10, 13-20.
Senevet, G., & Andarelli, L. (1955b). Races et variétés de l'Anopheles claviger Meigen, 1804. Archives de l'Institut Pasteur d'Algérie, 33, 128–137.
Service, M.W. (1973). The biology of Anopheles claviger (Mg.) (Dipt., Culicidae) in southern England. Bulletin of Entomological Research, 63(2), 347–359.
Shingarev, N.I. (1926). New information on Culicidae of U.S.S.R. Tropichenskai Meditsina I Veterinariia, 2, 47–48.
Stone, A., Knight, K.L., & Starcke, H. (1959). A synoptic catalog of the mosquitoes of the World (Diptera, Culicidae) (Vol. 6). Washington, D.C.: Entomological Society of America, The Thomas Say Foundation.
Torres Canamares, F. (1945). Contribucion al conocimiento del Anopheles claviger Mg. de España (Dip. Cul.). EOS Revista Española de Entomología, 20, 233–245.
Townsend, B.C. (1990). A catalogue of the types of bloodsucking flies. Culicidae. Occasional Papers of the Natural History Museum (London)(7), 35–152.
Trari, B., Dakki, M., Himmi, O., & El Agbani, M.A. (2002). Le moustiques (Diptera: Culicidae) du Maroc: Revue bibliographique (1916–2001) et inventaire des espèces. Bulletin de la Société de pathologie exotique (Paris), 96(4), 329–334.
Utrio, P. (1975). Anopheles claviger (Meig.), Aedes pionips Dyar and A. beklemishevi Den. (Diptera, Culicidae) found in Finland. Notulae Entomologicae, 55, 63–64.
Utrio, P. (1976). Identification key to Finnish mosquito larvae (Diptera, Culicidae). Annales Agriculturae Fenniae, 15, 128–136.
CITE THIS PAGE
Walter Reed Biosystematics Unit (Year). Anopheles claviger species page. Walter Reed Biosystematics Unit Website, http://wrbu.si.edu/vectorspecies/mosquitoes/claviger, accessed on [date (e.g. 03 February 2020) when you last viewed the site].
