Anopheles Meigen, 1818

GLOBAL DISTRIBUTION

Family
Subfamily

Generic abbreviation: An.

Type species: Anopheles maculipennis Meigen (ICZN 1959; Opinion 547)

Etymology: useless, unprofitable, hurtful (Gr)

With the exception of the polar icecaps and some isolated island groups in the Pacific and Atlantic Oceans, including Fiji and New Zealand, Anopheles species are found across the globe. Anopheles are the mosquitoes capable of transmitting the five Plasmodium parasites (P. falciparum, P. knowlesi, P. ovale, P. malariae and P. vivax) that cause human malaria. Of the 475 Anopheles species world-wide (<15%), only about 70 are of primary or secondary (local) importance as malaria vectors. Although often overlooked, the importance of Anopheles in arboviral transmission is substantial. O’nyong-nyong virus (ONNV) and Guaroa (GROV) viruses are spread only by Anopheles mosquitoesusually An. gambiae Giles and An. funestus Giles), and An. neivai Howard, Dyar & Knab, respectively.

The genus Anopheles is subdivided into eight subgenera: Anopheles (185 species; globally distributed), Baimaia (1 species; Oriental), Cellia (224 species; Old World), Christya (2 species; Afrotropical), Kerteszia (12 species; restricted to the Neotropics), Lophopodomyia (6 sp.; restricted to the Neotropics), Nyssorhynchus (40 species; restricted to the Neotropics) and Stethomyia (5 species; restricted to the Neotropics). Together with the small and geographically restricted genera Bironella (n = 8 species) and Chagasia (n = 5), Anopheles mosquitoes (n = 475) make up the distinctive subfamily Anophelinae.

 

DIAGNOSTIC CHARACTERS  (Click photo to zoom in.)

ADULT (illustrated): Head: Palpus about as long as proboscis, apically clubbed in males.  Thorax: Posterior margin of scutellum evenly rounded, with evenly distributed setae. Wing: Usually with regular pattern of pale and dark spots; wing scales all dark in a few species; veins M1+2 and M3+4 straight or curved, but not wavy. Abdomen: Largely devoid of scales or with only small scale-patches; setae present.

LARVA (not illustrated): Thorax: Seta 1-M not palmate. Abdomen: Seta 1 usually palmate, at least on abdominal segments III–VI. Terminal segments: Siphon absent.

 

TAXONOMIC KEYS

Azari-Hamidian & Harbach, 2009 (Iran)

Baba 1950:1 (China)

Cagampang-Ramos & Darsie 1970 (Philippines)

Causey, Deane & Deane 1946 (Amazon)

Chow 1949b (China)

Clark-Gil & Darsie 1983 (Guatemala)

Colless 1948 (Borneo)

Cova García & Sutil Oramas 1975a: 201 (Venezuela)

Cova García & Sutil Oramas 1976a: 13 (Venezuela)

Cova García & Sutil Oramas 1977 (Venezuela)

Darsie & Ward 1981 (North America)

Darsie & Ward 2005 (North America)

De Meillon 1947b (Ethiopian Region)

Do-Van-Quy 1963 (Vietnam)

Doucet 1951b (Madagascar)

DuBose & Curtin 1965 (Mediterranean area)

Evans 1938 (Ethiopian Region)

Faran & Linthicum 1981 (Nyssorhynchus, Amazonia)

Forattini 1961:169 (Neotropical Region)

Forattini 1962 (Neotropical Region)

Gelfand 1954 (Liberia)

Gillies & De Meillon 1968 (Ethiopian Region)

Glick 1992 (southwestern Asia & Egypt)

González & Carrejo 2007 (Colombia)

Gorham et al. 1973 (western South America)

Gunathilaka 2017 (Sri Lanka)

Gunathilaka et al. 2014 (Sri Lanka)

Guy 1959 (Morocco)

Hara 1959 (Japan)

Jepson et al. 1947 (Mauritius)

Lane 1953 (Neotropical Region)

Lee et al. 1987 (Australasian Region)

Linthicum 1988 (Argyritarsis Section, subgenus Nyssorhynchus)

Otsuru & Ohmori 1960 (Japan)

Peyton & Scanlon 1966 (Thailand)

Rao 1984 (India)

Gillies & Coetzee 1987 (Afrotropical Region)

Rattanarithikul & Harrison 1973 (Thailand)

Rattanarithikul & Harrison et al. 2005 (Thailand)

Rattanarithikul & Panthusiri 1994 (Thailand)

Rattanarithikul et al. 2006b (Thailand)

Reid 1968 (Malaysia & Borneo)

Ribeiro & da Cunha Ramos 1975 (Angola)

Pringle 1954 (Iraq)

Senevet & Andarelli 1955a (Mediterranean Region)

Senevet 1934 (review)

Service 1962 (W. Africa)

Suárez et al. 1988 (Colombia)

Vargas & Martínez-Palacíos 1956 (Mexico)

Vargas 1959 (New World)

Weyer 1954 (Palaearctic Region)

Wilkerson & Strickman 1990 (Mexico & Central America)

Wood et al. 1979 (Canada)

Zavortink 1973 (subgenus Kerteszia)

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 – Anopheles – Afrotropical – Adult

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WRBU – Anopheles – Afrotropical – Larva

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WRBU – Genera – Australasia – Adult

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WRBU – Genera – Australasia – Larva

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WRBU – Anopheles (Ano.) – Myzorhynchus Series – Australasia – Adult

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WRBU – Anopheles (Cel.) – Neocellia Series – Australasia – Adult

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WRBU – Anopheles (Cel.) – Neomyzomyia Series – Australasia – Adult

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WRBU – Anopheles (Cel.) – Neomyzomyia Series – Australasia – Larva

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WRBU – Anopheles – Subgenera and Series – Australasia – Adult

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WRBU – Genera – IndoMalaya – Adult

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WRBU – Genera – IndoMalaya – Larva

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WRBU – Anopheles (Ano.) – Myzorhynchus Series – IndoMalaya – Adult

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WRBU – Anopheles (Ano.) – Myzorhynchus Series – IndoMalaya – Larva

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WRBU – Anopheles (Cel.) – Myzomyia Series – IndoMalaya – Adult

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WRBU – Anopheles (Cel.) – Myzomyia Series – IndoMalaya – Larva

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WRBU – Anopheles (Cel.) – Neocellia Series – IndoMalaya – Adult

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WRBU – Anopheles (Cel.) – Neocellia Series – IndoMalaya – Larva

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WRBU – Anopheles (Cel.) – Neomyzomyia Series – IndoMalaya – Adult

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WRBU – Anopheles (Cel.) – Neomyzomyia Series – IndoMalaya – Larva

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WRBU – Anopheles (Cel.) – Pyretophorus Series – IndoMalaya – Adult

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WRBU – Anopheles (Cel.) – Pyretophorus Series – IndoMalaya – Larva

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WRBU – Anopheles – Subgenera and Series – IndoMalaya – Adult

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WRBU – Anopheles – Subgenera and Series – IndoMalaya – Larva

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WRBU – Genera – Oriental – Adult

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WRBU – Genera – Oriental – Larva

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WRBU – Anopheles (Ano.) – Myzorhynchus Series – Oriental – Adult

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WRBU – Anopheles (Ano.) – Myzorhynchus Series – Oriental – Larva

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WRBU – Anopheles (Cel.) – Neocellia Series – Oriental – Adult

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WRBU – Anopheles (Cel.) – Neocellia Series – Oriental – Larva

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WRBU – Anopheles (Cel.) – Neomyzomyia Series – Oriental – Adult

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WRBU – Anopheles (Cel.) – Neomyzomyia Series – Oriental – Larva

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WRBU – Anopheles (Cel.) – Pyretophorus Series – Oriental – Adult

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WRBU – Anopheles (Cel.) – Pyretophorus Series – Oriental – Larva

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WRBU – Anopheles – Subgenera and Series – Oriental – Adult

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WRBU – Anopheles – Subgenera and Series – Oriental – Larva

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WRBU – Genera – Eastern Palearctic – Adult

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WRBU – Genera – 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 – Anopheles – Western Palearctic – Adult

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WRBU – Anopheles – Republic of Georgia – Adult

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WRBU – Genera – Nearctic – Adult

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WRBU – Genera – Nearctic – Larva

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WRBU – Genera – Neotropical – Adult

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WRBU – Genera – Neotropical – Larva

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WRBU – Anopheles – Mexico and Central America – Adult

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WRBU – Anopheles – Mexico and Central America – Larva

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Exemplar DNA sequences

All Anopheles DNA sequences

Subgenus Anopheles (whole genomes)

An. atroparvus (11555)

An. sinensis (12170)

Subgenus Cellia (whole genomes)

An. arabiensis (11544)

An. christyi (575528)

An. coluzzii (41035)

An. cracens (17028)

An. culicifacies (11552)

An. dirus (11554)

An. epiroticus (11546)

An. farauti (11553)

An. farauti no. 4 (17208)

An. funestus (2697)

An. gambiae (46)

An. koliensis (36653)

An. maculatus (11550)

An. melas (11549)

An. merus (11548)

An. minimus (11551)

An. nili (17525)

An. punctulatus (36652)

An. quadriannulatus (11545)

An. stephensi (2653)

Subgenus Nyssorhynchus (whole genomes)

An. aquasalis (66449)

An. albimanus (11556)

An. darlingi (2624)

 

BIONOMICS

Immatures

Immatures Anopheles habitats are typically clean, fresh, still or slow-moving waters with plentiful floating vegetation, but there are many exceptions. In primary forests in northern Vietnam, An. cucphuongensis Vu et al., 1990 individually occupy egg-cup-sized pools in limestone outcrops. Immature An. litoralis King tolerate the highly saline waters (≥ 2.5% to 3%) of the salt flats in the Philippines, and An. bwambae White are endemic to the hot springs in Bwambae County, Uganda.

Adults

Anopheles females are typically active at night, although several species will feed during the day in heavily shaded sites. Helium balloon captures over the Sahel in Mali showed that several African Anopheles, including An. coluzzii Coetzee & Wilkerson, An. gambiae s.s. and An. coustani Laveran, escape the harsh arid conditions of the Sahara Desert as fully gravid females by migrating long distances at altitudes of at least 240m, presumably to find more suitable conditions. That potentially infected Anopheles mosquitoes can disperse over large distances greatly impacts the accuracy of current malaria models and may go some way to explaining ongoing malaria in sub-Saharan African countries despite intensive malaria eradication programs.

Associated Pathogens

 

IMPORTANT REFERENCES (full citations below)

Meigen 1818: 10 (as genus)

Christophers 1915 (M genitalia*)

Senevet 1934 (P*; key, review)

Evans 1938 (M*, F*, P*, L*, E*; keys to F & L, Ethiopian Region)

Causey, Deane & Deane 1946 (M genitalia; key; Amazon Anophelini)

De Meillon 1947b (M*, F*, P*, L*; keys, Ethiopian Region)

Jepson et al. 1947 (bionomics, keys, Mauritius)

Colless 1948 (keys, distribution; Borneo)

Chow 1949b (keys, distribution; China)

Baba 1950:1 (keys, distribution; China)

Doucet 1951b (keys; Madagascar)

Romeo Viamonte & Castro 1951 (F*; buccopharyngeal armature)

Bonne-Wepster & Swellengrebel 1953: 1 (distribution; Indo-Australian Region)

Lane 1953 (revision, keys, Neotropical Region)

Gelfand 1954 (M, F; keys, Liberia)

Pringle 1954 (keys, Iraq)

Weyer 1954 (key, Palearctic Region)

Senevet & Andarelli 1955a (keys, Mediterranean Region)

Senevet & Andarelli 1955c (antepalmate hairs)

Mattingly & Knight 1956 (Arabia)

Vargas & Martínez-Palacíos 1956 (keys; Mexico)

Guy 1959 (keys; Morocco)

Hara 1959 (M genitalia, keys; Japan)

Vargas 1959 (keys, New World)

Otsuru & Ohmori 1960 (keys; Japan)

Cova García 1961 (Venezuela)

Forattini 1961:169 (M, F, L, keys; Neotropical Region)

Belkin 1962 (tax. distribution; South Pacific)

Forattini 1962 (review, keys, Neotropical Region)

Service 1962 (keys, W. Africa)

Do-Van-Quy 1963 (keys, Vietnam)

Various authors in Russell et al. 1963 (keys)

DuBose & Curtin 1965 (keys, Mediterranean area)

Grjebine 1966 (Madagascar)

Peyton & Scanlon 1966 (F key; Thailand)

Gillies & De Meillon 1968 (F*, L*; keys, review, Ethiopian Region)

Hinton 1968a (E*; bionomics)

Reid 1968 (M*, F*, P*, L*; keys, revision, Malaysia & Borneo)

Scanlon et al. 1968 (checklist, Thailand); Mattingly 1969c (E*, taxonomy)

Belkin et al. 1970 (Jamaica)

Cagampang-Ramos & Darsie 1970 (F, L; keys, distribution; Philippines)

Zavortink 1970b (New World treehole species)

Gorham et al. 1973 (keys, western South America)

Postiglione et al. 1973 (Turkey)

Rattanarithikul & Harrison 1973 (L keys, Thailand)

Zavortink 1973 (M*, F*, L*, P*; keys, subgenus Kerteszia)

Gutsevich et al. 1974 (former USSR)

Coluzzi & Kitzmiller 1975: 285 (genetics)

Cova García & Sutil Oramas 1975a: 201 (F*; key, Venezuela)

Harrison & Klein 1975 (Cambodia)

Harrison & Scanlon 1975 (subgenus Anopheles, Thailand)

Ribeiro & da Cunha Ramos 1975 (F, L; keys, Angola)

Vargas 1975 (F key, Costa Rica)

Cova García & Sutil Oramas 1977 (M genitalia*, F*, L*, E*; keys, Venezuela)

Kitzmiller 1977 (chromosomes)

Klein 1977b (review, Cambodia)

White 1977 (tax. morph. Anopheles species complexes)

Tanaka et al. 1979 (revision, Japan)

Wood et al. 1979 (keys, review, Canada)

Faran 1980 (Albimanus Section, subgenus Nyssorhynchus)

Harrison 1980 (Revision, Myzomyia Series, subgenus Cellia, Thailand)

Faran & Linthicum 1981 (M*, F*, P, L*; keys, subgenus Nyssorhynchus, Amazonia)

Lu & Li 1982a (China)

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

Rao 1984 (India); Gillies & Coetzee 1987 (F, L; keys, Afrotropical Region)

Lee et al. 1987b (review, key, Australasian Region)

Linthicum 1988 (M*, F*, P*, L*; keys, revision, Argyritarsis Section, subgenus Nyssorhynchus)

Suárez et al. 1988 (keys, Colombia)

Darsie & Pradhan 1990 (Nepal)

Das et al. 1990 (keys, India)

Wilkerson & Peyton 1990 (Anopheles wing spots)

Wilkerson & Strickman 1990 (illustrated key, Mexico & Central America)

Glick 1992 (F key, southwestern Asia & Egypt)

Peyton et al. 1992 (tax. subgenus Kerteszia & Nyssorhynchus)

Wilkerson et al. 1993 (1990 Central America Spanish key translation)

Harbach 1994a (rev. genus Anopheles internal classification)

Rattanarithikul & Panthusiri 1994 (keys, medical importance species;  Thailand)

Lambert & Spencer 1995 (genetics, evolution)

Nagpal & Sharma 1995 (India)

Rubio-Palis & Zimmerman 1997 (review, bionomics, malaria vectors)

Lu et al. 1997 (China)

Harbach 2004 (classification, phylogeny)

Darsie & Ward 1981 (keys, North America)

Darsie & Ward 2005 (keys, North America)

Sallum et al. 2000 (phylogeny)

Krzywinski et al. 2001a (molecular phylogeny)

Krzywinski et al. 2001b (molecular phylogeny)

Sallum et al. 2002 (phylogeny)

Krzywinski & Besansky 2003 (review)

Harbach et al. 2005 (Baimaia, new subgenus)

Harbach & Kitching 2005 (phylogeny)

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

Rattanarithikul et al. 2006b (F*, L*; keys, bionomics, review, Thailand)

Corbel et al. 2007 (insecticide resistance)

González & Carrejo 2007 (keys, Colombia)

Harbach 2007 (phylogeny)

Azari-Hamidian & Harbach, 2009 (keys, distribution; Iran)

Morgan et al. 2009 (molecular phylogeny; Neocellia Series)

Service 2010 (review, Anopheles names)

Sinka et al. 2010a (vector distribution; Africa, Europe, Middle East)

Sinka et al. 2010b (vector distribution; Americas)

Sinka et al. 2011 (vector distribution;  Asia-Pacific)

Antonio-Nikondjio & Simard 2013 (taxonomic review; nili, moucheti)

Beebe et al. 2013 (systematics, bionomics; SW Pacific)

Boete & Beisel 2013 (transgenic mosquitoes)

Choochote & Saeung 2013 (systematics methods)

Conn et al. 2013 (phylogeography, malaria; S America)

Dev & Sharma 2013 (review, malaria vectors; India)

Dia et al. 2013 (tax. Funestus Group)

Drame et al. 2013 (salivary biomarkers)

Duchet et al. 2013 (control issues)

Durnez & Coosemans 2013 (malaria transmission)

Garros 2013 (systematics, genetics, morphology)

Gendrin & Christophides 2013 (mosquito microbiota)

Harbach 2013b (phylogeny classification)

Lahondere & Lazzari 2013 (bionomics)

Lanzaro & Lee 2013 (taxonomy, genetics, distribution, Gambiae Complex)

Manguin et al. 2013 (Anopheles bacterial fauna)

Obsomer et al. 2013 (surveillance methods)

Rejmankova et al. 2013 (bionomics; larvae)

Roberts et al. 2013 (control issues)

Sinka 2013 (distribution bionomics, niche models)

Suwonkerd et al. 2013 (bionomics, malaria; SE Asia)

Vythilingam & Hii 2013 (monkey malaria; SE Asia)

Freitas et al. 2015 (molecular phylogeny)

Harbach & Kitching 2015 (Anopheles phylogeny; Christya to subgenus)

Neafsey et al. 2015 (molecular phylogeny; 16 genomes)

Gunathilaka et al. 2014 (key, larvae; Sri Lanka)

Gunathilaka 2017 (key, adults; Sri Lanka)

Foster et al. 2017 (phylogeny)

Kyalo et al. 2017 (distribution subsaharan Africa)

Brack et al. 2018 (virus vectors, Africa)

Bourke et al. 2018 (mtDNA phylogeny Neotropical Anopheles)

Harbach 2018a (Anopheles genera comment)

Minkeu & Vernick 2018 (review arbovirus vectors)

 

VALID SUBGENERA

Anopheles Meigen [Ano.]

Baimaia Harbach, Rattanarithikul & Harrison [Bmi.]

Cellia Theobald [Cel.]

Christya Theobald [Chs.]

Kerteszia Theobald [Ker.]

Lophopodomyia Antunes [Lph.]

Nyssorhynchus Blanchard [Nys.]

Stethomyia Theobald [Ste.]

 

CURRENT GENERIC SYNONYMS

syn. Cycloleppteron Theobald 1901b: 234 (July 15); 1901c: 312 (Nov. 23) (as genus). Type species: Anopheles grabhamii Theobald.

syn. Cyclolepidopteron Blanchard 1901: 1046 (Dec. 6) (as genus). Type species: Anopheles grabhamii Theobald.

syn. Rossia Theobald 1902b: 183 (not Owen 1835; not Bonaparte 1838) (as genus). Type species: Anopheles sinensis Wiedemann.

syn. Myzorhynchus Blanchard 1902: 795 (as genus). Type species: Anopheles sinensis Wiedemann.

syn. Arribalzagia Theobald 1903a:13, 81 (as genus). Type species: Arribalzgia maculipes Theobald.

syn. Lophoscelomyia Theobald 1904c: 12 (January) (as genus). Type species: Lophoscelomyia asiatica Leicester.

syn. Lophomyia Giles 1904a: 366 (as genus) (not Fitzinger 1863) (Dec.). Type species: Lophomyia asiatica Giles.

syn. Arribalzagaia Blanchard 1905: 624 (emendment of Arribalzagia).

syn. Nototricha Coquillett 1906d: 13 (as genus). Type species: Cycloleppteron mediopunctatus Theobald.

syn. Coelodiazesis Dyar & Knab 1906d: 177 (as genus). Type species: Anopheles barberi Coquillett.

syn. Neostethopheles James 1910b: 98 (as genus). Type species: Anopheles aitkenii James.

syn. Patagiamyia James 1910b: 98 (as genus). Type species: Anopheles gigas Giles.

syn. Cyclophorus Eysell 1912: 422 (not Montfort 1910, not Agassiz 1846, not Pfeiffer 1847, not Kraatz 1880) (as genus). Type species: Anopheles nigripes Staeger.

syn. Proterorhynchus Brèthes 1912: 10 14 (as genus). Type species: Proterorhynchus argentinus Brèthes.

syn. Memnemyia Strickland 1915: 204 (as genus). Type species: Anopheles brevipalpis Roper.

syn. Anopheles subgenus Shannoniella da Fonseca & da Silva Ramos 1940 (March)(1939): 384. Type species: Shannoniella limai da Fonseca & da Silva Ramos.

syn. Anopheles subgenus Shannonesia da Fonseca & da Silva Ramos (as subgenus of genus Anopheles) in Neiva 1940b: 966. Type species: Shannonesia limai da Fonseca & da Silva Ramos.

syn. Anopheles subgenus Ayrozamyia Coutinho & Farias 1942: 343 (as genus). Type species: Ayrozamyia tibiamaculatus Neiva.

syn. Anopheles subgenus Russellia Vargas 1943a: 60, 63, 68-70. Type species: Anopheles xelajuensis Leon.

syn. species group, Maculipennia Buonomini & Mariani 1946:105. (Not valid since no type was designated; not absolute tautonymy).

syn. Anopheles subgenus Missirolimyia Vargas 1950a: 80. Type species: Missirolymyia algeriensis Theobald.

syn. Anopheles subgenus Maculipennia Buonomini & Mariani 1953: 178 (A, L, E; tax.) Type species: Maculipennia maculipennis Meigen.

 

CITED REFERENCES

Agassiz, L. (1846). Nomenclatoris zoologici index universalis, continens nomina systematica classium, ordinum, familiarum et generum animalium omnium, tam viventium quam fossilium, secundum ordinem alphabeticum unicum disposita, adjectis homonymiis plantarum, nec non variis adnotionibus et emendationibus. Jent et Gassmann, Soloduri [= Solothrun, Switzerland], 393 pp.

Antonio-Nkondjio, C., & Simard, F. (2013). Highlights on Anopheles nili and Anopheles moucheti, malaria vectors in Africa. In S. Manguin (Ed.), Anopheles mosquitoes: new insights into malaria vectors (pp. 221–238). Janeza Trdine 9, 51000 Rijeka, Croatia: InTech.

Azari-Hamidian, S., & Harbach, R.E. (2009). Keys to the adult females and fourth-instar larvae of the mosquitoes of Iran (Diptera: Culicidae). Zootaxa, 2078, 1–33.

Baba, K. (1950). A study of anopheline mosquitoes in relation to the epidemiology of malaria in Canton delta, South China. I. Species of anopheline mosquitoes recorded from Kwangtong Province, South China and their breeding habits. Transactions of the Kansai Entomological Society (China), 15(2), 1–11.

Beebe, N. W., Russell, T. L., Burkot, T. R., Lobo, N. F., & Cooper, R. D. (2013). The systematics and bionomics of malaria vectors in the Southwest Pacific. In S. Manguin (Ed.), Anopheles mosquitoes - New insights into malaria vectors (pp. 357–394). Janeza Trdine 9, 51000 Rijeka, Croatia: InTech.

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

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.

Blanchard, R. (1901). Observations sur quelques moustiques. Comptes Rendus des Seances de la Société de Biologie, 53, 1045–1046.

Blanchard, R. (1902). Nouvelle note sur les moustiques. Société de Biologie, 54, 793–795.

Blanchard, R. (1905) Les moustiques, histoire naturelle et médicale. F. R. de Rudeval, Imprimeur-Editeur, Paris. 673 pp.

Boete, C., & Beisel, U. (2013). Transgenic mosquitoes for malaria control: from the bench to the public opinion survey. In S. Manguin (Ed.), Anopheles mosquitoes - New insights into malaria vectors (pp. 797–813). Janeza Trdine 9, 51000 Rijeka, Croatia: InTech.

Bonne-Wepster, J., & Swellengrebel, N.H. (1953). The anopheline mosquitoes of the Indo-Australian Region. Amsterdam: J. H. de Bussy.

Bourke, B P., Foster, P.G., Bergo, E.S., Calado, D.C., & Sallum, M.A.M. (2018). Phylogenetic relationships among species of Anopheles (Nyssorhynchus) (Diptera, Culicidae) based on nuclear and mitochondrial gene sequences. Acta Tropica, 114, 88–96.

Brèthes, J. (1912). Los mosquitos de la República Argentina. Bol Inst ent Pat veg Buenos Aires, 1–48.

Buonomini, G. & Mariani, M. (1953). World anophelines belonging to the subgenus Maculipennia Buonomini e Mariani. Rivista di Malariologia, 32, 173–188.

Buonomini, G., & Mariani, M. (1946). Qual'e il valore sistematico del nome maculipennis Meig. Rivista di Malariologia, 25, 101–107.

Cagampang-Ramos, A., & Darsie Jr., R.F. (1970). Illustrated keys to the Anopheles mosquitoes of the Philippine Islands. US Air Force 5TH Epidemiology Flight, PACAF, Technical Report, 1–49.

Causey, O.R., Deane, L.M., & Deane, M.P. (1946). An illustrated key by male genitalic characteristics for the identification of thirty-four species of Anophelini from the northeast and Amazon regions of Brazil, with a note on dissection technique. American Journal of Hygiene Monograph Series, 18, 21–31.

Choochote, W., & Saeung, A. (2013). Genetic and phenetic approaches to Anopheles systematics. In S. Manquin (Ed.), Anopheles mosquitoes - New insights into malaria vectors (pp. 57–79). Janeza Trdine 9, 51000 Rijeka, Croatia: InTech.

Chow, C.Y. (1949b). The identification and distribution of Chinese anopheline mosquitoes. Journal of the National Malaria Society, 8(2), 121–131.

Christophers, S.R. (1915). The male genitalia of Anopheles. Indian Journal of Medical Research, 3, 371–394.

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.

Colless, D.H. (1948). The anopheline mosquitoes of north-west Borneo. Proceedings of the Linnean Society of New South Wales, 73, 71–119.

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