Taxonomy of Anopheles

Anopheles is a genus of mosquitoes (Culicidae). Of about 484 recognised species, over 100 can transmit human malaria, but only 30–40 commonly transmit parasites of the genus Plasmodium that cause malaria, which affects humans in endemic areas. Anopheles gambiae is one of the best known, because of its predominant role in the transmission of the deadly species Plasmodium falciparum.

Anopheles
Anopheles stephensi
Scientific classification
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Order:	Diptera
Superfamily:	Culicoidea
Family:	Culicidae
Subfamily:	Anophelinae
Genus:	Anopheles
Subgenera
Anopheles Baimaia Cellia Kerteszia Lophopodomyia Nyssorhynchus Stethomyia

Classification

Range map of selected Anopheles species

The classification of this genus began in 1901 with Frederick Vincent Theobald.[1] Despite the passage of time, the taxonomy remains incompletely settled.[2][3][4] Classification into species is based on morphological characteristics - wing spots, head anatomy, larval and pupal anatomy, and chromosome structure, and more recently on DNA sequences.[5][6][7][8]

The genus Anopheles belongs to a subfamily Anophelinae with three genera: Anopheles Meigen (nearly worldwide distribution), Bironella Theobald (Australia only: 11 described species) and Chagasia Cruz (Neotropics: four described species). The genus Bironella has been divided into three subgenera: Bironella Theobald (two species), Brugella Edwards (three species) and Neobironella Tenorio (three species). Bironella appears to be the sister taxon to the Anopheles, with Chagasia forming the outgroup in this subfamily.

The type species of the genus is Anopheles maculipennis.[9]

Subgenera

The genus has been subdivided into seven subgenera based primarily on the number and positions of specialized setae on the gonocoxites of the male genitalia. The system of subgenera originated with the work of Christophers, who in 1915 described three subgenera: Anopheles (widely distributed), Myzomyia (later renamed Cellia) (Old World) and Nyssorhynchus (Neotropical). Nyssorhynchus was first described as Lavernia by Theobald. Frederick Wallace Edwards in 1932 added the subgenus Stethomyia (Neotropical distribution). Kerteszia was also described by Edwards in 1932, but then was recognised as a subgrouping of Nyssorhynchus. It was elevated to subgenus status by Komp in 1937; this subgenus is also found in the Neotropics. Two additional subgenera have since been recognised: Baimaia (Southeast Asia only) by Harbach et al. in 2005 and Lophopodomyia (Neotropical) by Antunes in 1937.

One species within each subgenus has been identified as the type species of that particular subgenus:

• Subgenus Anopheles - Anopheles maculipennis Meigen 1918[9]
• Subgenus Baimaia - Anopheles kyondawensis Abraham 1947[10]
• Subgenus Cellia - Anopheles pharoensis Giles 1899
• Subgenus Kerteszia - Anopheles boliviensis Theobald 1905
• Subgenus Lophopodomyia - Anopheles squamifemur Antunes 1937
• Subgenus Nyssorhynchus - Anopheles argyritarsis Robineau-Desvoidy 1827
• Subgenus Stethomyia - Anopheles nimbus Theobald 1902[11]

Within the genus Anopheles are two main groupings, one formed by the Cellia and Anopheles subgenera and a second by Kerteszia, Lophopodomyia, and Nyssorhynchus. Subgenus Stethomyia is an outlier with respect to these two taxa. Within the second group, Kerteszia and Nyssorhynchus appear to be sister taxa. Cellia appears to be more closely related to the Kerteszia-Lophopodomyia-Nyssorhynchus group than to Anopheles or Stethomyia, tentatively suggesting the following branching order: ( Stethomyia ( Anopheles ( Cellia ( Lophopodomyia ( Kerteszia, Nyssorhynchus))))).

The number of species currently recognised within the subgenera is given here in parentheses: Anopheles (206 species), Baimaia (one), Cellia (239), Kerteszia (12), Lophopodomyia (six), Nyssorhynchus (34) and Stethomyia (five).

The subgenus Baimaia may be elevated to genus level, as it appears to be a sister group to Bironella and all other Anopheles.[12]

The ancestors of Drosophila and Anopheles diverged 260 million years ago. The Old and New World Anopheles species subsequently diverged between 80 and 95 million years ago.[13][14]

Divisions below subgenus

Taxonomic units between subgenus and species are not currently recognised as official zoological names. In practice, a number of taxonomic levels have been introduced. The larger subgenera (Anopheles, Cellia, and Nyssorhynchus) have been subdivided into sections and series, which in turn have been divided into groups and subgroups. Below subgroup but above species level is the species complex. Taxonomic levels above species complex can be distinguished on morphological grounds. Species within a species complex are either morphologically identical or extremely similar and can only be reliably separated by microscopic examination of the chromosomes or DNA sequencing. The classification continues to be revised.

The first species complex was described in 1926 when the problem of nontransmission of malaria by Anopheles gambiae was solved by Falleroni, who recognised that An. gambiae was a complex of six species, of which only four could transmit malaria. This complex has subsequently been revised to a total of seven species of which five transmit malaria.

Subgenus Nyssorhynchus has been divided in three sections: Albimanus (19 species), Argyritarsis (11 species) and Myzorhynchella (four species). The Argyritarsis section has been subdivided into Albitarsis and Argyritarsis groups.

The Anopheles group was divided by Edwards into four series: Anopheles (worldwide), Myzorhynchus (Palearctic, Oriental, Australasian and Afrotropical), Cycloleppteron (Neotropical) and Lophoscelomyia (Oriental); and two groups, Arribalzagia (Neotropical) and Christya (Afrotropical). Reid and Knight (1961) modified this classification by subdividing the subgenus Anopheles into two sections, Angusticorn and Laticorn and six series. The division was based on the shape of their pupal trumpets. The Laticorn section was created for those species with wide, funnel-shaped trumpets having the longest axis transverse to the stem, and the Angusticorn section for species with semitubular trumpets having the longest axis vertical more or less in line with the stem. The earlier Arribalzagia and Christya groups were considered to be series. The Angusticorn section includes members of the Anopheles, Cycloleppteron, and Lophoscelomyia series, and the Laticorn section includes the Arribalzagia (24 species), Christya, and Myzorhynchus series.

Cellia is the largest subgenus: all species within this subgenus are found in the Old World. It has been divided into six series - Cellia (eight species), Myzomyia (69 species), Neocellia (33 species), Neomyzomyia (99 species), Paramyzomyia (six species) and Pyretophorus (22 species). This classification was developed by Grjebine (in 1966), Reid (in 1968), and Gillies & de Meillon (also in 1968)[15] based on the work by Edwards in 1932. Series definition within this subgenus is based on the cibarial armature - a collection of specialized spicules borne ventrally at the posterior margin of the cibarium - which was first used as a taxonomic method by Christophers in 1933.

Kerteszia is a small subgenus found in South America whose larvae have specific ecological requirements; these can only develop within water that accumulates at the base of the follicular axis of the epiphytic Bromeliaceae. Unlike the majority of mosquitoes, species in this subgenus are active during the day.

Within a number of species, separate subspecies have been identified. The diagnostic criteria and characteristic features of each subgenus are discussed on the own page.

Species complexes

Anopheles nuneztovari is a species complex with at least one occurring in Colombia and Venezuela and another occurring in the Amazon Basin.[16] These clades appear to have diverged and expanded in the Pleistocene.

Medical and veterinary importance

The first demonstration that mosquitoes could act as vectors of disease was by Patrick Manson, a British physician working in China, who showed that a Culex species could transmit filariasis in 1878. This was then followed in 1897 by Ronald Ross, who showed avian malaria could also be transmitted by a species of Culex. Grassi in Italy showed that the species causing human malaria were transmitted by species of the genus Anopheles in 1898. Anopheles gambiae (then Anopheles coastalis), the most important of the vectors transmitting human malaria, was first recognised as such in 1899 at Freetown, Sierra Leone.[17] It was later realised that only a small number of species of mosquitoes were responsible for the vast majority of human malaria and other diseases. This generated a considerable interest in the taxonomy of this and other mosquito genera.

The species of the subgenera Baimaia, Lophopodomyia, and Stethomyia are not of medical importance.

All species within the subgenus Anopheles known to carry human malaria lie within either the Myzorhynchus or the Anopheles series. Anopheles maculipennis s.l. is a known vector of West Nile virus.

Six species in the subgenus Kerteszia can carry human malaria. Of these, only An. bellator and An. cruzii are of importance. Anopheles bellator can also transmit Wuchereria bancrofti.

Several species of the subgenus Nyssorhynchus are of medical importance.

All series of the subgenus Cellia contain vectors of malarial protozoa and microfilariae.

Five species of anopheline mosquitoes (An. arabiensis, An. funestus, An. gambiae, An. moucheti, An. nili) all belonging to the subgenus Cellia are responsible for over 95% of total malaria transmission for Plasmodium falciparum in continental sub-Saharan Africa.

Anopheles sundaicus and An. subpictus are important vectors of Plasmodium vivax.

Species evolution

The Anopheles gambiae complex has a number of important malaria vectors. A chromosomal study suggests that An. merus is the basal member of this complex and is sister species to An. gambiae.[18] The two species An. quadriannulatus A and An. quadriannulatus B - neither of whom are vectors for malaria - are derived from An. gambiae.

The subgenera Anopheles and Cellia appear to be sister clades as do Kerteszia and Nyssorhynchus.[19]

Species listing

Species that have been shown to be vectors of human malaria are marked with a star (*) after the name.

Subgenus Anopheles

Anopheles anthropophagus* Xu & Feng 1975

Anopheles confusa[20]

Anopheles derooki Soesilo & Van Slooten 1931

Anopheles gracilis Theobald 1905

Anopheles hollandi Taylor 1934

Anopheles obscura Tenorio 1975

Anopheles papuae Swellengrebel & Swellengrebel de Graaf 1919

Anopheles simmondsi Tenorio 1977

Anopheles travestita Brug 1928

Section Angusticorn

Series Anopheles

Anopheles algeriensis Theobald 1903

Anopheles concolor Edwards 1938

Anopheles marteri[21]

subspecies marteri

subspecies sogdianus Keshishian

Complex Claviger (Coluzzi et al. 1965)

Anopheles claviger* Meigen 1804

Anopheles petragnani Del Vecchio 1939

Group Aitkenii (Reid & Knight, 1961)

Anopheles aberrans Harrison & Scanlon 1975

Anopheles acaci Baisas 1946

Anopheles aitkenii James 1903

Anopheles bengalensis Puri 1930

Anopheles borneensis McArthur 1949

Anopheles fragilis Theobald 1903

Anopheles insulaeflorum Swellengrebel & Swellengrebel de Graaf 1919

Anopheles palmatus Rodenwaldt 1926

Anopheles peytoni Kulasekera Harrison & Amerasinghe 1989

Anopheles pilinotum Harrison & Scanlon 1974

Anopheles pinjaurensis Barraud 1932

Anopheles stricklandi Reid 1965

Anopheles tigertti Scanlon & Peyton 1967

Group Alongensis (Phan et al. 1991)

Anopheles alongensis Evenhuis 1940

Anopheles cucphuongensis Phan, Manh, Hinh & Vien 1991

Group Atratipes (Lee et al. 1987)

Anopheles atratipes Skuse 1889

Anopheles tasmaniensis Dobrowtorsky 1966

Group Culiciformis (Reid & Knight 1961)

Anopheles culiciformis Cogill 1903

Anopheles sintoni Puri 1929

Anopheles sintonoides Ho 1938

Group Lindesayi (Reid & Knight 1961)

Anopheles mengalengensis Ma 1981

Anopheles nilgiricus Christophers 1924

Anopheles wellingtonianus Alcock 1912

Complex Gigas (Harrison et al. 1991)

Anopheles baileyi Edwards 1923

Anopheles gigas Giles 1901

subspecies crockeri Colless

subspecies danaubento Mochtar & Walandouw

subspecies formosus Ludlow

subspecies gigas Giles

subspecies oedjalikalah Nainggolan

subspecies pantjarbatu Waktoedi

subspecies refutans Alcock

subspecies simlensis James

subspecies sumatrana Swellengrebel & Rodenwaldt

Complex Lindesayi (Harrison et al. 1991)

Anopheles lindesayi Giles 1900

subspecies benguetensis King

subspecies cameronensis Edwards

subspecies japonicus Yamada

subspecies lindesayi Giles

subspecies pleccau Koidzumi

Group Maculipennis (Reid & Knight 1961)

Anopheles atropos Dyar & Knab 1906

Anopheles aztecus Hoffmann 1935

Anopheles lewisi Ludlow 1920

Anopheles walkeri[1]

Complex Quadrimaculatus (Linton 2004)

Anopheles diluvialis Reinert 1997

Anopheles inundatus Reinert 1997

Anopheles maverlius Reinert 1997

Anopheles smaragdinus Reinert 1997

Anopheles quadrimaculatus* Say 1824

Subgroup Freeborni (Linton 2004)

Anopheles earlei Vargas 1943

Anopheles freeborni* Aitken 1939

Anopheles hermsi Barr & Guptavanij 1989

Subgroup Maculipennis (Linton 2004)

Anopheles artemievi Gordeyev, Zvantsov, Goryacheva, Shaikevich & Yezhov

Anopheles atroparvus* Van Thiel 1927

Anopheles beklemishevi Stegnii & Kabanova 1976

Anopheles daciae Linton, Nicolescu & Harbach 2004

Anopheles labranchiae* Falleroni 1926

Anopheles maculipennis[9]

Anopheles martinius Shingarev 1926

Anopheles melanoon* Hackett 1934

Anopheles messeae* Falleroni 1926

Anopheles occidentalis Dyar & Knab 1906

Anopheles persiensis Linton, Sedaghat & Harbach 2003

Anopheles sacharovi* Favre 1903

Anopheles sicaulti Roubaud 1935

Anopheles subalpinus Hackett & Lewis 1935

Group Plumbeus (Reid & Knight 1961)

Anopheles arboricola Zavortink 1970

Anopheles barberi Coquillett 1903

Anopheles barianensis James 1911

Anopheles fausti Vargas 1943

Anopheles judithae Zavortink 1969

Anopheles omorii Sakakibara 1959

Anopheles plumbeus* Stegnii & Kabanova 1828

Anopheles powderi Zavortink 1970

Anopheles xelajuensis De Leon 1938

Group Pseudopunctipennis (Reid & Knight 1961)

Anopheles chiriquiensis Komp 1936

Anopheles franciscanus McCracken 1904

Anopheles hectoris Giaquinto-Mira 1931

Anopheles tibiamaculatus Neiva 1906

Anopheles eiseni Coquillett 1902

subspecies eiseni Coquillett

subspecies geometricus Corrêa

Anopheles parapunctipennis* Martini 1932

subspecies guatemalensis de Leon

subspecies parapunctipennis Martini

Anopheles pseudopunctipennis[1]

subspecies levicastilloi Levi-Castillo

subspecies neghmei Mann

subspecies noei Mann

subspecies patersoni Alvarado & Heredia

subspecies pseudopunctipennis Theobald

subspecies rivadeneirai Levi-Castillo

Group Punctipennis (Reid & Knight 1961)

Anopheles perplexens Ludlow 1907

Anopheles punctipennis Say 1823

Complex Crucians (Wilkerson et al. 2004)

Anopheles bradleyi King 1939

Anopheles crucians Wiedemann 1828

Anopheles georgianus King 1939

Group Stigmaticus (Reid & Knight 1961)

Anopheles colledgei Marks 1956

Anopheles corethroides Theobald 1907

Anopheles papuensis Dobrowtorsky 1957

Anopheles powelli Lee 1944

Anopheles pseudostigmaticus Dobrowtorsky 1957

Anopheles stigmaticus Skuse 1889

Series Cycloleppteron (Edwards 1932)

Anopheles annulipalpis Lynch 1878

Anopheles grabhamii[1]

Series Lophoscelomyia (Edwards 1932)

Anopheles bulkleyi Causey 1937

Group Asiaticus (Reid 1968)

Anopheles annandalei Prashad 1918

Anopheles noniae Reid 1963

Subgroup Asiaticus (Rattanarithikul et al. 2004)

Anopheles asiaticus Leicester 1903

Subgroup Interruptus (Rattanarithikul et al. 2004)

Anopheles interruptus Puri 1929

Section Laticorn (Reid & Knight 1961)

Series Arribalzagia (Root 1922)

Anopheles anchietai Correa & Ramalho 1968

Anopheles apicimacula Dyar & Knab 1906

Anopheles bustamentei Galvao 1955

Anopheles calderoni Wilkerson 1991

Anopheles costai Da Fonseca & Da Silva Ramos 1939

Anopheles evandroi Da Costa Lima 1937

Anopheles fluminensis Root 1927

Anopheles forattinii Wilkerson 1999

Anopheles gabaldoni Vargas 1941

Anopheles guarao Anduze & Capdevielle 1949

Anopheles intermedius* Peryassu 1908

Anopheles maculipes Theobald 1903

Anopheles malefactor Dyar & Knab 1907

Anopheles mattogrossensis Lutz & Neiva 1911

Anopheles mediopunctatus Lutz 1903

Anopheles minor Da Costa Lima 1929

Anopheles neomaculipalpis Curry 1931

Anopheles peryassui Dyar & Knab 1908

Anopheles pseudomaculipes Peryassu 1908

Anopheles punctimacula Dyar & Knab 1906

Anopheles mediopunctatus Lutz 1903

Anopheles rachoui Galvao 1952

Anopheles shannoni Davis 1931

Anopheles veruslanei Vargas 1979

Anopheles vestitipennis Dyar & Knab 1906

Series Christya (Christophers 1924)

Anopheles implexus Theobald 1903

Anopheles okuensis Brunhes, le Goff & Geoffroy 1997

Series Myzorhynchus (Edwards 1932)

Anopheles obscurus Grunberg 1905

Anopheles bancroftii Giles 1902

Anopheles barbirostris* Van der Wulp 1884

Anopheles pollicaris Reid 1962

Group Albotaeniatus (Reid & Knight 1961)

Anopheles albotaeniatus Theobald 1903

Anopheles balerensis Mendoza 1947

Anopheles ejercitoi Mendoza 1947

Anopheles montanus Stanton & Hacker 1917

Anopheles saperoi Bohart & Ingram 1946

subspecies ohamai Ohama

subspecies saperoi Bohart & Ingram

Group Bancroftii (Reid & Knight 1961)

Anopheles pseudobarbirostris Ludlow 1935

Anopheles bancroftii Giles 1902

subspecies bancroftii Giles

subspecies barbiventris Brug

Group Barbirostris (Reid & Knight 1961)

Anopheles freyi Meng 1957

Anopheles koreicus Yamada & Watanabe 1918

Subgroup Barbirostris (Reid 1968)

Anopheles barbirostris van der Wulp 1884

Anopheles campestris Reid 1962

Anopheles donaldi Reid 1962

Anopheles franciscoi Reid 1962

Anopheles hodgkini Reid 1962

Anopheles pollicaris Reid 1962

Subgroup Vanus (Reid 1968)

Anopheles ahomi Chowdhury 1929

Anopheles barbumbrosus Strickland & Chowdhury 1927

Anopheles manalangi Mendoza 1940

Anopheles reidi Harrison 1973

Anopheles vanus Walker 1859

Group Coustani (Reid & Knight 1961)

Anopheles caliginosus De Meillon 1943

Anopheles coustani Laveran 1900

Anopheles crypticus Coetzee 1994

Anopheles fuscicolor Van Someren 1947

Anopheles namibiensis Coetzee 1984

Anopheles paludis Theobald 1900

Anopheles symesi Edwards 1928

Anopheles tenebrosus Donitz 1902

Anopheles ziemanni Grunberg 1902

Group Hyrcanus (Reid 1953)

Anopheles anthropophagus Xu and Feng 1975

Anopheles argyropus Swellengrebel 1914

Anopheles belenrae Rueda 2005

Anopheles changfus Ma 1981

Anopheles chodukini Martini 1929

Anopheles dazhaius Ma 1981

Anopheles engarensis Kanda & Oguma 1978

Anopheles hailarensis Xu JinJiang & Luo XinFu 1998

Anopheles heiheensis Ma 1981

Anopheles hyrcanus* Pallas 1771

Anopheles junlianensis Lei 1996

Anopheles kiangsuensis Xu and Feng 1975

Anopheles kleini Rueda 2005

Anopheles kummingensis Dong & Wang 1985

Anopheles kweiyangensis Yao & Wu 1944

Anopheles liangshanensis Kang Tan Cao Cheng Yang & Huang 1984

Anopheles nimpe Nguyen, Tran & Harbach

Anopheles pseudopictus Graham 1899

Anopheles pullus Yamada 1937

Anopheles sinensis* Wiedemann 1828

Anopheles sineroides Yamada 1924

Anopheles xiaokuanus Ma 1981

Anopheles xui Dong, Zhou, Dong & Mao 2007

Anopheles yatsushiroensis Miyazaki 1951

Subgroup Lesteri (Harrison 1972)

Anopheles crawfordi Reid 1953

Anopheles kiangsuensis Xu & Feng 1975

Anopheles lesteri de Meillon 1931

Anopheles paraliae Sandosham 1959

Anopheles peditaeniatus Leicester 1908

Anopheles vietnamensis Manh Hinh & Vien 1993

Subgroup Nigerrimus (Harrison 1972)

Anopheles nigerrimus* Giles 1900

Anopheles nitidus Harrison, Scanlon & Reid 1973

Anopheles pseudosinensis Baisas 1935

Anopheles pursati Laveran 1902

Group Umbrosus (Reid 1950)

Anopheles brevipalpis Roper 1914

Anopheles brevirostris Reid 1950

Anopheles hunteri Strickland 1916

Anopheles samarensis Rozeboom 1951

Anopheles similissimus Strickland & Chowdhury 1927

Subgroup Baezai (Rattanarithikul et al. 2004)

Anopheles baezai Gater 1934

Subgroup Letifer (Reid 1968)

Anopheles collessi Reid 1963

Anopheles letifer* Sandosham 1944

Anopheles roperi Reid 1950

Anopheles whartoni Reid 1963

Subgroup Separatus (Rattanarithikul et al. 2004)

Anopheles separatus Leicester 1908

Subgroup Umbrosus (Rattanarithikul et al. 2004)

Anopheles umbrosus Theobald 1903

Subgenus Baimaia

Anopheles kyondawensisAbraham 1947[10]

Subgenus Cellia

Anopheles rageaui Mattingly and Adam

Series Cellia (Christophers 1924)

Anopheles argenteolobatus Gough 1910

Anopheles brumpti Hamon & Rickenbach 1955

Anopheles carnevalei Brunhes le Goff & Geoffroy 1999

Anopheles cristipalpis Service 1977

Anopheles murphyi Gillies 1968[15]

Anopheles pharoensis Theobald 1901[1]

Anopheles swahilicus Gillies 1964

Group Squamosus (Grjebine 1966)

Anopheles cydippis de Meillon 1931

Anopheles squamosusTheobald 1901[1]

Series Myzomyia

Anopheles apoci Marsh 1933

Anopheles azaniae Bailly-Choumara 1960

Anopheles barberellus Evans 1932

Anopheles brunnipes Theobald 1910

Anopheles domicola Edwards 1916

Anopheles dthali Patton 1905

Anopheles erythraeus Corradetti 1939

Anopheles ethiopicus Gillies & Coetzee 1987

Anopheles flavicosta Edwards 1911

Anopheles fontinalis Gillies 1968[15]

Anopheles majidi Young & Majid 1928

Anopheles moucheti* Evans 1925

subspecies bervoetsi D'Haenans 1961

subspecies moucheti Evans 1925

subspecies nigeriensis

Anopheles schwetzi Evans 1934

Anopheles tchekedii de Meillon & Leeson 1940

Anopheles walravensi Edwards 1930

Group Demeilloni (Gillies & De Meillon 1962)

Anopheles carteri Evans & de Meillon 1933

Anopheles demeilloni Evans 1933

Anopheles freetownensis Evans 1925

Anopheles garnhami Edwards 1930

Anopheles keniensis Evans 1931

Anopheles lloreti Gil Collado 1936

Anopheles sergentii* Theobald 1907

subspecies macmahoni Evans 1936

subspecies sergentii Theobald 1907

Group Funestus (Garros et al 2004)

Anopheles jeyporiensis James 1902

Subgroup Aconitus (Chen et al. 2003)

Anopheles aconitus Dönitz 1902

Anopheles filipinae Manalang 1930

Anopheles mangyanus Banks 1906

Anopheles pampanai Buttiker & Beales 1959

Anopheles varuna Iyengar 1924

Subgroup Culicifacies (Garros et al. 2004)

Anopheles culicifacies* Giles 1901

Subgroup Funestus (Garros et al. 2004)

Anopheles aruni Sobti 1968

Anopheles confusus Evans & Leeson 1935

Anopheles funestus* Giles 1900

Anopheles funestus-like* Spillings et al 2009[22]

Anopheles longipalpis Type C Koekemoer et al. 2009

Anopheles parensis Gillies 1962

Anopheles vaneedeni Gillies & Coetzee 1987

Subgroup Minimus (Chen et al. 2003)

Anopheles flavirostris* Ludlow 1914

Anopheles leesoni Evans 1931

Anopheles longipalpis Type A Koekemoer et al 2009

Complex Fluviatilis (Salara et al. 1993)

Anopheles fluviatilis* (species S, T, U, V) James 1902[23]

Complex Minimus (Green et al. 1990)

Anopheles harrisoni Harbach & Manguin 2007

Anopheles minimusTheobald 1901*[1]

Subgroup Rivulorum (Garros et al 2004)

Anopheles brucei Service 1960

Anopheles fuscivenosus Leeson 1930

Anopheles rivulorum* Leeson 1935

Group Marshallii[15]

Anopheles austenii Theobald 1905

Anopheles berghei Vincke & Leleup 1949

Anopheles brohieri Edwards 1929

Anopheles gibbinsi Evans 1935

Anopheles hancocki Edwards 1929

Anopheles hargreavesi Evans 1927

Anopheles harperi Evans 1936

Anopheles mortiauxi Edwards 1938

Anopheles mousinhoi de Meillon & Pereira 1940

Anopheles njombiensis Peters 1955

Anopheles seydeli Edwards 1929

Complex Marshalli (Gillies & Coetzee 1987)

Anopheles hughi Lambert & Coetzee 1982

Anopheles kosiensis Coetzee, Segerman & Hunt 1987

Anopheles letabensis Lambert & Coetzee 1982

Anopheles marshallii Theobald 1903

Group Wellcomei[15]

Anopheles distinctus Newstead & Carter 1911

Anopheles erepens Gillies 1958

Anopheles theileri Edwards 1912

Anopheles wellcomei Theobald 1904

subspecies ugandae Evans 1934

subspecies ungujae White 1975

subspecies wellcomei Theobald 1904

Series Neocellia (Christophers 1924)

Anopheles ainshamsi Gad, Harbach & Harrison 2006

Anopheles dancalicus Corradetti 1939

Anopheles hervyi Brunhes, le Goff & Geoffroy 1999

Anopheles jamesiiTheobald 1901[1]

Anopheles karwari* James 1903

Anopheles maculipalpis Giles 1902

Anopheles moghulensis Christophers 1924

Anopheles paltrinierii Shidrawi & Gillies 1988

Anopheles pattoni Christophers 1926

Anopheles pretoriensis Theobald 1903

Anopheles pulcherrimusTheobald 1902*[11]

Anopheles rufipes Gough 1910

subspecies broussesi Edwards 1929

subspecies rufipes Gough 1910

Anopheles salbaii Maffi & Coluzzi 1958

Anopheles splendidus Koidzumi 1920

Anopheles theobaldi Giles 1901

Complex Stephensi

Anopheles stephensi* Liston 1901

Complex Superpictus

Anopheles superpictus* Grassi 1899

Group Annularis (Reid 1968)

Anopheles pallidus Theobald 1901[1]

Anopheles philippinensis* Ludlow 1902

Anopheles schueffneri Stanton 1915

Complex Annularis (Reid 1968)

Anopheles annularis* van der Wulp 1884

Complex Nivipes (Green et al. 1985)

Anopheles nivipes Theobald 1903

Group Jamesii (Rattanarithikul et al. 2004)

Anopheles jamesii Theobald 1901[1]

Anopheles pseudojamesi Strickland & Chowdhury 1927

Anopheles splendidus Koidzumi 1920

Group Maculatus (Rattanarithikul & Green 1987)

Anopheles dispar Rattanarithikul & Harbach 1991

Anopheles greeni Rattanarithikul & Harbach 1991

Anopheles pseudowillmori Theobald 1910

Anopheles rampae

Anopheles willmori James 1903

Subgroup Maculatus (Rattanarithikul et al 2004)

Anopheles dravidicus Christophers 1924

Anopheles maculatus*[1]

Subgroup Sawadwongporni (Rattanarithikul et al 2004)

Anopheles notanandai Rattanarithikul & Green 1987

Anopheles sawadwongporni Rattanarithikul & Green 1987

Series Neomyzomyia (Christophers 1924)

Anopheles amictus Edwards 1921

Anopheles annulatus Haga 1930

Anopheles aurirostris Watson 1910

Anopheles dualaensis Brunhes le Goff & Geoffroy 1999

Anopheles hilli Woodhill & Lee 1944

Anopheles incognitus Brug 1931

Anopheles kochi Dönitz 1901

Anopheles kokhani Vythilingam, Jeffery & Harbach 2007

Anopheles kolambuganensis Baisas 1932

Anopheles longirostris Brug 1928

Anopheles mascarensis de Meillon 1947

Anopheles meraukensis Venhuis 1932

Anopheles novaguinensis Venhuis 1933

Anopheles saungi Colless 1955

Anopheles stookesi Colless 1955

Anopheles watsonii Leicester 1908

Complex Annulipes

Anopheles annulipes Walker 1856

Complex Lungae

Anopheles lungae Belkin & Schlosser 1944

Anopheles nataliae Belkin 1945

Anopheles solomonis Belkin, Knight & Rozeboom 1945

Complex Punctulatus

Anopheles clowi Rozeboom & Knight 1946

Anopheles farauti* Laveran 1902

Anopheles hinesorum Schmidt 2001

Anopheles irenicus Schmidt 2003

Anopheles koliensis Owen 1945

Anopheles punctulatus Dönitz 1901

Anopheles torresiensis Schmidt 2001

Group Ardensis[15]

Anopheles ardensis Theobald 1905

Anopheles buxtoni Service 1958

Anopheles cinctus Newstead & Carter 1910

Anopheles deemingi Service 1970

Anopheles eouzani Brunhes le Goff & Bousses 2003

Anopheles kingi Christophers 1923

Anopheles machardyi Edwards 1930

Anopheles maliensis Bailly-Choumara & Adam 1959

Anopheles millecampsi Lips 1960

Anopheles multicinctus Edwards 1930

Anopheles natalensis Hill & Haydon 1907

Anopheles vernus Gillies 1968[15]

Anopheles vinckei de Meillon 1942

Complex Nili[15]

Anopheles carnevalei Brunhes, le Geoff & Geoffrey 1999

Anopheles nili* Theobald 1904

Anopheles ovengensis Awono-Ambene Simard Antonio-Nkonkjio & Fontenille 2004

Anopheles somalicus Rivola & Holstein 1957

Group Kochi (Rattanarithikul et al 2004)

Anopheles kochi Donitz 1901

Group Leucosphyrus

Anopheles baisasi Colless 1957

Anopheles cristatus King & Baisas

Subgroup Elegans

Anopheles elegans James 1903

Subgroup Hackeri

Anopheles hackeri Edwards 1921

Anopheles mirans Sallum & Peyton 2005

Anopheles pujutensis Colless 1948

Anopheles recens Sallum & Peyton 2005

Anopheles sulawesi* Waktoedi 1954

Subgroup Leucosphyrus

Anopheles baimaii* Sallum & Peyton 2005

Anopheles cracens Sallum & Peyton 2005

Anopheles scanloni Sallum & Peyton 2005

Complex Dirus

Anopheles dirus* Peyton & Harrison 1979

Anopheles nemophilous Peyton & Ramalingam 1988

Anopheles takasagoensis Morishita 1946

Complex Leucosphyrus (Peyton 1990)

Anopheles balabacensis* Baisas 1936

Anopheles introlatus Colless 1957

Anopheles latens* Sallum & Peyton 2005

Anopheles leucosphyrus* Dönitz 1901

Subgroup Riparis (Peyton 1990)

Anopheles cristatus King & Baisas

Anopheles macarthuri Colless 1956

Anopheles riparis King & Baisas 1936

Group Tessellatus (Rattanarithikul et al 2004)

Anopheles tessellatus Theobald

subspecies A. t. kalawara Stoker & Waktoedi

subspecies A. t. orientalis Swellengrebel & Swellengrebel de Graaf

subspecies A. t. tessellatus Theobald

Series Paramyzomyia (Christophers & Barraud 1931)

Group Cinereus[15]

Anopheles azevedoi Ribeiro 1969

Anopheles cinereus Theobald 1901[1]

subspecies cinereus Theobald 1901[1]

subspecies hispaniola Theobald 1903

Complex Turkhudi (Liston)

Anopheles turkhudi Liston 1901

subspecies telamali Saliternik & Theodor 1942

subspecies turkhudi Liston 1901

Group Listeri[15]

Anopheles listeri de Mellion 1931

Anopheles multicolor* Cambouliu 1902

Anopheles seretsei Abdulla-Chan Coetzee & Hunt 1998

Series Pyretophorus (Blanchard 1902)

Anopheles christyi Newstead & Carter 1911

Anopheles daudi Coluzzi 1958

Anopheles indefinitus Ludlow 1904

Anopheles limosus King 1932

Anopheles litoralis King 1932

Anopheles ludlowae Theobald 1903

subspecies ludlowae Theobald 1903

subspecies torakala Stoker & Waktoedi 1949

Anopheles parangensis Ludlow 1914

Anopheles vagus* Dönitz 1902

Complex Gambiae (White 1985)

Anopheles amharicus Hunt, Wilkerson & Coetzee 2013

Anopheles arabiensis* Patton 1905

Anopheles bwambae White 1985

Anopheles coluzzii* Coetzee & Wilkerson 2013 [24]

Anopheles comorensis Brunhes le Goff & Geoffroy 1997

Anopheles fontenillei Barrón et al 2019

Anopheles gambiae* Giles 1902

Anopheles melas* Theobald 1903

Anopheles merus Dontiz 1902

Anopheles quadriannulatus Theobald 1911

Complex Subpictus (Sugana et al. 1994)

Anopheles subpictus* Grassi 1899

Complex Sundaicus (Sukowati 1999)

Anopheles epiroticus Linton & Harbach 2005

Anopheles sundaicus* Rodenwaldt 1925

Subgenus Kerteszia

Anopheles auyantepuiensis Harbach & Navarro 1996

Anopheles bambusicolus Komp 1937

Anopheles bellator* Dyar & Knab 1906

Anopheles boliviensis Theobald 1905

Anopheles cruzii* Dyar & Knab 1908

Anopheles gonzalezrinconesi Cova Garcia, Pulido & de Ugueto, 1977

Anopheles homunculus* Komp 1937

Anopheles laneanus Corrêa & Cerqueira 1944

Anopheles lepidotus Zavortink 1973

Anopheles neivai Howard, Dyar & Knab 1913

Anopheles pholidotus Zavortink 1973

Anopheles rollai Cova Garcia, Pulido & de Ugueto 1977

Note: Anopheles cruzii is known to be a species complex,[25] but the number species in this complex has yet to be finalised.

Subgenus Lophopodomyia

Anopheles gilesi Peryassu 1928

Anopheles gomezdelatorrei Levi-Castillo 1955

Anopheles oiketorakras Osorno-Mesa 1947

Anopheles pseudotibiamaculata Galvao & Barretto 1941

Anopheles squamifemur Antunes 1937

Anopheles vargasi Gabaldon Cova Garcia & Lopez 1941

Subgenus Nyssorhynchus

Anopheles dominicanus Zavortinkb and Poinarab 2000

Section Albimanus

Anopheles noroestensis Galvao and Lane 1937

Series Albimanus (Faran 1980)

Anopheles albimanus* Weidemann 1820

Series Oswaldoi (Faran 1980)

Group Oswaldoi (Faran 1980)

Subgroup Oswaldoi (Faran 1980)

Anopheles anomalphyllus Komp

Anopheles aquasalis* Curry 1932

Anopheles dunhamii Causey 1945

Anopheles evansae Brethes 1926

Anopheles galvaoi Causey, Deane and Deane 1943

Anopheles ininii Sevenet & Abonnenc 1938

Anopheles konderi Galvfio & Damasceno 1942

Anopheles oswaldoi Peryassú 1922

Anopheles rangeli Galabadon, Cova-Garcia & Lopez 1941

Anopheles sanctielii Sevenet & Abonnenc 1938

Anopheles trinkae Faran 1980

Complex Nuneztovari (Conn et al. 1993)

Anopheles geoeldii Rozeboom and Gabaldón 1941

Anopheles nuneztovari* Galbadón 1940

Subgroup Strodei (Faran 1980)

Anopheles alberto

Anopheles arthuri*

Anopheles benarrochi Galabadon, Cova-Garcia & Lopez

Anopheles rondoni Neiva & Pinto 1922

Anopheles strodei Root 1926

Group Triannulatus

Anopheles halophylus do Nascimento & de Oliveira 2002

Anopheles triannulatus Neiva & Pinto 1922

Section Argyritarsis (Levi Castillo 1949)

Series Albitarsis[26]

Anopheles rooti Brethes 1926

Group Albitarsis[26]

Anopheles albitarsis[26]

Anopheles deaneorum Rosa-Freitas 1989

Anopheles janconnae Wilkerson & Sallum 2009

Anopheles oryzalimnetes Wilkerson & Motoki 2009

Anopheles marajoara* Galvao & Damesceno 1942

Group Braziliensis[26]

Anopheles braziliensis Chagas 1907

Series Argyritarsis[26]

Group Argyritarsis[26]

Anopheles argyitarsis Robineau-Desvoidy 1827

Anopheles sawyeri Causey, Deane, Deane & Sampaio 1943

Group Darlingi[26]

Anopheles darlingi* Root

Group Lanie[26]

Anopheles lanei Galvao & Amaral 1938

Group Pictipennis[26]

Anopheles pictipennis Phillippi 1865

Section Myzorhynchella (Peyton et al. 1992)

Anopheles antunesi Galvao & Amaral 1940

Anopheles lutzii Cruz 1901

Anopheles nigritarsis Chagas 1907

Anopheles parvus Chagas 1907

Subgenus Stethomyia

Anopheles acanthotorynus Komp 1937

Anopheles canorii Floch & Abonnenc 1945

Anopheles kompi Edwards 1930

Anopheles nimbus[11]

Anopheles thomasi Shannon 1933

Notes

• Anopheles anthropophagus Xu and Feng is considered to be a junior synonym of Anopheles lesteri de Meillon 1931.
• Anopheles bonneorum Fonseca & Ramos is considered to be a synonym of Anopheles costai.
• Anopheles lewisi Ludlow 1920 is a synonym of Anopheles thomasi Shannon 1933.
• Anopheles lineata Lutz is a synonym of Anopheles nimbus Theobald.
• Anopheles mesopotamiae is considered to be a synonym of Anopheles hyrcanus.
• Anopheles rossii Giles 1899 was originally described as Anopheles subpictus Grassi 1899.
• Bironella derooki is a synonym of Anopheles soesiloi.

The following are currently regarded as nomina nuda:

• Anopheles (Anopheles) solomonensis Cumpston 1924
• Anopheles (Cellia) melanotarsis Woodhill & Lee

A subgroup of Anopheles gambiae sensu stricto has been reported and given the name Goundry. This subgroup has not yet been elevated to species status.[27]

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23. Nanda, Nutan; Singh, Om P; Dua, Virendra K; Pandey, Akhilesh C; Nagpal, Bhupender N; Adak, Tridibes; Dash, Aditya P; Subbarao, Sarala K (2013). "Population cytogenetic and molecular evidence for existence of a new species in Anopheles fluviatilis complex (Diptera: Culicidae)". Infection, Genetics and Evolution. 13: 218–23. doi:10.1016/j.meegid.2012.09.018. PMID 23146830.
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External links

• Mosquito Taxonomic Inventory page on Anopheles
• The Walter Reed Biosystematics Unit