Taxonomy of Anopheles

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Anopheles
Anopheles stephensi.jpeg
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

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.

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), Edwards (three species) and 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 .[9]

Subgenera[]

Key to the morphology of female Anopheles

The genus has been subdivided into seven subgenera based primarily on the number and positions of specialized setae on the 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: (Southeast Asia only) by Harbach et al. in 2005 and (Neotropical) by Antunes in 1937.

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

  • Subgenus Anopheles - Meigen 1918[9]
  • Subgenus Baimaia - Abraham 1947[10]
  • Subgenus Cellia - Giles 1899
  • Subgenus Kerteszia - Theobald 1905
  • Subgenus Lophopodomyia - Antunes 1937
  • Subgenus Nyssorhynchus - Robineau-Desvoidy 1827
  • Subgenus Stethomyia - 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 - 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[]

* Xu & Feng 1975
[20]
Soesilo & Van Slooten 1931
Theobald 1905
Taylor 1934
Tenorio 1975
Swellengrebel & Swellengrebel de Graaf 1919
Tenorio 1977
Brug 1928
Section
Series Anopheles
Theobald 1903
Edwards 1938
[21]
subspecies marteri
subspecies sogdianus Keshishian
Complex Claviger (Coluzzi et al. 1965)
Anopheles claviger* Meigen 1804
Del Vecchio 1939
Group Aitkenii (Reid & Knight, 1961)
Harrison & Scanlon 1975
Baisas 1946
Anopheles aitkenii James 1903
Puri 1930
McArthur 1949
Theobald 1903
Anopheles insulaeflorum Swellengrebel & Swellengrebel de Graaf 1919
Rodenwaldt 1926
Kulasekera Harrison & Amerasinghe 1989
Harrison & Scanlon 1974
Barraud 1932
Reid 1965
Scanlon & Peyton 1967
Group Alongensis (Phan et al. 1991)
Evenhuis 1940
Phan, Manh, Hinh & Vien 1991
Group Atratipes (Lee et al. 1987)
Skuse 1889
Dobrowtorsky 1966
Group Culiciformis (Reid & Knight 1961)
Cogill 1903
Puri 1929
Ho 1938
Group Lindesayi (Reid & Knight 1961)
Ma 1981
Christophers 1924
Alcock 1912
Complex Gigas (Harrison et al. 1991)
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)
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
Hoffmann 1935
Ludlow 1920
Anopheles walkeri[1]
Complex Quadrimaculatus (Linton 2004)
Reinert 1997
Reinert 1997
Reinert 1997
Reinert 1997
Anopheles quadrimaculatus* Say 1824
Subgroup Freeborni (Linton 2004)
Anopheles earlei Vargas 1943
Anopheles freeborni* Aitken 1939
Barr & Guptavanij 1989
Subgroup Maculipennis (Linton 2004)
Gordeyev, Zvantsov, Goryacheva, Shaikevich & Yezhov
Anopheles atroparvus* Van Thiel 1927
Stegnii & Kabanova 1976
Anopheles daciae Linton, Nicolescu & Harbach 2004
* Falleroni 1926
[9]
Shingarev 1926
* Hackett 1934
* Falleroni 1926
Dyar & Knab 1906
Linton, Sedaghat & Harbach 2003
* Favre 1903
Roubaud 1935
Hackett & Lewis 1935
Group Plumbeus (Reid & Knight 1961)
Zavortink 1970
Anopheles barberi Coquillett 1903
Anopheles barianensis James 1911
Vargas 1943
Zavortink 1969
Sakakibara 1959
* Stegnii & Kabanova 1828
Zavortink 1970
De Leon 1938
Group Pseudopunctipennis (Reid & Knight 1961)
Komp 1936
Anopheles franciscanus McCracken 1904
Giaquinto-Mira 1931
Neiva 1906
Coquillett 1902
subspecies eiseni Coquillett
subspecies geometricus Corrêa
* Martini 1932
subspecies guatemalensis de Leon
subspecies parapunctipennis Martini
[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)
Ludlow 1907
Anopheles punctipennis Say 1823
Complex Crucians (Wilkerson et al. 2004)
King 1939
Anopheles crucians Wiedemann 1828
King 1939
Group Stigmaticus (Reid & Knight 1961)
Marks 1956
Theobald 1907
Dobrowtorsky 1957
Lee 1944
Dobrowtorsky 1957
Skuse 1889
Series (Edwards 1932)
Lynch 1878
[1]
Series (Edwards 1932)
Causey 1937
Group Asiaticus (Reid 1968)
Prashad 1918
Reid 1963
Subgroup Asiaticus (Rattanarithikul et al. 2004)
Leicester 1903
Subgroup Interruptus (Rattanarithikul et al. 2004)
Anopheles interruptus Puri 1929
Section (Reid & Knight 1961)
Series (Root 1922)
Correa & Ramalho 1968
Dyar & Knab 1906
Galvao 1955
Wilkerson 1991
Da Fonseca & Da Silva Ramos 1939
Da Costa Lima 1937
Root 1927
Wilkerson 1999
Vargas 1941
Anduze & Capdevielle 1949
* Peryassu 1908
Theobald 1903
Dyar & Knab 1907
Lutz & Neiva 1911
Lutz 1903
Da Costa Lima 1929
Curry 1931
Dyar & Knab 1908
Peryassu 1908
Dyar & Knab 1906
Lutz 1903
Galvao 1952
Davis 1931
Vargas 1979
Dyar & Knab 1906
Series Christya (Christophers 1924)
Theobald 1903
Brunhes, le Goff & Geoffroy 1997
Series (Edwards 1932)
Grunberg 1905
Giles 1902
Anopheles barbirostris* Van der Wulp 1884
Reid 1962
Group Albotaeniatus (Reid & Knight 1961)
Theobald 1903
Mendoza 1947
Mendoza 1947
Stanton & Hacker 1917
Bohart & Ingram 1946
subspecies ohamai Ohama
subspecies saperoi Bohart & Ingram
Group Bancroftii (Reid & Knight 1961)
Ludlow 1935
Giles 1902
subspecies bancroftii Giles
subspecies barbiventris Brug
Group Barbirostris (Reid & Knight 1961)
Meng 1957
Yamada & Watanabe 1918
Subgroup Barbirostris (Reid 1968)
Anopheles barbirostris van der Wulp 1884
Reid 1962
Reid 1962
Reid 1962
Reid 1962
Reid 1962
Subgroup Vanus (Reid 1968)
Chowdhury 1929
Anopheles barbumbrosus Strickland & Chowdhury 1927
Mendoza 1940
Anopheles reidi Harrison 1973
Walker 1859
Group Coustani (Reid & Knight 1961)
De Meillon 1943
Laveran 1900
Coetzee 1994
Van Someren 1947
Coetzee 1984
Theobald 1900
Edwards 1928
Donitz 1902
Grunberg 1902
Group Hyrcanus (Reid 1953)
Xu and Feng 1975
Swellengrebel 1914
Anopheles belenrae Rueda 2005
Ma 1981
Martini 1929
Ma 1981
Kanda & Oguma 1978
Xu JinJiang & Luo XinFu 1998
Ma 1981
* Pallas 1771
Lei 1996
Xu and Feng 1975
Anopheles kleini Rueda 2005
Dong & Wang 1985
Yao & Wu 1944
Kang Tan Cao Cheng Yang & Huang 1984
Nguyen, Tran & Harbach
Graham 1899
Yamada 1937
Anopheles sinensis* Wiedemann 1828
Yamada 1924
Ma 1981
Dong, Zhou, Dong & Mao 2007
Miyazaki 1951
Subgroup Lesteri (Harrison 1972)
Reid 1953
Xu & Feng 1975
de Meillon 1931
Sandosham 1959
Anopheles peditaeniatus Leicester 1908
Manh Hinh & Vien 1993
Subgroup Nigerrimus (Harrison 1972)
Anopheles nigerrimus* Giles 1900
Harrison, Scanlon & Reid 1973
Baisas 1935
Laveran 1902
Group Umbrosus (Reid 1950)
Roper 1914
Reid 1950
Strickland 1916
Rozeboom 1951
Strickland & Chowdhury 1927
Subgroup Baezai (Rattanarithikul et al. 2004)
Gater 1934
Subgroup Letifer (Reid 1968)
Reid 1963
* Sandosham 1944
Reid 1950
Reid 1963
Subgroup Separatus (Rattanarithikul et al. 2004)
Leicester 1908
Subgroup Umbrosus (Rattanarithikul et al. 2004)
Theobald 1903

Subgenus []

Abraham 1947[10]

Subgenus Cellia[]

Mattingly and Adam

Series Cellia (Christophers 1924)

Gough 1910
Hamon & Rickenbach 1955
Brunhes le Goff & Geoffroy 1999
Service 1977
Gillies 1968[15]
Theobald 1901[1]
Gillies 1964
Group Squamosus (Grjebine 1966)
de Meillon 1931
Theobald 1901[1]

Series

Marsh 1933
Bailly-Choumara 1960
Evans 1932
Theobald 1910
Edwards 1916
Patton 1905
Corradetti 1939
Gillies & Coetzee 1987
Edwards 1911
Gillies 1968[15]
Young & Majid 1928
* Evans 1925
subspecies bervoetsi D'Haenans 1961
subspecies moucheti Evans 1925
subspecies nigeriensis
Evans 1934
de Meillon & Leeson 1940
Edwards 1930
Group Demeilloni (Gillies & De Meillon 1962)
Evans & de Meillon 1933
Evans 1933
Evans 1925
Edwards 1930
Evans 1931
Gil Collado 1936
* Theobald 1907
subspecies macmahoni Evans 1936
subspecies sergentii Theobald 1907
Group Funestus (Garros et al 2004)
James 1902
Subgroup Aconitus (Chen et al. 2003)
Anopheles aconitus Dönitz 1902
Manalang 1930
Banks 1906
Buttiker & Beales 1959
Anopheles varuna Iyengar 1924
Subgroup Culicifacies (Garros et al. 2004)
Anopheles culicifacies* Giles 1901
Subgroup Funestus (Garros et al. 2004)
Sobti 1968
Evans & Leeson 1935
Anopheles funestus* Giles 1900
* Spillings et al 2009[22]
Type C Koekemoer et al. 2009
Gillies 1962
Gillies & Coetzee 1987
Subgroup Minimus (Chen et al. 2003)
* Ludlow 1914
Evans 1931
Type A Koekemoer et al 2009
Complex Fluviatilis (Salara et al. 1993)
* (species S, T, U, V) James 1902[23]
Complex Minimus (Green et al. 1990)
Harbach & Manguin 2007
Theobald 1901*[1]
Subgroup Rivulorum (Garros et al 2004)
Service 1960
Leeson 1930
* Leeson 1935
Group Marshallii[15]
Theobald 1905
Vincke & Leleup 1949
Edwards 1929
Evans 1935
Edwards 1929
Evans 1927
Evans 1936
Edwards 1938
de Meillon & Pereira 1940
Peters 1955
Edwards 1929
Complex Marshalli (Gillies & Coetzee 1987)
Lambert & Coetzee 1982
Coetzee, Segerman & Hunt 1987
Lambert & Coetzee 1982
Theobald 1903
Group Wellcomei[15]
Newstead & Carter 1911
Gillies 1958
Edwards 1912
Theobald 1904
subspecies ugandae Evans 1934
subspecies ungujae White 1975
subspecies wellcomei Theobald 1904

Series (Christophers 1924)

Gad, Harbach & Harrison 2006
Corradetti 1939
Brunhes, le Goff & Geoffroy 1999
Anopheles jamesiiTheobald 1901[1]
Anopheles karwari* James 1903
Giles 1902
Christophers 1924
Shidrawi & Gillies 1988
Anopheles pattoni Christophers 1926
Theobald 1903
Theobald 1902*[11]
Gough 1910
subspecies broussesi Edwards 1929
subspecies rufipes Gough 1910
Maffi & Coluzzi 1958
Koidzumi 1920
Giles 1901
Complex Stephensi
Anopheles stephensi* Liston 1901
Complex Superpictus
* Grassi 1899
Group Annularis (Reid 1968)
Anopheles pallidus Theobald 1901[1]
* Ludlow 1902
Stanton 1915
Complex Annularis (Reid 1968)
Anopheles annularis* van der Wulp 1884
Complex Nivipes (Green et al. 1985)
Theobald 1903
Group Jamesii (Rattanarithikul et al. 2004)
Anopheles jamesii Theobald 1901[1]
Strickland & Chowdhury 1927
Koidzumi 1920
Group Maculatus (Rattanarithikul & Green 1987)
Rattanarithikul & Harbach 1991
Rattanarithikul & Harbach 1991
Theobald 1910
James 1903
Subgroup Maculatus (Rattanarithikul et al 2004)
Christophers 1924
*[1]
Subgroup Sawadwongporni (Rattanarithikul et al 2004)
Rattanarithikul & Green 1987
Rattanarithikul & Green 1987

Series (Christophers 1924)

Edwards 1921
Haga 1930
Watson 1910
Brunhes le Goff & Geoffroy 1999
Woodhill & Lee 1944
Brug 1931
Dönitz 1901
Vythilingam, Jeffery & Harbach 2007
Baisas 1932
Brug 1928
de Meillon 1947
Venhuis 1932
Venhuis 1933
Colless 1955
Colless 1955
Leicester 1908
Complex Annulipes
Walker 1856
Complex Lungae
Belkin & Schlosser 1944
Belkin 1945
Belkin, Knight & Rozeboom 1945
Complex Punctulatus
Rozeboom & Knight 1946
* Laveran 1902
Schmidt 2001
Schmidt 2003
Owen 1945
Dönitz 1901
Schmidt 2001
Group Ardensis[15]
Theobald 1905
Service 1958
Newstead & Carter 1910
Service 1970
Brunhes le Goff & Bousses 2003
Christophers 1923
Edwards 1930
Bailly-Choumara & Adam 1959
Lips 1960
Edwards 1930
Hill & Haydon 1907
Gillies 1968[15]
de Meillon 1942
Complex Nili[15]
Brunhes, le Geoff & Geoffrey 1999
Anopheles nili* Theobald 1904
Awono-Ambene Simard Antonio-Nkonkjio & Fontenille 2004
Rivola & Holstein 1957
Group Kochi (Rattanarithikul et al 2004)
Donitz 1901
Group Leucosphyrus
Colless 1957
King & Baisas
Subgroup Elegans
Anopheles elegans James 1903
Subgroup Hackeri
Edwards 1921
Sallum & Peyton 2005
Colless 1948
Sallum & Peyton 2005
* Waktoedi 1954
Subgroup Leucosphyrus
* Sallum & Peyton 2005
Sallum & Peyton 2005
Sallum & Peyton 2005
Complex Dirus
Anopheles dirus* Peyton & Harrison 1979
Peyton & Ramalingam 1988
Morishita 1946
Complex Leucosphyrus (Peyton 1990)
Anopheles balabacensis* Baisas 1936
Anopheles introlatus Colless 1957
Anopheles latens* Sallum & Peyton 2005
* Dönitz 1901
Subgroup Riparis (Peyton 1990)
King & Baisas
Colless 1956
King & Baisas 1936
Group Tessellatus (Rattanarithikul et al 2004)
Anopheles tessellatus Theobald
subspecies Stoker & Waktoedi
subspecies Swellengrebel & Swellengrebel de Graaf
subspecies Theobald

Series (Christophers & Barraud 1931)

Group Cinereus[15]
Ribeiro 1969
Theobald 1901[1]
subspecies cinereus Theobald 1901[1]
subspecies hispaniola Theobald 1903
Complex Turkhudi (Liston)
Liston 1901
subspecies telamali Saliternik & Theodor 1942
subspecies turkhudi Liston 1901
Group Listeri[15]
de Mellion 1931
* Cambouliu 1902
Abdulla-Chan Coetzee & Hunt 1998

Series (Blanchard 1902)

Newstead & Carter 1911
Coluzzi 1958
Ludlow 1904
King 1932
King 1932
Theobald 1903
subspecies ludlowae Theobald 1903
subspecies torakala Stoker & Waktoedi 1949
Ludlow 1914
Anopheles vagus* Dönitz 1902
Complex Gambiae (White 1985)
Hunt, Wilkerson & Coetzee 2013
* Patton 1905
White 1985
Anopheles coluzzii* Coetzee & Wilkerson 2013 [24]
Brunhes le Goff & Geoffroy 1997
Barrón et al 2019
Anopheles gambiae* Giles 1902
* Theobald 1903
Dontiz 1902
Theobald 1911
Complex Subpictus (Sugana et al. 1994)
Anopheles subpictus* Grassi 1899
Complex Sundaicus (Sukowati 1999)
Linton & Harbach 2005
* Rodenwaldt 1925

Subgenus Kerteszia[]

Harbach & Navarro 1996
Komp 1937
* Dyar & Knab 1906
Theobald 1905
* Dyar & Knab 1908
Cova Garcia, Pulido & de Ugueto, 1977
* Komp 1937
Corrêa & Cerqueira 1944
Zavortink 1973
Howard, Dyar & Knab 1913
Zavortink 1973
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 []

Peryassu 1928
Levi-Castillo 1955
Osorno-Mesa 1947
Galvao & Barretto 1941
Antunes 1937
Gabaldon Cova Garcia & Lopez 1941

Subgenus []

Zavortinkb and Poinarab 2000
Section
Galvao and Lane 1937
Series Albimanus (Faran 1980)
Anopheles albimanus* Weidemann 1820
Series (Faran 1980)
Group Oswaldoi (Faran 1980)
Subgroup Oswaldoi (Faran 1980)
Komp
* Curry 1932
Causey 1945
Brethes 1926
Causey, Deane and Deane 1943
Sevenet & Abonnenc 1938
Galvfio & Damasceno 1942
Peryassú 1922
Galabadon, Cova-Garcia & Lopez 1941
Sevenet & Abonnenc 1938
Faran 1980
Complex Nuneztovari (Conn et al. 1993)
Rozeboom and Gabaldón 1941
Anopheles nuneztovari* Galbadón 1940
Subgroup Strodei (Faran 1980)
*
Galabadon, Cova-Garcia & Lopez
Neiva & Pinto 1922
Root 1926
Group Triannulatus
do Nascimento & de Oliveira 2002
Neiva & Pinto 1922
Section (Levi Castillo 1949)
Series [26]
Brethes 1926
Group Albitarsis[26]
[26]
Rosa-Freitas 1989
Wilkerson & Sallum 2009
Wilkerson & Motoki 2009
* Galvao & Damesceno 1942
Group Braziliensis[26]
Chagas 1907
Series Argyritarsis[26]
Group Argyritarsis[26]
Robineau-Desvoidy 1827
Causey, Deane, Deane & Sampaio 1943
Group Darlingi[26]
Anopheles darlingi* Root
Group Lanie[26]
Galvao & Amaral 1938
Group Pictipennis[26]
Phillippi 1865
Section (Peyton et al. 1992)
Galvao & Amaral 1940
Cruz 1901
Chagas 1907
Chagas 1907

Subgenus Stethomyia[]

Komp 1937
Floch & Abonnenc 1945
Edwards 1930
[11]
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]

References[]

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