Taxonomy of Anopheles

Genus of flies

Taxonomy of Anopheles

Anopheles stephensi
Scientific classification
Domain:  | Eukaryota
Kingdom:  | Animalia
Phylum:  | Arthropoda
Class:  | Insecta
Order:  | Diptera
Family:  | Culicidae
Subfamily:  | Anophelinae
Genus:  | Anopheles
Subgenera

* Anopheles
* Baimaia
* Cellia
* Kerteszia
* Lophopodomyia
* Nyssorhynchus
* Stethomyia

Anopheles is a genus of mosquitoes (Culicidae) comprising over 500 recognized species.[1] These mosquitoes are the primary vectors responsible for transmitting malaria to humans.[2]

The genus is taxonomically divided into several subgenera, including Anopheles (206 species), Baimaia (one species), Cellia (239 species), Kerteszia (12 species), Lophopodomyia (six species), Nyssorhynchus (34 species), and Stethomyia (five species).[3] Notably, only about 30 to 40 of these species are commonly involved in malaria transmission.[4]

## Classification

[edit]

The classification of this genus began in 1901 with Frederick Vincent Theobald.[5] Despite the passage of time, the taxonomy remains incompletely settled.[6][7][8] 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.[9][10][11][12]

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.[13]

### Subgenera

[edit]

Morphology of female Anopheles

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[13]
* Subgenus Baimaia \- Anopheles kyondawensis Abraham 1947[14]
* 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[15]

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.[16]

### Divisions below subgenus

[edit]

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)[17] 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

[edit]

Anopheles nuneztovari is a species complex with at least one occurring in Colombia and Venezuela and another occurring in the Amazon Basin.[18]

## Species listing

[edit]

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

### Subgenus Anopheles

[edit]

Anopheles anthropophagus Xu & Feng 1975
Anopheles confusa[19]
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 (Edwards, 1932a)

Anopheles algeriensis Theobald 1903
Anopheles concolor Edwards 1938
Anopheles marteri[20]

subspecies marteri
subspecies sogdianus Keshishian
Claviger Complex (Coluzzi et al., 1965)

Anopheles claviger* Meigen 1804
Anopheles petragnani Del Vecchio 1939
Aitkenii Group (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
Alongensis Group (Phan et al., 1991)

Anopheles alongensis Venhuis 1940
Anopheles cucphuongensis Vu, Nguyen, Tran & Nguyen 1991
Atratipes Group (Lee et al., 1987b)

Anopheles atratipes Skuse 1889
Anopheles tasmaniensis Dobrotworsky 1966
Culiciformis Group (Reid & Knight, 1961)

Anopheles culiciformis Cogill 1903
Anopheles sintoni Puri 1929
Anopheles sintonoides Ho 1938
Lindesayi Group (Reid & Knight, 1961)

Gigas Subgroup (Harrison et al., 1991, as Gigas Complex)

Baileyi Complex (Somboon et al., 2020)

Anopheles baileyi Edwards (species A)
Anopheles inthanonensis Somboon & Harbach, 2024 (species D)
Anopheles bhutanensis Somboon, Namgay & Harbach (species C)
Anopheles monticola Somboon, Namgay & Harbach (species B)
Anopheles simlensis James
Gigas Complex (redefined by Somboon et al., 2020)

Anopheles gigas Giles

subspecies crockeri Colless
subspecies danaubento Mochtar & Walandouw
subspecies formosus Ludlow
subspecies gigas Giles
subspecies oedjalikalah Nainggolan
subspecies pantjarbatu Waktoedi Koesoemawinangoen
subspecies refutans Alcock
Anopheles prachongae Rattanarithikul & Harrison
Anopheles sumatrana Swellengrebel & Rodenwaldt
Lindesayi Subgroup (Harrison et al., 1991, as Lindesayi Complex)

Anopheles menglangensis Ma 1981
Anopheles nilgiricus Christophers 1924
Anopheles wellingtonianus Alcock 1912
Lindesayi Complex (Namgay et al., 2020)

Anopheles benguetensis King
Anopheles cameronensis Edwards
Anopheles druki Somboon, Namgay & Harbach (lindesayi species C)
Anopheles himalayensis Somboon, Namgay & Harbach (lindesayi species D)
Anopheles japonicus Yamada
Anopheles lindesayi Giles (species A)
Anopheles lindesayi (species B)
Anopheles pleccau Koidzumi
Anopheles thimphuensis Somboon, Namgay & Harbach (lindesayi species E)
Maculipennis Group (Reid & Knight, 1961)

Anopheles atropos Dyar & Knab 1906
A. atropos

Anopheles aztecus Hoffmann 1935
Anopheles lewisi Ludlow 1920
Anopheles walkeri Theobald
Maculipennis Subgroup (Linton, 2004)

Anopheles artemievi Gordeyev, Zvantsov, Goryacheva, Shaikevich & Yezhov
Anopheles atroparvus* van Thiel 1927
Anopheles daciae Linton, Nicolescu & Harbach 2004
Anopheles labranchiae* Falleroni 1926
Anopheles maculipennis Meigen
Anopheles martinius Shingarev 1926
Anopheles melanoon* Hackett 1934
Anopheles messeae* Falleroni 1926
Anopheles persiensis Linton, Sedaghat & Harbach 2003
Anopheles sacharovi* Favre 1903
Quadrimaculatus Subgroup (Linton, 2004)

Anopheles beklemishevi Stegnii & Kabanova 1976
A. beklemishevi

Anopheles diluvialis Reinert 1997
Anopheles inundatus Reinert 1997
Anopheles maverlius Reinert 1997
Anopheles quadrimaculatus* Say 1824
Anopheles smaragdinus Reinert 1997
Freeborni Subgroup (Linton, 2004)

Anopheles earlei Vargas 1943
Anopheles freeborni* Aitken 1939
Anopheles hermsi Barr & Guptavanij 1989
Anopheles occidentalis Dyar & Knab 1906
A. occidentalis

Plumbeus Group (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* Stephens 1828
Anopheles powderi Zavortink 1970
Anopheles xelajuensis de Leon 1938
Pseudopunctipennis Group (Reid & Knight, 1961)

Anopheles chiriquiensis Komp 1936
Anopheles eiseni Coquillett 1902

subspecies eiseni Coquillett
subspecies geometricus Corrêa
Anopheles franciscanus McCracken 1904
Anopheles hectoris Giaquinto-Mira 1931
Anopheles parapunctipennis* Martini 1932

subspecies guatemalensis de Leon
subspecies parapunctipennis Martini
Anopheles pseudopunctipennis Theobald 1901
A. pseudopunctipennis

subspecies levicastilloi Levi-Castillo
subspecies neghmei Mann
subspecies noei Mann
subspecies patersoni Alvarado & Heredia
subspecies pseudopunctipennis Theobald
subspecies rivadeneirai Levi-Castillo
Anopheles tibiamaculatus Neiva 1906
Punctipennis Group (Reid & Knight, 1961)

Anopheles perplexens Ludlow 1907
A. perplexens

Anopheles punctipennis Say 1823
Crucians Complex (Wilkerson et al., 2004)

Anopheles bradleyi King 1939
Anopheles crucians Wiedemann 1828 (species A, B, C, D and E)
Anopheles georgianus King 1939
Stigmaticus Group (Reid & Knight, 1961)

Anopheles colledgei Marks 1956
Anopheles corethroides Theobald 1907
Anopheles papuensis Dobrotworsky 1957
Anopheles powelli Lee 1944
Anopheles pseudostigmaticus Dobrotworsky 1957
Anopheles stigmaticus Skuse 1889

Series Cycloleppteron (Edwards, 1932a)

Anopheles annulipalpis Lynch Arribálzaga 1878
Anopheles grabhamii Theobald 1901

Series Lophoscelomyia (Edwards, 1932a)

Anopheles bulkleyi Causey 1937
Asiaticus Group (Reid, 1968)

Anopheles annandalei Prashad 1918
Anopheles noniae Reid 1963
Asiaticus Subgroup (Rattanarithikul et al., 2006b)

Anopheles asiaticus Leicester 1903
Interruptus Subgroup (Rattanarithikul et al., 2006b)

Anopheles interruptus Puri 1929

Section Laticorn (Reid & Knight, 1961)

Series Arribalzagia (Root, 1922a)

Anopheles anchietai Corrêa & Ramalho 1968
Anopheles apicimacula Dyar & Knab 1906
Anopheles bustamentei Galvão 1955
Anopheles calderoni Wilkerson 1991
Anopheles costai Fonseca & Ramos 1939
Anopheles evandroi da Costa Lima 1937
Anopheles fluminensis Root 1927
Anopheles forattinii Wilkerson & Sallum 1999
Anopheles gabaldoni Vargas 1941
Anopheles guarao Anduze & Capdevielle 1949
Anopheles intermedius* Peryassú 1908
Anopheles maculipes Theobald 1903
Anopheles malefactor Dyar & Knab 1907
Anopheles mattogrossensis Lutz & Neiva 1911
Anopheles medialis Harbach
Anopheles mediopunctatus Lutz 1903
Anopheles minor da Costa Lima 1929
Anopheles neomaculipalpus Curry 1931
Anopheles peryassui Dyar & Knab 1908
Anopheles pseudomaculipes Chagas 1907
Anopheles punctimacula Dyar & Knab 1906
Anopheles rachoui Galvão 1952
Anopheles shannoni Davis 1931
Anopheles veruslanei Vargas 1979
Anopheles vestitipennis Dyar & Knab 1906

Series Christya (Christophers, 1924a)

Anopheles implexus Theobald 1903
Anopheles okuensis Brunhes, le Goff & Geoffroy 1997

Series Myzorhynchus (Edwards, 1932a)

Anopheles obscurus Grünberg 1905
Albotaeniatus Group (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
Bancroftii Group (Reid & Knight, 1961)

Anopheles bancroftii Giles 1902
A. bancroftii

subspecies bancroftii Giles
subspecies barbiventris Brug
Anopheles pseudobarbirostris Ludlow 1935
Barbirostris Group (Reid & Knight, 1961)

Anopheles freyi Meng 1957
Anopheles koreicus Yamada & Watanabe 1918
Barbirostris Subgroup (Reid, 1968)

Barbirostris Complex (Satoto, 2001)

Anopheles barbirostris* van der Wulp 1884
Anopheles campestris Reid 1962
Anopheles dissidens Taai & Harbach (species III of Paredes-Esquivel et al., 2009)
Anopheles saeungae Taai & Harbach (species IV of Paredes-Esquivel et al., 2009)
Anopheles vanderwulpi Townson & Harbach
Anopheles wejchoochotei Taai & Harbach
Anopheles sarpangensis Somboon, Namgay & Harbach, 2024
Anopheles donaldi Reid 1962
Anopheles franciscoi Reid 1962
Anopheles hodgkini Reid 1962
Anopheles pollicaris Reid 1962
Vanus Subgroup (Reid, 1968)

Anopheles ahomi Chowdhury 1929
Anopheles barbumbrosus Strickland & Chowdhury 1927
Anopheles manalangi Mendoza 1940
Anopheles reidi Harrison 1973
Anopheles vanus Walker 1859
Coustani Group (Reid & Knight, 1961)

Anopheles caliginosus de Meillon 1943
Anopheles coustani Laveran 1900
A. coustani

Anopheles crypticus Coetzee 1994
Anopheles fuscicolor van Someren 1947
Anopheles namibiensis Coetzee 1984
Anopheles paludis Theobald 1900
Anopheles symesi Edwards 1928
Anopheles tenebrosus Dönitz 1902
Anopheles ziemanni Grünberg 1902
Hyrcanus Group (Reid, 1953)

Anopheles anthropophagus Xu & 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 & Luo 1998
Anopheles heiheensis Ma 1981
Anopheles hyrcanus* Pallas 1771
A. hyrcanus

Anopheles hyrcanus spIR (Djadid et al., 2009)
Anopheles junlianensis Lei 1996
Anopheles kiangsuensis Xu & 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 Grassi 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
Lesteri Subgroup (Harrison, 1972)

Anopheles crawfordi Reid 1953
Anopheles lesteri Baisas & Hu 1936
Anopheles paraliae Sandosham 1959
Anopheles peditaeniatus Leicester 1908
Anopheles vietnamensis Nguyen, Tran & Nguyen 1993
Nigerrimus Subgroup (Harrison, 1972)

Anopheles nigerrimus* Giles 1900
Anopheles nitidus Harrison, Scanlon & Reid 1973
Anopheles pseudosinensis Baisas 1935
Anopheles pursati Laveran 1902
Umbrosus Group (Reid, 1950)

Anopheles brevipalpis Roper 1914
Anopheles brevirostris Reid 1950
Anopheles hunteri Strickland 1916
Anopheles samarensis Rozeboom 1951
Anopheles similissimus Strickland & Chowdhury 1927
Baezai Subgroup (Rattanarithikul et al., 2006b)

Anopheles baezai Gater 1934
Letifer Subgroup (Reid, 1968)

Anopheles collessi Reid 1963
Anopheles letifer* Sandosham 1944
Anopheles roperi Reid 1950
Anopheles whartoni Reid 1963
Separatus Subgroup (Rattanarithikul et al., 2006b)

Anopheles separatus Leicester 1908
Umbrosus Subgroup (Rattanarithikul et al., 2006b)

Anopheles umbrosus Theobald 1903

### Subgenus Baimaia

[edit]

Anopheles kyondawensis Abraham 1947[14]

### Subgenus Cellia

[edit]

Anopheles rageaui Mattingly & Adam

Series Cellia (Christophers, 1924a)

Anopheles argenteolobatus Gough 1910
Anopheles brumpti Hamon & Rickenbach 1955
Anopheles cristipalpis Service 1977
Anopheles murphyi Gillies & de Meillon 1968[17]
Anopheles pharoensis Theobald 1901[5]
Anopheles swahilicus Gillies 1964
Squamosus Group (Grjebine, 1966)

Anopheles cydippis de Meillon 1931
Anopheles squamosus Theobald 1901[5]

Series Myzomyia (Christophers, 1924a)

Anopheles apoci Marsh 1933
Anopheles azaniae Bailly-Choumara 1960
Anopheles barberellus Evans 1932
Anopheles brunnipes Theobald 1910
Anopheles domicolus Edwards 1916
Anopheles dthali Patton 1905
Anopheles erythraeus Corradetti 1939
Anopheles ethiopicus Gillies & Coetzee 1987
Anopheles flavicosta Edwards 1911
Anopheles fontinalis Gillies & de Meillon 1968[17]
Anopheles gabonensis Rahola, Makanga & Paupy 2014
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
Demeilloni Group (Gillies & de Meillon, 1968)

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
Funestus Group (Garros et al., 2005b)

Anopheles jeyporiensis James 1902
Aconitus Subgroup (Chen et al., 2003)

Anopheles aconitus Dönitz 1902
Anopheles filipinae Manalang 1930
Anopheles mangyanus Banks 1906
Anopheles pampanai Büttiker & Beales 1959
Anopheles varuna Iyengar 1924
Culicifacies Subgroup (Garros et al., 2005b)

Anopheles culicifacies* Giles 1901 (species A, B, C, D and E) (Kar et al., 1999)
Funestus Subgroup (Garros et al., 2005b)

Anopheles aruni Sobti 1968
Anopheles confusus Evans & Leeson 1935
Anopheles funestus* Giles 1900
Anopheles funestus-like* Spillings et al. 2009[21]
Anopheles longipalpis (Theobald) (Type C) (Koekemoer et al., 2009)
Anopheles parensis Gillies 1962
Anopheles vaneedeni Gillies & Coetzee 1987
Minimus Subgroup (Chen et al., 2003)

Anopheles flavirostris* Ludlow 1914
Anopheles leesoni Evans 1931
Anopheles longipalpis (Theobald) (Type A) (Koekemoer et al., 2009)
Fluviatilis Complex (Sarala et al., 1994)

Anopheles fluviatilis* James 1902 (species S, T and U)
Minimus Complex (Green et al., 1990)

Anopheles harrisoni Harbach & Manguin 2007 (species C)
Anopheles minimus Theobald 1901 (species A)[5]
Anopheles yaeyamaensis Somboon & Harbach
Rivulorum Subgroup (Garros et al., 2005b)

Anopheles brucei Service 1960
Anopheles fuscivenosus Leeson 1930
Anopheles rivulorum* Leeson 1935
Anopheles rivulorum-like (Cohuet et al., 2003)
Marshallii Group (Gillies & de Meillon, 1968)

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
Marshallii Complex (Gillies & Coetzee, 1987)

Anopheles hughi Lambert & Coetzee 1982
Anopheles kosiensis Coetzee, Segerman & Hunt 1987
Anopheles letabensis Lambert & Coetzee 1982
Anopheles marshallii Theobald 1903
Wellcomei Group (Gillies & de Meillon, 1968)

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, 1924a)

Anopheles ainshamsi Gad, Harbach & Harrison 2006
Anopheles dancalicus Corradetti 1939
Anopheles hervyi Brunhes, le Goff & Geoffroy 1999
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 pulcherrimus* Theobald 1902[15]
Anopheles rufipes Gough 1910

subspecies broussesi Edwards 1929
subspecies rufipes Gough 1910
Anopheles salbaii Maffi & Coluzzi 1958
Anopheles stephensi* Liston 1901 (potentially a species complex; see Firooziyan et al., 2018)
Anopheles superpictus* Grassi 1899 (species A and B) (Oshaghi et al., 2007; Oshaghi et al., 2008)
Anopheles theobaldi Giles 1901
Annularis Group (Reid, 1968)

Annularis Complex (Atrie et al., 1999)

Anopheles annularis* van der Wulp 1884 (species A and B)
Anopheles pallidus Theobald 1901[5]
Anopheles philippinensis* Ludlow 1902
Anopheles schueffneri Stanton 1915
Nivipes Complex (Green et al., 1985b; Harrison et al., 1991)

Anopheles nivipes Theobald 1903 (2 cytogenetic species in Thailand)
Jamesii Group (Rattanarithikul et al., 2006b)

Anopheles jamesii Theobald 1901[5]
Anopheles pseudojamesi Strickland & Chowdhury 1927
Anopheles splendidus Koidzumi 1920
Maculatus Group (Rattanarithikul & Green, 1987)

Anopheles dispar Rattanarithikul & Harbach 1991
Anopheles greeni Rattanarithikul & Harbach 1991
Anopheles pseudowillmori Theobald 1910
Anopheles willmori James 1903
Maculatus Subgroup (Rattanarithikul et al., 2006b)

Anopheles dravidicus Christophers 1924
Maculatus Complex (Ali et al., 2019b)

Anopheles maculatus* Theobald 1901[5]
Anopheles maculatus Javanese form (Ali et al., 2019a)
Sawadwongporni Subgroup (Rattanarithikul et al., 2006b)

Anopheles notanandai Rattanarithikul & Green 1987
Anopheles rampae Harbach & Somboon 2011
Anopheles sawadwongporni Rattanarithikul & Green 1987

Series Neomyzomyia (Christophers, 1924a)

Anopheles amictus Edwards 1921
A. amictus

Anopheles annulatus de Rook 1930
Anopheles aurirostris Watson 1910
Anopheles dualaensis Brunhes, le Goff & Geoffroy 1999
Anopheles hilli Woodhill & Lee 1944
Anopheles incognitus Brug 1931
Anopheles kokhani Vythilingam, Jeffery & Harbach 2007
Anopheles kolambuganensis Baisas 1932
Anopheles longirostris Brug 1928
Anopheles meraukensis Venhuis 1932
Anopheles novaguinensis Venhuis 1933
Anopheles saungi Colless 1955
Anopheles stookesi Colless 1955
Anopheles watsonii Leicester 1908
Annulipes Complex (Green, 1972)

Anopheles annulipes Walker 1856 (species A-Q) (Foley et al., 2007b)
A. annulipes

Kochi Group (Rattanarithikul et al., 2006b)

Anopheles kochi Dönitz 1901
Preserved specimens of A. kochi

Lungae Complex (Belkin, 1962)

Anopheles lungae Belkin & Schlosser 1944
Anopheles nataliae Belkin 1945
Anopheles solomonis Belkin, Knight & Rozeboom 1945
Ardensis Group (Gillies & de Meillon, 1968)

Anopheles ardensis Theobald 1905
Anopheles buxtoni Service 1958
Anopheles cinctus Newstead & Carter 1910
Anopheles deemingi Service 1970
Anopheles dureni Edwards 1930
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 & de Meillon 1968[17]
Anopheles vinckei de Meillon 1942
Nili Complex (Gillies & de Meillon, 1968)

Anopheles carnevalei Brunhes, le Goff & Geoffroy 1999
Anopheles nili* Theobald 1904
Anopheles ovengensis Awono-Ambene, Kengne, Simard, Antonio-Nkondjio & Fontenille 2004
Anopheles somalicus Rivola & Holstein 1957
Leucosphyrus Group (Reid, 1949)

Hackeri Subgroup (Sallum et al., 2005a)

Anopheles hackeri Edwards 1921
Anopheles mirans Sallum & Peyton 2005
Anopheles pujutensis Colless 1948
Anopheles recens Sallum & Peyton 2005
Anopheles sulawesi* Waktoedi Koesoemawinangoen 1954
Leucosphyrus Subgroup (Peyton, 1990; Sallum et al., 2005b)

Anopheles baisasi Colless 1957
Dirus Complex (Sallum et al., 2005b)

Anopheles aff. takasagoensis (Takano et al., 2010)
Anopheles baimaii* Sallum & Peyton 2005
Anopheles cracens Sallum & Peyton 2005
Anopheles dirus* Peyton & Harrison 1979
Anopheles elegans James 1903
Anopheles nemophilous Peyton & Ramalingam 1988
Anopheles scanloni Sallum & Peyton 2005
Anopheles takasagoensis Morishita 1946
Leucosphyrus Complex (Sallum et al., 2005a)

Anopheles balabacensis* Baisas 1936
Anopheles introlatus Colless 1957
Anopheles latens* Sallum & Peyton 2005
Anopheles leucosphyrus* Dönitz 1901
Riparis Subgroup (Peyton, 1990)

Anopheles cristatus King & Baisas 1936
Anopheles macarthuri Colless 1956
Anopheles riparis King & Baisas 1936
Mascarensis Group (Harbach, 1994a)

Anopheles mascarensis de Meillon 1947
Pauliani Group (Grjebine, 1966)

Anopheles grassei Grjebine
Anopheles grenieri Grjebine
Anopheles milloti Grjebine & Lacan
Anopheles pauliani Grjebine
Anopheles radama de Meillon
Punctulatus Group (Schmidt et al., 2001)

Anopheles clowi Rozeboom & Knight 1946
Anopheles koliensis Owen 1945
Anopheles punctulatus Dönitz 1901

Hosts: Bos taurus, Canis familiaris, Equus, Felis, Gallus[22]
Anopheles rennellensis Taylor & Maffi
Anopheles sp. near punctulatus (Foley et al., 1995)
Farauti Complex (Schmidt et al., 2003)

Anopheles farauti* Laveran 1902
Anopheles farauti 4 and 5 (Foley et al., 1993)
Anopheles farauti 8 (Bower et al., 2008)
Anopheles hinesorum Schmidt 2001
Anopheles irenicus Schmidt 2003
Anopheles oreios Bangs & Harbach 2014
Anopheles torresiensis Schmidt 2001
Ranci Group (Grjebine, 1966)

Anopheles griveaudi Grjebine
Ranci Subgroup (Grjebine, 1966)

Anopheles ranci Grjebine
Roubaudi Subgroup (Grjebine, 1966)

Anopheles lacani Grjebine
Anopheles notleyi van Someren
Anopheles roubaudi Grjebine
Rhodesiensis Group (Gillies & de Meillon, 1968)

Anopheles cameroni de Meillon & Evans
Anopheles lounibosi Gillies & Coetzee
Anopheles rhodesiensis Theobald

subspecies rhodesiensis Theobald
subspecies rupicolus Lewis
Anopheles rodhaini Leleup & Lips
Anopheles ruarinus Edwards
Smithii Group (Gillies & de Meillon, 1968)

Anopheles caroni Adam
Anopheles faini Leleup
Anopheles hamoni Adam
Anopheles jebudensis Froud
Anopheles lovettae Evans
Anopheles smithii Theobald
Anopheles vanhoofi Manson & Lebied
Anopheles wilsoni Evans
Tessellatus Group (Rattanarithikul et al., 2006b)

Tessellatus Complex (Bourke et al., 2021)

Anopheles tessellatus Theobald 1901 (species A, B, C, D, E and F)

subspecies kalawara Stoker & Waktoedi Koesoemawinangoen
subspecies orientalis Swellengrebel & Swellengrebel de Graaf
subspecies tessellatus Theobald

Series Paramyzomyia (Christophers & Barraud, 1931)

Cinereus Group (Gillies & de Meillon, 1968)

Anopheles azevedoi Ribeiro 1969
Anopheles cinereus Theobald 1901[5]

subspecies cinereus Theobald 1901[5]
subspecies hispaniola Theobald 1903
Anopheles turkhudi Liston 1901

subspecies telamali Saliternik & Theodor 1942
subspecies turkhudi Liston 1901
Listeri Group (Gillies & de Meillon, 1968)

Anopheles listeri de Meillon 1931
Anopheles multicolor* Cambouliu 1902
Anopheles seretsei Abdulla-Khan, Coetzee & Hunt 1998

Series Pyretophorus (Edwards, 1932a)

Anopheles christyi Newstead & Carter 1911
Anopheles comorensis Brunhes, le Goff & Geoffroy 1997
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 Koesoemawinangoen 1949
Anopheles parangensis Ludlow 1914
Anopheles pseudosundaicus Tyagi, Hiriyan, Tewari et al.
Anopheles vagus* Dönitz 1902
Gambiae Complex (White, 1985)

Anopheles amharicus Hunt, Wilkerson & Coetzee 2013
Anopheles arabiensis* Patton 1905
Anopheles bwambae White 1985
Anopheles coluzzii* Coetzee & Wilkerson 2013 [23]
Anopheles gambiae* Giles 1902
Anopheles melas* Theobald 1903
Anopheles merus Dönitz 1902
Anopheles quadriannulatus Theobald 1911
Subpictus Complex (Suguna et al., 1994)

Anopheles subpictus* Grassi 1899 (species A, B, C and D)
Sundaicus Complex (Sukowati et al., 1999)

Anopheles epiroticus Linton & Harbach 2005
Anopheles sundaicus* Rodenwaldt 1925
Anopheles sundaicus (species B, C, D and E) (Dusfour et al., 2007)

### Subgenus Kerteszia

[edit]

Anopheles auyantepuiensis Harbach & Navarro 1996
Anopheles bambusicolus Komp 1937
Bellator Complex (Bourke et al., 2023)

Anopheles bellator* Dyar & Knab 1906 (bellator 1)
Anopheles bellator 2
Boliviensis Complex (Bourke et al., 2023)

Anopheles boliviensis 1 (identity of boliviensis s.s. not determined)
Anopheles boliviensis 2
Anopheles boliviensis 3
Cruzii Complex (Ramirez & Dessen, 2000a)

Anopheles cruzii* Dyar & Knab 1908 (species A, B and C) (Ramirez & Dessen, 2000b)[24]
Homunculus Complex (Bourke et al., 2023)

Anopheles homunculus* Komp 1937 (homunculus 1?)
Anopheles homunculus 2
Laneanus Complex (Bourke et al., 2023)

Anopheles laneanus 1
Anopheles laneanus Corrêa & Cerqueira 1944 (laneanus 2)
Neivai Complex (Bourke et al., 2023)

Anopheles neivai Howard, Dyar & Knab 1913 (neivai 1)
Anopheles neivai 2
Anopheles neivai 3
Anopheles neivai 4
Anopheles neivai 5
Anopheles neivai 6
Anopheles neivai 7
Anopheles neivai 8 (neivai nr neivai 4 of Ahumada et al., 2016)
Pholidotus Complex (Bourke et al., 2023)

Anopheles pholidotus 1 (identity of pholidotus s.s. not determined)
Anopheles pholidotus 2
Anopheles pholidotus 3
Anopheles pholidotus 4
Rollai Complex (Bourke et al., 2023)

Anopheles rollai 1 (identity of rollai s.s. not determined)
Anopheles rollai 2
Anopheles rollai 3
Anopheles rollai 4
Anopheles rollai 5
Anopheles gonzalezrinconesi Cova García, Pulido & de Ugueto 1977
Anopheles lepidotus Zavortink 1973

### Subgenus Lophopodomyia

[edit]

Anopheles gilesi Peryassú 1928
Anopheles gomezdelatorrei Leví-Castillo 1955
Anopheles oiketorakras Osorno-Mesa 1947
Anopheles pseudotibiamaculatus Galvão & Barretto 1941
Anopheles squamifemur Antunes 1937
Anopheles vargasi Gabaldón, Cova García & Lopez 1941

### Subgenus Nyssorhynchus

[edit]

Section Albimanus (Levi Castillo, 1949)

Anopheles noroestensis Galvão & Lane 1937
Series Albimanus (Faran, 1980)

Anopheles albimanus* Wiedemann 1820
Series Oswaldoi (Faran, 1980)

Oswaldoi Group (Faran, 1980)

Oswaldoi Subgroup (Faran, 1980)

Anopheles anomalophyllus Komp
Anopheles aquasalis* Curry 1932
Anopheles evansae Brèthes 1926
Anopheles galvaoi Causey, Deane & Deane 1943
Anopheles ininii Senevet & Abonnenc 1938
Konderi Complex (Ruiz-Lopez et al., 2013)

Anopheles konderi Galvão & Damasceno 1942
Anopheles tadei Saraiva & Scarpassa (formerly sp. nr. konderi of Ruiz-Lopez et al., 2013)
Nuneztovari Complex (Mirabello & Conn, 2008; Foster et al., 2013; Scarpassa et al., 2016)

Anopheles dunhami Causey 1945
Anopheles goeldii Rozeboom & Gabaldón 1941
Anopheles jamariensis Sant'Ana & Sallum
Anopheles nuneztovari* Gabaldón 1940
Anopheles nuneztovari B and C
Oswaldoi Complex (Ruiz-Lopez et al., 2013)

Anopheles oswaldoi Peryassú 1922
Anopheles oswaldoi A (Ruiz-Lopez et al., 2013)
Anopheles oswaldoi B (Ruiz et al., 2005; Ruiz et al., 2010)
Anopheles rangeli Gabaldón, Cova García & Lopez 1941
Anopheles sanctielii Senevet & Abonnenc 1938
Anopheles trinkae Faran 1980
Strodei Subgroup (Faran, 1980)

Anopheles albertoi Unti
Anopheles CP Form (Sallum et al., 2010)
Anopheles rondoni Neiva & Pinto 1922
Anopheles striatus Sant'Ana & Sallum
Anopheles strodei Root 1926
Arthuri Complex (Bourke et al., 2013)

Anopheles arthuri*
Anopheles ibiapabaensis (Sant’Ana & Sallum, 2023)
Anopheles untii (Sant’Ana & Sallum, 2023)
Anopheles rondoniensis Sant'Ana & Sallum 2022
Benarrochi Complex (Ruiz et al., 2005)

Anopheles benarrochi Gabaldón, Cova García & Lopez 1941
Anopheles benarrochi B (Ruiz et al., 2005)
Triannulatus Group (Faran, 1980)

Anopheles halophylus Silva do Nascimento & Lourenço-de-Oliveira 2002
Anopheles triannulatus Neiva & Pinto 1922
Anopheles triannulatus (species C) (Silva-do-Nascimento & Lourenço-de-Oliveira, 2007)

Section Argyritarsis (Levi Castillo, 1949)

Series Albitarsis[25]

Anopheles rooti Brèthes 1926
Albitarsis Group (Linthicum, 1988)

Albitarsis Complex (Wilkerson et al., 1995)

Anopheles albitarsis Lynch Arribálzaga 1878
Anopheles albitarsis (species F, G, H and I) (Brochero et al., 2007; Ruiz-Lopez et al., 2012)
Anopheles deaneorum Rosa-Freitas 1989
Anopheles janconnae Wilkerson & Sallum 2009
Anopheles lineage nr janconnae (Gutiérrez et al., 2010)
Anopheles marajoara* Galvão & Damasceno 1942 (lineages 1 and 2) (McKeon et al., 2010)
Anopheles oryzalimnetes Wilkerson & Motoki 2009
Braziliensis Group (Linthicum, 1988)

Anopheles braziliensis Chagas 1907
Series Argyritarsis (Linthicum, 1988)

Argyritarsis Group (Linthicum, 1988)

Anopheles argyritarsis Robineau-Desvoidy 1827
Anopheles sawyeri Causey, Deane, Deane & Sampaio 1943
Darlingi Group (Linthicum, 1988)

Anopheles darlingi* Root 1926
Lanei Group (Linthicum, 1988)

Anopheles lanei Galvão & Franco do Amaral 1938
Pictipennis Group (Linthicum, 1988)

Anopheles atacamensis González & Sallum
Anopheles pictipennis Philippi 1865

Section Myzorhynchella (Peyton et al., 1992)

Anopheles antunesi Galvão & Franco do Amaral 1940
Anopheles guarani Shannon 1928
Anopheles lutzii Cruz 1901
Anopheles nigritarsis Chagas 1907
Anopheles parvus Chagas 1907
Anopheles pristinus Nagaki & Sallum

### Subgenus Stethomyia

[edit]

Anopheles acanthotorynus Komp 1937
Anopheles canorii Floch & Abonnenc 1945
Anopheles kompi Edwards 1930
Anopheles nimbus Theobald 1902[15]
Anopheles thomasi Shannon 1933

### Nomen dubium

[edit]

Anopheles africanus Roque, 1903
Anopheles allopha (Peryassú)
Anopheles arnoulti Grjebine
Anopheles brachypus Dönitz, 1902
Anopheles costalis Loew, 1866
Anopheles courdurieri Grjebine, 1966
Anopheles jacobi (Hill & Haydon)
Anopheles minutus Macquart
Anopheles nero (Doleschall)
Anopheles soalalaensis Grjebine
Anopheles upemba Lips
Anopheles vulgaris Hatori

### Extinct species

[edit]

†Anopheles (Nyssorhynchus) dominicanus Zavortink & Poinar 2000
†Anopheles rottensis Statz, 1944

## Notes

[edit]

* 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.[26]

## References

[edit]

1. ^ Harbach, Ralph E. (2020). "An updated classification of the Anopheles genus (Diptera: Culicidae)". Parasites & Vectors. 13 (1): 1–14\. doi:10.1186/s13071-020-04203-1. PMC 7329488.
2. ^ "Anopheles – Walter Reed Biosystematics Unit". Smithsonian Institution. Retrieved 1 April 2025.
3. ^ "Taxonomy of Anopheles". Wikipedia. Retrieved 1 April 2025.
4. ^ "Anopheles gambiae s.l.: Morphology, Life Cycle, and Ecology". Target Malaria. Retrieved 1 April 2025.
5. ^ a b c d e f g h i Theobald, Frederick Vincent (1901). A Monograph of the Culicidae, or Mosquitoes. Vol. 1\. London: British Museum (Natural History). ISBN 978-1178519037. `{{cite book}}`: ISBN / Date incompatibility (help)[page needed]
6. ^ Krzywinski, Jaroslaw; Besansky, Nora J (2003). "MOLECULARSYSTEMATICS OFANOPHELES: From Subgenera to Subpopulations". Annual Review of Entomology. 48: 111–39\. doi:10.1146/annurev.ento.48.091801.112647. PMID 12208816.
7. ^ Foley, Desmond H; Bryan, Joan H; Yeates, David; Saul, Allan (1998). "Evolution and Systematics of Anopheles:Insights from a Molecular Phylogeny of Australasian Mosquitoes". Molecular Phylogenetics and Evolution. 9 (2): 262–75\. Bibcode:1998MolPE...9..262F. doi:10.1006/mpev.1997.0457. PMID 9562985.
8. ^ Calle, D. A; Quiñones, M. L; Erazo, H. F; Jaramillo, N (2008). "Discriminación por morfometría geométrica de once especies de Anopheles (Nyssorhynchus) presentes en Colombia" [Differentiation by geometric morphometrics among 11 Anopheles (Nyssorhynchus) in Colombia]. Biomedica: Revista del Instituto Nacional de Salud (in Portuguese). 28 (3): 371–85\. PMID 19034360.
9. ^ Rattanarithikul, R; Harrison, B. A; Harbach, R. E; Panthusiri, P; Coleman, R. E; Panthusiri, P (2006). "Illustrated keys to the mosquitoes of Thailand. IV. Anopheles". The Southeast Asian Journal of Tropical Medicine and Public Health. 37 (Suppl 2): 1–128\. PMID 17262930.
10. ^ Sedaghat, M. M; Harbach, R. E (2005). "An annotated checklist of the Anopheles mosquitoes (Diptera: Culicidae) in Iran". Journal of Vector Ecology. 30 (2): 272–6\. PMID 16599162.
11. ^ Walton, C; Somboon, P; o'Loughlin, S.M; Zhang, S; Harbach, R.E; Linton, Y.-M; Chen, B; Nolan, K; Duong, S; Fong, M.-Y; Vythilingum, I; Mohammed, Z.D; Trung, Ho Dinh; Butlin, R.K (2007). "Genetic diversity and molecular identification of mosquito species in the Anopheles maculatus group using the ITS2 region of rDNA". Infection, Genetics and Evolution. 7 (1): 93–102\. Bibcode:2007InfGE...7...93W. doi:10.1016/j.meegid.2006.05.001. PMID 16782411.
12. ^ Garros, C; Harbach, R. E; Manguin, S (2005). "Morphological assessment and molecular phylogenetics of the Funestus and Minimus groups of Anopheles (Cellia)". Journal of Medical Entomology. 42 (4): 522–36\. doi:10.1603/0022-2585(2005)042[0522:maampo]2.0.co;2. PMID 16119539. S2CID 22776177.
13. ^ a b Meigen, J.W. (1818). Systematische Beschreibung der bekannten europäischen zweiflügeligen Insekten [Systematic description of the known European two-winged insects] (in German). Aachen. OCLC 13208765.[page needed]
14. ^ a b Abraham, O. K. (1947). "Anopheles kyondawensis n. sp. from Lower Burma". Medical Journal of Malaya. 1947 (1): 173–176.
15. ^ a b c Theobald, Frederick Vincent (1902). "The Classification of the Anophelina" (PDF). Journal of Tropical Medicine. 5: 181–183. Retrieved 13 September 2015.
16. ^ Harbach, Ralph E; Kitching, Ian J (2005). "Reconsideration of anopheline mosquito phylogeny (Diptera: Culicidae: Anophelinae) based on morphological data". Systematics and Biodiversity. 3 (4): 345–74\. Bibcode:2005SyBio...3..345H. doi:10.1017/S147720000500174X. S2CID 84208117.
17. ^ a b c d Gillies, M. T.; De Meillon, Botha (1968). The Anophelinae of Africa south of the Sahara (Ethiopian zoogeographical region) (2 ed.). Johannesburg: South African Institute for Medical Research. OCLC 9614034.[page needed]
18. ^ Scarpassa, Vera Margarete; Conn, Jan E (2011). "Mitochondrial DNA Detects a Complex Evolutionary History with Pleistocene Epoch Divergence for the Neotropical Malaria Vector Anopheles nuneztovari Sensu Lato". The American Journal of Tropical Medicine and Hygiene. 85 (5): 857–67\. doi:10.4269/ajtmh.2011.11-0150. PMC 3205631. PMID 22049039.
19. ^ Bonne-Wepster, J. (1951). "Anopheles venhuisi n. sp. [sic]". Documenta Neerlandica et Indonesica de Morbis Tropicis. 3: 24.
20. ^ Senevet, G.; Prunnelle, M. (1927). "Une nouvelle espèce d'anophèle en Algérie, Anopheles marteri n. sp" [A new species of anopheles in Algeria, Anopheles marteri n. sp]. Archives de l'Institut Pasteur d'Algérie (in French). 5 (4): 529–533.
21. ^ Spillings BL, Brooke BD, Koekemoer LL, Chiphwanya J, Coetzee M, Hunt RH (2009) A new species concealed by Anopheles funestus Giles, a major malaria vector in Africa. Am J Trop Med Hyg 81(3):510-5
22. ^ "Anopheles punctulatus". Invasive Species Compendium (ISC). CABI (Centre for Agriculture and Bioscience International). 2019-11-20. Retrieved 2022-01-09.
23. ^ Coetzee, M.; Hunt, R. H.; Wilkerson, R.; Della Torre, A.; Coulibaly, M. B.; Besansky, N. J. (2013). "Anopheles coluzzii and Anopheles amharicus, new members of the Anopheles gambiae complex". Zootaxa. 3619 (3): 246–74\. doi:10.11646/zootaxa.3619.3.2. PMID 26131476.
24. ^ Rona LD, Carvalho-Pinto CJ, Peixoto AA (2010) Molecular evidence for the occurrence of a new sibling species within the Anopheles (Kerteszia) cruzii complex in south-east Brazil. Malar J 26, 9(1):33
25. ^ Linthicum, Kenneth J. (1988). "A revision of Argyritarsis section of the subgenus Nyssorhynchus of Anopheles" (PDF). Mosquito Systematics. 20 (2): 98–271. Retrieved 13 September 2015.
26. ^ Riehle, M. M; Guelbeogo, W. M; Gneme, A; Eiglmeier, K; Holm, I; Bischoff, E; Garnier, T; Snyder, G. M; Li, X; Markianos, K; Sagnon, N; Vernick, K. D (2011). "A Cryptic Subgroup of Anopheles gambiae is Highly Susceptible to Human Malaria Parasites". Science. 331 (6017): 596–8\. Bibcode:2011Sci...331..596R. doi:10.1126/science.1196759. PMC 3065189. PMID 21292978.

## External links

[edit]

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

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