Anopheles is a genus of mosquitoes (Culicidae) with about 484 recognised species.
Classification
The classification of this genus began in 1901 with Frederick Vincent Theobald. Despite the passage of time, the taxonomy remains incompletely settled. 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.
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.
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 1818 Subgenus Baimaia - Anopheles kyondawensis Abraham 1947 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
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.
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) 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.
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 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 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
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 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
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