Subgenus Anopheles Meigen, 1818
Anopheles maculipennis Meigen, 1818.
Subfamily Anophelinae, genus Anopheles. Subgenus Anopheles includes 208 formally recognised species divided between two Sections that are further subdivided into a number of Series comprised of Groups and Subgroups believed to represent phylogenetically related assemblages of species based principally on morphological similarity (Harbach, 2004). Subgenus abbreviation – Ano.
Subgenus Anopheles includes species with and without spotted wings. Those with pale markings have no more than two pale areas on the costal margin. Pale scales of other veins, when present, usually do not form distinct pale areas. Cibarial armature is absent in females and the genitalia of males have 1‒3 (usually 2) specialised setae (generally one internal and two parabasal setae) on the gonocoxites, with at least the inner parabasal borne on a distinct prominence. Proepisternal setae are usually numerous and prespiracular setae are usually present. Larvae, with few exceptions, have seta 1-A branched and usually inserted on the mesal (inner) side of the antenna. Seta 2-C are closely approximated (closer, rarely equal, than the distance between seta 2-C and seta 3-C on either side) and setae of the pleural groups (setae 9‒12-P,M,T) are generally all simple. The subgenus is divided into two Sections based on the shape of the pupal trumpet. The Laticorn Section includes species with a wide funnel-shaped trumpet having the longest axis transverse to the stem (laticorn), and the Angusticorn Section includes species with a semi-tubular trumpet having the longest axis vertical more or less in line with the stem (angusticorn). Pupal seta 9 on segments V‒VII is usually short and blunt, and seta 1-Pa is usually short and straight, seldom long and bent. See genus Anopheles.
The morphological and molecular phylogenetic studies of Sallum et al. (2000, 2002) and Harbach & Kitching (2005, 2016) indicate that subgenus Anopheles is not demonstrably monophyletic with regard to genus Bironella and subgenera Lophopodomyia and Stethomyia, and the monophyly of the Sections and most Series are doubtful. In contrast, the results of a cladistic analysis of morphological data by Collucci & Sallum (2007) and relationships recovered in the phylogenetic analyses of COI and COII mtDNA and 2.5S rDNA sequences by Freitas et al. (2015) suggest that the subgenus is monophyletic. The results of Collucci & Sallum (2007), however, were biased by the selection of outgroup taxa whose interrelationships with the ingroup taxa were unresolved in the earlier studies of Sallum et al. (2000, 2002). A molecular phylogeny of 18 species constructed from aligned protein sequences of 1,085 single-copy orthologs indicates that subgenus Anopheles is sister to subgenus Cellia (Neafsey et al., 2015). Anopheles was also recovered as sister to Cellia in the phylogeny of Foster et al. (2017) based on analyses of mitochondrial protein coding genes obtained for four species of the Old World subgenus Cellia and nine predominantly Neotropical species of subgenus Anopheles (one species, An. quadrimaculatus, occurs in Central and North America).
Anopheles larvae are adapted to a variety of aquatic habitats, but occur predominantly in ground waters. Some species require aerated water, others brackish water and some inhabit cavities such as rock holes and tree holes (e.g. the Plumbeus Group). Adults are active at night and during twilight periods, but the females of some species will accost potential hosts in dimly lit places during daylight hours. Many species are known feed on humans and domestic animals, but the host preferences of most species are unknown.
Subgenus Anopheles is divided into six Series, but only females of the Myzorhynchus and Anopheles Series include vectors of human malarial parasites. The Hyrcanus and Maculipennis Groups include principal vectors of historical and contemporary importance in the Holarctic Region. Some members of the subgenus are effective vectors of microfilariae and some may be involved in the transmission of encephalitis viruses.
Subgenus Anopheles has an almost world-wide distribution. Species are found at elevations from coastal areas to mountainous terrain in temperate, subtropical and tropical areas, but are absent from the majority of the Pacific Islands, including the large ones of New Zealand, Fiji and New Caledonia.
Lane, 1953 (Neotropical Region); Mattingly & Knight, 1956 (Arabia); Cova-Garcia, 1961 (Venezuela); Forattini, 1962 (Neotropical Region); Gillies & de Meillon, 1968 (Afrotropical Region); Reid, 1968 (Malaysia, Borneo); Belkin et al., 1970 (Jamaica); Gutsevich et al., 1974 (former USSR); Harrison & Scanlon, 1975 (Thailand); Klein, 1977 (Cambodia); Tanaka et al., 1979 (Japan); Wood et al., 1979 (Canada); Darsie & Ward, 1981, 2005 (keys, North America); Lu & Li, 1982 (China); Clark-Gil & Darsie, 1983 (Guatemala); Rao, 1984 (India); Lee et al., 1987 (Australasian Region); Gillies & Coetzee, 1987 (Afrotropical Region); Das et al., 1990 (keys, India); Darsie & Pradhan, 1990 (Nepal); Wilkerson & Strickman, 1990 (keys, Central America and Mexico); Glick, 1992 (keys, southwestern Asia and Egypt); Rattanarithikul & Panthusiri, 1994 (keys, medically important species, Thailand); Nagpal & Sharma, 1995 (India); Lu Baolin et al., 1997 (China); Rattanarithikul et al., 2006 (keys, Thailand); Collucci & Sallum, 2007 (phylogeny); Gómez et al., 2015 (Arribalzagia Series, molecular taxonomy); Foster et al., 2017 (phylogenetic relationships); Namgay et al., 2020 (Lindesayi Complex); Coetzee, 2022 (Afrotropical Region, adult females, descriptions, biology, distributions).
aberrans Harrison & Scanlon, 1975
acaci Baisas, 1946
ahomi Chowdhury, 1929
aitkenii James, 1903 (in Theobald, 1903)
albotaeniatus (Theobald, 1903)
algeriensis Theobald, 1903
alongensis Venhuis, 1940
anchietai Corrêa & Ramalho, 1968
annandalei Prashad, 1918
annulipalpis Lynch Arribálzaga, 1878
apicimacula Dyar & Knab, 1906
arboricolus Zavortink, 1970
argentinus (Brèthes, 1912)
argyropus (Swellengrebel, 1914)
artemievi Gordeyev, Zvantsov, Goryacheva, Shaikevich & Yezhov, 2005
asiaticus Leicester, 1903
atratipes Skuse, 1889
atroparvus van Thiel, 1927
atropos Dyar & Knab, 1906
aztecus Hoffmann, 1935
baezai Gater, 1933
baileyi Edwards, 1929
balerensis Mendoza, 1947
bancroftii Giles, 1902
barberi Coquillett, 1903
barbirostris van der Wulp, 1884
barbiventris Brug, 1938
barbumbrosus Strickland & Chowdhury, 1927
barianensis James, 1911 (in James & Liston, 1911)
beklemishevi Stegnii & Kabanova, 1976
belenrae Rueda, 2005
bengalensis Puri, 1930
benguetensis King, 1931
bhutanensis Somboon, Namgay & Harbach (in Somboon et al., 2021)
borneensis McArthur, 1949
bradleyi King, 1939
brevipalpis Roper, 1914
brevirostris Reid, 1950
bulkleyi Causey, 1937
bustamentei Galvão, 1955
calderoni Wilkerson, 1991
caliginosus de Meillon, 1943
cameronensis Edwards, 1929
campestris Reid, 1962
chiriquiensis Komp, 1936
chodukini Martini, 1929
claviger (Meigen, 1804)
colledgei Marks, 1956
collessi Reid, 1963
concolor Edwards, 1938 (in Evans, 1938)
corethroides Theobald, 1907
costai da Fonseca & da Silva Ramos, 1940
coustani Laveran, 1900
crawfordi Reid, 1953
crockeri Colless, 1955
crucians Wiedemann, 1828
crypticus Coetzee, 1995
cucphuongensis Vu, Nguyen, Tran & Nguyen, 1991
culiciformis Cogill, 1903
daciae Linton, Nicolescu & Harbach, 2004 (in Nicolescu et al., 2004)
danaubento Mochtar & Walandouw, 1934
diluvialis Reinert, 1997 (in Reinert et al., 1997)
dissidens Taai & Harbach, 2015
donaldi Reid, 1962
druki Somboon, Namgay & Harbach, 2022 (in Somboon et al., 2022
earlei Vargas, 1943
eiseni Coquillett, 1902
ejercitoi Mendoza,1947
engarensis Kanda & Oguma, 1978
evandroi da Costa Lima, 1937
fausti Vargas, 1943
fluminensis Root, 1927
forattinii Wilkerson & Sallum, 1999
formosus Ludlow, 1909
fragilis (Theobald, 1903)
franciscanus McCracken, 1904
franciscoi Reid, 1962
freeborni Aitken, 1939
freyi Meng, 1957
fuscicolor van Someren, 1947
gabaldoni Vargas, 1941
geometricus Corrêa, 1944
georgianus King, 1939
gigas Giles, 1901
grabhamii Theobald, 1901
guarao Anduze & Capdevielle, 1949
hailarensis Xu & Luo, 1998
hectoris Giaquinto Mira, 1931
heiheensis Ma, 1981
hermsi Barr & Guptavanij, 1989
himalayensis Somboon, Namgay & Harbach, 2022 (in Somboon et al., 2022)
hodgkini Reid, 1962
hunteri (Strickland, 1916)
hyrcanus (Pallas, 1771)
insulaeflorum (Swellengrebel & Swellengrebel de Graaf, 1920)
interruptus Puri, 1929
inthanonensis Somboon & Harbach, 2024 (in Phanitchakun et al., 2024)
inundatus Reinert,1997 (in Reinert et al., 1997)
japonicus Yamada, 1918
judithae Zavortink, 1969
kleini Rueda, 2005
koreicus Yamada & Watanabe, 1918
kweiyangensis Yao & Wu, 1944
labranchiae Falleroni, 1926
lesteri Baisas & Hu, 1936
letifer Sandosham, 1944
levicastilloi Levi Castillo, 1944
lewisi Ludlow, 1920
liangshanensis Kang, Tan, Cao, Cheng, Yang & Huang, 1984
lindesayi Giles, 1900
maculipennis Meigen, 1818
maculipes (Theobald, 1903)
malefactor Dyar & Knab, 1907
manalangi Mendoza, 1940
marteri Senevet & Prunnelle, 1927
martinius Shingarev, 1926
mattogrossensis Lutz & Neiva, 1911
maverlius Reinert, 1997 (in Reinert et al., 1997)
medialis Harbach, 2018
mediopunctatus (Lutz, 1903) (in Theobald, 1903)
melanoon Hackett, 1934
mengalangensis Ma, 1981
messeae Falleroni, 1926
minor da Costa Lima, 1929
montanus Stanton & Hacker, 1917
monticola Somboon, Namgay & Harbach, 2021 (in Somboon et al., 2021)
namibiensis Coetzee, 1984
neghmei Mann, 1950
neomaculipalpus Curry, 1931
nigerrimus Giles, 1900
nilgiricus Christophers, 1924
nimpe Nguyen, Tran & Harbach, 2000 (in Nguyen et al., 2000)
nitidus Harrison, Scanlon & Reid, 1973
noei Mann, 1950
noniae Reid, 1963
obscurus (Grünberg, 1905)
occidentalis Dyar & Knab, 1906
omorii Sakakibara, 1959
palmatus (Rodenwaldt, 1926)
paludis Theobald, 1900
papuensis Dobrotworsky, 1957
parapunctipennis Martini, 1932
peditaeniatus (Leicester, 1908)
perplexens Ludlow, 1907
persiensis Linton, Sedaghat & Harbach, 2003 (in Sedaghat et al., 2003)
peryassui Dyar & Knab, 1908
petragnani del Vecchio, 1939
peytoni Kulasekera, Harrison & Amerasinghe, 1989
pilinotum Harrison & Scanlon, 1974
pinjaurensis Barraud, 1932
pleccau Koidzumi, 1924
plumbeus Stephens, 1828
pollicaris Reid, 1962
powderi Zavortink, 1970
powelli Lee, 1944
prachongae Rattanarithikul & Harrison, 2017 (in Harbach et al., 2017)
pseudobarbirostris Ludlow, 1902
pseudomaculipes (Chagas, 1908) (in Peryassú, 1908)
pseudopictus Grassi, 1899 (in Grassi et al., 1899)
pseudopunctipennis Theobald, 1901
pseudosinensis Baisas, 1935
pseudostigmaticus Dobrotworsky, 1957
pullus Yamada, 1937
punctimacula Dyar & Knab, 1906
punctipennis (Say, 1823)
pursati Laveran, 1902
quadrimaculatus Say, 1824
rachoui Galvao, 1952
refutans Alcock, 1913
reidi Harrison, 1973
rivadeneirai Levi-Castillo, 1945
roperi Reid, 1950
sacharovi Favre, 1903
saeungae Taai & Harbach, 2015
samarensis Rozeboom, 1951
saperoi Bohart & Ingram, 1946
sarpangensis Somboon, Namgay & Harbach, 2024 (in Somboon et al., 2024)
separatus (Leicester, 1908
shannoni Davis, 1931
simlensis (James, 1911) (in James & Liston, 1911
similissimus Strickland & Chowdhury, 1927
sinensis Wiedemann, 1828
sineroides Yamada, 1924
sintoni Puri, 1929
sintonoides Ho, 1938
smaragdinus Reinert, 1997 (in Reinert et al., 1997)
stigmaticus Skuse, 1889
stricklandi Reid, 1965
sumatranus Swellengrebel & Rodenwaldt, 1932
symesi Edwards, 1928
tasmaniensis Dobrotworsky, 1966
tenebrosus Dönitz, 1902
thimphuensis Somboon, Namgay & Harbach, 2022 (in Somboon et al., 2022)
tibiamaculatus (Neiva, 1906
tigertti Scanlon & Peyton, 1967
umbrosus (Theobald, 1903)
vanderwulpi Townson & Harbach, 2013 (in Townson et al., 2013)
vanus Walker, 1859
wejchoochotei Taai & Harbach, 2015
veruslanei Vargas, 1979
vestitipennis Dyar & Knab, 1906
vietnamensis Nguyen, Tran & Nguyen, 1993
walkeri Theobald, 1901
wellingtonianus Alcock, 1912
whartoni Reid, 1963
xelajuensis de Leon, 1938
xui Dong, Zhou, Dong & Mao, 2007
ziemanni Grünberg, 1902
