Subgenus Cellia Theobald, 1902
Anopheles pharoensis Theobald, 1901 [see Theobald, 1902].
Subfamily Anophelinae, genus Anopheles. Subgenus Cellia includes 233 formally recognised species and a growing number of unnamed members of sibling species complexes that are divided between six Series comprised of Groups and Subgroups (see Anopheles Classification, subgenus Cellia) believed to represent phylogenetically related assemblages based principally on morphological similarity (Harbach, 2004). Subgenus abbreviation – Cel.
Subgenus Cellia includes species with four or more pale spots involving the costa and veins R-R1, and the other veins usually also have distinct pale markings. With few exceptions, the cibarial armature is well developed in females and the genitalia of males have a cluster of 4‒6 (usually 5) parabasal setae (no internal seta) on the gonocoxites, usually not borne on raised tubercles. Larvae usually have a small, single seta 1-A inserted on the lateral (outer) side of the antenna. Seta 2-C are normally well separated, at least as far apart as the distance between 2-C and 3-C on either side. Each pleural setal groups (setae 9-12-P,M,T) is variable, but one or both long mesopleural and metapleural setae are often plumose. Pupae have a semi-tubular trumpet (angusticorn) with the longest axis vertical more or less in line with the stem, seta 9 on segments V-VII is usually relatively long and sharply pointed, paddle seta 1-Pa is commonly long and hooked or curved, and the genital lobe of male pupae has an apical pair of rounded points or knobs. See genus Anopheles.
Phylogenetic analyses of morphological and molecular data (Krzywinski et al., 2001a, 2001b; Sallum et al., 2000, 2002; Harbach & Kitching, 2005, 2016; Mohanty et al., 2009; Wang et al., 2014; Norris & Norris, 2015; Freitas et al., 2015) have shown that subgenus Cellia is a monophyletic group. Two unique, shared characters, cibarial armature of females with cones only and the genital lobe of pupae with a finger-like projection, support a sister-group relationship between Cellia (Old World) and Kerteszia + Nyssorhynchus (New World) (Harbach & Kitching, 2005). It is noteworthy, however, that a molecular phylogeny of 18 species constructed from aligned protein sequences of 1,085 single-copy orthologs indicated that the Nyssorhynchus lineage diverged prior to the lineages that gave rise to subgenera Anopheles and Cellia (Neafsey et al., 2015). Dixit et al. (2010) recently inferred the phylogeny and divergence times of the Myzomyia, Neocellia and Pyretophorus Series based on COII sequences of the mitochondrial genome. The molecular phylogeny of Neafsey et al. (2015) convincingly supports the monophyly of the Pyretophorus Series and its emergence about 30 Mya. The molecular studies of Sallum et al. (2007), Sum et al. (2014) and Wang et al. (2014) provide evidence that the Funestus Group (Myzomyia Series), Maculatus Group (Neocellia Series) and the Leucosphyrus Group (Neomyzomyia Series) are monophyletic assemblages; however, the results of morphological (Sallum et al., 2000; Harbach & Kitching, 2005, 2016) and other molecular studies (Sallum et al. 2002; Mohanty et al. 2009; Norris & Norris 2015; Freitas et al., 2015) indicate that a significant part of the current internal classification of Cellia does not reflect evolutionary relationships. The molecular phylogeny of Neafsey et al., 2015 indicated that subgenus Cellia is sister to subgenus Anopheles. Cellia was also recovered as sister to Anopheles in the phylogeny of Foster et al. (2017) based on analyses of mitochondrial protein coding genes obtained for four species of subgenus Cellia and nine predominantly Neotropical species of subgenus Anopheles (one species, An. quadrimaculatus, occurs in Central and North America).
The immature stages of species of subgenus Cellia are adapted to a wide variety of ground-water habitats, ranging from small collections of water such as animal footprints and puddles that lack vegetation to large bodies of water such as rice fields, swamps and river margins that have shelter provided by vegetation. A number of species thrive in both fresh- and brackish-water habitats and some normally inhabit only brackish-water pools. Whereas the bionomics of species involved in malaria transmission is generally well known, comparatively little is known about the biology of most species. See subfamily Anophelinae and genus Anopheles for general information concerning the bionomics of anopheline mosquitoes.
Subgenus Cellia is divided into six Series (Cellia, Neocellia, Myzomyia, Neomyzomyia, Paramyzomyia and Pyretophorus). Each series contains vectors of malarial protozoa and microfilariae. The principal vectors of malaria include members of sibling species complexes: the gambiae and funestus complexes in the Afrotropical Region; the culicifacies, dirus; fluviatilis, leucosphyrus, maculatus, minimus and sundaicus complexes in the Oriental Region; the farauti and punctulatus complexes in the Australasian Region. Anopheles sergentii (Myzomyia Series) and An. stephensi (Neocellia Series) are also major vectors in the Middle East and the Indian Subcontinent. Analysis of DNA sequence for the Ansteobp1 intron I gene (Firooziyan et al., 2018) suggests that An. stephensi could be a complex of three sibling species: An. stephensi sibling A (type form), An. stephensi sibling B (intermediate form) and An. stephensi sibling C (mysorensis form).
Most species of subgenus Cellia have distributions in the Afrotropical, Australian and Oriental Regions, but some species occur in southern areas of the Palaearctic. Species of Cellia are conspicuously absent from the majority of the islands of the Pacific, including the large islands of New Zealand, Fiji and New Caledonia.
Lane, 1953 (Neotropical Region); Mattingly & Knight, 1956 (Arabia); Cova-García, 1961 (Venezuela); Belkin, 1962 (taxonomy, South Pacific); DuBose & Curtin, 1965 (keys, Mediterranean area); Grjebine, 1966 (Madagascar); Gillies & de Meillon, 1968 (Afrotropical Region); Reid, 1968 (Malaysia, Borneo); Cagampang-Ramos & Darsie, 1970 (keys, Philippine Islands); Gutsevich et al., 1974 (former USSR); Klein, 1977 (Cambodia); Tanaka et al., 1979 (Japan); Harrison, 1980 (Myzomyia Series, Thailand); Lu & Li, 1982 (China); Rao, 1984 (India); Lee et al., 1987 (Australasian Region); Gillies & Coetzee, 1987 (Afrotropical Region); Das et al., 1990 (keys, India); Darsie & Pradhan, 1990 (Nepal); 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); Beebe et al., 2015 (Punctulatus Group, evolution, distribution); Foster et al., 2017 (phylogenetic relationships).
aconitus Dönitz, 1902
ainshamsi Gad, Harbach & Harrison, 2006
amharicus Hunt, Wilkerson & Coetzee, 2013 (in Coetzee et al., 2013)
amictus Edwards, 1921
annularis van der Wulp, 1884
annulatus Haga, 1930
annulipes Walker, 1856
apoci Marsh, 1933
arabiensis Patton, 1905
ardensis (Theobald, 1905)
argenteolobatus (Gough, 1910)
aruni Sobti, 1968
aurirostris (Watson, 1910)
austenii (Theobald, 1905)
azaniae Bailly-Choumara, 1960
azevedoi Ribeiro, 1969
baimaii Sallum & Peyton, 2005 (in Sallum et al., 2005)
baisasi Colless, 1957
balabacensis Baisas, 1936
barberellus Evans, 1932
basilewskyi Leleup, 1957
berghei Vincke & Leleup, 1949
bervoetsi D'Haenens, 1961
brohieri Edwards, 1929
broussesi Edwards, 1929
brucei Service, 1960
brumpti Hamon & Rickenbach, 1955
brunnipes (Theobald, 1910)
buxtoni Service, 1958
bwambae White, 1985
cameroni de Meillon & Evans, 1935
carnevalei Brunhes, Le Goff & Geoffroy, 1999
caroni Adam, 1961
carteri Evans & de Meillon, 1933
christyi (Newstead & Carter, 1911)
cinctus (Newstead & Carter, 1910)
cinereus Theobald, 1901
clowi Rozeboom & Knight, 1946
coluzzii Coetzee & Wilkerson, 2013 (in Coetzee et al., 2013)
comorensis Brunhes, Le Goff & Geoffroy, 1997
confusus Evans & Leeson, 1935
cracens Sallum & Peyton, 2005 (in Sallum et al., 2005)
cristatus King & Baisas, 1936
cristipalpis Service, 1977
culicifacies Giles, 1901
cydippis de Meillon, 1931
dancalicus Corradetti, 1939
daudi Coluzzi, 1958
deemingi Service, 1970
demeilloni Evans, 1933
dirus Peyton & Harrison, 1979
dispar Rattanarithikul & Harbach, 1991
distinctus (Newstead & Carter, 1911)
domicolus Edwards, 1916
dravidicus Christophers, 1924
dthali Patton, 1905
dualaensis Brunhes, Le Goff & Geoffroy, 1999
dureni Edwards, 1938 (in Evans, 1938)
elegans (James, 1903) (in Theobald, 1903)
eouzani Brunhes, Le Goff & Boussès, 2003
epiroticus Linton & Harbach, 2005 (in Linton et al., 2005)
erepens Gillies, 1958
erythraeus Corradetti, 1939
ethiopicus Gillies & Coetzee, 1987
faini Leleup, 1952
farauti Laveran, 1902
filipinae Manalang, 1930
flavicosta Edwards, 1911
flavirostris (Ludlow, 1914)
fluviatilis James, 1902
fontinalis Gillies & de Meillon, 1968
freetownensis Evans, 1925
funestus Giles, 1900
fuscivenosus Leeson, 1930
gabonensis Rahola, Makanga & Paupy, 2014 (in Rahola et al., 2014)
gambiae Giles, 1902
garnhami Edwards, 1930
gibbinsi Evans, 1935
grassei Grjebine, 1953
greeni Rattanarithikul & Harbach, 1991
grenieri Grjebine, 1964
griveaudi Grjebine, 1961
hackeri Edwards, 1921
hamoni Adam, 1962
hancocki Edwards, 1929
hargreavesi Evans, 1927
harperi Evans, 1936
harrisoni Harbach & Manguin, 2007 (in Harbach et al., 2007)
hervyi Brunhes, Le Goff & Geoffroy, 1999
hilli Woodhill & Lee, 1944
hinesorum Schmidt, 2001 (in Schmidt et al., 2001)
hispaniola Theobald, 1903
hughi Lambert & Coetzee, 1982
incognitus Brug, 1931
indefinitus (Ludlow, 1904)
introlatus Colless, 1957
irenicus Schmidt, 2003 (in Schmidt et al., 2003)
jamesii Theobald, 1901
jebudensis Froud, 1944
jeyporiensis James, 1902
karwari (James, 1903) (in Theobald, 1903)
keniensis Evans, 1931
kingi Christophers, 1923
kochi Dönitz, 1901
kokhani Vythilingam, Jeffery & Harbach, 2007 (in Vythilingam et al., 2007)
kolambuganensis Baisas, 1932
koliensis Owen, 1945
kosiensis Coetzee, Segerman & Hunt, 1987
lacani Grjebine, 1953
latens Sallum & Peyton, 2005 (in Sallum et al., 2005)
leesoni Evans, 1931
letabensis Lambert & Coetzee, 1982
leucosphyrus Dönitz, 1901
limosus King, 1932
listeri de Meillon, 1931
litoralis King, 1932
lloreti Gil Collado, 1936
longipalpis (Theobald, 1903)
longirostris Brug, 1928
lounibosi Gillies & Coetzee, 1987
lovettae Evans, 1934
ludlowae (Theobald, 1903)
lungae Belkin & Schlosser, 1944
macarthuri Colless, 1956
machardyi Edwards, 1930
macmahoni Evans, 1936
maculatus Theobald, 1901
maculipalpis Giles, 1902
majidi Young & Majid, 1928
maliensis Bailly-Choumara & Adam, 1959
mangyanus (Banks, 1906)
marshallii (Theobald, 1903)
mascarensis de Meillon, 1947
melas (Theobald, 1903)
meraukensis Venhuis, 1932
merus Dönitz, 1902
millecampsi Lips, 1960
milloti Grjebine & Lacan, 1953 (in Grjebine, 1953)
minimus Theobald, 1901
mirans Sallum & Peyton, 2005 (in Sallum et al., 2005)
moghulensis Christophers, 1924
mortiauxi Edwards, 1938 (in Evans, 1938)
moucheti Evans, 1925
subspecies moucheti Evans, 1925
subspecies nigeriensis Evans, 1931
mousinhoi de Meillon & de Carvalho Pereira, 1940
multicinctus Edwards, 1930
multicolor Cambouliu, 1902
murphyi Gillies & de Meillon, 1968
natalensis (Hill & Haydon, 1907)
nataliae Belkin, 1945
nemophilous Peyton & Ramalingam, 1988
nigeriensis Evans, 1931
nili (Theobald, 1904)
nivipes (Theobald, 1903)
njombiensis Peters, 1955
notanandai Rattanarithikul & Green, 1987
notleyi van Someren, 1949
novaguinensis Venhuis, 1933
oreios Bangs & Harbach, 2014 (in Bangs et al., 2014)
ovengensis Awono-Ambene, Simard, Antonio-Nkonkjio & Fontenille, 2004
pallidus Theobald, 1901
paltrinierii Shidrawi & Gillies, 1988
pampanai Büttiker & Beales, 1959
parangensis (Ludlow, 1914)
parensis Gillies, 1962
pattoni Christophers, 1926
pauliani Grjebine, 1953
pharoensis Theobald, 1901
philippinensis Ludlow, 1902
pretoriensis (Theobald, 1903)
pseudojamesi Strickland & Chowdhury, 1927
pseudosundaicus Tyagi, Hiriyan, Tewari, Ayanar, Samuel, Arunachalam, Paramasivan, Krishnamoorthy, Dhananjeyan, Leo & Rajendran, 2009
pseudowillmori (Theobald, 1910)
pujutensis Colless, 1948
pulcherrimus Theobald, 1902
punctulatus Dönitz, 1901
quadriannulatus (Theobald, 1911)
radama de Meillon, 1943
rageaui Mattingly & Adam, 1954
rampae Harbach & Somboon, 2011 (in Somboon et al., 2011)
ranci Grjebine, 1953
recens Sallum & Peyton, 2005 (in Sallum et al., 2005)
rennellensis Taylor & Maffi, 1991
rhodesiensis Theobald, 1901
riparis King & Baisas, 1936
rivulorum Leeson, 1935
rodhaini Leleup & Lips, 1950
roubaudi Grjebine, 1953
ruarinus Edwards, 1940
rufipes (Gough, 1910)
rupicolus Lewis, 1937
salbaii Maffi & Coluzzi, 1958
saungi Colless, 1955
sawadwongporni Rattanarithikul & Green, 1987
scanloni Sallum & Peyton, 2005 (in Sallum et al., 2005)
schueffneri Stanton, 1915
schwetzi Evans, 1934
seretsei Abdulla-Khan, Coetzee & Hunt, 1998
sergentii (Theobald, 1907)
seydeli Edwards, 1929
smithii Theobald, 1905
solomonis Belkin, Knight & Rozeboom, 1945
somalicus Rivola & Holstein, 1957
splendidus Koidzumi, 1920
squamosus Theobald, 1901
stephensi Liston, 1901
stookesi Colless, 1955
subpictus Grassi, 1899 (in Grassi et al., 1899)
sulawesi Waktoedi Koesoemawinangoen, 1954
sundaicus (Rodenwaldt, 1925)
superpictus Grassi, 1899
swahilicus Gillies, 1964
takasagoensis Morishita, 1946
tchekedii de Meillon & Leeson, 1940
tessellatus Theobald, 1901
theileri Edwards, 1912
theobaldi Giles, 1901
torakala Stoker & Waktoedi Koesoemawinangoen, 1949
torresiensis Schmidt, 2001 (in Schmidt et al., 2001)
turkhudi Liston, 1901
ugandae Evans, 1934
ungujae White, 1975
vagus Dönitz, 1902
vaneedeni Gillies & Coetzee, 1987
vanhoofi Wanson & Lebied, 1945
varuna Iyengar, 1924
vernus Gillies & de Meillon, 1968
vinckei de Meillon, 1942
walravensi Edwards, 1930
watsonii (Leicester, 1908)
wellcomei Theobald, 1904
willmori (James, 1903) (in Theobald, 1903)
wilsoni Evans, 1934
yaeyamaensis Somboon & Harbach, 2010 (in Somboon et al., 2010)
