Belongs within: Culicidae.
Anopheles is a genus of mosquitoes, distinguishable from others by their elongate palps. Many species in the genus are significant disease vectors, most notably of the malaria causative organism Plasmodium, and of various filarial nematodes. Examples include the Anopheles gambiae complex of Africa. Females of Anopheles generally lay their eggs in permanent bodies of water; the elongate eggs are laid singly, and have membranous folds on either side that act as floats (Liehne 1991). Several subgenera have been recognised in Anopheles; members of the subgenus Anopheles have an entirely dark hind tarsus whereas the hind tarsus is partially white in A. (Nyssorhynchus) albimanus (Stone 1981). Members of the subgenus Cellia have numerous spots of white scales on the wings as opposed to the more uniformly dark-scaled Anopheles sensu stricto (Liehne 1991).
The feared mosquito
Published 29 August 2021
It’s one of those standard pub-quiz “trick” questions. What animal kills the most people? The hope is that contestants will nominate the ‘obvious’—snakes, sharks, bears, whatever—before being blind-sided by the revelation that mosquitoes kill over a million people. They don’t kill them directly, of course; their victims die from the diseases they spread*. The statistic also glosses over the point that there are many hundreds of species of mosquito that vary significantly in the nature and severity of their role as disease vectors. Nevertheless, for this post I’m considering the group that includes some of the most notorious vectors: the genus Anopheles.
*For the record, if the question was confined to active killings, the most dangerous animal to humans is other humans. Dogs come a distant second.
Anopheles is one of the most divergent genera of mosquitoes, being placed in a distinct subfamily Anophelinae (along with a couple of small related genera) from the bulk of mosquitoes in the subfamily Culicinae. Adult Anopheles can be readily distinguished from culicine mosquitoes by their palps which are about as long as the proboscis (in other mosquitoes, the palps are distinctly shorter). Larvae of Anopheles lack the long respiratory siphons at the end of the abdomen found in other mosquito larvae so they rest parallel with the water surface rather than hanging below it. The genus is found around the world; over 450 named species are currently known (Harbach 2013) with many more waiting to be described. The genus is currently divided between seven subgenera though one of the largest of these, the cosmopolitan subgenus Anopheles, is not monophyletic. The remaining subgenera are better supported with the largest of these, Cellia, being found in the Old World. Between them, the subgenera Anopheles and Cellia account for over 400 of the known Anopheles species. The remaining small subgenera are mostly Neotropical with a single Oriental species being awarded its own subgenus.
Anopheles is of most concern to humans, of course, for its role as a disease vector. As with other mosquitoes, the transmission of disease is done entirely by females taking blood meals to provide nutrients for their developing eggs. Males are not blood feeders, instead feeding entirely on sugar sources such as nectar (females also feed on nectar for their own nutrition). The main disease spread by Anopheles is malaria, but they may also spread malaises such as filariasis and arboviruses (Krzywinski & Besansky 2003). As noted above, species may vary significantly in their importance as disease vectors, even between quite closely related taxa. Many historically recognised vector “species” have proved, on close inspection, to represent species complexes of which some may be vectors and others not. For instance, one of the most important transmitters of malaria, the African A. gambiae, has been divided between at least eight different species (Coetzee et al. 2013). Misidentification of vectors can be a significant issue. For instance, mosquito control regimes in central Vietnam during the 1990s focused on two species, A. dirus and A. minimus, that were each active at different times of year. However, Van Bortel et al. (2001) found that A. minimus was in fact very rare in this area, with specimens previously thought to be A. minimus proving to be another species, A. varuna. Anopheles varuna is not a significant malaria vector, feeding almost entirely on animals such as cattle rather than on humans. Large amounts of resources would have been wasted trying to control a mosquito that was of little concern. What is more, the fact that malaria was not being transmitted by A. minimus raises the possibility that it was being spread by yet another species, one that had managed to escape attention. Remember, kids: bad taxonomy kills.
Systematics of Anopheles
Characters (from Liehne 1991): Clypeus longer than broad, rounded anteriorly. Palps of female as long as proboscis. Scutellum evenly rounded, without distinct lobes. Abdomen usually without scales or with a few loosely applied scales. Larva with head longer than broad, rotating freely; seta 2-C close to apical margin of head; setae 5-C to 7-C pinnate and aligned in distinct row across middle of head; mentum generally long and narrow; abdomen with dorsal palmate tufts on all segments; siphon absent; pecten on posterior margin of distinct triangular chitinous plate; setae 2-X and 4-X irregularly branched.
<==Anopheles |--A. (Anopheles)L91 | |--A. (A.) atratipes Skuse 1889L91 | |--A. (A.) bancroftii Giles 1902L91 | |--A. (A.) chiriquiensis Komp 1936SK56 | |--A. (A.) earlei Vargas 1943S81, SK56 | |--A. (A.) fluminensis Root 1927SK56 | |--A. (A.) maculipennisDW84 [=Culex (A.) maculipennisG20] | |--A. (A.) powelli Lee 1944L91 | |--A. (A.) punctipennisS81 | |--A. (A.) samarensis Rozeboom 1951SK56 | |--A. (A.) saperoi Bohart & Ingram 1946SK56 | `--A. (A.) shannoni Davis 1931SK56 |--A. (Cellia)L91 | | i. s.: A. (C.) amharicus Coetzee, Hunt et al. 2013CH13 | | A. (C.) amictus Edwards 1921L91 | | A. (C.) annulipes Walker 1856 (see below for synonymy)L91 | | A. (C.) coluzzii Coetzee, Hunt et al. 2013CH13 | | A. (C.) farauti Laveran 1902 [incl. A. moluccensis Swellengrebel & Swellengrebel de Graf 1919]L91 | | A. (C.) hilli Woodhill & Lee 1944L91 [=A. amictus hilliW91] | | A. (C.) meraukensis Venhuis 1932L91 | | A. (C.) novaguinensis Venhuis 1933L91 | | A. (C.) punctulatusL91 | | A. (C.) quadriannulatusCH13 | |--ser. NeocelliaNNK72 | | |--A. (C.) maculatusNNK72 | | |--A. (C.) pulcherrimusNNK72 | | `--A. (C.) stephensiNNK72 | |--A. (C.) (ser. Neomyzomyia) tessellatusNNK72 | `--ser. PyretophorusNNK72 | |--A. (C.) gambiae Giles 1902NNK72, CH13 [incl. A. costalis Loew 1866 (n. d.)CH13, A. gracilis Dönitz 1902CH13] | `--A. (C.) subpictusNNK72 [incl. A. rossiA71; incl. Myzomyia rossi var. indefinita Ludlow 1904SK56] |--A. (Dendropaedium) bellator Dyar & Knab 1906 [incl. A. (D.) bellator race bromelicola Dyar & Knab 1925]SK56 |--A. (Kerteszia)SK56 | |--A. (K.) anoplus Komp 1937SK56 | |--A. (K.) bambusicolus Komp 1937SK56 | `--A. (K.) homunculus Komp 1937SK56 |--A. (Myzomyia)SK56 | |--A. (M.) clowi Rozeboom & Knight 1946SK56 | |--A. (M.) cristatus King & Baisas 1936SK56 | |--A. (M.) leucosphyrus [incl. A. (M.) leucosphyrus var. riparis King & Baisas 1936]SK56 | |--A. (M.) lungae Belkin & Schlosser 1944SK56 | |--A. (M.) nataliae Belkin 1945SK56 | `--A. (M.) solomonis Belkin, Knight & Rozeboom 1945SK56 |--A. (Nyssorhynchus)S81 | |--A. (N.) albimanusS81 | |--A. (N.) anomalophyllus Komp 1936SK56 | |--A. (N.) darlingi Root 1926SK56 | |--A. (N.) davisi Paterson & Shannon 1927SK56 | |--A. (N.) dunhami Causey 1945SK56 | |--A. (N.) emilianus Komp 1941SK56 | |--A. (N.) goeldii Rozeboom & Gabaldon 1941SK56 | |--A. (N.) guarani Shannon 1928SK56 | |--A. (N.) sawyeri Causey, Deane et al. 1943SK56 | `--A. (N.) strodei Root 1926SK56 `--A. (Stethomyia)SK56 |--A. (S.) acanthotorynus Komp 1937SK56 `--A. (S.) niveopalpis Ludlow 1919SK56 Anopheles incertae sedis: A. apicimacula Dyar & Knab 1906SK56 A. aquasalisC09 A. arabiensis [incl. A. quadriannulatus davidsoni]CH13 A. argenteolobatusCH13 A. argyritarsisC09 A. atroparvusBP00 [=A. maculipennis atroparvusA71] A. atropos Dyar & Knab 1906SK56 A. barberi Caquillett 1903SK56 A. bwambaeCH13 A. clavigerKK91 A. comorensis Brunhes, Le Goff & Geoffroy 1997CH13 A. crucians [incl. A. crucians var. bradleyi King 1939, A. crucians var. georgianus King 1939]SK56 A. cruziiEN20 A. culicifaciesD51 A. cydippisCH13 A. dirusB08 A. eiseni Coquillett 1902SK56 ‘Cellia’ flava Ludlow 1908SK56 ‘Myzomyia’ flavirostris Ludlow 1914SK56 A. fluviatilisRD77 A. formosus Ludlow 1909SK56 A. funestusRD77 A. gorgasi Dyar & Knab 1907SK56 A. hinesorumRK04 ‘Culex’ hyemalis Fitch 1847SK56 A. koliensis Owen 1945SK56 A. labranchiaeD51 A. lewisi Ludlow 1920SK56 ‘Stethomyia lewisi’ Shannon 1931 non Anopheles lewisi Ludlow 1920SK56 A. lindesayi [incl. A. lindesayi var. benguetensis King 1931]SK56 ‘Chagasia’ lineata Ludlow 1908SK56 A. litoralis King 1932SK56 A. lutziRD77 A. malefactor Dyar & Knab 1907SK56 A. melanoonBP00 A. melasCH13 A. merus [incl. A. gambiae litoralis Halcrow 1957 non A. litoralis King 1932, A. tangensis Kuhlow 1962]CH13 A. messaeBP00 A. minimusRD77 |--A. m. minimusB08 `--A. m. varunaB08 A. mouchetiKSM06 A. multicolorRD77 A. neivai Howard, Dyar & Knab 1917SK56 A. niliKSM06 A. nimbus [incl. A. nimbus var. kompi Edwards 1930]SK56 A. occidentalis Dyar & Knab 1906SK56 A. oiketorakras Osorno-Mesa 1947SK56 ‘Stethomyia’ pallida Ludlow 1905SK56 ‘Myzomyia’ parangensis Ludlow 1914SK56 A. perplexens Ludlow 1907SK56 A. pharoensisKSM06 A. philippinensis Ludlow 1902SK56 A. plumbeusKK91 A. pseudobarbirostris Ludlow 1902SK56 A. pseudopunctipennisC09 A. punctimacula Dyar & Knab 1906SK56 A. quadrimaculatusHF01 A. sacharoviD51 A. sanctielii Senevet & Abonnenc 1938S69 A. selengensis Ludlow 1920SK56 A. strigimacula Dyar & Knab 1906SK56 A. subalpinusD51 A. sundaicusRD77 ‘Myzomyia’ thorntonii Ludlow 1904SK56 A. torresiensisRK04 A. vagus [incl. A. vagus var. limosus King 1932]SK56 A. vestitipennis Dyar & Knab 1906SK56
Anopheles (Cellia) annulipes Walker 1856 [incl. A. derricki Taylor 1943, A. mastersi Skuse 1889, A. musivus Skuse 1889, A. perplexus Taylor 1943, A. persimilis Taylor 1943]L91
*Type species of generic name indicated
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