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Type: Article
Published: 2022-05-04
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Literature review of the systematics, biology and role in malaria transmission of species in the Afrotropical Anopheles subgenus Anopheles (Diptera: Culicidae)

Wits Research Institute for Malaria, School of Pathology, Faculty of Health Sciences, University of the Witwatersrand, 2001 Johannesburg, South Africa. Centre for Emerging Zoonotic and Parasitic Diseases, National Institute for Communicable Diseases, Johannesburg, South Africa.
Diptera caliginosus concolor coustani crypticus fuscicolor namibiensis obscurus paludis symesi tenebrosus ziemanni

Abstract

It has been 34 years since the last update of the subgenus Anopheles Meigen, 1818 in the Afrotropical Region. Eleven species occur in the region, and of these, eight occur only on the African continent, two in both Africa and Madagascar, and one exclusively in Madagascar. Three species are implicated in the transmission of malarial parasites in localised areas: An. coustani Laveran, 1900 (Cameroon, Central African Republic, Democratic Republic of the Congo, Kenya, Madagascar, Tanzania and Zambia), An. paludis Theobald, 1900 (Cameroon, Democratic Republic of the Congo) and An. ziemanni Grünberg, 1902 (Cameroon, Chad, Ethiopia and Rwanda). Several arboviruses have been isolated from An. coustani in Kenya, Madagascar and Senegal. Recent molecular studies indicate possible undescribed species within An. coustani that could be resolved with integrated molecular, morphological and cytogenetic methods.

 

References

  1. Abdullah, M.A.R. & Merdan, A.I. (1995) Distribution and ecology of the mosquito fauna in the southwestern Saudi Arabia. Journal of the Egyptian Society of Parasitology, 25 (3), 815–837.
    Abduselam, N., Zeynudin, A., Berens-Riha, N., Seyoum, D., Pritsch, M., Tibebu, H., Eba, K., Hoelscher, M., Wieser, A. & Yewhalaw, D. (2016) Similar trends of susceptibility in Anopheles arabiensis and Anopheles pharoensis to Plasmodium vivax infection in Ethiopia. Parasites & Vectors, 9, 552. https://doi.org/10.1186/s13071-016-1839-0
    Amvongo-Adjia, N., Wirsiy, E.L., Riveron, J.M., Chounna Ndongmo, W.P., Enyong, P.A., Njiokou, F., Wondji, C.S. & Wanji, S. (2018) Bionomics and vectorial role of anophelines in wetlands along the volcanic chain of Cameroon. Parasites & Vectors, 11, 471. https://doi.org/10.1186/s13071-018-3041-z
    Antonio-Nkondjio, C., Kerah, C.H., Simard, F., Awono-Ambene, P., Chouaibou, M., Tchuinkam, T. & Fontenille, D. (2006) Complexity of the malaria vectorial system in Cameroon: contribution of secondary vectors to malaria transmission. Journal of Medical Entomology, 43 (6), 1215–1221. https://doi.org/10.1093/jmedent/43.6.1215
    Aranda, C., Aponte, J.J., Saute, F., Casimiro, S., Pinto, J., Sousa, C., Rosario, V.D., Petrarca, V., Dgedge, M. & Alonso, P. (2005) Entomological characteristics of malaria transmission in Manhica, a rural area in southern Mozambique. Journal of Medical Entomology, 42 (2), 180–186. https://doi.org/10.1093/jmedent/42.2.180
    Ayala, D., Costantini, C., Ose, K., Kamdem, G.C., Antonio-Nkondjio, C., Agbor, J-P., Awono-Ambene, P., Fontenille, D. & Simard, F. (2009) Habitat suitability and ecological niche profile of major malaria vectors in Cameroon. Malaria Journal, 8, 307. https://doi.org/10.1186/1475-2875-8-307
    Bakker, J.W., Loy, D.E., Takken, W., Hahn, B.H. & Verhulst, N.O. (2020) Attraction of mosquitoes to primate odours and implications for zoonotic Plasmodium transmission. Medical & Veterinary Entomology, 34 (1), 17–26. https://doi.org/10.1111/mve.12402
    Bamou, R., Mbakop, L.R., Kopya, E., Ndo, C., Awono-Ambene, P., Tchuinkam, T., Rono, M.K., Mwangangi, J. & Antonio-Nkondjio, C. (2018) Changes in malaria vector bionomics and transmission patterns in the equatorial forest region of Cameroon between 2000 and 2017. Parasites & Vectors, 11, 464. https://doi.org/10.1186/s13071-018-3049-4
    Bigoga, J.D., Nanfack, F.M., Awono-Ambene, P.H., Patchoke, S., Atangana, J., Otia, V.S., Fondjo, E., Moyou, R.S. & Leke, R.G. (2012) Seasonal prevalence of malaria vectors and entomological inoculation rates in the rubber cultivated area of Niete, South Region of Cameroon. Parasites & Vectors, 5, 197. https://doi.org/10.1186/1756-3305-5-197
    Brunhes, J., Le Goff, G. & Geoffroy, B. (1997) Anophèles Afro-tropicaux. I.–Descriptions d’espèces nouvelles et changements de statuts taxonomiques (Diptera: Culicidae). Annales de la Société Entomologique de France, 33 (2), 173–183.
    Brunhes, J., Le Goff, G. & Geoffroy, B. (1998) Anophèles afrotropicaux. II—Mises au point sur les espèces de la sous-région malgache (Diptera, Culicidae). Bulletin de la Société Entomologique de France, 103 (2), 139–152. https://doi.org/10.3406/bsef.1998.17406
    Charlwood, J.D., Macia, G.A., Manhaca, M., Sousa, B., Cuamba, N. & Braganca, M. (2013) Population dynamics and spatial structure of human-biting mosquitoes, inside and outside of houses, in the Chockwe irrigation scheme, southern Mozambique. Geospatial Health, 7 (2), 309–320. https://doi.org/10.4081/gh.2013.89
    Christophers, S.R. (1924) Provisional list and reference catalogue of the Anophelini. Indian Medical Research Memoirs, 3, 1–105.
    Ciubotariu, I.I., Jones, C.M., Kobayashi, T., Bobanga, T., Muleba, M., Pringle, J.C., Stevenson, J.C., Carpi, G. & Norris, D.E. (2020) Genetic diversity of Anopheles coustani (Diptera: Culicidae) in malaria transmission foci in southern and central Africa. Journal of Medical Entomology, 57 (6), 1782–1792. https://doi.org/10.1093/jme/tjaa132
    Coene, J. (1993) Malaria in urban and rural Kinshasa: the entomological input. Medical and Veterinary Entomology, 7, 127–137. https://doi.org/10.1111/j.1365-2915.1993.tb00665.x
    Coetzee, M. (1983) Chromosomal and cross-mating evidence for two species within Anopheles (A.) coustani (Diptera: Culicidae). Systematic Entomology, 8 (2), 137–141. https://doi.org/10.1111/j.1365-3113.1983.tb00473.x
    Coetzee, M. (1984) A new species of Anopheles (Anopheles) from Namibia (Diptera: Culicidae). Systematic Entomology, 9 (1), 1–8. https://doi.org/10.1111/j.1365-3113.1984.tb00498.x
    Coetzee, M. (1995) Anopheles crypticus, new species from South Africa is distinguished from Anopheles coustani (Diptera: Culicidae). Mosquito Systematics, 26 (3), 125–131. [for 1994].
    Coetzee, M. (2020) Key to the females of Afrotropical Anopheles mosquitoes (Diptera: Culicidae). Malaria Journal, 19, 70. https://doi.org/10.1186/s12936-020-3144-9
    Coetzee, M. & Le Sueur, D. (1988) Effects of salinity on the larvae of some Afrotropical anopheline mosquitoes. Medical and Veterinary Entomology, 2 (4), 385–390. https://doi.org/10.1111/j.1365-2915.1988.tb00212.x
    de Grandpré, A.D. & de Charmoy, D. d’E. (1901) Les moustiques: Anatomie et biologie. Contribution à l’étude des culicidées et principalement des genres Culex et Anopheles de leur rôle dans la propagation de la malaria et de la filariose et des moyens de s’en préserver. Ile Maurice, Port Louis, iv + 59 pp., 5pls. [for 1900]
    de Meillon, B. (1936) Entomological studies. Studies on insects of medical importance in South Africa (Part III). Publications of the South African Institute for Medical Research, 7 (38), 125–215.
    de Meillon, B. (1943) New records and new species of Nematocera (Diptera) from the Ethiopian region. Journal of the Entomological Society of Southern Africa, 6 (1), 90–113.
    de Meillon, B. (1947) The Anophelini of the Ethiopian Geographical Region. Publications of the South African Institute for Medical Research, 10 (49), 1–272
    Diallo, D., Sall, A.A., Diagne, C.T., Faye, Oum., Faye, Ous., Ba, Y., Hanley, K.A., Buenemann, M., Weaver, S.C. & Diallo, M. (2014) Zika virus emergence in mosquitoes in southeastern Senegal, 2011. Plos One, 9 (10), e109442. https://doi.org/10.1371/journal.pone.0109442
    Dönitz, W. (1902) Beiträge zur Kenntniss der Anopheles. Zeitschrift für Hygiene und Infektionskrankheiten, 41, 15–88, 2 pls.  https://doi.org/10.1017/S0007485300019994
    Edwards, F.W. (1928) Mosquito notes.—VII. Bulletin of Entomological Research, 18 (3), 267–284.
    Evans, A.M. (1932) Notes on African mosquitoes. Annals of Tropical Medicine and Parasitology, 26 (1), 85–108.
    Evans, A.M. (1938) Mosquitoes of the Ethiopian Region. II.—Anophelini adults and early stages. British Museum (Natural History), London, x, 404 pp.
    Finney, M., McKenzie, B.A., Rabaovola, B., Sutcliffe, A., Dotson, E. & Zohdy, S. (2021) Widespread zoophagy and detection of Plasmodium spp. in Anopheles mosquitoes in southeastern Madagascar. Malaria Journal, 20, 25. https://doi.org/10.1186/s12936-020-03539-4
    Getachew, D., Balkew, M. & Tekie, H. (2020) Anopheles larval species composition and characterization of breeding habitats in two localities in the Ghibe River Basin, southwestern Ethiopia. Malaria Journal, 19, 65. https://doi.org/10.1186/s12936-020-3145-8
    Giaquinto-Mira, M. (1950) Notes on the geographical distribution and biology of “Anophelinae” and “Culicinae” in Etiopia [sic]. Rivista di Malariologia, 29 (5), 281–313.
    Gillies, M.T. & Coetzee, M. (1987) A supplement to the Anophelinae of Africa south of the Sahara (Afrotropical Region). Publications of the South African Institute for Medical Research, 55, 1–143.
    Gillies, M.T. & de Meillon, B. (1968) The Anophelinae of Africa south of the Sahara (Ethiopian Zoogeographical Region). Publications of the South African Institute for Medical Research, 54, 1–343.
    Glick, J.I. (1992) Illustrated key to the female Anopheles of southwestern Asia and Egypt (Diptera: Culicidae). Mosquito Systematics, 24 (2), 125–153.
    Goupeyou-Youmsi, J., Rakotondranaivo, T., Puchot, N., Peterson, I., Girod, R., Vigan-Womas, I., Paul, R., Ndiath, M.O. & Bourgouin, C. (2020) Differential contribution of Anopheles coustani and Anopheles arabiensis to the transmission of Plasmodium falciparum and Plasmodium vivax in two neighbouring villages of Madagascar. Parasites & Vectors, 13, 430. https://doi.org/10.1186/s13071-020-04282-0
    Grjebine, A. (1954) Observations sur les Nematoceres vulnerants de Madagascar. Regions de Majunga et de la Mandraka. Memoires de l’Institut Scientifique de Madagascar, 4, 443–503.
    Grjebine, A. (1966) Faune de Madagascar. XXII. Insectes Diptéres Culicidae Anophelinae. Centre National de la Recherche Scientifique, Office de la Recherche Scientifique et Technique Outre-Mer, Paris, 487 pp., table, map, 8 pls.
    Grünberg, K. (1902) Ein neuer Anopheles aus Westafrika, Anopheles ziemanni nov. spec. Zoologischer Anzeiger, 25 (677), 550–551.
    Grünberg, K. (1905) Zur Kenntnis der Culicidenfauna von Kamerun und Togo. Zoologischer Anzeiger, 29 (12), 377–390.
    Habtewold, T., Prior A., Torr, S.J. & Gibson, G. (2004) Could insecticide-treated cattle reduce Afrotropical malaria transmission? Effects of deltamethrin-treated Zebu on Anopheles arabiensis behaviour and survival in Ethiopia. Medical and Veterinary Entomology, 18 (4), 408–417. https://doi.org/10.1111/j.0269-283X.2004.00525.x
    Hamon, J., Adam, J.P. & Grjebine, A. (1956) Observations sur la répartition et le comportement des anophèles de l’Afrique-Equatoriale Francaise, du Cameroun et de l’Afrique Occidentale. Bulletin of the World Health Organization, 15 (3–5), 549–591.
    Harbach, R.E. (1994) Review of the internal classification of the genus Anopheles (Diptera: Culicidae): the foundation for comparative systematics and phylogenetic research. Bulletin of Entomological Research, 84 (3), 331–342. https://doi.org/10.1017/S0007485300032454
    Harbach, R.E. (2004) The classification of genus Anopheles (Diptera: Culicidae): a working hypothesis of phylogenetic relationships. Bulletin of Entomological Research, 94 (6), 537–553. https://doi.org/10.1079/BER2004321
    Harbach, R.E. (2018) An Anopheles by any other name …? Journal of Medical Entomology, 55 (5), 1069–1070. https://doi.org/10.1093/jme/tjy108
    Harbach, R.E. & Kitching, I.J. (2016) The phylogeny of Anophelinae revisited: inferences about the origin and classifications of Anopheles (Diptera: Culicidae). Zoologica Scripta, 45 (1), 34–47. https://doi.org/10.1111/zsc.12137
    Harbach, R.E. & Knight, K.L. (1980) Taxonomists’ Glossary of Mosquito Anatomy. Plexus Publishing, Inc., New Jersey, Medford, xi + 413 pp.
    Harbach, R.E. & Knight, K.L. (1982) Corrections and additions to Taxonomists’ Glossary of Mosquito Anatomy. Mosquito Systematics, 13 (2), 201–217. [for 1981]
    Irish, S.R., Kyalo, D., Snow, R.W. & Coetzee, M. (2020) Updated list of Anopheles species (Diptera: Culicidae) by country in the Afrotropical Region and associated islands. Zootaxa, 4747 (3), 401–449. https://doi.org/10.11646/zootaxa.4747.3.1
    Kamau, L., Mulaya, N. & Vulule, J.M. (2006) Evaluation of potential role of Anopheles ziemanni in malaria transmission in western Kenya. Journal of Medical Entomology, 43 (4), 774–776. https://doi.org/10.1093/jmedent/43.4.774
    Karch, S., Asidi, N., Manzambi, Z.M. & Salaun, J.J. (1992) La faune anophélienne et la transmission du paludisme humain a Kinshasa (Zaïre). Bulletin de la Société de Pathologie Exotique, 85 (4), 304–309.
    Karch, S. & Mouchet, J. (1992) Anopheles paludis: vecteur important du paludisme au Zaïre. Bulletin de la Societe de Pathologie Exotique, 85 (5), 388–389.
    Kerah-Hinzoumbe, C., Peka, M., Antonio-Nkondjio, C., Donan-Gouni, I., Awono-Ambene, P., Same-Ekobo, A. & Simard, F. (2009) Malaria vectors and transmission dynamics in Goulmoun, a rural city in south-western Chad. BMC Infectious Diseases, 9, 71. https://doi.org/10.1186/1471-2334-9-71
    Knight, K.L. & Stone, A. (1977) A catalog of the mosquitoes of the world. The Thomas Say Foundation. Vol.VI. 2nd Edition. Entomological Society of America, College Park, Maryland, xi + 611 pp.
    Kyalo, D., Amratia, P., Mundia, C.W., Mbogo, C.M., Coetzee, M. & Snow, R.W. (2017) A geo-coded inventory of anophelines in the Afrotropical Region south of the Sahara: 1898−2016. Wellcome Open Research, 2, 57. https://doi.org/10.12688/wellcomeopenres.12187.1
    Laveran, [A.] (1900) Sur un Anopheles provenant de Madagascar. Comptes Rendus Hebdomadaires des Seìances et Meìmoires de la Socieìteì de Biologie, 52, 109–110.
    Lobo, N.F., St. Laurent, B., Sikaala, C.H., Hamainza, B., Chanda, J., Chinula, D., Krishnankutty, S.M., Mueller, J.D., Deason, N.A., Hoang, Q.T., Boldt, H.L., Thumloup, J., Stevenson, J., Seyoum, A. & Collins, F.H. (2015) Unexpected diversity of Anopheles species in eastern Zambia: implications for evaluating vector behavior and interventions using molecular tools. Scientific Reports, 5, 17952.
    https://doi.org/10.1038/srep17952
    Makanga, B., Costantini, C., Rahola, N., Yangari P., Rougeron, V., Ayala, D., Prugnolle, F. & Paupy, C. (2017) “Show me which parasites you carry and I will tell you what you eat”, or how to infer the trophic behaviour of hematophagous arthropods feeding on wildlife. Ecology and Evolution, 7 (19), 7578–7584. https://doi.org/10.1002/ece3.2769
    Manga, L., Mbingue, S., Nkou Etoundi, M. & Ngollo, M. (1997) Anopheles namibiensis is anthropophilic and widespread in Cameroon. Medical and Veterinary Entomology, 11 (4), 409. https://doi.org/10.1111/j.1365-2915.1997.tb00432.x
    Meigen, J.W. (1818) Systematische Beschreibung der bekannten europaischen zweiflugeligen Insekten. Vol. 1. Aachen, xxxvi + 332 pp., errata (1 p.), 11 pls. https://doi.org/10.5962/bhl.title.13731
    Morsy, T.A., El Kadry, A.A., Salama, M.M., Sabry, A.H. & El Sharkawy, I.M. (1995) Studies on the bionomics and vector competence of adult anopheline mosquitoes in El Faiyum Governorate, Egypt. Journal of the Egyptian Society of Parasitology, 25 (1), 213–244.
    Mwangangi, J.M., Muturi, E.J., Muriu, S.M., Nzovu, J., Midega, J.T. & Mbogo, C. (2013) The role of Anopheles arabiensis and Anopheles coustani in indoor and outdoor malaria transmission in Taveta District, Kenya. Parasites & Vectors, 6, 114. https://doi.org/10.1186/1756-3305-6-114
    Ndiath, M.O., Eiglmeier, K., Olé Sangba, M.L., Holm, I., Kazanji, M. & Vernick, K.D. (2016) Composition and genetics of malaria vector populations in the Central African Republic. Malaria Journal, 15, 387. https://doi.org/10.1186/s12936-016-1431-2
    Ndiath, M.O., Mazenot, C., Gaye, A., Konate, L., Bouganali, C., Faye, O., Sokhna, C. & Trape, J-F. (2011) Methods to collect Anopheles mosquitoes and evaluate malaria transmission: A comparative study in two villages in Senegal. Malaria Journal, 10, 270. https://doi.org/10.1186/1475-2875-10-270
    Nepomichene, T.N., Tata, E. & Boyer, S. (2015) Malaria case in Madagascar, probable implication of a new vector, Anopheles coustani. Malaria Journal, 14, 475. https://doi.org/10.1186/s12936-015-1004-9
    Nyirakanani, C., Chibvongodze, R., Kariuki, L., Habtu, M., Masika, M., Mukoko, D. & Njunwa, K.J. (2017) Characterization of malaria vectors in Huye District, southern Rwanda. Tanzania Journal of Health Research, 19 (3), 1–10. https://doi.org/10.4314/thrb.v19i3.8
    Ogola, E., Villinger, J., Mabuka, D., Omondi, D., Orindi, B., Mutunga, J., Owino, V. & Masiga, D.K. (2017) Composition of Anopheles mosquitoes, their blood-meal hosts, and Plasmodium falciparum infection rates in three islands with disparate bed net coverage in Lake Victoria, Kenya. Malaria Journal, 16, 360. https://doi.org/10.1186/s12936-017-2015-5
    Osman, A.A., Gafer, A.A., Abdalmagid, M.A., Jamal, A.E., Bashir, A., Elnaeim, H., Abdalgadir, O.M., Brair, M. & Toto, H.K. (2014) Up to date anopheline mosquitoes distribution in Sennar and South Darfur states during dry season, (Sudan) 2009. Sudanese Journal of Public Health, 9 (1), 53–58.
    Paupy, C., Makanga, B., Ollomo, B., Rahola, N., Durand, P., Magnus, J., Willaume, E., Renaud, F., Fontenille, D. & Prugnolle, F. (2013) Anopheles moucheti and Anopheles vinckei are candidate vectors of ape Plasmodium parasites, including Plasmodium falciparum in Gabon. PLoS One, 8, e57294. https://doi.org/10.1371/journal.pone.0057294
    Peffly, R.L. (1959) Insecticide resistance in anophelines in eastern Saudi Arabia. Bulletin of the World Health Organization, 20, 757–776.
    Ratovonjato, J., Olive, M.M., Tantely, L.M., Andrianaivolambo, L., Tata, E., Razainirina, J., Jeanmaire, E., Reynes, J.M. & Elissa, N. (2011) Detection, isolation, and genetic characterization of Rift Valley fever virus from Anopheles (Anopheles) coustani, Anopheles (Anopheles) squamosus, and Culex (Culex) antennatus of the Haute Matsiatra region, Madagascar. Vector-Borne Zoonotic Diseases, 11 (6), 753–759. https://doi.org/10.1089/vbz.2010.0031
    Reid, J.A & Knight, K.L. (1961) Classification within the subgenus Anopheles (Diptera: Culicidae). Annals of Tropical Medicine & Parasitology, 55 (4), 474–488. https://doi.org/10.1080/00034983.1961.11686077
    Ribeiro, H. & da Cunha Ramos, H. (1975) Research on the mosquitoes of Angola. VI―The genus Anopheles Meigen, 1818 (Diptera, Culicidae) [sic]. Check-list with new records, keys to the females and larvae, distribution and bioecological notes. Garcia de Orta, Série de Zoologia, 4 (1), 1– 39, 17 pls.
    Sallum, M.A.M., Obando, R.G., Carrejo, N. & Wilkerson, R.C. (2020) Identification keys to the Anopheles mosquitoes of South America (Diptera: Culicidae). IV. Adult females. Parasites & Vectors, 13, 584. https://doi.org/10.1186/s13071-020-04301-0
    Stevenson, J.C., Simubali, L., Mbambara, S., Musonda, M., Mweetwa, S., Mudenda, T., Pringle, J.C., Jones, C.M. & Norris, D.E. (2016) Detection of Plasmodium falciparum infection in Anopheles squamosus (Diptera: Culicidae) in an area targeted for malaria elimination, southern Zambia. Journal of Medical Entomology, 53 (6), 1482–1487. https://doi.org/10.1093/jme/tjw091
    St. Laurent, B., Cooke, M., Krishnankutty, S.M., Asih, P., Mueller, J.D., Kahindi, S., Ayoma, E., Oriango, R.M., Thumloup, J., Drakeley, C., Cox, J., Collins, F.H., Lobo, N.F. & Stevenson, J.C. (2016) Molecular characterization reveals diverse and unknown malaria vectors in the western Kenyan highlands. American Journal of Tropical Medicine and Hygiene, 94 (2), 327–335. https://doi.org/10.4269/ajtmh.15-0562
    Stone, A., Knight, K.L. & Starcke, H. (1959) A synoptic catalog of the mosquitoes of the world (Diptera, Culicidae). The Thomas Say Foundation. Volume VI. Entomological Society of America, College Park, Maryland, vi + 358 pp.
    Tabue, R.N., Awono-Ambene, P., Etang, J., Atangana, J., Antonio-Nkondjio, C., Toto, J.C., Patchoke, S., Leke, R.G., Fondjo, E., Mnzava, A.P., Knox, T.B., Tougordi, A., Donnelly, M.J. & Bigoga, J.D. (2017) Role of Anopheles (Cellia) rufipes (Gough, 1910) and other local anophelines in human malaria transmission in the northern savannah of Cameroon: a cross-sectional survey. Parasites & Vectors, 10, 22. https://doi.org/10.1186/s13071-016-1933-3
    Tabue, R.N., Nem, T., Atangana, J., Bigoga, J.D., Patchoke, S., Tchouine, F., Fodjo, B.Y., Leke, R.G. & Fondjo, E. (2014) Anopheles ziemanni a locally important malaria vector in Ndop health district, north west region of Cameroon. Parasites & Vectors, 7, 262. https://doi.org/10.1186/1756-3305-7-262
    Theobald, F.V. (1900) A new Anopheles (A. paludis) from Sierra Leone. Reports to the Malaria Committee of the Royal Society, 1, 75–76.
    Theobald, F.V. (1901) A monograph of the Culicidae or mosquitoes. Vol. 1. British Museum (Natural History), London, xviii + 424 pp., pls. viii + I‒XXXVII + A‒E under separate cover.
    Theobald, F.V. (1903) Report on a collection of mosquitoes and other flies from Equatorial East Africa and the Nile provinces of Uganda. Reports to the Sleeping Sickness Committee of the Royal Society, 7 (3), 33–42, map.
    Theobald, F.V. (1907) A monograph of the Culicidae or mosquitoes. Volume 4. British Museum (Natural History), London, xix + 639 pp, 16 pls.
    van Someren, E.C.C. (1947) The description of a new Anopheles of the Myzorhynchus series from Madagascar with notes on its systematic position in relation to the Ethiopian species of this group. East African Medical Journal, 24 (1), 42–46.
    Villinger, J., Mbaya, M.K., Ouso, D., Kipanga, P.N., Lutomiah, J. & Masiga, D.K. (2017) Arbovirus and insect-specific virus discovery in Kenya by novel six genera multiplex high-resolution melting analysis. Molecular Ecology Resources, 17 (3), 466–480. https://doi.org/10.1111/1755-0998.12584
    Wanji, S., Mafo, F.F., Tendongfor, N., Tanga, M.C., Tchuente, F., Bilong Bilong, C.F. & Njine, T. (2009) Spatial distribution, environmental and physicochemical characterization of Anopheles breeding sites in the Mount Cameroon region. Journal of Vector-borne Diseases, 46 (1), 75–80.
    White, G.B. (1980) Family Culicidae. In: Crosskey, R.W. (Ed.), Catalogue of the Diptera of the Afrotropical Region. British Museum (Natural History), London, pp. 114–148.
    Wilkerson, R.C., Linton, Y.-M. & Strickman, D. (2021) Mosquitoes of the world. Vols. 1 & 2. Johns Hopkins University Press, Baltimore, Maryland, 1332 pp. 
    https://doi.org/10.1186/s13071-021-04848-6
    Wilkerson, R.C. & Peyton, E.L. (1990) Standardized nomenclature for the costal wing spots of the genus Anopheles and other spotted-wing mosquitoes (Diptera: Culicidae). Journal of Medical Entomology, 27 (2), 207–224. https://doi.org/10.1093/jmedent/27.2.207
    Zeru, M.A., Shibru, S. & Massebo, F. (2020) Exploring the impact of cattle on human exposure to malaria mosquitoes in the Arba Minch area district of southwest Ethiopia. Parasites & Vectors, 13, 322. https://doi.org/10.1186/s13071-020-04194-z

  2.