Anopheles (Cellia) rufipes (Gough)

Harbach, Ralph E. & Wilkerson, Richard C., 2023, The insupportable validity of mosquito subspecies (Diptera: Culicidae) and their exclusion from culicid classification, Zootaxa 5303 (1), pp. 1-184 : 71-72

Anopheles (Cellia) rufipes (Gough)

subspecies broussesi Edwards, 1929a —original combination: Anopheles (Myzomyia) broussesi (subspecific status by Rioux 1961). Distribution: Algeria, Burkina Faso, Chad, Egypt, Ghana, Libya, Mali, Niger, Nigeria ( Wilkerson et al. 2021) .

subspecies rufipes ( Gough, 1910) —original combination: Nyssorhynchus pretoriensis var. rufipes (specific status by Edwards 1912c). Distribution: Algeria, Angola, Benin, Botswana, Burkina Faso, Cameroon, Central African Republic, Chad, Côte d’Ivoire, Democratic Republic of the Congo, Eritrea, Eswatini, Ethiopia, Gabon, Gambia, Ghana, Guinea, Guinea-Bissau, Kenya, Madagascar, Malawi, Mali, Mauritania, Mozambique, Namibia, Niger, Nigeria, Republic of the Congo, Republic of South Africa, Senegal, Seychelles, Sierra Leone, South Sudan, Sudan, Tanzania, Togo, Uganda, Zambia, Zimbabwe ( Wilkerson et al. 2021).

The taxa discussed here have very similar larval, pupal and male genitalic characters. Most are illustrated or discussed in Gillies & de Meillon (1968). The adult females are distinguished from other species of the subgenus Cellia by the following combination of characters: Lack of shaggy palpus; uniformly brownish leg coloration; various extent of white banding on the hindtarsomeres; medium to broad pale scutal scales either in a regular pattern or grading to evenly distributed; usually with 6 distinct pale wing spots and pale fringe spots at the ends of all veins except the anal vein; abdomen uniformly without scales; gonocoxite of male with 4 parabasal setae and aedeagus with several pairs of unmodified leaflets; larval setae 2,3-C long with sparse branches; blades of palmate setae with very short stumpy filaments, the setae resembling blunt-looking fans.

Gough (1910) described rufipes as a variety of Nyssorhynchus pretoriensis Theobald, 1903a (not pretoriensis Gough, 1910 , a synonym of An. squamosus Theobald, 1901a ) from collections made in Onderstepoort, Transvaal, Republic of South Africa. Gough’s description is brief but he noted that variety rufipes has “uniformly ruddy brown” legs, quite different from pretoriensis , which has speckled femora and tibiae. Other distinctive adult and larval characters that separate pretoriensis from rufipes are also evident. Without comment, in a key, Edwards (1912c) recognized rufipes as a species and also placed An. watsoni Edwards, 1911a , from Katagumm Northern Provinces, Nigeria, in synonymy with it.

Edwards (1929a) described broussesi from “Djanet, the most southerly post in the Algerian Sahara… on the southern extremity of the Tasili n Ajjer [sic] mountains.” Djanet is the capital of Djanet Province, an oasis city in a hot desert climate in southeastern Algeria. The adult female has a combination of distinguishing characters: “Palpi of ♀ slender, dark, with three very narrow white rings on joints, last segment almost entirely dark…”; scutum with well-defined lines of supraalar, dorsocentral and acrostichal scales; vein R 4 +5 mostly dark with a pair of small subbasal pale spots (illustrated but not noted by Edwards); legs with “extremely faint pale rings present at tips of first four segments [tarsomeres 1–4] of hind tarsi; last hind tarsal segment [tarsomere 5] in several specimens entirely cream-coloured, but in others apparently dark like remainder of tarsi.”

Evans (1938) described the nominotypical form in detail but did not compare it with broussesi . She did, however, recognize rufipes var. ingrami Edwards, 1929b , a current synonym of ssp. broussesi , as listed by Harbach (2018) and Wilkerson et al. (2021) or as ssp. rufipes per Gillies & de Meillon (1968). Evans noted several features that can be contrasted with Edwards’s (1929a) characters for broussesi : Female palpus with 3 pale bands, the first 2 broad; scutum nearly evenly covered with somewhat broad pale scales; vein R 4+5 with a single small subbasal pale spot and long median pale spot (illustrated but not noted by Evans); hindtarsomere 2 pale on apical 0.4 and on all of tarsomeres 3–5; larval seta 2-C single or with fine aciculae.

De Meillon (1947) treated An. rufipes and An. rufipes var. ingrami in keys, descriptions and illustrations, but did not mention broussesi , perhaps because he considered it to be a Saharan taxon. Senevet & Andarelli (1956), in a treatise on species of North Africa and the Mediterranean basin, used Edwards’s (1929a) characters and illustration of the wing, and considered broussesi to be a species. It is to be noted that broussesi and rufipes both were listed as separate species in Stone et al. (1959).

Rioux (1961) discussed geographic distribution, variation in the extent of hindtarsal pale scaling, and the extent and distribution of pale scutal scales, to determine that rufipes consisted of three subspecies: An. rufipes rufipes , An. rufipes broussesi and An. rufipes seneveti Rioux, 1959 . This was the first time broussesi was considered as a subspecies of rufipes . The taxonomic treatment of Gillies & Coetzee (1987) and catalog listings ( Knight & Stone 1977; White 1980; Harbach 2018; Wilkerson et al. 2021) followed the recognition of rufipes rufipes and rufipes broussesi . Hamon et al. (1961) named a new variety, An. rufipes var. brucechwatti , and also recognized An. broussesi as a species and An. rufipes as a species with varieties ingrami and seneveti. Dubose & Curtin (1965), who stated they used the collection of the U. S. National Museum extensively, presumably with Alan Stone’s input, retained broussesi as a species. Variety seneveti was later placed in synonymy with An. rufipes broussesi by Gillies & de Meillon (1968), as was brucechwatti. We note that the illustration of the wing of seneveti differs from illustrated wings of the other nominal forms in having a nearly entirely pale-scaled vein CuA, suggesting its possible validity. Gillies & de Meillon (1968) recognized only An. rufipes rufipes and An. rufipes broussesi and included ingrami as a synonym of subspecies rufipes and not broussesi as indicated by Harbach (2018) and Wilkerson et al. (2021). We do not know the source of this placement, or which is correct.

Ribeiro & da Cunha Ramos (1975) also recognized subspecies rufipes rufipes and rufipes broussesi . They stated: “ A. rufipes rufipes is a widespread subspecies of African savannas, while A. rufipes broussesi is restricted to the western portion of the Northern Savanna of Moreau.” Julvez et al. (1998) reported that subspecies rufipes rufipes and rufipes broussesi are sympatric in Chad.

We believe that the above provides evidence of two species, rufipes and broussesi . Anopheles rufipes is a variable, widespread Afrotropical species, or species complex, and An. broussesi is a hot-dry climate Saharan and Mediterranean species. The observed variability, mostly in the pattern of palpal and hindtarsal pale scales, has been expressed by authors using varietal or subspecies names, which have been used over time in many different combinations. We see no empirical evidence of genetic independence other than between these two species, and therefore return broussesi to species status: Anopheles (Cellia) broussesi Edwards, 1929a . Anopheles broussesi is currently listed as a species in the Encyclopedia of Life.