Abstract
Several house bat specimens superficially resembling the white-bellied house bat Scotophilus leucogaster (Cretzschmar, 1830), were recently captured in southwestern Ethiopia and southern South Sudan. These S. cf. leucogaster differed from typical S. leucogaster by their slightly smaller size and ventral coloration, conforming instead with the original description of S. altilis Allen, 1914. Scotophilus altilis is an overlooked taxon known from the Blue Nile region in Sudan that is currently considered a junior synonym of S. leucogaster. Phylogenetic analysis of mitochondrial cytochrome b gene (cytb) sequences revealed S. cf. leucogaster as a sister clade to S. leucogaster with a genetic distance of ca. 10%. Comparative specimens of questionable S. nigritellus de Winton, 1899 from northwestern Ethiopia and a wing biopsy sample of another S. cf. leucogaster from western Kenya also fell within this clade. Sequence data from two nuclear markers (zfy and fgb7) corroborated the distinction of S. cf. leucogaster from S. leucogaster. Likewise, morphometric analysis of cranial data largely supported this distinction, as well as taxonomic affiliation with S. altilis based on comparison with the only available paratype specimen. The position of this paratype specimen within the new Scotophilus clade, inferred from analysis of a short fragment of cytb, confirmed its taxonomic identity. Based on the presented evidence, the overlooked East African taxon S. altilis should be resurrected as a full species within the genus Scotophilus.
References
Aellen, V. (1952) Contribution à l’étude des chiroptères du Cameroun. Mémoires de la Société neuchâteloise des sciences naturelles, 8, 1–121.
Allen, G.M. (1914) Mammals from the Blue Nile Valley. Bulletin of the Museum of Comparative Zoology at Harvard College, 58, 305–357.
Allen, J.A., Lang, H. & Chapin, J.P. (1917) The American Museum Congo Expedition collection of bats. Bulletin of the American Museum of Natural History, 37, 405–563.
Allen, G.M. (1939) A checklist of African mammals. Bulletin of the Museum of Comparative Zoology at Harvard College, 83, 1–763.
Baker, R.J. & Bradley, R.D. (2006) Speciation in mammals and the genetic species concept. Journal of Mammalogy, 87 (4), 643–662.
https://doi.org/10.1644/06-MAMM-F-038R2.1
Bandelt, H.-J., Forster, P. & Röhl, A. (1999) Median-joining networks for inferring intraspecific phylogenies. Molecular Biology and Evolution, 16, 37–48.
https://doi.org/10.1093/oxfordjournals.molbev.a026036
Brooks, D.M. & Bickham, J.W. (2014) New species of Scotophilus (Chiroptera: Vespertilionidae) from Sub-Saharan Africa. Occasional Papers Museum of Texas Tech University, 326, 1–21.
Cretzschmar, P.J. (1830) Säugethiere. In: Atlas zu der Reise im nördlichen Afrika von Eduard Rüppell. Senckenberghische Naturforschende Gesellschaft, Frankfurt am Main, pp. 71–72.
Demos, T.C., Webala, P.W., Bartonjo, M. & Patterson, B.D. (2018) Hidden diversity of African yellow house bats (Vespertilionidae, Scotophilus): Insights from multilocus phylogenetics and lineage delimitation. Frontiers in Ecology and Evolution, 6, 1–13.
https://doi.org/10.3389/fevo.2018.00086
De Winton, W.E. (1899) On mammals collected by Lieut. -Colonel W. Giffard in the Northern Territory of the Gold Coast. Annals and Magazine of Natural History, Series 7, 4, 353–359.
https://doi.org/10.1080/00222939908678212
Goodman, S.M., Jenkins, R.K.B. & Ratrimomanarivo, F.H. (2005) A review of the genus Scotophilus (Chiroptera: Vespertilionidae) on Madagascar, with the description of a new species. Zoosystema, 27, 867–882.
Helgen, K.M. & McFadden, T.L. (2001) Type specimens of recent mammals in the Museum of Comparative Zoology. Bulletin of the Museum of Comparative Zoology, 157, 93–181.
Heuglin von, M.T. (1861) Beiträge zur Fauna der Säugethiere N.O.—Afrikas. Nova Acta Academiae Caesareae Leopoldino-Carolinae Halle, 29, 1–18.
Jacobs, D.S., Eick, G.N., Schoeman, M.C. & Mathee, C.A. (2006) Cryptic species in an insectivorous bat, Scotophilus dinganii. Journal of Mammalogy, 87 (1), 161–170.
https://doi.org/10.1644/04-MAMM-A-132R2.1
Kock, D. (1969) Die Fledermaus-Fauna des Sudan (Mammalia, Chiroptera). Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft, 521, 1–238.
Koopman, K.F. (1965) Status of forms described or recorded by J.A. Allen in "The American Museum Congo Expedition collection of bats." American Museum Novitates, 2219, 1–34.
Koopman, K.F. (1975) Bats of the Sudan. Bulletin of the American Museum of Natural History, 154, 353–444.
Koopman, K.F. (1994) Chiroptera: systematics. In: Niethammer, J., Schliemann, H. & Starck, D. (Eds.), Handbook of zoology: a natural history of the phyla of the animal kingdom, Vol. VIII. Mammalia. Part 60. Walter de Gruyter, Berlin, pp. 1–217.
Kruskop, S.V., Benda, P., Vasenkov, D.A. & Lavrenchenko, L.A. (2016) First records of bats from the Alatish National Park, north-western Ethiopia (Chiroptera). Lynx, New Series, 47, 51–69.
Lanza, B., Funaioli, U. & Riccucci, M. (2015) The bats of Somalia and neighbouring areas. Editions Chimaira, Frankfurt am Main, 566 pp.
Monadjem, A., Taylor, P.J., Cotterill, F.D.P. & Schoeman, M.C. (2010) Bats of Southern and Central Africa. A biogeographic and taxonomic synthesis. Wits University Press, Johannesburg, 596 pp.
Robbins, C.B., De Vree, F. & Van Cakenberghe, V. (1985) A systematic revision of the African bat genus Scotophilus (Vespertilionidae). Annales Musée Royal de l'Afrique Centrale, Sciences Zoologiques, 246, 51–84.
Ronquist, F. & Huelsenbeck, J.P. (2003) MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics, 19, 1572–1574.
https://doi.org/10.1093/bioinformatics/btg180
Simmons, N.B. (2005) Order Chiroptera. In: Wilson, D.E. & Reeder, D.M. (Eds.), Mammal species of the world: a taxonomic and geographic reference. 3rd Edition. Johns Hopkins University Press, Baltimore, pp. 312–529.
Thomas, O. (1904) On new bats from British Africa and Cameroons. Annals and Magazine of Natural History, Series 7, 13, 207.
Trujillo, R.G., Patton, J.C., Schlitter, D.A. & Bickham, J.W. (2009) Molecular phylogenetics of the bat genus Scotophilus (Chiroptera: Vespertilionidae): Perspectives from paternally and maternally inherited genomes. Journal of Mammalogy, 90, 548–560.
https://doi.org/10.1644/08-MAMM-A-239R2.1
Van Cakenberghe, V. & Happold, M. (2013) Scotophilus leucogaster. White-bellied House-bat. In: Happold, M. & Happold, D. (Eds.), Mammals of Africa. Volume IV. Hedgehogs, shrews and bats. Bloomsbury, London, New Delhi, New York and Sydney, pp. 676–678.
https://doi.org/10.1515/mammalia-2013-0137
Vallo, P., Benda, P. & Reiter, A. (2011) Yellow-bellied or white-bellied? Identity of Arabian house bats (Vespertilionidae: Scotophilus) revealed from mitochondrial DNA and morphology. African Zoology, 46 (2), 350–361.
https://doi.org/10.3377/004.046.0214
Vallo, P., Benda, P., Červený, J. & Koubek, P. (2013) Conflicting mitochondrial and nuclear paraphyly in small-sized West African house bats (Vespertilionidae). Zoologica Scripta, 42, 1–12.
https://doi.org/10.1111/j.1463-6409.2012.00563.x
Vallo, P., Benda, P., Červený, J. & Koubek, P. (2015) Phylogenetic position of the giant house bat Scotophilus nigrita (Chiroptera, Vespertilionidae). Mammalia, 79, 225–231.
https://doi.org/10.1515/mammalia-2013-0137
Vallo, P. & Van Cakenberghe, V. (2017) Advances in taxonomy of African house bats (Scotophilus, Vespertilionidae). African Bat Conservation News, 46, 4–9.
Wettstein von, O. (1918) Wissenschaftliche Ergebnisse der mit Unterstützung der Kaiserlichen Akademie der Wissenschaften in Wien aus der Erbschaft Treitl von F. Werner unternommenen zoologischen Expedition nach dem Anglo-Ägyptischen Sudan (Kordofan) 1914. II. Bearbeitung der auf der Expedition gesamelten Vögel und Säugetiere. Denkschrifte der Kaiserlichen Akademie der Wissenschaften Wien, mathematisch-naturwissenschaftliche Klasse, 94, 555–691.