Abstract
The taxonomy of the small, sub-Saharan, insectivorous bat, Afronycteris helios (Heller, 1912), has been unresolved for decades. The name A. cf. helios was introduced in the literature to recognise bats found in east and southern Africa that were like A. helios but had glands on the uropatagium. Cranio-dental morphology, bacular morphology, and molecular genetics (albeit the latter two being without representation of A. helios), provided evidence to formally describe “A. cf. helios”, which is currently known from Kenya, Mozambique, and South Africa. Bayesian analyses based on cytochrome oxidase b, cytochrome c oxidase subunit 1, and 12S rRNA confirm that it belongs to the genus Afronycteris with A. nanus (Peter, 1852) and A. helios. These analyses also revealed genetic, bacular, and cranio-dental morphological differences within A. nanus, which are described here. Pending a more thorough geographic analysis, including all existing synonyms, A. cf. nanus has been introduced to refer to bats that were smaller than A. nanus. Although co-occurring in north-eastern parts of South Africa, A. cf. nanus has a more westerly distribution extending to west Africa, relative to the more easterly distribution of A. nanus. Afronycteris sp. nov. showed at least a 6.9%, 3.4% and 2.9% nucleotide difference to its nearest relative based on cytochrome oxidase b, cytochrome c oxidase subunit 1, and 12S rRNA respectively. It is cranio-dentally smaller than A. helios, slightly smaller than A. nanus, and larger than A. cf. nanus, with a distinct baculum, and a unique pair of glands on the uropatagium.
References
- ACR (2024) African Chiroptera Report. 2024. African Bats NPC, Pretoria, xv + 1397 pp. https://doi.org/10.13140/RG.2.2.15903.27045
- Benda, P., Uvizl, M., Šklíba, J., Mazoch, V. & Červený, J. (2022) African bats in the collection of the National Museum, Prague (Chiroptera). I. Bats from Zambia. Lynx, series nova, 53 (1), 291–323. https://doi.org/10.37520/lynx.2022.021
- Bickham, J.W., Wood, C.C. & Patton, J.C. (1995) Biogeographic implications of cytochrome b sequences and allozymes in sockeye (Oncorhynchus nerka). Journal of Heredity, 86, 140–144. https://doi.org/10.1093/oxfordjournals.jhered.a111544
- Bickham, J.W., Patton, J.C., Schlitter, D.A., Rautenbach, I.L. & Honeycutt, R.L. (2004) Molecular phylogenetics, karyotypic diversity, and partition of the genus Myotis (Chiroptera, Vespertilionidae). Molecular Phylogenetics and Evolution, 33 (2), 333–338. https://doi.org/10.1016/j.ympev.2004.06.012
- Darriba, D., Taboada, G.L., Doallo, R. & Posada, D. (2012) JModelTest 2: more models, new heuristics and parallel computing. Nature Methods, 9, 772. https://doi.org/10.1038/nmeth.2109
- Demos, T.C., Webala, P.W., Monadjem, A. & Patterson, B.D. (2025) Nuclear introns support the subtribe Laephotina and recently proposed genera of African Vespertilionidae. Acta Chiropterologica, 26 (2), 143–152. https://doi.org/10.3161/15081109ACC2024.26.2.001
- Drummond, A.J. & Rambaut, A. (2007) BEAST: Bayesian evolutionary analysis by sampling trees. BMC Evolutionary Biology, 7, 214. https://doi.org/10.1186/1471-2148-7-214
- Folmer, O., Black, M., Hoeh, W., Lutz, R. & Vrijenhoek, R. (1994) DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology, 3, 294–299.
- Geldenhuys, M., Mortlock, M., Weyer, J., Bezuidt, O., Seamark, E.C., Kearney, T., Gleasner, C., Erikkil, T.H., Cui, H. & Markotter, W. (2018) A metagenomic viral discovery approach identifies potential zoonotic and novel mammalian viruses in Neoromicia bats within South Africa. PloS ONE, 13 (3), e0194527. https://doi.org/10.1371/journal.pone.0194527
- Goodman, S.M., Rakotondramananus, C.F., Ramasindrazana, B., Kearney, T., Monadjem, A., Schoeman, M.C., Taylor, P.J., Naughton, K. & Appleton, B. (2015) An integrative approach to characterize Malagasy bats of the subfamily Vespertilioninae. Zoological Journal of the Linnean Society, 173 (4), 988–1018. https://doi.org/10.1111/zoj.12223
- Goodman, S.M., Kearney, T., Ratsimbazafy, M.M. & Hassanin, A. (2017) Description of a new species of Neoromicia (Chiroptera: Vespertilionidae) from southern Africa: a name for ‘N. cf. melckorum’. Zootaxa, 4236 (2), 351–374. https://doi.org/10.11646/zootaxa.4236.2.10
- Hall, T.A. (1999) BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic Acids Symposium Series, 41, 95–98.
- Hammer, O., Harper, D.A.T. & Ryan, P.D. (2001) PAST: Paleontological Statistics software package for education and data analysis. Palaeontologia Electronica, 4 (1), 1–9.
- Happold, D.C.D. & Happold, M. (1996) The social organization and population dynamics of leaf-roosting banana bats, Pipistrellus nanus (Chiroptera, Vespertilionidae), in Malawi, east-central Africa. Mammalia, 60 (4), 517–544. https://doi.org/10.1515/mamm.1996.60.4.517
- Happold, M. & Van Cakenberghe, V. (2013) Pipistrellus cf. helios Samburu Pipistrelle. In: Happold, M. & Happold, D.C.D. (Eds.), Mammals of Africa. Vol. IV. Hedgehogs, Shrews and Bats. Bloomsbury Publishing, London, pp. 627–629.
- Harima, H., Qiu, Y., Yamagishi, J., Kajihara, M., Changula, K., Okuya, K., Isono, M., Yamaguchi, T., Ogawa, H., Nao, N., Sasaki, M., Simulundu, E., Mweene, A.S., Sawa, H., Ishihara, K., Hang’ombe, B.M. & Takada, A. (2023) Surveillance, Isolation, and Genetic Characterization of Bat Herpesviruses in Zambia. Viruses, 15 (6), 1369. https://doi.org/10.3390/v15061369
- Hassanin, A., Colombo, R., Gembu, G.-C., Merle, M., Tu, V.T., Gorfol, T., Akawa, P.M., Csorba, G., Kearney, T., Monadjem, A. & Ing, R.K. (2017) Multilocus phylogeny and species delimitation within the genus Glauconycteris (Chiroptera, Vespertilionidae), with the description of a new bat species from the Tshopo Province of the Democratic Republic of the Congo. Journal of Zoological Systematics and Evolutionary Research, 56, 1–22. https://doi.org/10.1111/jzs.12176
- Hassanin, A., Delsuc, F., Ropiquet, A., Hammer, C., Jansen van Vuuren, B., Matthee, C., Ruiz-Garcia, M., Catzeflis, F., Areskoug, V., Nguyen, T.T. & Couloux, A. (2012) Pattern and timing of diversification of Cetartiodactyla (Mammalia, Laurasiatheria), as revealed by by a comprehensive analysis of mitochondrial genomes. Comptes Rendus Biologies, 335, 32–50. https://doi.org/10.1016/j.crvi.2011.11.002
- Hill, J.E. & Harrison, D.L. (1987) The baculum in the Vespertilioninae (Chiroptera: Vespertilionidae) with a systematic review, a synopsis of Pipistrellus and Eptesicus, and the descriptions of a new genus and subgenus. Bulletin of the British Museum (Natural History), Zoology Series, 52 (7), 225–305.
- Heller, E. (1912) New races of insectivores, bats and lemurs from British East Africa. Smithsonian Miscellaneous Collections, 60 (12), 1–13.
- Kearney, T.C. (2005) Systematic revision of southern African species in the genera Eptesicus, Neoromicia, Hypsugo and Pipistrellus (Chiroptera: Vespertilionidae). Ph.D. Thesis, The University of KwaZulu-Natal, Durban, 580 pp.
- Kearney, T.C. & Taylor, P.J. (2011) Selection of cranial and mandible measurements for traditional morphometric analyses of southern African vesper bats of the genera Eptesicus, Hypsugo, Neoromicia and Pipistrellus (Mammalia: Chiroptera: Vespertilionidae). Annals of the Ditsong National Museum of Natural History, 1, 53–61.
- https://hdl.handle.net/10520/EJC83700
- Kearney, T.C., Volleth, M., Contrafatto, G. & Taylor, P.J. (2002) Systematic implications of chromosome GTG-band and bacula morphology for Southern African Eptesicus and Pipistrellus and several other species of Vespertilioninae (Chiroptera: Vespertilionidae). Acta Chiropterologica, 4 (1), 55–76. https://doi.org/10.3161/001.004.0107
- Koopman, K.F. (1993) Order Chiroptera. In: Wilson, D.E. & Reeder, D.M. (Eds.), Mammal species of the world: a taxonomic and geographic reference. Smithsonian Institution Press, Washington, D.C., pp. 137–241.
- Kumar, S., Stecher, G., Li, M., Knyaz, C. & Tamura, K. (2018) MEGA X: molecular evolutionary genetics analysis across platforms. Molecular Biology and Evolution, 35, 1547–1549. https://doi.org/10.1093/molbev/msy096
- LaVal, R.K. & LaVal, M.L. (1977) Reproduction and behaviour of the African Banana Bat, Pipistrellus nanus. Journal Of Mammalogy, 58 (3), 403–410. https://doi.org/10.2307/1379339
- Meester, J.A.J., Rautenbach, I.L., Dippenaar, N.J. & Baker, C.M. (1986) Classification of Southern African mammals. Transvaal Museum Monograph. No. 5. Transvaal Museum, Pretoria, 359 pp.
- Miller, M.A., Pfeiffer, W. & Schwartz, T. (2010) Creating the CIPRES Science Gateway for inference of large phylogenetic trees. In: 2010 Gateway Computing Environments workshop (GCE). IEEE, New Orleans, Louisiana, pp. 1–8. https://doi.org/10.1109/GCE.2010.5676129
- Monadjem, A., Richards, L., Taylor, P.J. & Stoffberg, S. (2013) High diversity of pipistrelloid bats (Vespertilionidae: Hypsugo, Neoromicia and Pipistrellus) in a West African rainforest with the description of a new species. Zoological Journal of the Linnean Society, 167 (1), 191–207. https://doi.org/10.1111/j.1096-3642.2012.00871
- Monadjem, A., Demos, T.C., Dalton, D.L., Webala, P.W., Musila, S., Kerbis Peterhans, J.C. & Patterson, B.D. (2021) A revision of pipistrelle-like bats (Mammalia: Chiroptera: Vespertilionidae) in East Africa with the description of new genera and species. Zoological Journal of the Linnean Society, 191 (4), 1114–1146. https://doi.org/10.1093/zoolinnean/zlaa087
- O’Shea, T.J. (1980) Roosting, social organization and the annual cycle in a Kenya population of the bat Pipistrellus nanus. Zeitschrift für Tierpsychologie, 53 (2), 171–195. https://doi.org/10.1111/j.1439-0310.1980.tb01048.x
- Peters, W.C. (1852) Naturwissenschaftliche Reise nach Mossambique: 1. Säugethiere. Mit sechs und vierzig Taflen. Zoologie. Druck und Verlag von Georg Reimer, 670 pp.
- Rautenbach, I.L., Fenton, M.B. & Braack, L.E.O. (1985) First records of five species of insectivorous bats from the Kruger National Park. Koedoe, 28, 73–80. https://doi.org/10.4102/koedoe.v28i1.537
- Rautenbach, I.L., Fenton, M.B. & Whiting, M.J. (1996) Bats in riverine forests and woodlands: a latitudinal transect in southern Africa. Canadian Journal of Zoology, 74, 312–322. https://doi.org/10.1139/z96-039
- Rosevear, D.R. (1965) The bats of West Africa. British Museum (Natural History), London, 418 pp.
- Rutherford, M.C., Mucina, L. & Powrie, L.W. (2006) Biomes and bioregions of southern Africa. The vegetation of South Africa, Lesotho and Swaziland, 19, 30–51.
- Simmons, N.B. (2005) Order Chiroptera. In: Wilson, D.E. & Reeder, D.M. (Eds.), Mammal species of the world: a taxonomic and geographic reference. John Hopkins University Press, Baltimore, Maryland, pp. 312–529.
- Torrent, L., Juste, J., Garin, I., Aihartza, J., Dalton, D.L., Mamba, M., Tanshi, I., Powell, L.L., Padidar, S., Garcia Mudarra, J.L., Richards, L. & Monadjem, A. (2025) Taxonomic revision of African pipistrelle-like bats with a new species from the West Congolean rainforest. Zoological Journal of the Linnean Society, 204 (2), zlaf020. https://doi.org/10.1093/zoolinnean/zlaf020
