African Bat Conservation News 4 August 2017 vol. 46 ISSN 1812-1268

Advances in taxonomy of African house bats (Scotophilus, Vespertilionidae)

Peter VALLO1,2 and Victor VAN CAKENBERGHE3 1Institute of Evolutionary Ecology and Conservation Genomics, Ulm University, Albert Einstein Alle 11, 89069 Ulm, Germany. 2Institute of Vertebrate Biology, v.v.i., Academy of Sciences of the Czech Republic, Květná 8, 60365 Brno, Czech Republic. [email protected]. 3Functional Morphology, Department of Biology, University of Antwerp (CDE), Universiteitsplein 1, B-2610 Antwerpen (Wilrijk), Belgium.

Bats of the vespertilionid genus Scotophilus Leach, 1821, in a complicated issue, less euphemistically best labelled as "a mess" English called house bats or, rather misleadingly, yellow bats, are (see ROBBINS et al., 1985). Some progress, however, has been commonly encountered throughout sub-Saharan Africa and southern achieved in recent years. An overview of all available African names Asia. Their strong dentition has left its marks on mistnets and and their current assignment is presented in Appendix 1. fingers of many a bat researcher, and contributed so to the general For a considerable part of the twentieth century, bat taxonomists familiarity of this genus. In external appearance and morphological had been satisfied with calling most of the African medium-sized traits, these bats look very similar, with size and pelage colouration Scotophilus simply S. nigrita, a name that actually pertains to the usually the most striking difference between species. For over a giant Scotophilus species known at those times under its junior century, this overall similarity has made the taxonomy of Scotophilus synonym S. gigas Dobson, 1875 (ROBBINS, 1978). Although today

Offshore observation Eidolon helvum Gulf of Guinea / Pakula et al. (2017) / Advances in taxonomy on African Scotophilus Scientific contributions Vallo and Van Cakenberghe (2017) African Bat Conservation News ISSN 1812-1268 August 2017 vol. 46 5 this taxonomic confusion has been corrected, the ghost of former designation to one of them and S. nigritellus (de Winton, 1899), times is still haunting many museum collections over the world, previously considered a subspecies of S. viridis in West Africa, to where quite a few specimens are still mislabelled as S. nigrita. Within another. Two other evolutionary units of S. dinganii remained without the former content of S. nigrita (see KOCK, 1969 and KOOPMAN, any taxonomic treatment, although their specific distinctness 1975), many currently acknowledged taxa had been hidden, which appeared obvious. These comprised populations from East and are familiar to contemporary bat researchers. The tribute for the West Africa, respectively, with the population from Kakamega, current taxonomic arrangement of African Scotophilus must be southwestern Kenya, clustering together with the latter. For the given to ROBBINS et al. (1985), who revised and substantiated sake of simplification, these clades will be further denoted as East recognition of the currently acknowledged species of continental African and West African. In S. viridis, two evolutionary units from Africa: S. dinganii (A. Smith, 1833), S. leucogaster (Cretzschmar, East and Southern Africa, respectively, seem to have been implicitly 1826), S. viridis (Peters, 1852), S. nigrita (Schreber, 1774), S. nux recognized as S. viridis s. str. and a so far undescribed species. Thomas, 1904, and S. nucella Robbins, 1983. After this crucial work by TRUJILLO et al. (2009), several Since the revision by ROBBINS et al. (1985), the taxonomic further studies on African Scotophilus appeared. In their study on situation in African Scotophilus has not changed much until Scotophilus from Yemen, VALLO et al. (2011) showed one of the 2005, when GOODMAN et al. (2005, 2006) described two new two Scotophilus species occurring there to be closely related to species from Madagascar, S. tandrefana Goodman, Jenkins Ethiopian S. dinganii, and thus belonging to the East African clade and Ratrimomanarivo, 2005 and S. marovaza Goodman, (Figure 1). Because of its unresolved internal arrangement, the Ratrimomanarivo and Randrianandrianina, 2006. Their descriptions East African evolutionary unit of the S. dinganii morphogroup was were based on morphological differences from the only traditionally considered as one - although highly variable - species and the name recognised Madagascan species S. robustus Milne-Edwards, 1881, S. colias (Thomas, 1904) was provisionally suggested for it. This was as well as species from adjacent continental Africa. At about the an available name for a yellow-bellied medium-sized Scotophilus same time, JACOBS et al. (2006) published the existence of a cryptic form from central Kenya suggested by SIMMONS (2005) for the species within the Southern African S. dinganii, clearly distinct in vaguely substantiated and geographically delimited East African morphology, echolocation and genetics from the latter species. It subspecies of S. dinganii. Two years later, a study on small-sized was provisionally named S. mhlanganii, but it was never formally Scotophilus in West Africa (VALLO et al., 2013) reported evidence described. Although it has been mentioned in two ecological studies for a yet unknown white-bellied form provisionally identified as S. (JACOBS et al., 2007; JACOBS and BARCLAY 2009), it is currently a aff. nigritellus bearing mitochondrial haplotypes closely related to nomen nudum (see MONADJEM et al., 2010; VAN CAKENBERGHE sympatric medium-sized white-bellied S. leucogaster (Figure 1). and HAPPOLD, 2013). This Scotophilus species has not yet been properly revised, but Eventually, molecular genetic approach to taxonomy has been known facts indicate that none of the available names apply to applied to the genus Scotophilus. TRUJILLO et al. (2009) published it, hence probably representing an undescribed species. Another a molecular phylogeny of Scotophilus based on sequences of the taxon surveyed was the giant S. nigrita (VALLO et al. 2015), whose mitochondrial (mt) gene for cytochrome b (cytb) and an additional Southern African population may represent a distinct species, for nuclear, male-only gene for zinc finger protein on the Y chromosome. which the name S. alvenslebeni Dalquest, 1965 would be available. While their study largely corroborated delimitation of species However, a thorough revision of additional material is needed before proposed by ROBBINS et al. (1985), they also found evidence for a definite decision is made. Finally, the sister relationship between polyphyly in the widely distributed yellow-bellied (morpho)species S. the rainforest species S. nucella and S. nux was confirmed recently dinganii and S. viridis (Figure 1). Three independent evolutionary by VALLO et al. (2016). units have been shown in both taxa, respectively, with S. dinganii Until recently, there were eight named Scotophilus species s. str. from South Africa being the only straightforward taxonomic recognized on the African mainland, excluding S. alvenslebeni,

Figure 1: Phylogenetic relationships of African Scotophilus species based on phylogenies published by TRUJILLO et al. (2009) and VALLO et al. (2011, 2013, 2015, 2016). All nodes are supported. Positions of South African S. viridis, S. nigrita and S. robustus are depicted in an unresolved multifurcation. Polyphyletic evolutionary units within S. dinganii and S. viridis morphospecies are labelled according to geographical region and recognized or presumed names. Numbers in brackets identify clades as delimited by TRUJILLO et al. (2009) and used by BOOKS and BICKHAM (2014). Shaded area includes species not discussed in detail in the text.

Vallo and Van Cakenberghe (2017) Scientific contributions Advances in taxonomy African Scotophilus African Bat Conservation News 6 August 2017 vol. 46 ISSN 1812-1268

but including S. colias, and three species on Madagascar, making a reasonable provisional solution. Since S. colias was described it eleven in total for the Afrotropics. Furthermore, there was one from central Kenya and the population from the Kakamega forest in distinctive West African species not yet described. Another taxon, southwestern Kenya belongs to the West African S. dinganii clade, Scotophilus borbonicus (Geoffroy, 1803), into which many African there is also a certain possibility that S. colias might represent a species were included at one time or another (see DOBSON, 1878), senior synonym to the newly introduced S. livingstonii. Anyway, a is not included into this account, given its obscure and unidentifiable more thorough analysis including at least the type specimen of S. content, making it a nomen dubium (see also ROBBINS et al., 1985; colias should have been undertaken before introducing these new GOODMAN et al., 2005). In addition to these species, a handful of East African names. The available candidate names also should not new ones were introduced by BROOKS and BICKHAM (2014), largely have been ignored completely. based on the molecular phylogeny by TRUJILLO et al. (2009). They The last species introduced by BROOKS and BICKHAM (2014), revised morphological traits of four of the clades of the S. dinganii S. trujilloi, harbours another problem. A major caveat in the and S. viridis morphospecies highlighted in the latter paper, and identification of any bat specimen as S. viridis is that reference respectively numbered 8, 9, 11, 12 in the original study (Figure 1). specimens in museum collections labelled S. viridis may in fact be The following names have been given to these clades: Scotophilus erroneously identified. Historically, there has actually been a large ejetai (8), Scotophilus andrewreborii (9), Scotophilus livingstonii misunderstanding in distinguishing S. viridis from S. leucogaster (11), and Scotophilus trujilloi (12). Unfortunately, the authors only damarensis (Thomas, 1906), which is another Southern African used univariate analyses of five external and eight skull dimensions form of similar size. Many museum specimens of S. viridis do rather than multivariate comparisons such as principal components not conform to the original description of S. viridis, which clearly and discriminant function analyses. These latter techniques usually states yellow colouration of belly (see MONADJEM et al. 2010). provide convincing delimitation of distinct morphotypes if these Furthermore, the presumption on the comparative specimens of are present in the dataset. In the S. dinganii morphospecies, it S. viridis s. str. from Mozambique to be topotypic cannot be really would have been intriguing to see four distinct morphogroups corroborated. S. viridis was not described from the inland Tete corresponding to S. dinganii s.str. and the three new species, as this province, where the comparative specimens originated, but from would have helped to support the diagnostic features discriminating the Island of Mozambique, which is an island off the coast in the among them. The genetic distinction alone of the West and East Nampula province, some five hundred kilometres to the east. Thus, African clades from S. dinganii s.str. obviously substantiates their it would have been much more appropriate to simply compare the recognition as separate species, given the known interspecific Kenyan specimens to the type specimen of S. viridis than to rely variation in Scotophilus and generally in bats (BAKER and BRADLEY on non-revised museum specimens. It cannot be excluded that 2006). On the other hand, BROOKS and BICKHAM (2014) did not the specimens from Kenya are actually S. viridis s. str. and the demonstrate morphological differences distinguishing the three introduction of the new name S. trujilloi for this yellow-bellied Kenyan new species S. ejetai, S. andrewreborii, and S. livingstonii from each form may thus have been premature. other, making their diagnosis unconvincing. As indicated in this short note, African Scotophilus continues Especially between the two proposed East African sister species, to be a controversial bat group despite advances in its taxonomy the Ethiopian S. ejetai and Kenyan S. andrewreborii, a careful in the last decade. Molecular data have immensely improved argumentation should have been provided in order to exclude perception of largely acknowledged species and created a basis for phenotypic plasticity as a possible explanation of the presented the recognition of other, yet unknown or unsuspected ones. On the morphological differences. Interestingly, based on cytb sequences other hand, interpretation of these data may not always be simple. A certain specimens within S. andrewreborii differ more from their rigorous taxonomic approach and caution must be applied, in order presumed conspecifics (up to 4.4% uncorrected genetic distance) not to increase confusion. Regarding the latter four newly introduced than from individuals of the sister species S. ejetai, as the genetic species, it is advisable to restrict their use until thoroughly revised. distance between these two species varies 3.8–5.4%. Moreover, Definitely, once they are validated, taxonomy of African Scotophilus monophyly of S. andrewreborii may not be regarded as convincingly will become a bit less confusing. supported (see TRUJILLO et al., 2009, VALLO et al., 2011). In no way would the authors of this note doubt the existence of distinct Acknowledgements evolutionary units, possible species, in the heterogeneous We would like to thank Ernest Seamark, Jakob Fahr and Petr Benda environment of East Africa. The genetic difference between Southern for their critical comments and suggestions to this manuscript, as well as encouraging and inspiring discussions on African Scotophilus. African S. dinganii s. str. and West African S. livingstonii reaches similar values (4.8–5.4%), so genetic distinction of these two East Literature cited African clades seems substantiated. Nevertheless, the question ACR 2016. African Chiroptera Report 2016. AfricanBats, Pretoria. i -xvii, remains whether the East Africa clade could not be regarded as just 1—7380 pp. url: http://www.africanbats.org/Documnets/ACR/2016/ one species with a high internal variability. This would be another ACR_2016.pdf. plausible option given the currently available data. BAKER, R. J. and BRADLEY, R. D. 2006. Speciation in mammals and the Another, more serious issue with regards to taxonomy by BROOKS genetic species concept. Journal of Mammalogy 87 (4): 643—662. doi: 10.1644/06-MAMM-F-038R2.1. and BICKHAM (2014), is the introduction of new species names BROOKS, D. M. and BICKHAM, J. W. 2014. New species of Scotophilus without reviewing available names already given to local forms. (Chiroptera: Vespertilionidae) from sub-Saharan Africa. Occasional In East Africa, three relevant names are readily available for the papers of the Museum of Texas Tech University 326: 1—21. url: http:// medium-sized yellow-bellied Scotophilus: colias Thomas, 1904, www.nsrl.ttu.edu/publications/opapers/ops/OP326.pdf. murino-flavus von Heuglin, 1861 and flavigaster von Heuglin, DOBSON, G. E. 1878. Catalogue of the Chiroptera of the collection of the 1861. The latter two names, originating from Eritrea, remain British Museum. Trustees of the British Museum, London. i–xlii, 1—567, somewhat obscure and type specimens are not easily accessible plates i–xxx. for comparison. Their synonymisation with S. leucogaster, however, GOODMAN, S. M., JENKINS, R. K. B. and RATRIMOMANARIVO, F. H. as suggested by KOOPMAN (1975) has not been validated (see 2005. A review of the genus Scotophilus (Mammalia, Chiroptera, Vespertilionidae) on Madagascar, with the description of a new species. ROBBINS et al. 1985) and therefore they still remain as possible Zoosystema 27 (4): 867—882. url: http://sciencepress.mnhn.fr/sites/ candidate names. On the other hand, the former name S. colias default/files/articles/pdf/z2005n4a8.pdf. is easily comparable. The morphological similarity of Ethiopian S. GOODMAN, S. M., RATRIMOMANARIVO, F. H. and RANDRIANANDRIANINA, dinganii and the type specimen of S. colias presented in VALLO et F. H. 2006. A new species of Scotophilus (Chiroptera: Vespertilionidae) al. (2011) seems supportive of the taxonomic affiliation of the East from western Madagascar. Acta Chiropterologica 8 (1): 21—37. doi: 10.3161/1733-5329(2006)8[21:ANSOSC]2.0.CO;2. African clade with the latter named form, and currently represents JACOBS, D. S. and BARCLAY, R. M. R. 2009. Niche differentiation in two

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sympatric sibling bat species, Scotophilus dinganii and Scotophilus systematic_revision_of_the_African_bat_genus_Scotophilus_ mhlanganii. Journal of Mammalogy 90 (4): 879—887. doi: (Vespertilionidae)/links/00b7d522ae89d7d687000000.pdf. 10.1644/08-MAMM-A-235.1. SIMMONS, N. B. 2005. Order Chiroptera. In: WILSON, D. E. and REEDER, JACOBS, D. S., EICK G. N., SCHOEMAN M. C. and MATHEE C. A. 2006. D. M. (eds), Mammal Species of the World, pp. 312—529. The Johns Cryptic species in an insectivorous bat, Scotophilus dinganii. Journal Hopkins University Press, Baltimore, MD, USA. of Mammalogy 87 (1): 161—170. doi: 10.1644/04-MAMM-A-132R2.1. TRUJILLO, R. G., PATTON, J. C., SCHLITTER, D. A. and BICKHAM, J. W. 2009. JACOBS, D. S., KELLY, E. J., MASON, M. and STOFFBERG, S. 2007. Molecular phylogenetics of the bat genus Scotophilus (Chiroptera: Thermoregulation in two free-ranging subtropical insectivorous bat Vespertilionidae): Perspectives from paternally and maternally species: Scotophilus species (Vespertilionidae). Canadian Journal of inherited genomes. Journal of Mammalogy 90 (3): 548—560.doi: Zoology 85: 883—890. doi: 10.1139/Z07-067. 10.1644/08-MAMM-A-239R2.1. KOCK, D. 1969. Die Fledermaus-Fauna des Sudan (Mammalia, Chiroptera). VALLO, P., BENDA, P. and REITER, A. 2011. Yellow-bellied or white-bellied? Abhandlungen der Senckenbergischen naturforschenden Gesellschaft Identity of Arabian house bats (Vespertilionidae: Scotophilus) revealed 521: 1—238. from mitochondrial DNA and morphology. African Zoology 46 (2): 350— KOOPMAN, K. F. 1975. Bats of the Sudan. Bulletin of the American Museum 361. doi: 10.1080/15627020.2011.11407508. of Natural History 154 (4): 353—444. VALLO P., BENDA, P., ČERVENÝ, J. and KOUBEK, P. 2013. Conflicting LANZA, B., FUNAIOLI, U. and RICCUCCI, M. 2015. The Bats of Somalia and mitochondrial and nuclear paraphyly in small-sized West African neighbouring areas. Edition Chimaira, Frankfurt am Main: 566 pp. house bats (Vespertilionidae). Zoologia Scripta 42: 1—12. doi: 10.1111/j.1463-6409.2012.00563.x. MONADJEM, A., TAYLOR, P. J., COTTERILL, F. P. D. and SCHOEMAN, M. C. 2010. Bats of Southern and Central Africa. A Biogeographic and VALLO P., BENDA, P., ČERVENÝ, J. and KOUBEK, P. 2015. Phylogenetic position Taxonomic synthesis. Wits University Press, Johannesburg: 596 pp. of the giant house bat Scotophilus nigrita (Chiroptera, Vespertilionidae). Mammalia 79 (2): 225—231. doi: 10.1515/mammalia-2013-0137. ROBBINS, C.B. 1978. Taxonomic identification and history of Scotophilus nigrita (Schreber) (Chiroptera:Vespertilionidae). Journal of Mammalogy VALLO, P., NKRUMAH, E. E., TEHODA, P., BENDA, P., BADU, E. K. and 59 (1): 212—213. doi: 10.2307/1379901. DECHER, J. 2016: Nutlet is a little nut: disclosure of the phylogenetic position of Robbins’ house bat Scotophilus nucella (Vespertilionidae). ROBBINS, C. B., DE VREE, F. and VAN CAKENBERGHE, V. 1985. A systematic Folia zoologica 65: 302—309. url: https://www.researchgate.net/ revision of the African bat genus Scotophilus (Vespertilionidae). publication/313574523_Nutlet_is_a_little_nut_Disclosure_of_the_ Annalen van het Koninklijk Museum voor Midden Afrika: Zoölogische phylogenetic_position_of_Robbins%27_house_bat_Scotophilus_ Wetenschappen 246: 51—84. url: https://www.researchgate.net/ nucella_Vespertilionidae. profile/Victor_Van_Cakenberghe/publication/256453727_A_

Appendix 1. Overview of the African Scotophilus names according to current state of knowledge, including the sources of description and type localities (see ACR, 2016). Scotophilus andrewreborii Brooks and Bickham, 2014 Occ. Pap. Mus. Texas Tech Univ., 326: 1, 8, fig. 2. Kenya: Rift Valley province, Nakuru district, 12 km S, 4 km E Nakuru [00°24‘S 36°07‘E]. Holotype: CM 98049: ad ♂, skin and skull. Collected by: Dr. Kimberlyn (Kim) Nelson; collection date: 7 October 1985; original number: 217. Muscle tissue sample TK 33143. Paratype: CM 98048: ad ♀, skin and skull. Collected by: Dr. Kimberlyn (Kim) Nelson; collection date: 7 October 1985; original number: 214. Muscle tissue sample TK 33140. Paratype: CM 98050: ad ♂, skin and skull. Collected by: Dr. Kimberlyn (Kim) Nelson; collection date: 8 October 1985; original number: 219. Muscle tissue sample TK 33149. Scotophilus borbonicus (E. Geoffroy Saint-Hilaire, 1803) Vespertilio borbonicus E. Geoffroy Saint-Hilaire, 1803 Cat. Mamm. Mus. nat. Hist. nat., Paris, 55. France: Ile Bourbon [=Réunion Island]. Lectotype: RMNH MAM.28508: ad ♂, mounted skin (skull not removed). Scotophilus colias Thomas, 1904 Scotophilus nigrita colias Thomas, 1904 Ann. Mag. nat. Hist., ser. 7, 13 (75): 207. Kenya: Fort Hall [=Murango] [00°43‘S 37°09‘E]. Holotype: BMNH 1902.7.6.11: ad ♂. Collected by: Mrs. Hildegarde Hinde; collection date: 25 January 1902; original number: 107. Presented/Donated by: Mrs. Hildegarde Hinde. Scotophilus dinganii (A. Smith, 1833) Vespertilio dinganii A. Smith, 1833 S. Afr. Quart. J., ser. 2, 1 (2): 59. South Africa: Between Port Natal (= Durban) and Delagoa Bay (=Maputo). Lectotype: NHRM 1065: mounted skin. Collected by: J. Wahlberg, collection date: 18 April 1841, from Caffer. Vespertilio epichrysus Temminck, 1832 [in Smuts] Enumer. Mamm. Capensium, 106. South Africa: Cape Town [33°56‘S 18°28‘E]. No type specimen known. Scotophilus nigrita herero Thomas, 1906 Ann. Mag. nat. Hist., ser. 7, 17 (98): 174. Namibia: Ovamboland, Olifants Vlei [approx. 18°00‘S 17°30‘E]. Holotype: BMNH 1907.1.1.449: ad ♀, skin and skull. Collected by: Mr. C.J. Andersson; collection date: 8 August 1859; original number: 260, originally in the Tomes collection. Nycticejus planirostris Peters, 1852 Naturw. Reise Mossamb., Säugethiere, p. 65, pl. 17, fig. 1, a-e. Mozambique: Zambezi, Tete [16°10’S 33°35’E] Holotype: ZMB 527: ad ♂, skin and skull. Collected by: Prof. Wilhelm Carl Hartwig Peters; collection date: March 1845. Scotophilus nigrita pondoensis Roberts, 1946 Ann. Transv. Mus., 20 (4): 304. South Africa: Transkei, Pondoland, Port St.Johns [31°27‘S 29°33‘E].

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Holotype SAMC 6858, collected by Captain G. C. Shortridge at Port St Johns, Pondoland, 22 August 1902. Scotophilus ejetai Brooks and Bickham, 2014 Occ. Pap. Mus. Texas Tech Univ., 326: 1, 14, fig. 5. Ethiopia: Orimaya Region, Dogy River Bridge [08°21‘43“N 35°53‘02“E]. Holotype: BRTC 57950: ad ♂, skin and skull. Collected by: Dr. Duane Albert Schlitter; collection date: 23 February 2001; original number: 10310. Muscle tissue sample AK 21235. Paratype: BRTC 57954: ad ♀, skin and skull. Collected by: Dr. Duane Albert Schlitter; collection date: 21 February 2001; original number: 10292. Tissue sample AK 21213. Paratype: CM 114043: ad ♀, skin and skull. Collected by: Dr. Duane Albert Schlitter; collection date: 2 April 1995; original number: 9986. Tissue sample SP 13027. Scotophilus leucogaster (Cretzschmar, 1826) Nycticejus leucogaster Cretzschmar, 1826 [in Rüppell] Atlas Reise Nördl. Afr., Zool. Säugeth., 1: 71, pl. 28a, 1, 2. Sudan: Kordofan, Brunnen Nedger (Nedger Well) [=Bir Nedger]. Lectotype: SMF 4309: Collected by: Wilhem Peter Edward Simon Rüppell. Mertens (1925: 22) old catalog II.O.1.b. Type: RMNH MAM.27432: ad. Scotophilus altilis G.M. Allen, 1914 Bull. Mus. comp. Zool., 58 (7): 350. Sudan: Blue Nile, north of [Er] Roseires, Aradeiba [12°25‘N 34°20‘E]. Holotype: MCZ 14463: ad ♂, skin and skull. Collected by: Glover Morrill Allen and The Phillips Sudan Expedition; collection date: 22 January 1913; original number: 73. Paratype: MCZ 14462: ad ♂, skin and skull (exchanged to FMNH). Paratype: MCZ 14610: ad ♂, alcoholic. Paratype: MCZ 14611: ad ♂, alcoholic. Scotophilus damarensis Thomas, 1906 Ann. Mag. nat. Hist., ser. 7, 17 (98): 175. Namibia: Ovamboland, Olifants Vlei [approx. 18°00‘S 17°30‘E]. Holotype: BMNH 1907.1.1.463: ad ♂, skin and skull. Collected by: Mr. C.J. Andersson; collection date: 10 August 1859; original number: 276, originally in the Tomes collection. Nycticejus flavigasterHeuglin, 1861 Nova Acta Acad. Cæs. Leop.-Carol., Halle, 29 (8): 5, 14. Eritrea: Kérén [approx. 15°45‘N 38°20‘E]. Syntype: SMNS 983: ad ♂, skull and alcoholic. Collected by: Martin Theodor von Heuglin; collection date: 1861. Syntype: SMNS 983a: ad ♀, skull and alcoholic. Collected by: Martin Theodor von Heuglin; collection date: 1861. Nycticejus murino-flavus Heuglin, 1861 Nova Acta Acad. Cæs. Leop.-Carol., Halle, 29 (8): 5, 15. Eritrea: near Massaua, M'kullu [15°38‘N 39°24‘E]. Type: [Unknown] ad ♂. Scotophilus livingstonii Brooks and Bickham, 2014 Occ. Pap. Mus. Texas Tech Univ., 326: 1, 11, fig. 4. Kenya: Western province, Kakamega district, Ikuywa River Bridge, 6.5 km S, 19 km E Kakamega [00°13‘N 34°55‘ E]. Holotype: CM 98051: ad ♂, skin and skull. Collected by: Dr. Duane Albert Schlitter; collection date: 8 November 1985; original number: 7394. Muscle tissue sample TK 33534. Paratype: CM 98053: ad ♀, skin and skull. Collected by: Dr. Duane Albert Schlitter; collection date: 8 November 1985; original number: 7396. Muscle tissue sample TK 33536. Paratype: USNM 412150: ad ♀, skin and skull. Collected by: Bruce Joliffe Hayward; collection date: 22 July 1967; original number: 4135. Collected at University of Ghana Botanical Garden (5º40'N, 0º12'W), Legon, Eastern (Greater Accra) Region, Ghana. Scotophilus marovaza Goodman, Ratrimomanarivo and Randrianandrianina, 2006 Acta Chiropt., 8 (1): 23, figs 2 - 5. Madagascar: Mahajanga province, Marovaza [14°56‘S 47°16‘E]. Holotype: FMNH 184050: ad ♂, skull and alcoholic. Collected by: Steven M. Goodman and Fania H. Ratrimomanarivo; collection date: 11 December 2004. Paratype: FMNH 184051: Collected by: Steven M. Goodman and Fania H. Ratrimomanarivo; collection date: 21 - 22 April 2005. locality: Madagascar: Province de Mahajanga - Marovaza [14 56 S 47 16 E, 5 m asl]. Paratype: FMNH 184052: Collected by: Steven M. Goodman and Fania H. Ratrimomanarivo; collection date: 21 - 22 April 2005. Locality: Madagascar: Province de Mahajanga - Marovaza [14 56 S 47 16 E, 5 m asl]. Paratype: FMNH 184085: Collected by: Steven M. Goodman and Fania H. Ratrimomanarivo; collection date: 21 - 22 April 2005. Locality: Madagascar: Province de Mahajanga - Marovaza [14 56 S 47 16 E, 5 m asl]. Paratype: FMNH 184086: Collected by: Steven M. Goodman and Fania H. Ratrimomanarivo; collection date: 21 - 22 April 2005. Locality: Madagascar: Province de Mahajanga - Marovaza [14 56 S 47 16 E, 5 m asl]. Paratype: MNHN ZM-MO-1984-433: collection date: 29 April 1869. Paratype: UADBA 46965: Collected by: Richard K.B. Jenkins; collection date: 24 November 2004; original number: RBJ 215. Locality: Madagascar: Antafinimihakatra, Parc National d' Ankarafantsika [16 16 S 46 48 E]. Scotophilus nigritellus de Winton, 1899 Ann. Mag. nat. Hist., ser. 7, 4 (23): 355. Ghana: Gambaga [10°32‘N 00°26‘W]. Holotype: BMNH 1899.6.15.9, ad ♂, collected and presented by Captain W. Giffard, collection date: 9 January 1899. Scotophilus nigrita (Schreber, 1774) Vespertilio Nigrita Schreber, 1774 Die Säugethiere, 1 (8): pl. 58 + 1 (9): 171. Senegal: Senegal. No type specimen known. Type specimen of V. nigrita originated from Senegal, Senegal River. Lanza et al., 2015 suggested emendation for the species name nigrita, which is an adjective in femininum form, to masculinum nigritus, in order to conform to the gender of both genus names. Why J.C.D. Schreber at his time, and his classically educated scholarly followers for further two centuries, should have used a gramatically wrong name combination, remains unclear. An alternative explanation of the species name nigrita given by Lanza et al. (2015) would be misspelling of the Greek substantive nigritia used in aposition. Scotophilus alvenslebeni Dalquest, 1965 J. Mamm., 46 (2): 254, 258. Mozambique: S bank Save River, Zinave National Park [approx. 21°26‘S 33°52‘E]. Holotype: KU 105222: ad ♂, skin and skull. Collected by: Prof. Dr. Walter Woelberg Dalquest; collection date: 9 October 1963; original number: MWUC

Advances in taxonomy African Scotophilus Scientific contributions Vallo and Van Cakenberghe (2017) African Bat Conservation News ISSN 1812-1268 August 2017 vol. 46 9

18767. Scotophilus gigas Dobson, 1875 Ann. Mag. nat. Hist., ser. 4, 16 (92): 122. Nigeria: Lagos [06°27‘N 03°23‘E]. Type: BMNH 1872.10.24.5: ad ♀, skull and alcoholic. Scotophilus nucella Robbins, 1983 Ann. Kon. Mus. Mid. Afr. Zool. Wetensch., 273: 19. Ghana: Eastern region, 1 mi N Nkawkaw [06°33‘N 00°44‘W]. Type: USNM 412141: ad ♂, skin and skull. Collected by: Bruce Joliffe Hayward; collection date: 5 August 1967; original number: 4256. Scotophilus nux Thomas, 1904 Scotophilus nigrita nux Thomas, 1904 Ann. Mag. nat. Hist., ser. 7, 13 (75): 208. Cameroon: Efulen [02°46‘N 10°42‘E]. Holotype: BMNH 1903.2.4.5: ad ♂. Collected by: George Latimer Bates. Scotophilus robustus A. Milne-Edwards, 1881 C. R. Acad. Sci., Paris, 91: 1035. Madagascar: Madagascar. Paratype: MNHN ZM-MO-1997-1883b: ad ♀, alcoholic (skull not removed). Collected by: Léon Humblot. Number 218a in Rode (1941: 248). Goodman et al. (2005: 873) additionally mention that the jar with the type specimen contains two specimens (1997-1883a and 1997-1883b), but that not can be determined which of the two is the holotype. They also mention the locality as "entre Foulpointe et le lac d'Alaoutre" in the central portion of het eastern humid forest. Type: MNHN ZM-MO-1997-1883a: ad ♀, alcoholic (skull not removed). Collected by: Léon Humblot. Number 218 in Rode (1941: 248). Scotophilus tandrefana Goodman, Jenkins and Ratrimomanarivo, 2005 Zoosystema, 27 (4): 867, 875, figs 3 - 5. Madagascar: Mahajange province, just outside the limit of the Parc national de Bemaraha, 1.8 km SE from Bekopaka, and 0.6 km NE from Andadoany [19°08.454‘S 44°48.732‘E]. Holotype: UADBA 46923: ad ♂, skull and alcoholic. Collected by: Richard K.B. Jenkins and Fania H. Ratrimomanarivo; collection date: 27 July 2003; original number: RBJ 161. Scotophilus trujilloi Brooks and Bickham, 2014 Occ. Pap. Mus. Texas Tech Univ., 326: 1, 14, fig. 5. Kenya: Coast province, Kwale district, Moana Marine Station, 1 km S, 2 km E Ukunda [04°18‘S 39°35‘E] Holotype: CM 98038: ad ♂, skin and skull. Collected by: Dr. Duane Albert Schlitter; collection date: 18 October 1985; original number: 7086. Muscle tissue sample TK 33263. Paratype: CM 98040: ad ♀, skin and skull. Collected by: Dr. Duane Albert Schlitter; collection date: 18 October 1985; original number: 7089. Tissue sample TK 33266. Paratype: CM 98041: ad ♀, skin and skull. Collected by: Dr. Duane Albert Schlitter; collection date: 18 October 1985; original number: 7090. Tissue sample: TK 33267. Scotophilus viridis (Peters, 1852) Nycticejus viridis Peters, 1852 Naturw. Reise Mossamb., Säugethiere, 667, pl. 17, fig. 2, a-e. Mozambique: Mozambique Island [15°00‘S 40°42‘E]. Holotype: ZMB 73610: ad ♂, skin and skull. Collected by: Prof. Wilhelm Carl Hartwig Peters; collection date: March 1845. Formerly ZMB 526 [=An 16362]. S[cotophilus] mhlanganii Jacobs, Kelly, Mason and Stoffberg, 2007 Can. J. Zool., 85: 883. Submitted: 29 June 2015 Accepted: 08 August 2017 Managing Editor: E.C.J. Seamark

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