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NEOROMICIA HLANDZENI SP. NOV. LOWVELD SEROTINE Pipistrellus (Hypsugo) anchietae Hill & Harrison (1987) (In part): Zambia? Pipistrellus (Hypsugo) anchietae Koopman (1993) (In part): South Africa. Hypsugo anchietae Cotterill (1996): Zimbabwe. Hypsugo anchietae Kearney et al. (2002): KwaZuluNatal Province, South Africa. Hypsugoanchietae Kearney (2005): Zimbabwe, Limpopo and KwaZulu-Natal provinces, South Africa. Hypsugo anchietae Simmons (2005) (In part): Zimbabwe, South Africa. Hypsugo anchietae Monadjem et al. (2010) (In part): Zimbabwe, Mozambique, Eswatini, South Africa. Neoromicia anchietae Monadjem et al. (2020) (In part): Zimbabwe, Mozambique, Eswatini, South Africa. Laephotis anchietae Simmons & Cirranello (2022) (In part): Zimbabwe, Mozambique, Eswatini, South Africa. Holotype: Durban Natural Science Museum (DM) No. 8423. The adult male specimen collected by Ara Monadjem (field number AM 2066) on 5 September 2005, has muscle tissue in 90% ethanol and the body preserved in ethanol, with skull (Fig. 6) and baculum (Fig. 5) extracted and cleaned. cytb sequences are available from GenBank (No. KM 886057). Type locality: Eswatini (formerly Swaziland), Mlawula Nature Reserve, near the Siweni train siding in the north-eastern lowveld savanna region of the country (latitude 26.17998°S; longitude 32.04871°E). The specimen was netted in riparian vegetation over a small side channel of the Mbuluzi River at an elevation of 110 m a.s.l. Paratypes: DM No. 8422; Ditsong National Museum of Natural History (TM) No. 47718. Both adult males, with bacula, from Mlawula Nature Reserve, Eswatini. Etymology: eHlandzeni means the ‘lowveld’ or wilderness in the SiSwati language indicating the specific lowveld (low-lying savanna) habitat of the new species which differentiates it from N. anchietae known from higher-lying (> 1000 m a.s.l.) elevations. With SiSwati names, the prefix is typically dropped when a noun is turned into a name. Diagnosis: The species is distinguished genetically (11% divergence in cytb sequences) from its sister species, N. anchietae from Angola. When compared with craniometric data of positively identified (from either mtDNA sequences or baculum) N. hlandzeni from South Africa and Eswatini (N = 17; Table 2; Fig. 3), specimens of N. anchietae s.s. from Angola (N = 20) have distinctly smaller-sized skulls (Fig. 3; Table 2). Greatest skull length varies from 11.2 to 12.9 mm (mean 12.1 mm) in Angolan N. anchietae and from 12.7 to 13.5 mm (mean 13.1 mm) in N. hlandzeni. Based on material examined in this study, there is a definite and clear dental feature that distinguishes both N. anchietae and N. hlandzeni from P. hesperidus. Unless the teeth are worn, the inner, anterior (larger of two) upper incisor is clearly bifid in N. anchietae and N. hlandzeni, although sometimes this is only manifested as a ‘step’ in the tooth. Neither the step, nor the bifid condition was found in any P. hesperidus examined in this study. This condition of the inner upper incisor is confirmed byVan Cakenberghe & Happold (2013). The original description by Seabra (1900) refers to the inner upper incisor as ‘tricuspid’. The baculum is> 1.30 mm long and bears a bilobed tip and base, with one of the basal lobes more enlarged than the other. Eswatini and South African individuals of N. hlandzeni can be distinguished from Angolan and Zambia samples of N. anchietae (also> 1.30 mm long) by their lack of well-defined lateral projections of the tip lobes (Fig. 5). Description: In pelage coloration and general body size and appearance, the new species is indistinguishable from N. anchietae (see description of Seabra, 1900 in Van Cakenberghe & Seamark, 2020; Van Cakenberghe & Happold, 2013). The new bat represents a relatively small-sized vespertilionid species with relatively large ears. The thumb is relatively long and has a distinct white marking at the base, on the thumb pad (Fig. 7). The tragus is about half the length of the ear, with a rounded and convex posterior surface and basal lobe. The antitragus is clearly visible and triangular in shape. The pelage is longish (up to 6 mm in length), bicoloured (hairs darker below), dark brown to yellowish brown dorsally and light brown, cream or white ventrally, darker in the pelvic area. The skull is relatively small and fragile for an African vespertilionid with a ‘notably depressed head’, i.e. concave lateral profile (Seabra, 1900; Fig. 6). There are five upper cheek teeth (including a minute anterior premolar), five lower cheek teeth and one canine and two upper and three lower incisors on each side. The anterior (inner) upper incisor is typically bifid or retains a ‘step’ in individuals with worn teeth. The posterior (outer) incisor is much smaller, about half the length of the anterior incisor or less. In cranial shape, the new species, together with N. anchietae, have an upturned end to the skull in the region of the incisors and canines relative to P. rusticus and P. hesperidus giving a concave lateral profile, whereas the others have a straighter profile. Furthermore, the zygomatic arch is not straight but has a wavy profile in N. anchietae and N. hlandzeni, whereas in P. rusticus and P. hesperidus it is straight. Based on eight genotyped and released individuals in the Eastern Cape Province of South Africa (Moir et al., 2020), of which mtDNA sequences of five were included in our molecular study (Fig. 2), the echolocation call parameters of this species are almost indistinguishable from those of P. hesperidus: duration = 2.73 ± 0.32 ms; maximum frequency = 87.09 ± 8.74 kHz; minimum f r e q u e n c y = 4 6.0 6 ± 1.2 6 k H z; c h a r a c t e r i s t i c f r e q u e n c y = 4 6. 6 3 ± 1. 2 9 k H z; f r e q u e n c y a t knee = 50.5 ± 2.08 kHz. Biology: The species seems to be associated with coastal and scarp forests in the Eastern Cape and KwaZuluNatal provinces of South Africa (Moir et al., 2020), the lowveld savanna of Eswatini and well wooded riparian areas in Zimbabwe and Mozambique (Monadjem et al., 2020). These habitats contrast somewhat with the higher-elevation habitat of N. anchietae based on the type specimen from Cahata in western Angola (c. 1300 m a.s.l.) and the localities in central Angolan represented in the current study (1000–1300 m a.s.l.). Further molecular sampling is required to determine whether previous records from Zambia, Botswana and the Democratic Republic of Congo (DRC) represent N. anchietae or N. hlandzeni. Preliminary bacular analyses and comparison with published records indicate that Zambian samples represent N. anchietae s.s., Published as part of Taylor, Peter John, Strydom, Erika, Richards, Leigh, Markotter, Wanda, Toussaint, Dawn Cory, Kearney, Teresa, D., F. P., Woody, Cotterill, Howard, Alexandra, Weier, Sina Monika, Keith, Mark, Neef, Gӧtz, Mamba, Mnqobi L., Magagula, Siphesihle & Monadjem, Ara, 2022, Integrative taxonomic analysis of new collections from the central Angolan highlands resolves the taxonomy of African pipistrelloid bats on a continental scale, pp. 1570-1590 in Zoological Journal of the Linnean Society 196 (4) on pages 1583-1586, DOI: 10.1093/zoolinnean/zlac071, http://zenodo.org/record/7390987, {"references":["Hill JE, Harrison DL. 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: 225 - 305.","Koopman KF. 1993. Chiroptera. In: Wilson DE, Reeder DM, eds. Mammal species of the world. A taxonomic and geographic reference, 2 nd edn. Washington DC: Smithsonian Institution Press, 137 - 241.","Cotterill FPD. 1996. New distribution records of insectivorous bats of the families Nycteridae, Rhinolophidae and Vespertilionidae (Microchiroptera: Mammalia) in Zimbabwe. Arnoldia Zimbabwe 10: 71 - 89.","Kearney TC, Volleth M, Contrafatto G, Taylor PJ. 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: 55 - 76.","Simmons NB. 2005. Order Chiroptera. In: Wilson DE, Reeder DM, eds. Mammal species of the world: a taxonomic and geographic reference, 3 rd edn. Washington DC: Smithsonian Institution Press / Johns Hopkins University Press, 312 - 529.","Monadjem A, Taylor PJ, Cotterill FPD, Schoeman MC. 2010. Bats of southern and central Africa: a biogeographic and taxonomic synthesis. Johannesburg: Wits University Press.","Monadjem A, Taylor PJ, Cotterill FPD, Schoeman MC. 2020. Bats of southern and central Africa: a biogeographic and taxonomic synthesis, 2 nd edn. Johannesburg: Wits University Press.","Simmons NB, Cirranello AL. 2022. Bat species of the world: a taxonomic and geographic database. Available at: https: // www. batnames. org /. Accessed on 23 August 2022.","Van Cakenberghe V, Happold M. 2013. Genus Pipistrellus. In: Happold M, Happold D, eds. Mammals of Africa. Volume IV: hedgehogs, shrews and bats. London: Bloomsbury Publishing, 600 - 608.","Seabra AF. 1900. Sobre um caracter importante para a determinacao dos generos e especies dos Microchiropteros e lista das especies d'este grupo existentes nas colleccoes do museu nacional. Jornal de Sciencias, Mathematicas, Physicas e Naturaes. Lisboa, Ser. 2 5: 16 - 35.","Van Cakenberghe V, Seamark ECJ, eds. 2020. African Chiroptera report 2020. Pretoria: AfricanBats NPC.","Moir MI, Richards LR, Rambau RV, Cherry MI. 2020. Bats of eastern Cape and southern KwaZulu-Natal forests, South Africa: diversity, call library and range extensions. Acta Chiropterologica 22: 365 - 381."]}

Taylor, Peter John, Strydom, Erika, Richards, Leigh, Markotter, Wanda, Toussaint, Dawn Cory, Kearney, Teresa, D., F. P., Woody, Cotterill, Howard, Alexandra, Weier, Sina Monika, Keith, Mark, Neef, Gӧtz, Mamba, Mnqobi L., Magagula, Siphesihle, Monadjem, Ara (2022): Integrative taxonomic analysis of new collections from the central Angolan highlands resolves the taxonomy of African pipistrelloid bats on a continental scale. Zoological Journal of the Linnean Society 196 (4): 1570-1590, DOI: 10.1093/zoolinnean/zlac071, URL: http://dx.doi.org/10.1093/zoolinnean/zlac071

Monadjem, Ara, Richards, Leigh R., Decher, Jan, Hutterer, Rainer, Mamba, Mnqobi L., Guyton, Jen, Naskrecki, Piotr, Markotter, Wanda, Wipfler, Benjamin, Kropff, Anna S., Dalton, Desire L. (2021): A phylogeny for African Pipistrellus species with the description of a new species from West Africa (Mammalia: Chiroptera). Zoological Journal of the Linnean Society 191 (191): 548-574, DOI: 10.1093/zoolinnean/zlaa068/5901436

PIPISTRELLUS SIMANDOUENSIS MONADJEM, RICHARDS, DECHER AND HUTTERER, SP . NOV . SIMANDOU SEROTINE LSID: http://zoobank.org/urn:lsid:zoobank.org:pub:A1F9BAC5-40BA-42FD-8D76-5284691656F4 Holotype: ZMFK-MAM-2008.0302, field number: JD 614. The bat was collected by Jan Decher and team. It is an adult male fixed in formalin and currently preserved in 70% alcohol, with the skull extracted and cleaned. Photographs of the head and tragus are illustrated in Figures 4 and 5, while the mandible and skull of the holotype are illustrated in Figure 6. Images of the entire skeleton of the holotype, including the skull, are presented in Supporting Information (Fig. S 1). The glans penis is illustrated in Figure 8 and the dorsal, ventral and lateral views of the baculum are presented in Figure 9. Type locality: Guinea, Macenta, Simandou Range, “Whiskey 1” as mapped in Decher et al. (2015) (Fig. 1). The bat was netted on 25 February 2008 across a small creek in a forested ravine on the east slope of the Simandou Range (08°32’53.84”N, 08°53’48.07”W) at an elevation of 950 m above sea level. Paratypes: T w o o t h e r b a t s, a f e m a l e (ZMFK MAM2008.0300; field number: JD 656) and a male (ZMFK-MAM-2008.0301; field number: JD 661) identified as belonging to this species were captured 5.9 km to the south-west of the type locality at “Foko confluence” on the west slope of the Simandou Range, on 8 and 9 March 2008, respectively, as mapped in Decher et al. (2015), in more humid forest habitat and at 737 m above sea level. Both these specimens have been sequenced and clearly group with the holotype (Fig. 2A) and can be considered as paratypes. Etymology: This species is named after the type region, the Simandou mountain range in eastern Guinea. Diagnosis: A m e d i u m - s i z e d p i p i s t r e l l o i d b a t, assignable to the genus Pipistrellus on the basis of the presence of a small anterior upper premolar (Hill & Harrison, 1987) and phylogeny (Fig. 2). Some members of the genus Parahypsugo may also have this anterior upper premolar but differ in skull morphology and shape of rhinarium (Hutterer et al., 2019), the cranium being more inflated in Pi. simandouensis than in any Parahypsugo species. Pi. simandouensis is readily distinguishable from Pi. hesperidus (its sister taxon) by its unicoloured pelage (Fig. 4) and shape of baculum (Fig. 5); however, these two species are indistinguishable in craniodental or external measurements. It is significantly larger in external and cranial features (see below) than Pi. nanulus Thomas, 1904, Pi. rusticus (Tomes, 1861) and Pipistrellus deserti Thomas, 1902. It differs from Pipistrellus inexspectatus Aellen, 1959 by its unicoloured pelage (bicoloured in the latter) and lack of white on the trailing edge of the wing membrane. Pi. (Vansonia) rueppellii is readily distinguished from all other Pipistrellus species by its pure white underparts and is currently placed in a separate genus Vansonia Roberts, 1946 (Moratelli & Burgin, 2019). Finally, the poorly diagnosed Pi. musciculus Thomas, 1913 (which may not even be a Pipistrellus; Hill & Harrison, 1987), is far smaller in external and cranial measurements. Description: External characters: Pi. simandouensis is a medium-sized pipistrelloid bat (similar in size to that of Pi. hesperidus), but large for the genus Pipistrellus (in fact it is the largest within the genus in Africa), with a total length of 80–86 mm and forearm length 31–34 mm (Table 3). The pelage is bright yellowishbrown and paler below than above (Fig. 4), with the individual hairs being unicoloured on both the upper and under parts. The patagium and uropatagium are both dark brown. Typically for the genus Pipistrellus, the ears are subtriangular in shape, rounded at the tip, and dark brown in colour (Fig. 4C). The tragus is also typical of the genus and is moderately long (roughly half of the length of the ear), relatively broad with a straight leading edge and convex outer edge (Fig. 9). It bears a markedly pointed projection, situated near the base and lying immediately below the indentation/notch of the outer margin. The rhinarium is as illustrated for the genus Pipistrellus in Hutterer et al. (2019), with the nostrils rounded in shape and obviously protruding from the snout. The external measurements of the holotype and other specimens of Pi. simandouensis are shown in Table 3. Craniodental characters: The skull is relatively robust for a Pipistrellus, while the rostrum is neither particularly broad nor narrow. The brain case is moderately inflated and rises distinctly and sharply above the level of the rostrum (Fig. 6) in contrast to the relatively flatter skulls of Parahypsugo spp. (Hutterer et al., 2019). The posterior of the skull does not end in an extended parietal/supraoccipital crest and the sagittal and lambdoid crests are poorly developed. The zygomatic arch is moderately robust. Cranial measurements for the holotype and other specimens of Pi. simandouensis are shown in Table 4. The dentition of Pi. simandouensis is I 2/3, C 1/1, P 2/3, M 2/3, which is typical of the genus Pipistrellus (Van Cakenberghe & Happold, 2013a). In the upper tooth row, I 1 is not bifid and I 2 is relatively smallsized, not reaching half the length of I 1 (Fig. 7A). P 1 is relatively small in all specimens examined, and in the toothrow, creating an obvious gap between C and P 2 (Fig. 7B). Dental measurements for the holotype and other specimens of Pi. simandouensis are shown in Table 5. The penis of Pi. simandouensis is long (8.4 mm in the holotype), straight and covered by long hairs all over its length (Fig. 8). These hairs are mostly white, with only a few on the dorsal side are brown. The terminal glans is wider than the shaft (Fig. 8). Total length of the penis is about 10% of total body length (Table 3). In its long and straight shape, the penis of Pi. simandouensis is similar to other species of the genus, such as Pipistrellus abramus (Temminck, 1840) and other Asian (Francis, 2019) and African Pipistrellus species (Benda et al., 2004). The bacula of Pi. simandouensis, Pi. grandidieri, Pi. hesperidus, Pi. nanulus and Pi. rusticus are presented in Figure 9. The baculum of Pi. simandouensis was notably shorter than the other species, with a total length of 1.32 mm. Baculum total length for the remaining species was: 2.00 mm (Pi. grandidieri), 1.47 mm (Pi. hesperidus), 2.98 mm (Pi. nanulus) and 1.52 mm (Pi. rusticus). Pi. simandouensis, Pi. hesperidus and Pi. rusticus were overall similar in morphology, presenting a slender-shafted baculum with a bilobed base and an expanded tip with two distinct prongs. However, Pi. simandouensis can be differentiated from Pi. hesperidus and Pi. rusticus on the basis of its robust, triangular base measuring 0.32 mm. Similarly, the projections of the tip in Pi. simandouensis are more pointed than in Pi. hesperidus and Pi. rusticus. The full suite of bacula measurements for all five species are provided in Supporting Information (Table S4). Biology: Pipistrellus simandouensis is currently known from two localities in West Africa, based on the four sequenced specimens (appearing in Fig. 2A), one specimen from Mount Nimba in northern Liberia (Monadjem et al., 2013) and three from the Simandou Range in south-eastern Guinea (Decher et al., 2015). This species has also been reported from the Fouta Djallon in central Guinea (Decher et al., 2015); however, this has not been confirmed genetically and we have not been able to examine these specimens. However, all these sites are from upland regions of West Africa, ranging in elevation from the foothill slopes at around 450 m a.s.l., to the summit of some of the peaks at over 1200 m a.s.l. and higher. Therefore, we suspect that this is a species closely associated with these upland forested habitats. Because such upland habitats are relatively restricted in West Africa, and few are legally protected (Monadjem et al., 2016), we furthermore predict that this is a species under threat. Hence, we recommend that the conservation status of this species be assessed as a matter of urgency., Published as part of Monadjem, Ara, Richards, Leigh R., Decher, Jan, Hutterer, Rainer, Mamba, Mnqobi L., Guyton, Jen, Naskrecki, Piotr, Markotter, Wanda, Wipfler, Benjamin, Kropff, Anna S. & Dalton, Desire L., 2021, A phylogeny for African Pipistrellus species with the description of a new species from West Africa (Mammalia: Chiroptera), pp. 548-574 in Zoological Journal of the Linnean Society 191 (191) on pages 559-563, DOI: 10.1093/zoolinnean/zlaa068/5901436, http://zenodo.org/record/4451311, {"references":["Decher J, Hoffmann A, Schaer J, Norris RW, Kadjo B, Astrin J, Monadjem A, Hutterer R. 2015. Bat diversity in the Simandou Mountain Range of Guinea, with the description of a new white-winged vespertilionid. Acta Chiropterologica 17: 255 - 282.","Tamura K, Stecher G, Peterson D, Filipski A, Kumar S. 2013. MEGA 6: molecular evolutionary genetics analysis version 6.0. Molecular Biology and Evolution 30: 2725 - 2729.","Hutterer R, Decher J, Monadjem A, Astrin J. 2019. A new genus and species of vesper bat from West Africa, with notes on Hypsugo, Neoromicia, and Pipistrellus (Chiroptera: Vespertilionidae). Acta Chiropterologica 21: 1 - 22.","Moratelli R, Burgin CJ. 2019. Family Vespertilionidae (vesper bats). In: Wilson DE, Mittermeier RA, eds. Handbook of the mammals of the world - Vol. 9. Barcelona: Lynx Edicions.","Thomas O. 1913. On African bats and shrews. Annals and Magazine of Natural History 11: 314 - 321.","Van Cakenberghe V, Happold M. 2013 a. Genus Pipistrellus. Pipistrelles. In: Happold M, Happold DCD, eds. The mammals of Africa. Vol. IV: hedgehogs, shrews and bats. London: Bloomsbury Publishing, 600 - 604.","Francis CM. 2019. Field guide to the mammals of South-East Asia. London: Bloomsbury Publishing.","Benda P, Hulva P, Gaisler J. 2004. Systematic status of African populations of Pipistrellus pipistrellus complex (Chiroptera: Vespertilionidae), with a description of a new species from Cyrenaica, Libya. Acta Chiropterologica 6: 193 - 217.","Monadjem A, Richards L, Taylor PJ, 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: 191 - 207.","Monadjem A, Richards L, Denys C. 2016. An African bat hotspot: the exceptional importance of Mount Nimba for bat diversity. Acta Chiropterologica 18: 359 - 375."]}

AFRONYCTERIS MONADJEM, PATTERSON & DEMOS GEN . NOV. LSID: http://zoobank.org/ urn:lsid:zoobank.org:pub: 71737F08-2938-4403-8385-5438B2E5EABE Synonymy Vespertilio Peters, 1852 (part, not Linnaeus, 1758). Hypsugo Kolenati, 1860 (part, not Kolenati, 1856). Vesperugo Dobson, 1875 (part, not Keyserling & Blasius, 1839). Pipistrellus Miller, 1900 (part, not Kaup, 1829). Myotis Matschie, 1907 (part, not Kaup, 1829). Neoromicia Shortridge, 1934 (part, not Roberts, 1926). Eptesicops Roberts, 1951 (part, not Roberts, 1926). Complete synonymic histories for the species placed herein in Afronycteris are given in the African Chiroptera report (AfricanBats NPC, 2019). as Neo. tenuipinnis). However, its distribution beyond western Kenya is not known. It seems to be associated with the high plateau of western Kenya, which extends into eastern Uganda; presumably, it also occurs there. Thorn & Kerbis Peterhans (2009) recorded ‘ Pipistrellus tenuipinnis ’ as occurring widely in Uganda. The cranial measurements of specimens from Budongo, Entebbe and Sango Bay (at elevations similar to those we report from Kenya) all fall neatly within the range of Pse. nyanza and are generally larger than those for Pse. tenuipinnis. It would be instructive to re-examine these specimens (in the collections of the BMNH and LACM) to confirm their identities and help to determine the western limits of the distribution of Pse. nyanza. However, records from the eastern Democratic Republic of the Congo apparently refer to true Pse. tenuipinnis, owing to their small size, with total length ‘about 72 mm’ (Allen et al., 1917). We speculate that, despite the limited geographical range of Pse. nyanza (even if Uganda is included), this species is currently not threatened because it survives in human-altered habitats, and therefore we recommend the IUCN conservation status of ‘Least Concern’. The type specimen echolocated at a peak frequency (start and end frequencies) of 40.4 kHz (56.4–39.3 kHz). The mean (± SD) peak frequency for 16 individuals at the type locality was 40.4 ± 0.84 kHz (55.1 ± 7.91 to 39.5 ± 0.68 kHz). Type species: Afronycteris nana (Peters, 1852). Included species: Afronycteris helios (Heller, 1912). Etymology: From the Greek word νυχτερίδα, bat, and the prefix Afro- referring to the African continent, referring to the wide distribution of the type species A. nana. This species ranges, without obvious breaks in distribution, from Senegal in the west, east to Ethiopia and south to South Africa, being absent only from the more arid desert and semi-desert environments associated with the Sahara, Sahel and Chalbi Desert in the north and the Namib and Kalahari deserts in the south-west (Happold, 2013a). Diagnosis: Small-sized vespertilionids with the simple muzzle characteristic of this family. The cranium in lateral view is distinctly inflated, more so than any other member of the tribe Vespertilionini. The tragus is characteristically hatchet shaped, with the posterior margin having an abrupt angle and lacking a notch at its base, as illustrated by Van Cakenberghe & Happold (2013). The tragi of Laephotis, Neoromicia and Pseudoromicia all have a notch at the base of the posterior margin. The pelage of the upper and under parts is bicoloured, with the basal portion of each hair darker than the terminal portion. There is a distinct thumbpad at the base of the thumb, thought to be useful in climbing on smooth leaves. The outer incisor I 2 is well developed, reaching almost the same length as the I 1, the latter being slightly bicuspid or unicuspid; in Laephotis, Neoromicia and Pseudoromicia, I 2 is typically half the length of I 1 or shorter. The P 1 is present and relatively large, whereas this tooth is absent in Laephotis, Neoromicia (except Neo. bemainty and Neo. anchietae) and Pseudoromicia. The baculum (~2.0 mm in length) is shorter than in Pseudoromicia and similar in length to that of Laephotis and Neoromicia. It has a distinctly and deeply bilobed base and a gently curved shaft leading to a spatulate tip (Fig. 5D). Distribution: This genus is endemic to sub-Saharan Africa, probably occurring in suitable habitats across its wide range. It occurs throughout the Upper Guinea rainforest zone, extending northward into Sudanian savanna, possibly extending into the Sahel along major rivers and wetlands (Happold, 2013a). It occurs throughout mesic portions of Central and East Africa, but records are sparser in the Horn of Africa (Lanza et al., 2015). It is widespread in the wetter parts of southern Africa, avoiding the dry southwestern region of South Africa, much of Botswana and Namibia (Monadjem et al., 2010). Systematic relationships: Afronycteris is sister to Pseudoromicia, but the two genera can be distinguished easily by external characteristics, cranial features and the shape of the baculum (see ‘Diagnosis’ above for details)., Published as part of Monadjem, Ara, Demos, Terrence C, Dalton, Desire L, Webala, Paul W, Musila, Simon, Kerbis Peterhans, Julian C & Patterson, Bruce D, 2020, A revision of pipistrelle-like bats (Mammalia: Chiroptera: Vespertilionidae) in East Africa with the description of new genera and species, pp. 1-33 in Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) (Zool. J. Linn. Soc.) 190 on pages 25-26, DOI: 10.1093/zoolinnean/zlaa087, http://zenodo.org/record/4451420, {"references":["Roberts A. 1926. Some new S. African mammals and some changes in nomenclature. Annals of the Transvaal Museum 11: 245 - 263.","AfricanBats NPC. 2019. African Chiroptera report 2019. Pretoria: AfricanBats NPC. Available at: https: // africanbats. org / publication / african-chiroptera-report- 2019 /","Thorn E, Kerbis Peterhans J. 2009. Small mammals of Uganda: bats, shrews, hedghog, golden-moles, ottertenrec, elephant-shrews, and hares. Bonner Zoologische Monographien 55: 1 - 164.","Allen JA, Lang H, Chapin JP. 1917. The American Museum Congo Expedition collection of bats. Bulletin of the American Museum of Natural History 37: 405 - 563.","Happold M. 2013 a. Pipistrellus nanus. Banana pipistrelle (banana bat). In: Happold M, Happold D, eds. The mammals of Africa. Volume IV: hedgehogs, shrews and bats. London: Bloomsbury Publishing, 639 - 642.","Van Cakenberghe V, Happold M. 2013. Genus Pipistrellus. Pipistrelles. In: Happold M, Happold DCD, eds. The mammals of Africa. Volume IV: hedgehogs, shrews and bats. London: Bloomsbury Publishing, 600 - 604.","Lanza B, Funaioli U, Riccucci M. 2015. The bats of Somalia and neighbouring areas. Frankfurt am Main: Chimaira.","Monadjem A, Taylor PJ, Cotterill FPD, Schoeman MC. 2010. Bats of southern and central Africa: a biogeographic and taxonomic synthesis. Johannesburg: University of the Witwatersrand Press."]}

LAEPHOTIS THOMAS, 1901 Synonymy Vespertilio A. Smith, 1829 (part, not Linnaeus, 1758). Hypsugo Kolenati, 1860 (part, not Kolenati, 1856). Scotophilus Thomas, 1861 (part, not Leach, 1821). Vesperugo Dobson, 1878 (part, not Keyserling & Blasius, 1839). Vesperus Jentink, 1887 (part, not Keyserling & Blasius, 1839). Eptesicus Matschie, 1897 (part, not Rafinesque,1820). Scabrifer G.M. Allen, 1908. Rhinopterus G.M. Allen, 1939 (part, not Miller, 1906). Pipistrellus Heller & Volleth, 1984 (part, not Kaup, 1829). Nycterikaupius (part, not Menu, 1987). Neoromicia Volleth et al., 2001 (part, not Roberts, 1926). Complete synonymic histories for the species of Laephotis are given in the African Chiroptera report (AfricanBats NPC, 2019). Description: This genus was originally created for the species Laephotis wintoni Thomas, 1901, with the name referring to the large ‘sail-like’ ears of that species. A second, closely related species with large ears was described a quarter of a century later, Lae. angolensis Monard 1935, and two more species by Setzer in 1971: Lae. botswanae and Lae. namibensis. The baculum (1.5–2.0 mm in length) of Laephotis as defined herein is shorter than in Pseudoromicia and similar in length to that of Neoromicia and Afronycteris. It has a characteristic shape, with a bilobed base, straight shaft and a spatulate tip that is at an angle of ~45° to the shaft (Fig. 5A). Based on our genetic and morphometric analyses presented above, we have expanded further this genus to include the following species: Lae. capensis (A. Smith, 1829), Lae. matroka (Thomas & Schwann, 1905), Lae. robertsi (Goodman et al., 2012), Lae. malagasyensis (Peterson et al., 1995) and Lae. stanleyi (Goodman et al., 2017). Laephotis is readily distinguished by its bacular morphology (Hill & Harrison, 1987). It is easily separated from Afronycteris based on external features (for details, see the account of Afronycteris). This genus may also be distinguished from Neoromicia by its larger size. Furthermore, the cranium is more robust in Laephotis and obviously flattened compared with Neoromicia and Pseudoromicia. Laephotis also lacks the white wings of Pseudoromicia and is mostly associated with arid savannas and grasslands. Of the nine species that we recognize in this genus, all except the one we describe here are restricted to eastern and southern Africa and Madagascar, and none is associated with rainforests of tropical Africa., Published as part of Monadjem, Ara, Demos, Terrence C, Dalton, Desire L, Webala, Paul W, Musila, Simon, Kerbis Peterhans, Julian C & Patterson, Bruce D, 2020, A revision of pipistrelle-like bats (Mammalia: Chiroptera: Vespertilionidae) in East Africa with the description of new genera and species, pp. 1-33 in Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) (Zool. J. Linn. Soc.) 190 on pages 14-15, DOI: 10.1093/zoolinnean/zlaa087, http://zenodo.org/record/4451420, {"references":["Roberts A. 1926. Some new S. African mammals and some changes in nomenclature. Annals of the Transvaal Museum 11: 245 - 263.","AfricanBats NPC. 2019. African Chiroptera report 2019. Pretoria: AfricanBats NPC. Available at: https: // africanbats. org / publication / african-chiroptera-report- 2019 /","Goodman SM, Taylor PJ, Ratrimomanarivo F, Hoofer SR. 2012. The genus Neoromicia (Family Vespertilionidae) in Madagascar, with the description of a new species. Zootaxa 3250: 1 - 25.","Goodman SM, Kearney T, Ratsimbazafy MM, Hassanin A. 2017. Description of a new species of Neoromicia (Chiroptera: Vespertilionidae) from southern Africa: a name for ' N. cf. melckorum '. Zootaxa 4236: 351 - 374.","Hill JE, Harrison DL. 1987. The baculum in the Vespertilioninae (Chiroptera: Vespertilionidae) with a systematic review, a synopsis of Pipistrellus and Eptesicus, and the description of a new genus and subgenus. Bulletin of the British Museum (Natural History): Zoology 52: 225 - 305."]}

Monadjem, Ara, Demos, Terrence C, Dalton, Desire L, Webala, Paul W, Musila, Simon, Kerbis Peterhans, Julian C, Patterson, Bruce D (2020): 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 (Zool. J. Linn. Soc.) 190: 1-33, DOI: 10.1093/zoolinnean/zlaa087, URL: http://dx.doi.org/10.1093/zoolinnean/zlaa087

LAEPHOTIS KIRINYAGA MONADJEM, PATTERSON, WEBALA & DEMOS SP.NOV. EAST AFRICAN SEROTINE LSID: http://zoobank.org/ urn:lsid:zoobank.org:pub: 71737F08-2938-4403-8385-5438B2E5EABE Synonymy Eptesicus capensis Kingdon (1974). Pipistrellus capensis garambe Thorn & Kerbis Peterhans (2009) (in part). Neoromicia capensis Patterson & Webala (2012). Neoromicia somalica Benda et al. (2016) (in part). Holotype: FMNH 234558, field number BDP 7516. This specimen was collected by Bruce D. Patterson, Paul W. Webala, Carl W. Dick and Beryl Makori. It is an adult male, with muscle tissue in liquid nitrogen, the body fixed in formalin and preserved in ethanol, now with skull extracted and cleaned. Type locality: Marsabit National Park, 1.3 km SE of campground near Headquarters, Marsabit County, Kenya (2.3090°N, 38.0001°E; Fig. 1). The type specimen was netted on 27 July 2015 at an elevation of 1280 m above sea level. Paratypes: One other male (FMNH 234559) was captured at the same location and on the same night as the type specimen and is considered a paratype. Seven other individuals (FMNH 234546, FMNH 234549– 234553, FMNH 234556–234557, four males and three females), were collected close to the type locality at elevations ranging from 1157 to 1356 m from 16 to 26 July 2015 (Supporting Information, Table S1); they closely resemble the holotype genetically (Fig. 3C) and morphologically (Tables 2–4) and are also considered paratypes. Etymology: The specific epithet is a Kikuyu word for Mount Kenya and reflects the distribution of the species in the northern highlands of Kenya. It is a noun in apposition. Diagnosis: This species is similar in size and appearance to its sister species Lae. capensis. It is easily distinguished from the long-eared Laephotis species by its shorter ears. Of the short-eared Laephotis species, Lae. stanleyi and Lae. robertsi are significantly larger in forearm length and most craniodental measurements (Tables 2–4). In contrast, Lae. malagasyensis is smaller, especially in cranial measurements (Table 3). Laephotis matroka is similar in external and craniodental measurements but is typically darker brown above and medium brown below (Goodman, 2011). In any case, Lae. robertsi, Lae. malagasyensis and Lae. matroka are all endemic to Madagascar (Goodman et al., 2012, 2017) and genetically distinct from Lae. kirinyaga (Fig. 3C). Laephotis kirinyaga closely resembles Lae. capensis, from which it differs by 8.3% on the Cytb gene (Supporting Information, Table S4). Externally, the two species are alike and broadly overlap in size, but Lae. kirinyaga is on average smaller in most measurements, particularly total length and forearm length (Table 2). Likewise, Lae. capensis is on average larger for all craniodental measurements (but with significant overlap), except greatest skull height, which is greater in Lae. kirinyaga. This is borne out in the lateral profile of the skull (Fig. 7), which is visibly flatter in Lae. capensis. These two species occupy mostly separate regions in multivariate space (Fig. 6), but again with some overlap. In contrast, the three specimens assigned to Lae. kirinyaga from Ethiopia and Guinea (labelled ‘cf. capensisW’ in Fig. 6), for which genetic data are lacking, fall completely within the multivariate space of the Lae. kirinyaga specimens (labelled ‘cf. capensis’) that have been sequenced. Measurements are presented as the mean ± SD, range and sample size (N). Measurements are of the holotype, other individuals of the new species and other ‘short-eared’ species of Laephotis. Measurements for the three Malagasy endemics, Lae. robertsi, Lae. matroka and Lae. malagasyensis, are taken from Goodman et al. (2017). Description: Exernal characters: Laephotis kirinyaga is a medium-sized pipistrelle-like bat, with strongly contrasting fur dorsally and ventrally. The dorsal pelage is medium brown, with most individual hairs being tipped light yellowish brown, giving the bat a brightly coloured appearance. The ventral pelage is cream–white to light cream–brown, with a dark base. The ears are short and rounded, and the tragus is curved distally on both anterior and posterior margins, Measurements are presented as the mean ± SD, range and sample size (N). Measurements are of the holotype, other individuals of the new species and other ‘short-eared’ species of Laephotis. Measurements for the three Malagasy endemics, Lae. robertsi, Lae. matroka and Lae. malagasyensis, are taken from Goodman et al. (2017). ending in a rounded tip, as in Lae. capensis (Monadjem et al., 2020b). The ears and muzzle are dark brown in colour, and the skin around the eyes is dark brown in the type specimen (Fig. 8A) but mostly pinkish in the paratypes. Craniodental characters: The skull is relatively robust, as in Lae. capensis, but less so than in Lae. stanleyi. In lateral profile, the cranium is distinctly straight, rising only gently up from the rostrum to the top of the braincase. An occipital ‘helmet’ is present but poorly developed, and the sagittal and lambdoidal crests are visible. The zygomatic arches are relatively robust (Fig. 7), as in Lae. capensis. The dentition in Lae. kirinyaga is typical of the genus, with I 2/3, C 1/1, P 1/3, M 2/3. In the upper tooth row, I 1 is unicuspid and I 2 is small, not reaching halfway up the length of I 1. The P 1 is absent, putting C in contact with P 2. The m3 is myotodont sensu Van Cakenberghe & Happold (2013). Biology: This species has been captured infrequently across the highlands of Kenya on both sides of the Rift Valley. It is present in wet tropical forest (e.g. Kakamega forest, with ~ 1900 mm of rainfall per annum), less mesic montane forest (Marsabit National Park) and relatively dry savanna woodlands (e.g. Lolldaiga Hills conservancy ~ 600 mm), hence aridity per se does not seem to be an important variable in its distribution. However, it has been recorded only at elevations> 1000 m (current records are all between 1160 and 1700 m), and this might be an important limit in its geographical distribution. We also include two specimens (FMNH 233035, 233036) from Murchison Falls National Park, Uganda (1180 m above sea level) in this new species. Two specimens from Ethiopia (identified as ‘ Neoromicia somalica ’ by Benda et al., 2016) also group with Lae. kirinyaga in the phylogeny, as does a specimen from Senegal (Koubínová et al., 2013), suggesting that this newly described species has a wide distribution north of the equator. We recommend, based on its relatively large distribution range and habitat preference, that it be listed as ‘Least Concern’ in the IUCN red list. However, we did not examine the specimens from Ethiopia and Senegal and therefore recommend a detailed morphological investigation before our hypothesis concerning the geographical range of this species is accepted. The type specimen echolocated at a peak frequency (start and end frequencies) of 44.9 kHz (74.3–41.6 kHz). The mean (± SD) peak frequency for 14 other individuals at the type locality was 43.9 ± 0.91 kHz (73.9 ± 9.43 to 41.8 ± 1.64 kHz)., Published as part of Monadjem, Ara, Demos, Terrence C, Dalton, Desire L, Webala, Paul W, Musila, Simon, Kerbis Peterhans, Julian C & Patterson, Bruce D, 2020, A revision of pipistrelle-like bats (Mammalia: Chiroptera: Vespertilionidae) in East Africa with the description of new genera and species, pp. 1-33 in Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) (Zool. J. Linn. Soc.) 190 on pages 15-18, DOI: 10.1093/zoolinnean/zlaa087, http://zenodo.org/record/4451420, {"references":["Kingdon J. 1974. East African mammals: an atlas of evolution in Africa. Volume IIA, insectivores and bats. New York: Academic Press Inc.","Thorn E, Kerbis Peterhans J. 2009. Small mammals of Uganda: bats, shrews, hedghog, golden-moles, ottertenrec, elephant-shrews, and hares. Bonner Zoologische Monographien 55: 1 - 164.","Patterson BD, Webala PW. 2012. Keys to the bats (Mammalia: Chiroptera) of East Africa. Fieldiana: Life and Earth Sciences 1563: 1 - 60.","Benda P, Reiter A, Uhrin M, Varadinova Z. 2016. A new species of pipistrelle bat (Chiroptera: Vespertilionidae) from southern Arabia. Acta Chiropterologica 18: 301 - 323.","Goodman SM. 2011. Les chauves-souris de Madagascar: guide de leur distribution, biologie et identification. Antananarivo: Association Vahatra.","Goodman SM, Taylor PJ, Ratrimomanarivo F, Hoofer SR. 2012. The genus Neoromicia (Family Vespertilionidae) in Madagascar, with the description of a new species. Zootaxa 3250: 1 - 25.","Goodman SM, Kearney T, Ratsimbazafy MM, Hassanin A. 2017. Description of a new species of Neoromicia (Chiroptera: Vespertilionidae) from southern Africa: a name for ' N. cf. melckorum '. Zootaxa 4236: 351 - 374.","Monadjem A, Taylor PJ, Cotterill FPD, Schoeman MC. 2020 b. Bats of southern and central Africa: a biogeographic and taxonomic synthesis, 2 nd edn. Johannesburg: Wits University Press.","Van Cakenberghe V, Happold M. 2013. Genus Pipistrellus. Pipistrelles. In: Happold M, Happold DCD, eds. The mammals of Africa. Volume IV: hedgehogs, shrews and bats. London: Bloomsbury Publishing, 600 - 604.","Koubinova D, Irwin N, Hulva P, Koubek P, Zima J. 2013. Hidden diversity in Senegalese bats and associated findings in the systematics of the family Vespertilionidae. Frontiers in Zoology 10: 48."]}

PSEUDOROMICIA MONADJEM, PATTERSON, WEBALA & DEMOS GEN.NOV. LSID: http://zoobank.org/ urn:lsid:zoobank.org:pub: 71737F08-2938-4403-8385-5438B2E5EABE Synonymy Vesperus Peters 1872 (part, not Keyserling & Blasius, 1839). Vesperugo Dobson 1878 (part, not Keyserling & Blasius, 1839). Eptesicus Matschie, 1897 (part, not Rafinesque,1820). Vespertilio Miller, 1900 (part, not Linnaeus, 1758). Pipistrellus Monard, 1935 (part, not Kaup, 1829). Nycterikaupius (part, not Menu, 1987). Neoromicia Kearney et al., 2002 (part, not Roberts, 1926). Complete synonymic histories for the species placed herein in Pseudoromicia are given in the African Chiroptera report (AfricanBats NPC, 2019). Type species: Pseudoromicia tenuipinnis (Peters, 1872). Included species: Pseudoromicia brunnea (Thomas, 1880); Pseudoromicia isabella (Decher, Hutterer & Monadjem, 2015); Pseudoromicia rendalli (Thomas, 1889); Pseudoromicia roseveari (Monadjem et al., 2013); Pseudoromicia tenuipinnis (Peters, 1872); and two newly described species (see below). Etymology: This feminine name is derived from the Greek prefix ψευδο-, false, and the genus Romicia Gray, 1838, in turn derived from the Ancient Greek word ρóμιξα, meaning a ‘kind of javelin or huntingspear’. It also hints at the genus Neoromicia, to which members of Pseudoromicia were previously assigned. Members of this new genus resemble and have in the past been confused with Neoromicia species. Diagnosis: T h e s e a r e s m a l l t o m e d i u m - s i z e d vespertilionids with a simple muzzle. The tragus is typically curved anteriorly, with a notch at the base of the posterior margin. The pelage of the upper and under parts is variably coloured, but in most species tends to be unicoloured dorsally and bicoloured ventrally. In contrast, dorsal pelage is bicoloured in Afronycteris, Laephotis and Neoromicia. Four of the seven species in this genus have translucent white wing membranes, whereas membranes are dark brown or blackish in colour in the remaining three species. The cranium is slightly inflated to relatively flattish in lateral profile; in contrast, it is highly inflated in Afronycteris and moderately inflated in Neoromicia s.s., whereas it is flattened in Laephotis. The outer incisors are usually half the length or less of the inner incisors, the latter being weakly bicuspid or unicuspid. The P 1 is absent, contrasting with Afronycteris, in which it is present and relatively large. The baculum (~3.0 mm in length) is distinctly longer than that of any of the other three genera previously included in Neoromicia, with a robust trilobed base and strongly arched shaft leading to a bilobed tip (Fig. 5C). Distribution: This genus is widely distributed across sub-Saharan Africa. However, all but one of the species is associated with equatorial tropical forest and woodland belt. One species, Pse. rendalli, extends far into savanna habitats, ranging from 13°N to 28°S. Systematic relationships: The genera Pseudoromicia and Afronycteris are sister to the genera Laephotis and Neoromicia as now understood (see below)., Published as part of Monadjem, Ara, Demos, Terrence C, Dalton, Desire L, Webala, Paul W, Musila, Simon, Kerbis Peterhans, Julian C & Patterson, Bruce D, 2020, A revision of pipistrelle-like bats (Mammalia: Chiroptera: Vespertilionidae) in East Africa with the description of new genera and species, pp. 1-33 in Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) (Zool. J. Linn. Soc.) 190 on pages 19-20, DOI: 10.1093/zoolinnean/zlaa087, http://zenodo.org/record/4451420, {"references":["Roberts A. 1926. Some new S. African mammals and some changes in nomenclature. Annals of the Transvaal Museum 11: 245 - 263.","AfricanBats NPC. 2019. African Chiroptera report 2019. Pretoria: AfricanBats NPC. Available at: https: // africanbats. org / publication / african-chiroptera-report- 2019 /","Decher J, Hoffmann A, Schaer J, Norris RW, Kadjo B, Astrin J, Monadjem A, Hutterer R. 2015. Bat diversity in the Simandou Mountain Range of Guinea, with the description of a new white-winged vespertilionid. Acta Chiropterologica 17: 255 - 282.","Monadjem A, Richards L, Taylor PJ, 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: 191 - 207."]}

PSEUDOROMICIA NYANZA MONADJEM, PATTERSON, WEBALA & DEMOS SP.NOV. NYANZA SEROTINE LSID: http://zoobank.org/ urn:lsid:zoobank.org:pub: 71737F08-2938-4403-8385-5438B2E5EABE Neoromicia tenuipinnis Patterson & Webala (2012). Neoromicia tenuipinnis Musila et al. (2019). Neoromicia tenuipinnis Rydell et al. (2020). Holotype: FMNH 215626, field number BDP 5719. This specimen was collected on 8 January 2012 by Bruce D. Patterson, Paul W. Webala and Carl W. Dick. It is an adult male, formalin-fixed and preserved in ethanol. Its skull has been extracted and cleaned, its glans penis removed and the baculum stained and extracted. Muscle tissue was also preserved in liquid nitrogen at the time of capture. Type locality: Kisumu Impala Sanctuary, State Lodge Campsite, Kisumu County (formerly Nyanza province), Kenya, at an elevation of 1130 m above sea level; geographical coordinates: 0.10961°S, 34.74593°E (Fig. 1). The sanctuary borders both Lake Victoria and Kenya’s fifth largest city, Kisumu, and is only 0.34 km 2 in area. Vegetation consisted of open parkland, shortstatured trees and shrubs. Paratypes: Four other individuals (FMNH 215625, FMNH 215627, FMNH 215628 and FMNH 215629), all females, were collected at the same location and on the same night as the holotype and closely resemble it genetically (Fig. 3B) and morphologically (Tables 5–7), qualifying them as paratypes. Etymology: This species is named after the region where it was found, Nyanza, which derives from the Bantu word for ‘large body of water’. Covering nearly 60 000 km 2, Lake Victoria surely qualifies. The name is used as a noun in apposition. Diagnosis: This is a medium-sized member of the genus Pseudoromicia, with a mean forearm length of 31.2 mm (Table 5) and greatest skull length of 12.96 mm (Table 6). It is genetically distinct from all other Pseudoromicia species (Fig. 3B). Furthermore, it is readily distinguished from the dark-winged members of this genus (Pse. roseveari, Pse. brunnea and Pse. kityoi) by its white wings. It can be distinguished from Pse. rendalli by its smaller size (mostly non-overlapping forearm length and craniodental measurements (Tables 5–7) and weakly bicuspid I 1 (unicuspid in Pse. rendalli). It is significantly larger than Pse. tenuipinnis, with hardly any overlapping external and craniodental measurements (Tables 5–7); furthermore, its dorsal fur is medium brown and bicoloured (dark brown and unicoloured in Pse. tenuipinnis). It is most like Pse. isabella in size and external appearance, but that species has rusty tips to the fur on its upper parts, whereas Pse. nyanza has white-tipped hairs. The taxon Eptesicus ater J. A. Allen, 1917, which was described from north-eastern Democratic Republic of the Congo, is currently considered a synonym of Pse. tenuipinnis (Simmons, 2005) and is far smaller than Pse. nyanza, with a reported total length of 68 mm. Furthermore, Pse. tenuipinnis has ‘brownish black’ fur on its back (Allen et al., 1917), contrasting with the light-tipped fur of Pse. nyanza. Description: External characters: Pseudoromicia nyanza is a medium-sized pipistrelle-like bat with white patagial and uropatagial membranes (Fig. 8B). The dorsal pelage is medium brown with white-tipped hairs over most of the back. The ventral hairs are pure white with a dark base. The ears are short and rounded, and the tragus is broad and truncated, as in Pse. tenuipinnis (Monadjem et al., 2013). Craniodental characters: The skull is relatively gracile, as in Pse. tenuipinnis and Pse. isabella. In lateral profile, the cranium slopes gently up from the rostrum to the top of the braincase. There is no occipital ‘helmet’, and the sagittal and lambdoidal crests are absent. The zygomatic arches are fragile, as in Pse. tenuipinnis and Pse. isabella (Fig. 10). The dentition in Pse. nyanza is typical of the genus, with I 2/3, C 1/1, P 1/3, M 2/3. In the upper tooth row, I 1 is weakly but distinctly bicuspid and I 2 is moderate in size, slightly more than half the length of I 1. The P 1 is absent, putting C in contact with P 2. The m3 is myotodont sensu Van Cakenberghe & Happold (2013). Biology: Judging by how frequently this species is captured, it is common west of the Rift Valley in Kenya (B. D. Patterson & P. W. Webala, personal observation). It seems to prefer forest-edge habitats and avoids the forest interior (Rydell et al., 2020

PSEUDOROMICIA KITYOI MONADJEM, KERBIS PETERHANS, NALIKKA, WASWA, DEMOS & PATTERSON SP.NOV. KITYO’ S SEROTINE LSID: http://zoobank.org/ urn:lsid:zoobank.org:pub: 71737F08-2938-4403-8385-5438B2E5EABE Holotype: FMNH 223211, field number JCK 7436. This specimen was collected by Betty Nalikka and Sadic Waswa Babyesiza during a field training exercise with Julian Kerbis Peterhans. It is an adult male preserved in ethanol, with skull extracted and cleaned, and tissue taken from breast muscle and preserved in dimethyl sulfoxide. Type locality: Mabira Forest Reserve, 0.79 km northeast of Nagojje Station, Mukono District of the Central Region, Uganda; geographical coordinates: 0.4451°N, 32.88876°E (Fig. 1). The type specimen was netted on 19 October 2012 in cultivated gardens directly adjacent (for a photograph of the type locality, see Fig. S2) to Mabira forest at an elevation of 1130 m above sea level. Paratype: One other male (FMNH 223555) was netted at the same location and on the same night as the holotype, and closely resembles it genetically (Fig. 3B) and morphologically (Tables 5–7) and can therefore be considered a paratype. Etymology: This species is named in honour of Dr. Robert M. Kityo, mammalogist, mentor and longserving curator at the Museum of Zoology, Makerere University, in recognition of his valuable contributions to bats and small mammal research in the region. His welcoming nature, curiosity, hospitality and support have facilitated numerous and diverse research agendas over the decades for both national and international researchers. Diagnosis: This is the largest member of the genus Pseudoromicia, with forearm length of 37 and 38 mm (Table 5) and greatest skull length of 14.70 and 14.99 mm for the two known specimens (Table 6). In comparison, the maximum greatest skull length in Pse. roseveari (which is the second largest member of the genus) is 14.5 mm (Table 6). Pseudoromicia brunnea is smaller in forearm length and in most craniodental measurements. Therefore, this species is readily diagnosable by size alone. It can easily be distinguished from the white-winged members of this genus (Pse. rendalli, Pse. isabella and Pse. tenuipinnis) by its dark wings. Description: External characters: Pseudoromicia kityoi is a large-sized pipistrelle-like bat, similar in size to the largest members of the Nycticeinops group, specifically Nyc. macrocephalus and Nyc. happoldorum, which were both described in the genus Paraphypsugo (Hutterer & Kerbis Peterhans, 2019; Hutterer et al., 2019). Despite its large size, this species is similar in external features to other black-winged members of Pseudoromicia. The pelage is medium brown above and slightly paler below. The individual hairs are unicoloured on the upper parts and bicoloured on the under parts, with the proximal half darker than the distal half. Like Pse. brunnea and Pse. roseveari, the patagium and uropatagium are both dark in colour. The ears are short and rounded, and the tragus has a curved outer margin as is typical of the genus (Monadjem et al., 2013). Craniodental characters: The skull is robust for a Pseudoromicia, even more so than in Pse. roseveari. The rostrum has a shallow depression, and the brain case is moderately inflated as in other members of the genus. There is no occipital ‘helmet’ as seen in the cranium of Lae. capensis (Monadjem et al., 2020b). The sagittal and lambdoidal crests are visible, and the zygomatic arches are robust for a pipistrelle-like bat (Fig. 9). The dentition in Pse. kityoi is typical of the genus, with I 2/3, C 1/1, P 1/3, M2/3. In the upper tooth row, I 1 is unicuspid and I 2 is tiny, extending slightly beyond the cingulum of I 1. The P 1 is absent, putting C in contact with P 2. The m 3 is myotodont sensu Van Cakenberghe & Happold (2013). Measurements are presented as the mean ± SD, range and sample size (N). Measurements are of the holotypes, other individuals of the two new species and other species of Pseudoromicia. The three species listed above the horizontal black line are dark winged, the four below are white winged (see main text for more details). Measurements are presented as the mean ± SD, range and sample size (N). Measurements are of the holotypes, other individuals of the two new species and other species of Pseudoromicia. The three species listed above the horizontal black line are dark winged, the four below are white winged (see main text for more details). Measurements are presented as the mean ± SD, range and sample size (N). Measurements are of the holotypes, other individuals of the two new species and other species of Pseudoromicia. The three species listed above the horizontal black line are dark winged, the four below are white winged (see main text for more details). Biology: Owing to the paucity of specimens, almost nothing can be said about the biology of this species. The only two known specimens were captured within 200 m from the edge of Mabira Forest in a domestic garden (Supporting Information, Fig. S2). However, considering that most members of this genus are restricted to tropical rainforest habitats, and that the two known specimens of this species were captured in a remnant patch of rainforest, its global distribution might be both fragmented and limited in extent. Urgent surveys are required to assess the status of this species at Mabira Forest Reserve, which has been steadily losing habitat to agriculture over the past few decades (Boffa et al., 2008). We suggest that this species might be present in other Congo Basin forest patches in Uganda (e.g. Semliki, Kibale, Kashyoha-Kitomi) and Kenya (Kakamega), although extensive surveys at Kakamega forest have failed to locate this species there (Webala et al., 2019). Owing to the limited information available on this species, we recommend that it be given the IUCN conservation status of ‘Data Deficient’, but we note that because of its presumed close association with rapidly disappearing forest habitat, this species is probably of conservation concern. Its closest known relative is Pse. roseveari, recently described from Mount Nimba and with a limited distribution in the borderland zone between Liberia and Guinea (Monadjem et al., 2013; Decher et al., 2015; Mamba et al., in press), some 4700 km to the west. Whether either species occurs in the vast tropical rainforests between these two sites is unknown and deserves investigation., Published as part of Monadjem, Ara, Demos, Terrence C, Dalton, Desire L, Webala, Paul W, Musila, Simon, Kerbis Peterhans, Julian C & Patterson, Bruce D, 2020, A revision of pipistrelle-like bats (Mammalia: Chiroptera: Vespertilionidae) in East Africa with the description of new genera and species, pp. 1-33 in Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) (Zool. J. Linn. Soc.) 190 on pages 20-23, DOI: 10.1093/zoolinnean/zlaa087, http://zenodo.org/record/4451420, {"references":["Hutterer R, Kerbis Peterhans J. 2019. A further new species of vesper bat from Central Africa (Chiroptera: Vespertilionidae). Lynx, n. s. (Praha) 50: 51 - 59.","Hutterer R, Decher J, Monadjem A, Astrin J. 2019. A new genus and species of vesper bat from West Africa, with notes on Hypsugo, Neoromicia and Pipistrellus (Chiroptera: Vespertilionidae). Acta Chiropterologica 21: 1 - 22.","Monadjem A, Richards L, Taylor PJ, 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: 191 - 207.","Monadjem A, Taylor PJ, Cotterill FPD, Schoeman MC. 2020 b. Bats of southern and central Africa: a biogeographic and taxonomic synthesis, 2 nd edn. Johannesburg: Wits University Press.","Van Cakenberghe V, Happold M. 2013. Genus Pipistrellus. Pipistrelles. In: Happold M, Happold DCD, eds. The mammals of Africa. Volume IV: hedgehogs, shrews and bats. London: Bloomsbury Publishing, 600 - 604.","Boffa JM, Kindt R, Katumba B, Jourget JG, Turyomurugyendo L. 2008. Management of tree diversity in agricultural landscapes around Mabira Forest Reserve, Uganda. African Journal of Ecology 46: 24 - 32.","Webala PW, Mwaura J, Mware JM, Ndiritu GG, Patterson BD. 2019. Effects of habitat fragmentation on the bats of Kakamega Forest, western Kenya. Journal of Tropical Ecology 35: 260 - 269.","Decher J, Hoffmann A, Schaer J, Norris RW, Kadjo B, Astrin J, Monadjem A, Hutterer R. 2015. Bat diversity in the Simandou Mountain Range of Guinea, with the description of a new white-winged vespertilionid. Acta Chiropterologica 17: 255 - 282."]}

Parahypsugo crassulus (Thomas, 1904) Broad-headed pipistrelle Pipistrellus crassulus Thomas, 1904: 206. Pipistrellus (Pipistrellus) crassulus: Hill and Harrison, 1987: 242, 272. Hypsugo crassulus: Heller et al., 1994: 3, 4, 5. Pipistrellus (Vansonia) crassulus: Koopman, 1994: 116. Hypsugo eisentrauti: Volleth and Heller, 1994: 13, 20, 26. Material examined SMF 79.441 (Rwanda), SMF 79.442 (DR Congo, Kivu region, Centre de Recherche en Sciences Naturelles, 850 m a.s.l.); FMNH 167775- 776, male (Gabon), FMNH 108168, male (Sudan), FMNH 83599, male (Angola), FMNH 165170-171, male (Uganda), FMNH 152767, male (Kenya), BMNH 1904.2.8.1 (holotype), SMF 79441 -442 (DR Congo). Comments The species occurs from Cameroon and Angola eastwards to Kenya (Fahr, 2013). Chromosome number of a specimen from DR Congo was: 2n = 30, aFN = 56 (Volleth et al., 2001). This is the smallest species of the genus (Tables 3 and 4)., Published as part of Hutterer, Rainer, Decher, Jan, Monadjem, Ara & Astrin, Jonas, 2019, A new genus and species of vesper bat from West Africa, with notes on Hypsugo, Neoromicia, and Pipistrellus (Chiroptera: Vespertilionidae), pp. 1-22 in Acta Chiropterologica 21 (1) on page 15, DOI: 10.3161/15081109ACC2019.21.1.001, http://zenodo.org/record/3752833, {"references":["THOMAS, O. 1904. New bats from British East Africa collected by Mrs. Hinde, and from the Cameroons by Mr. G. L. Bates. Annals and Magazine of Natural History, 7, 13: 206 - 210.","HILL, J. E., and D. L. HARRISON. 1987. The baculum in the Vespertilioninae (Chiroptera: Vespertilionidae) with a systematic review, a synopsis of Pipistrellus and Eptesicus, and the description of a new genus and subgenus. Bulletin of the British Museum of Natural History (Zoology), 72: 225 - 305.","HELLER, K. - G., M. VOLLETH, and D. KOCK. 1994. Notes on some verspertilionid bats from the Kivu region, Central Africa (Mammalia: Chiroptera). Senckenbergiana biologica, 74: 1 - 8.","KOOPMAN, K. F. 1994. Chiroptera: systematics. Pp. 1 - 217, in Handbook of Zoology. Volume 8: Mammalia, Part 60 (J. NIETHAMMER, H. SCHLIEMANN, and D. STARCK, eds). Walter de Gruyter, Berlin, vii + 224 pp.","VOLLETH, M., and K. - G. HELLER. 1994. Phylogenetic relationships of vespertilionid genera (Mammalia: Chiroptera) as revealed by karyological analysis. Zeitschrift fur Zoologische Systematik und Evolutionsforschung, 32: 11 - 34.","FAHR, J. 2013. Pipistrellus crassulus Broad-headed pipistrelle.. The mammals of Africa. Volume IV: Hedgehogs, shrews and bats (M. HAPPOLD and D. C. D. HAPPOLD, eds.). Bloomsbury Publishing, London, 800 pp.","VOLLETH, M., G. BRONNER, M. C. GOPFERT, K. - G. HELLER, O. VON HELVERSEN, and H. - S. YONG. 2001. Karyotype comparison and phylogenetic relationships of Pipistrellus - like bats (Vespertilionidae; Chiroptera; Mammalia). Chromosome Research, 9: 25 - 46."]}

Parahypsugo, gen. nov. Type species Parahypsugo happoldorum, sp. nov. Other species included in the new genus Parahypsugo crassulus (Thomas, 1904), P. eisentrauti (Hill, 1968) and P. bellieri (De Vree, 1972). Etymology A combination of the Greek prefix para (beside, near) with Hypsugo, emphasizing the close genetic and morphological relationships between the two genera. Distribution Tropical Africa from Guinea in the west to Sudan and Somalia in the east, and south to Congo, DRC, Uganda, Rwanda and Kenya (Patterson and Webala, 2012; Bates et al., 2013; Monadjem et al., 2013; Decher et al., 2016; Van Cakenberghe et al., 2017). This genus is probably widely distributed in the rainforest zone of Africa, and occurs from 70 m to over 2,200 m a.s.l. (Happold and Happold, 2013). Diagnosis Medium-sized bats of the Vespertilionidae family forming a clade between Hypsugo and Pipistrellus (Fig. 1) with a broad and simple rhinarium (Fig. 2), second incisor small with traces of second cusp (Fig. 6), smooth rostrum and strongly pointed margins (Fig. 4), external penis bent with long terminal hairs (Fig. 3a), and baculum straight with on the cingulum of the second incisor (not present in Hypsugo). Scale is 1 mm larger structure (Fig. 3b). on basal part; diameter curved Morphological comparisons Based on the phylogenetic tree, Parahypsugo is closely related to the genera Hypsugo, Neoromicia and Pipistrellus (Fig. 1) and hence we compare these genera morphologically. Externally, Parahypsugo is characterized by its broad and flat rhinarium (Fig. 2), where the Pars supranarica and internarica cover most of the Pars supralabialis. The skull of Parahypsugo has a smooth rostrum, while Hypsugo has two large and deep pits on either side of the rostrum. Neoromicia has two straight rims, while the skull of Pipistrellus shows a simple smooth rostrum (Fig. 4). Furthermore, in Parahypsugo the lateral margin of the rostrum is pointed, while the margin is straight in the other genera. Nycticeinops Hill and Harrison, 1987 is cranio-dentally somewhat similar but differs by the lack of a second upper incisor (Hill and Harrison, 1987). The new bat species During the survey of the Simandou Mountains in Guinea (Decher et al., 2016), we captured seven individuals of a small bat, four of which were preserved and three released. We first assigned them to the genus Hypsugo, after tentatively naming them Neoromicia cf. capensis: two in the canopy net over the creeks at Foko Confluence (8��29���48.62���N, 8��54���48.22���W; 737 m a.s.l.), one in a forest net at 2.3 km east Tourela (08��30���31.99���N, 008��56���59.60���W; 621 m a.s.l.), three over the creek in the Western Spur Valley (08��33���26.44���N, 08��55���16.50���W; 900 m a.s.l.) and one over the creek in gallery forest at Canga Moribadou Road (8��35���35.65���E, 8��51���25.85���W; 750 m a.s.l.). See Decher et al. (2016) for a map with these localities. The bats ranged in body mass from 7 to 14 g. Female ZFMK-MAM-2008.0295 carried two embryos. In the molecular analysis (Fig. 1) the four preserved specimens group with a specimen from Mount Nimba (DM 13225) previously labeled Neoromicia sp. 1 (Monadjem et al., 2013), which was collected in forest at an altitude of 1,400 m above sea level in the East Nimba Nature Reserve (Monadjem et al., 2016). This new species lacks the ���much reduced, small��� anterior premolar described by Hill and Harrison (1987: 244) as a characteristic of the genus Hypsugo, but all the remaining Parahypsugo species have this character (Fig. 7). Scale is 1 mm, Published as part of Hutterer, Rainer, Decher, Jan, Monadjem, Ara & Astrin, Jonas, 2019, A new genus and species of vesper bat from West Africa, with notes on Hypsugo, Neoromicia, and Pipistrellus (Chiroptera: Vespertilionidae), pp. 1-22 in Acta Chiropterologica 21 (1) on pages 7-9, DOI: 10.3161/15081109ACC2019.21.1.001, http://zenodo.org/record/3752833, {"references":["THOMAS, O. 1904. New bats from British East Africa collected by Mrs. Hinde, and from the Cameroons by Mr. G. L. Bates. Annals and Magazine of Natural History, 7, 13: 206 - 210.","HILL, J. E. 1968. Bats from the Cameroons, with the description of a new species of Pipistrellus. Bonner zoologische Beitrage, 19: 43 - 48.","DE VREE, F. 1972. Description of a new form of Pipistrellus from Ivory Coast. Revue de Zoologie et Botanique Africaine, 85: 412 - 416.","PATTERSON, B. D., and P. W. WEBALA. 2012. Keys to the bats (Mammalia: Chiroptera) of East Africa. Fieldiana: Life and Earth Sciences, 1563: 1 - 60.","BATES, P. J. J., K. CAMERON, M. J. PEARCH, and B. HAYES. 2013. A review of the bats (Chiroptera) of the Republic of Congo, including eight species new to the country. Acta Chiropterologica, 15: 313 - 340.","MONADJEM, A., L. R. RICHARDS, P. J. TAYLOR, and S. STOFFBERG. 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: 191 - 207.","DECHER, J., A. HOFFMANN, J. SCHAER, B. KADJO, A. MONADJEM, and R. HUTTERER. 2016. Bat diversity in the Simandou Mountain Range of Guinea, with the description of a new white-winged vespertilionid. Acta Chiropterologica, 17: 255 - 282.","VAN CAKENBERGHE, V., G. C. GEMBU TUNGALUNA, AKAWA MUSABA, P. SEAMARK, and E. VERHEYEN. 2017. The bats of the Congo and of Rwanda and Burundi revisited (Mammalia: Chiroptera). European Journal of Taxonomy, 382: 1 - 327.","HAPPOLD, M., and D. C. D. HAPPOLD (eds.). 2013. Mammals of Africa Volume IV: Hedgehogs, shrews, and bats. Bloomsbury Publishing, London, 800 pp.","HILL, J. E., and D. L. HARRISON. 1987. The baculum in the Vespertilioninae (Chiroptera: Vespertilionidae) with a systematic review, a synopsis of Pipistrellus and Eptesicus, and the description of a new genus and subgenus. Bulletin of the British Museum of Natural History (Zoology), 72: 225 - 305.","MONADJEM, A., L. R. RICHARDS, and C. DENYS. 2016. An African bat hotspot: the exceptional importance of Mount Nimba for bat diversity. Acta Chiropterologica, 18: 359 - 375."]}

Parahypsugo bellieri (De Vree, 1972) Bellier���s Pipistrelle Pipistrellus eisentrauti bellieri De Vree, 1972: 412. Hypsugo eisentrauti bellieri: Heller et al., 1994: 2���3. Nycticeinops eisentrauti: Hoofer and Van Den Busche, 2003: 14, 19, 27, 53; Hoofer et al., 2006: 982, 984. Hypsugo (crassulus) bellieri: Weber and Fahr, 2007: 92 /93. Pipistrellus crassulus: Fahr, 2013: 617. Hypsugo crassulus: Decher et al., 2016: 266. Material examined Holotype, MRAC 35686, female, skin and skull (C��te d���Ivoire, Adiopodoum��; coll. F. De Vree, 19 November 1969); ZFMK-MAM-2008.0299 (ZFMK-TIS-4905, -4906, -25943; ZFMK-DNA- 0100417869), female (Guinea, Simandou, Foko Confluence, 8 March 2008); ZFMK-MAM- 2009.0030 (ZFMK-TIS-4545, -4546, -25947; ZFMK-DNA-0100417618), male (Guinea, Simandou, Western Spur Valley, 12 December 2008). DM 12616, male (Liberia, East Nimba Nature Reserve, coll. Ara Monadjem 22 December 2010, altitude 1,200 m a.s.l.); DM 13221, male (Liberia, East Nimba Nature Reserve, coll. A. M. 31 December 2011, altitude 900 m a.s.l.); DM 13222, male (Liberia, 10 km west of Mount Nimba, coll. A. M. 8 January 2012, altitude 420 m a.s.l.). Comments This species was originally described as a subspecies of Pipistrellus eisentrauti (De Vree 1972), and subsequently assigned to Hypsugo crassulus (Heller et al., 1994), although Koopman et al. (1995) argued to keep both species in Pipistrellus and retain bellieri as a subspecies of P. eisentrauti. The holotype compares well in external and cranium dimensions with other specimens from Guinea and C��te d���Ivoire (Table 4). It is one of two smaller species of Parahypsugo which is known only from the Upper Guinea forest zone in Guinea, Liberia and C��te d���Ivoire. Fahr (2013) discussed it as a subspecies of Pipistrellus crassulus with specimens from seven localities from these countries, but noted that it may represent a distinct species, as did Monadjem et al. (2016). Happold and Happold (2013) published a drawing of the skull (Fig. 137d) of the holotype of P. bellieri., Published as part of Hutterer, Rainer, Decher, Jan, Monadjem, Ara & Astrin, Jonas, 2019, A new genus and species of vesper bat from West Africa, with notes on Hypsugo, Neoromicia, and Pipistrellus (Chiroptera: Vespertilionidae), pp. 1-22 in Acta Chiropterologica 21 (1) on page 16, DOI: 10.3161/15081109ACC2019.21.1.001, http://zenodo.org/record/3752833, {"references":["DE VREE, F. 1972. Description of a new form of Pipistrellus from Ivory Coast. Revue de Zoologie et Botanique Africaine, 85: 412 - 416.","HELLER, K. - G., M. VOLLETH, and D. KOCK. 1994. Notes on some verspertilionid bats from the Kivu region, Central Africa (Mammalia: Chiroptera). Senckenbergiana biologica, 74: 1 - 8.","HOOFER, S. R., R. A. VAN DEN BUSSCHE, and I. HORACEK. 2006. Generic status of the American pipistrelles (Vespertilionidae) with description of a new genus. Journal of Mammalogy, 87: 981 - 992.","WEBER, N., and J. FAHR. 2007. A rapid survey of small mammals from the Atewa Range Forest Reserve, Eastern Region, Ghana, Pp. 90 - 98, in A rapid biological assessment of the Atewa Range Forest Reserve, Eastern Ghana (J. MC- CULLOUGH, L. E. ALONSO, P. NASKRECKI, H. E. WRIGHT, and Y. OSEI- OWUSU, eds.). RAP Bulletin of Biological Assessment, 47: 1 - 194.","FAHR, J. 2013. Pipistrellus crassulus Broad-headed pipistrelle.. The mammals of Africa. Volume IV: Hedgehogs, shrews and bats (M. HAPPOLD and D. C. D. HAPPOLD, eds.). Bloomsbury Publishing, London, 800 pp.","DECHER, J., A. HOFFMANN, J. SCHAER, B. KADJO, A. MONADJEM, and R. HUTTERER. 2016. Bat diversity in the Simandou Mountain Range of Guinea, with the description of a new white-winged vespertilionid. Acta Chiropterologica, 17: 255 - 282.","KOOPMAN, K. F., C. F. KOFRON, and A. CHAPMAN. 1995. The bats of Liberia: systematics, ecology, and distribution. American Museum Novitates, 3148: 1 - 24.","MONADJEM, A., L. R. RICHARDS, and C. DENYS. 2016. An African bat hotspot: the exceptional importance of Mount Nimba for bat diversity. Acta Chiropterologica, 18: 359 - 375.","HAPPOLD, M., and D. C. D. HAPPOLD (eds.). 2013. Mammals of Africa Volume IV: Hedgehogs, shrews, and bats. Bloomsbury Publishing, London, 800 pp."]}

Parahypsugo happoldorum sp. nov. Happolds’ pipistrelle Neoromicia sp. 1: Monadjem et al., 2013: 194, 196, 199, 200, 205. Hypsugo sp.: Decher et al., 2016: 266; Monadjem et al., 2016: 369. Holotype ZFMK-MAM-2009.0029, field number JD 700, collected by Jan Decher, Blaise Kadjo and Juliane Schaer on 12 December 2008. Adult male preserved in 70% ethanol, skull extracted. Frozen tissue and DNA preserved (ZFMK-TIS-4547, -4548, and -25937; ZFMK-DNA-0100417629); COI and Cyt-b sequences deposited in GenBank (accessions KT598198 and MK188525). Type locality Guinea, Préfécture Macenta, Simandou Mountains, Western Spur Valley, 08°33‘26.44”N, 08°55‘ 16.50”W, ca. 737 m a.s.l., in net placed across a fastflowing creek in a ravine in submontane forest. Paratype ZFMK-MAM-2008.0295, ♀, with two embryos preserved (Guinea, Préfécture Macenta, Foko Confluence, 8 March 2008) (ZFMK-TIS-25935; ZFMK-DNA-0100417627). Referred specimens ZFMK-MAM-2008.0296 (ZFMK-TIS-4549, -25936; ZFMK-DNA-0100417628), ♀ (Guinea, 2.3 km E Tourela, 14 March 2008); ZFMK-MAM- 2009.0032 (ZFMK-TIS-4550, -4551, -5938; ZFMK- DNA-0100417630), ♀ (Guinea, gallery forest on the Canga-Moribadou Road, 18 December 2008); DM 13225 (Liberia, Mount Nimba, submontane forest, 26 December 2011); reported as Neoromicia sp. 1 by Monadjem et al. (2013) who suggested that it may refer to an undescribed new species; COI sequence GenBank accession: JX508832. All preserved in alcohol and skulls extracted. Diagnosis Largest species of Parahypsugo (Tables 4 and 5), as defined above. Total length 92–97 mm, tail length 28–37 mm, forearm 34.6–36.5 mm, ear length 9–12 mm, body mass 7.3–9.5 g (Table 4). Free tail tip 1.2 to 1.9 mm long (Table 1; longer in Hypsugo, shorter in Pipistrellus). Thumb long (Fig. 8). Length of penis 6.4 mm, baculum 4.4 mm, thumb 7.2 mm, tragus 4.0 mm (Table 2). Greatest length of skull 14.96 mm, length of C–M 3 15.20 mm (Table 5). Etymology We are pleased to dedicate this new bat to Meredith and David Happold (Canberra, Australia) (Fig. 9) in recognition of their studies on African bats, and particularly of their monumental work for the Mammals of Africa volume IV on hedgehogs, shrews, and bats (Happold and Happold, 2013). As a vernacular name we propose ‘Happolds’ Pipistrelle’. Description and comparison Parahypsugo happoldorum gen. nov., sp. nov. is the largest species currently recognized in the genus, with a total length of 90–97 mm. The tail length (28–37 mm), however, is longer in P. eisentrauti (36–40 mm), while forearm (34.6–36.5 mm) and body mass (7.3–9.5 g) are larger in P. happoldorum than in the other three species (Table 4). The short free tail tip measures 1.2–1.9 mm, and is shorter than in most Hypsugo species (1.5–5.5 mm — Table 1). Specimens from Guinea and the Nimba specimen are unicolored and medium brown on the dorsal and ventral surfaces. Dorsal hairs are 5 mm long on back and 3 mm at venter. Thumb, pad and lip are paler (Figs. 8 and 10). Parahypsugo happoldorum has a broad and simply built rhinarium (Fig. 2), and the face bears two round glandular areas surrounded by longer hairs which are missing in P. bellieri (Fig. 10). Ear length is 10 mm in the holotype (9–12 mm, n = 4), and its tip is rounded; the inner ear conch bears 3 folds. The tragus (Fig. 8) of the holotype is 4.0 mm long (Table 2) and represents 40% of the ear length. The thumb of the holotype is 7.2 mm long (Table 2 and Fig. 8), the longest in the genus. The forearm of the holotype is 35.8 mm long in P. happoldorum, slightly smaller but overlapping in P. eisentrauti, but significantly shorter in P. bellieri and P. crassulus (Table 4). The penis of the holotype specimen is 6.4 mm long and covered by short whitish hairs (Fig. 3 left); the proximal part is bent and not visible from the outside. A digital x-ray of its baculum measures 4.0 mm which is straight with a larger structure on its basal part and a slightly curved cross-section (Fig. 3 middle). The bacula of P. eisentrauti and P. bellieri are much shorter (Table 2), while the baculum of P. crassulus (Heller et al., 1994; Bates et al., 2013) is only slightly shorter. The cranium of P. happoldorum is illustrated in Figs. 4 and 11. The species is distinguished from other species (Fig. 12) by larger cranial and dental measurements, particularly in the greatest skull the eye. Photographs by J. Decher length and the upper and lower molar rows (Table 5). The dorsal profile is relatively flat (slightly more rounded in P. eisentrauti and P. bellieri). The large skull in P. happoldorum is mainly due to a different anterior design of the skull (Fig. 12). The rostrum is smooth and shows no traces of pits (as in Hypsugo sensu stricto — Table 6). The zygomatic arches are wide and extend posteriorly (Figs. 4 and 11). The I 1 has a second cusp, which is shorter than the first (P. happoldorum) or almost equal in length (P. eisentrauti, P. bellieri). The second upper incisor is small, about half the length of the first, and bears traces of a second cusp (Fig. 6). In Hypsugo, Neoromicia and Pipistrellus, the I 2 almost never bears traces of a second cusp (Fig. 6). Furthermore, P 1 is absent in the few P. happoldorum specimens that have been examined to date, but is present in all other species of the genus (Fig. 7). The upper molars are widest in P. happoldorum, and this is also true for P 2. Discussion Based on our molecular analyses, Parahypsugo happoldorum is split into two clusters (Fig. 1; p -distances of up to 6.7% in COI and up to 5.6% in Cyt-b) but we are unable to recognize reliable morphological characters, and we therefore refrain from describing a further new species. However, cryptic species may occur within this new species which is a topic for future studies. Specimen ZFMK-MAM-2009.0032, the only specimen caught in gallery forest on the more arid eastern side of the Simandou Ridge, is lighter on the belly, but the other specimens are brown. Cranially, the four species are quite similar. Differences are mainly due to the length of the rostrum (Fig. 9). Other species in the genus Currently we recognize three further species in the new genus, based on genetics (Fig. 1) and morphology (Figs. 7 and 9). However, we expect that further new species will be described in the future, particularly from the central and eastern parts of its range (see below).

Parahypsugo eisentrauti (Hill, 1968) Eisentraut���s Pipistrelle Pipistrellus eisentrauti Hill, 1968: 45. Pipistrellus (Hypsugo) eisentrauti: Hill and Harrison, 1987: 246, 274. Hypsugo eisentrauti: Hoofer et al., 2006: 991. Material examined ZFMK 1968.0005, holotype (Cameroon, Rumpi Hills); ZFMK 1968.0006, paratype (Cameroon, Mt. Kupe); ZFMK 1999.0676 (ZFMK-DNA- FC19442872 ��� Cameroon, Mt. Nlonako). Comments The history of the taxon eisentrauti is rather complicated. It was originally described in the genus Pipistrellus based on specimens from Cameroon (Rumpi Hills and Mt Kupe) (Hill, 1968). This was followed by the description of the taxon bellieri that was originally assigned to P. eisentrauti (see above). Both these taxa and P. crassulus were then moved to the genus Hypsugo and bellieri removed from P. eisentrauti to P. crassulus (Heller et al., 1994), although this was disputed by Koopman et al. (1995). In addition, new specimens of the taxon eisentrauti were reported from Kenya, Rwanda, Somalia and Uganda (Heller et al., 1994; Varty and Hill, 1988; Thorn and Kerbis Peterhans, 2009; see also Van Cakenberghe et al., 2017). However, specimens collected east of Cameroon probably represent one (or more) new species, which have not yet been studied in detail. Also a specimen (ROM 100532) used by Hoofer and Van Den Bussche (2003) to characterize P. eisentrauti in their tree represents a different species, most probably P. bellieri (Van Cakenberghe and Happold, 2013). Hoofer et al. (2006) listed the holotype (with misspelled type locality ���Rumysi Hilus��� = Rumpi Hills) and the paratype of P. eisentrauti, but in their trees they used the wrong sequence that had been published previously (Hoofer and Van den Busche, 2003). The same (but wrong) sequence was recently used again by Amador et al. (2018). Our tree (Fig. 1) includes a new specimen of P. eisentrauti from Mt. Nlonako, a mountain neighboring Mt. Cameroon, the type locality for this species., Published as part of Hutterer, Rainer, Decher, Jan, Monadjem, Ara & Astrin, Jonas, 2019, A new genus and species of vesper bat from West Africa, with notes on Hypsugo, Neoromicia, and Pipistrellus (Chiroptera: Vespertilionidae), pp. 1-22 in Acta Chiropterologica 21 (1) on page 15, DOI: 10.3161/15081109ACC2019.21.1.001, http://zenodo.org/record/3752833, {"references":["HILL, J. E. 1968. Bats from the Cameroons, with the description of a new species of Pipistrellus. Bonner zoologische Beitrage, 19: 43 - 48.","HILL, J. E., and D. L. HARRISON. 1987. The baculum in the Vespertilioninae (Chiroptera: Vespertilionidae) with a systematic review, a synopsis of Pipistrellus and Eptesicus, and the description of a new genus and subgenus. Bulletin of the British Museum of Natural History (Zoology), 72: 225 - 305.","HOOFER, S. R., R. A. VAN DEN BUSSCHE, and I. HORACEK. 2006. Generic status of the American pipistrelles (Vespertilionidae) with description of a new genus. Journal of Mammalogy, 87: 981 - 992.","HELLER, K. - G., M. VOLLETH, and D. KOCK. 1994. Notes on some verspertilionid bats from the Kivu region, Central Africa (Mammalia: Chiroptera). Senckenbergiana biologica, 74: 1 - 8.","KOOPMAN, K. F., C. F. KOFRON, and A. CHAPMAN. 1995. The bats of Liberia: systematics, ecology, and distribution. American Museum Novitates, 3148: 1 - 24.","VARTY, N., and J. E. HILL. 1988. Notes on a collection of bats from the riverine forests of the Jubba Valley, southern Somalia. Mammalia 52: 533 - 540.","THORN, E., and J. KERBIS PETERHANS. 2009. Small mammals of Uganda. Bonner zoologische Monographien, 55: 1 - 164.","VAN CAKENBERGHE, V., G. C. GEMBU TUNGALUNA, AKAWA MUSABA, P. SEAMARK, and E. VERHEYEN. 2017. The bats of the Congo and of Rwanda and Burundi revisited (Mammalia: Chiroptera). European Journal of Taxonomy, 382: 1 - 327.","HOOFER, S. R., and R. A. VAN DEN BUSSCHE. 2003. Molecular phylogenetics of the chiropteran family Vespertilionidae. Acta Chiropterologica, 5 (Supplement 1): 1 - 63.","VAN CAKENBERGHE, V., and M. HAPPOLD. 2013. Pipistrellus eisentrauti Eisentraut's pipistrelle. Pp. 621 - 622, in The mammals of Africa. Volume IV. Hedgehogs, shrews and bats (M. HAPPOLD and D. C. D. HAPPOLD, eds). Bloomsbury Publishing, London, 800 pp.","AMADOR, L. I., R. L. MOYERS AREVALO, F. C. ALMEIDA, S. A. CATALANO, and N. P. GIANNINI. 2018. Bat systematics in the light of unconstrained analyses of a comprehensive molecular supermatrix. Journal of Mammalian Evolution, 25: 37 - 70."]}

Hutterer, Rainer, Decher, Jan, Monadjem, Ara, Astrin, Jonas (2019): A new genus and species of vesper bat from West Africa, with notes on Hypsugo, Neoromicia, and Pipistrellus (Chiroptera: Vespertilionidae). Acta Chiropterologica 21 (1): 1-22, DOI: 10.3161/15081109ACC2019.21.1.001

Monadjem, Ara, Shapiro, Julie T., Richards, Leigh R., Karabulut, Hatice, Crawley, Wing, Nielsen, Ida Broman, Hansen, Anders, Bohmann, Kristine, Mourier, Tobias (2019): Systematics of West African Miniopterus with the description of a new species. Acta Chiropterologica 21 (2): 237-256, DOI: 10.3161/15081109ACC2019.21.2.001

Miniopterus nimbae sp. nov. Nimba long-fingered bat Holotype DM 12621 (field no. AM2010_12_18_1), an adult male, was collected by Ara Monadjem. The specimen was fixed in formalin and then transferred to 70% alcohol. The skull has been extracted and cleaned. Photograph of the skull and drawing of the tragus of the holotype are illustrated in Figs. 6 and 7. Type locality Liberia, Nimba Province, Mount Gangra, 10 km to the west of Mount Nimba (Fig. 1). The bat was netted on 18 December 2010 exiting from a mine adit mid-way up Mount Gangra (7.55434°N, 8.62902°W) at 720 m a.s.l, in secondary forest. Paratypes No other specimens of this species were captured or collected on the same day at the same site. However, the previous night an adult female (DM 12614) was collected at the Yiti River 9 km to the south-east of Mount Gangra. Photograph of the paratype is illustrated in Fig. 8. Etymology This species is named after Mount Nimba, one of just three localities from which it is known, further highlighting the critical importance of this mountain for bat conservation in Africa (Monadjem et al., 2016). Diagnosis This is a large-sized Miniopterus from Mount Nimba, Liberia, with a mean forearm length of 47.4 mm (n = 26 individuals). The large size of this bat (particularly its forearm length) readily distinguishes M. nimbae from all other African Miniopterus taxa except the M. inflatus / M. africanus group. In external and craniodental measurements, M. nimbae is similar in size to other members of the M. inflatus group (M. inflatus s.s., M. cf. inflatus and M. africanus) (Tables 3–5); however, in multi-dimensional morphospace based on craniodental measurements, it overlaps only with M. inflatus s.s. The taxon M. cf. inflatus from eastern and southern Africa tends to have a light pelage, being more reddish-brown in colour (compared with a deep chocolate brown in M. nimbae). It is not possible, at present to distinguish M. nimbae from M. inflatus s.s. on external characters. However, they can be readily distinguished by cranial features. In particular, the 1st upper premolar has an additional lingual cusp, posterior to the main cusp, that is present in M. nimbae but absent in M. inflatus s.s. (Fig. 9); this feature being clearly visible, even with a low magnification hand lens, and consistently present in all the specimens examined in this study. The lower tooth row (i 1 –m 3) and lower molar (LWMOLS) lengths are also slightly larger in M. nimbae than M. inflatus s.s. In terms of cranial geometry, M. nimbae differs from M. inflatus s.s. as it bears a slightly more gracile skull, less ‘inflated’ braincase, and the point of maximum curvature along the occiput is more elevated in the newly described taxon than in the nominate form. Furthermore, these two taxa, are also distinguishable on molecular grounds (K2P pairwise genetic distance = 1.6%). Additionally, the ranges of the two taxa do not appear to overlap. Miniopterus africanus appears to be restricted to north-eastern Africa and is genetically distinct (K2P pairwise distance = 16.7%) from M. nimbae. Description External characters.— Miniopterus nimbae is large-sized for the genus, but showing typical generic features including a rounded head, an elongated second phalanx of the third digit, rounded ears, and a relatively long and straight tragus. The tail is slightly less than half that of the total length. The pelage is dark chocolate brown above and slightly paler below. Individual hairs are unicoloured. The mass and standard external measurements of the holotype compared with a small sample of other individuals are shown in Table 3. Craniodental characters.—The skull is robust for a Miniopterus species. The rostrum is broad and the braincase is rounded and high, typical for the genus of Miniopterus. The dentition of M. nimbae is I 2/3, C 1/1, P 2/3, M 3/3, which is typical of the genus. In the upper tooth row, the inner incisor is larger than the outer one, and the anterior premolar is relatively well developed. The cranial and dental measurements of the holotype compared with a small sample of other individuals are shown in Tables 4 and 5. Distribution Analysis of genetic samples taken from five specimens with large forearm lengths (range 46.9– 48.2 mm) at Mount Gangra (Nimba, Liberia), group them all as M. nimbae (Figs. 2 and 3). Although not genetically analysed, it is likely that specimens previously collected from northern Liberia or southeastern Guinea and identified as M. inflatus are instead referable to this new species, M. nimbae. Based on this assumption, this taxon is known from just three localities: Mount Nimba (and surrounding uplands), Liberia; Wonegizi Mountains, Liberia; and Mount Béro, Guinea (Wolton et al., 1982; Koopman et al., 1995; Fahr et al., 2006; Monadjem et al., 2016). The closest locality for M. inflatus s.s. is at least 2,000 km away in eastern Nigeria (Happold, 2013 b), and no other large Miniopterus specimens have been collected in the intermediate area, despite extensive surveys in a number of localities e.g., Thaï and Comoé National Parks in Côte d’Ivoire (Fahr and Kalko, 2011), the Simandou Range in eastern Guinea (Decher et al., 2015), the Fouta Djallon mountains in central Guinea (Weber and Fahr, 2007) or anywhere in Ghana or Sierra Leone (Grubb et al., 1998; Happold, 2013 b, 2013 c). Therefore, M. nimbae is probably an endemic to the upland areas of northern Liberia and south-eastern Guinea, and may be shown to also occur in the upland area of western Côte d’Ivoire at and around Mount Nimba. Biology: Practically nothing is known about the biology of this Upper Guinea forest endemic. It has been recorded roosting in mine adits at 720– 970 m above sea level (a.s.l.) at Mount Gangra and Mount Yuelliton (both within 10 km of Mount Nimba). The size of one roosting colony at Mount Gangra was estimated at between 20–30 individuals; the site was shared with large numbers of Myonycteris angolensis and Hipposideros cf. ruber, and a few H. marisae. A second roosting colony was estimated to include> 200 individuals of this species at Mount Yuelliton (which was not inhabited by any other bat species — A. Monadjem, personal observation). It has been netted at various localities in the foothills of Mount Nimba and in the low-lying rainforest region between these three upland areas. This suggests that this species roosts in upland areas (700 m a.s.l.), and then descends to forage in lower lying forested areas (about 500 m a.s.l.). In a sample of 13 females captured at Nimba between late December and end-March, five were pregnant. During the same period, three out of 10 males had scrotal testes. The mean frequency of the knee of handreleased M. nimbae captured at Mount Nimba was 48.4 kHz (range: 47.2–48.9 kHz, n = 4). Other Taxonomic Considerations In addition to the description of the new species, M. nimbae, the phylogeny presented here (Figs. 2 and 3) also identifies two other distinct taxa. The first is the taxon M. villiersi, which does not appear to be closely related to M. schreibersii s.s. and should therefore not be considered a subspecies of the latter mentioned taxon. In fact, M. villiersi is sister to M. nimbae (Figs. 2 and 3), from which it is readily distinguishable based on genetics and size (see Tables 2–5); the pairwise genetic distance between the two species is 9.0%, and there is no overlap between these two species in forearm length or any of the craniodental measurements presented here. The echolocation calls also differ, with the frequency of the knee of M. villiersi calls (based on hand-released individuals that were captured at Mount Nimba — x = 51.6 kHz, range 51.1–52.4 kHz, n = 5) being distinctly higher than that of M. nimbae (x = 48.4 kHz, range 47.2–48.9 kHz, n = 4) with no overlap between the two species. The second taxon refers to the M. inflatus group, which appears to be paraphyletic. Miniopterus inflatus s.s. (based on sequenced specimens from Gabon, and close to the type locality in southern Cameroon) is sister to the taxon M. nimbae and these two are sister to M. villiersi (Figs. 2 and 3). By contrast, the M. cf. inflatus specimens (from Malawi and Mozambique) are sister to M. fraterculus and M. minor. This suggests that the taxon M. inflatus s.l. comprises, in addition to the newly recognised M. nimbae, two distinct and not closely related taxa which we refer to as M. inflatus s.s. (from Gabon), and M. cf. inflatus (from Malawi and Mozambique)., Published as part of Monadjem, Ara, Shapiro, Julie T., Richards, Leigh R., Karabulut, Hatice, Crawley, Wing, Nielsen, Ida Broman, Hansen, Anders, Bohmann, Kristine & Mourier, Tobias, 2019, Systematics of West African Miniopterus with the description of a new species, pp. 237-256 in Acta Chiropterologica 21 (2) on pages 245-248, DOI: 10.3161/15081109ACC2019.21.2.001, http://zenodo.org/record/3944920, {"references":["MONADJEM, A., L. RICHARDS, and C. DENYS. 2016. An African bat hotspot: the exceptional importance of Mount Nimba for bat diversity. Acta Chiropterologica, 18: 359 - 375.","WOLTON, R. J., P. A. ARAK, H. C. J. GODFRAY, and R. P. WILSON. 1982. Ecological and behavioural studies of the Megachiroptera at Mount Nimba, Liberia, with notes on Microchiroptera. Mammalia, 18: 359 - 375.","KOOPMAN, K. F., C. P. KOFRON, and A. CHAPMAN. 1995. The bats of Liberia: systematics, ecology, and distribution. American Museum Novitates, 3148: 1 - 24.","FAHR, J., B. A. DJOSSA, and H. VIERHAUS. 2006. Rapid assessment of bats (Chiroptera) in Dere, Diecke and Mt. Bero classified forests, southeastern Guinea; including a review of the distribution of bats in Guinee Forestiere. Pp. 168 - 180, 245 - 247, in A rapid biological assessment of three classified forests in Southeastern Guinea (H. E. WRIGHT, J. MCCULLOUGH, L. E. ALONSO, and M. S. DIALLO, eds.). RAP Bulletin of Biological Assessment, 40: 1 - 248.","HAPPOLD, M. 2013 b. Miniopterus inflatus Greater long-fingered bat. Pp. 714 - 716, in The mammals of Africa. Volume IV: Hedgehogs, shrews and bats (M. HAPPOLD and D. C. HAP- POLD, eds.). Bloomsbury Publishing, London, 800 pp.","FAHR, J., and E. K. V. KALKO. 2011. Biome transitions as centres of diversity: habitat heterogeneity and diversity patterns of West African bat assemblages across spatial scales. Ecography, 34: 177 - 195.","DECHER, J., A. HOFFMANN, J. SCHAER, R. W. NORRIS, B. KADJO, J. ASTRIN, A. MONADJEM, and R. HUTTERER. 2015. Bat diversity in the Simandou Mountain Range of Guinea, with the description of a new white-winged vespertilionid. Acta Chiropterologica, 17: 255 - 282.","WEBER, N., and J. FAHR. 2007. Survey of endemic and globally threatened bat species in the Fouta Djallon Plateau for","GRUBB, P., T. S. JONES, A. G. DAVIES, E. EDBERG, E. D. STARIN, and J. E. HILL. 1998. Mammals of Ghana, Sierra Leone and The Gambia. Trendine Press, Cornwall, UK, 265 pp.","HAPPOLD, M. 2013 c. Miniopterus schreibersii Schreibers's long-fingered bat. Pp. 721 - 722, in The mammals of Africa. Volume IV: Hedgehogs, shrews and bats (M. HAPPOLD and D. C. D. HAPPOLD, eds.). Bloomsbury Publishing, London, 800 pp."]}

RHINOLOPHUS LOBATUS PETERS, 1852 PETERS’ S HORSESHOE BAT Synonyms: R. l. angolensis (Seabra, 1898), Angola, Hahna. Type locality: Mozambique, Tete (= Tete Province), Sena village, south bank of the Zambesi River, - 15.679° S, 33.809° E. Syntypes: ZMB 375 (female, poor skin only), ZMB 2496 (female, skin in alcohol, skull extracted, broken), ZMB 24922 (adult male, complete skeleton), ZMB 24927 (female, complete skeleton), collected between 1843 and 1847 by W. C. H. Peters from Mozambique, Tette (= Tete Province), Sena on the south bank of the Zambesi River, - 15.679° S, 33.809° E. Referred specimens having molecular identifications: FMNH 228936 (SMG 18959), male, 228937 (SMG 18988), female, 228938 (SMG 18989), male, 228939 (SMG 18990), male, 228940 (SMG 18991), female, 228941 (SMG 18992), male, all collected on 2 May 2015 by S. M. Goodman, M. C. Schoeman and G. le Minter from Mozambique, Chihalatan Cave, 38.2 km Eof Inhassoro, - 21,671° S, 34,864° E. FMNH: 228947 (SMG 19022), female, 228954 (SMG 19029), female, 228965 (SMG 19040), female, all collected on 2 May 2015 by S. M. Goodman, M. C. Schoeman and G. le Minter from Mozambique, Mozambique, Inhambane Province, Malashane Cave, 39.1 km Eof Inhassoro, - 21,668° S, 34,847° E, and situated Referred specimens having non - molecular identifications: DM 13894 (female), 13927 (male), 13938 (unknown sex), 13942 (unknown sex), 13943 (unknown sex), 13944 (unknown sex), 13945 (male), 14529 (male), 14531 (male), 14532 (female), collected by A. Monadjem, G. le Minter and E. Lagadec in July 2015 from Mozambique, Sofala Province, Gorongosa National Park. TMSA 49114 (female), TMSA 49117 (male), collected from Mozambique, Sofala Province, Gorongosa National Park by A. J. Guyton in July 2015 from Chitengo Camp and Cheringoma Plateau respectively. TMSA 14653 (female), 14655 (unknown sex), 14656 (female), 14657 (female), 14662 (male), 14663 (male), 14664 (female), collected 28 – 29 July 1964 from Mozambique, Tete Province, Muchena, - 15.679° S, 33.809° E. Incertae sedis: DM 8575 (male) from Mozambique, Cabo Delgado Province, Pemba Island (- 13.006° S, 40.524° E) and DM 8574 (female), from Mozambique, Sofala Province, Chinizuia Forest (- 18.977° S, 35.052° E), both collected by A. Monadjem in June 2006. TMSA 49115 (male), TMSA 49118 (male), collected from Mozambique, Sofala Province, Gorongosa National Park by A. J. Guyton in July 2015 from Chitengo Camp and Cheringoma Plateau, respectively. Etymology: The Latin word lobatus means lobed, perhaps referring to the general shape of the noseleaf (Fig. 13). Re-diagnosis and comparisons: This taxon is clearly phylogenetically distinct on Cytb gene sequences (see also: Dool et al., 2016) from topotypic landeri from West Africa and seems to be more closely affiliated with the capensis group of Csorba et al. (2003), i.e. R. capensis, R. denti, R. simulator and R. swinnyi (Fig. 2). Specimens from Chihalatan and Malashane (N = 4) had a mean echolocation call peak frequency of 106.8 ± 0.4 kHz (Table 2), close to the 107 kHz generally reported for southern Africa ‘ landeri ’ (Monadjem et al., 2010). This contrasts with a mean maximum frequency of 104.3 ± 0.42 kHz (N = 2) recorded for topotypic West African (Liberian) R. landeri, making a peak frequency of c.107 kHz a possible diagnostic criterion for R. lobatus. However, a wider range (102–108 kHz) and slightly lower mean frequency of 105 kHz was reported for 20 individuals at Gorongosa (Table 2). This variation could be indicative of further undetected cryptic speciation in this taxon, as also indicated by the very wide variation in baculum and noseleaf characters, discussed in the following paragraph. Although not easily distinguished on body or skull size (Table 2; Fig. 4), this taxon differs from topotypic R. landeri by displaying a prognathic rostrum (LMs 1–3), a shortened braincase that extends posteriorly (LMs 10–14) and a broader nasal inflation (LMs 16–18) (Figs 5, 6). Specimens of cf. lobatus from Pemba (Mozambique) had a shorter foramen magnum (LMs 9–10), a narrower jugal process (LM 3–4) and a slightly broader nasal inflation (LMs 16–18), than other Mozambique animals. The same individual from Pemba possessed a distinctive robust baculum (Fig. 8) and noseleaf (Fig. 9) having a markedly hastate lancet and pronounced posterior lobe of the connecting process. However, pending molecular data, we here refer them to lobatus incertae sedis. Although genetically clearly assigned to lobatus, individuals from Gorongosa National Park are somewhat smaller in external (but not cranial) measurements (Table 2), and also display unique noseleaf and bacular morphologies. The noseleaf of a Gorongosa individual has a small hastate lancet and bluntly pointed tip, and a relatively small and more erect connecting process with semi-symmetrical lobe (Figs 9, 13), while its baculum is highly divergent from all other forms considered in our study (Fig. 8), having a remarkable short baculum with wide shaft and very small base compared with landeri from West Africa, as well as cf. lobatus from Malashane and Chihalatan Caves. Given the clear evidence for close genetic identity in our study between Malashane and Chihalatan Caves and Gorongosa National Park for animals referred to R. lobatus, we provisionally regard this variation in noseleaf and baculum structure to represent polymorphic traits in this species. It is possible that these morphological types represent good species between which introgression has occurred. Further analysis with additional nuclear sequences is necessary to test this hypothesis. There is also the possibility that these could represent subadults with bacula that are not fully ossified, as detailed in R. adami described by Kock et al. (2000). Male R. lobatus from Chihalatan and Malashane Caves, as well as Gorongosa National Park, typically possessed a dark brown apical tuft of stiff hairs characteristic of R. landeri (Fig. 14; Csorba et al., 2003; Monadjem et al., 2010). This convergent character could be one of the contributing factors that have led to the historical misidentification of the species with R. landeri. Distribution and biology: As anticipated by Monadjem et al. (2010), who considered that the southern (and possibly eastern) African savannah-occurring lobatus might prove to be distinct from the West African forest-occurring landeri, we here refer all southern African specimens to lobatus (Fig. 11) [see appendix of Monadjem et al. (2010) for a full list of localities]. We agree with Monadjem et al. (2010) that R. angolensis Seabra, 1989 from western Angola may merit specific status. Nevertheless, in the absence of the type series that was destroyed in the Lisbon fire of 1978, resolution of its status must await a detailed revision, including molecular evidence with designation of new type material. Likewise, the status of east and central African populations must await further studies, including molecular data. Until further evidence becomes available, we suggest that it is prudent to use the name lobatus for all savannah populations in southern and east Africa., Published as part of Taylor, Peter J., Macdonald, Angus, Goodman, Steven M., Kearney, Teresa, Cotterill, Fenton P. D., Stoffberg, Sam, Monadjem, Ara, Schoeman, M. Corrie, Guyton, Jennifer, Naskrecki, Pitor & Richards, Leigh R., 2018, Integrative taxonomy resolves three new cryptic species of small southern African horseshoe bats (Rhinolophus), pp. 1249-1276 in Zoological Journal of the Linnean Society 184 on pages 1270-1272, DOI: 10.1093/zoolinnean/zly024, http://zenodo.org/record/4253044, {"references":["Dool SE, Puechmaille SJ, Foley NM, Allegrini B, Bastian A, Mutumi GL, Maluleke TG, Odendaal LJ, Teeling EC, Jacobs DS. 2016. Nuclear introns outperform mitochondrial DNA in inter-specific phylogenetic reconstruction: lessons from horseshoe bats (Rhinolophidae: Chiroptera). Molecular Phylogenetics and EVolution 97: 196 - 212.","Csorba G, Ujhelyip P, Thomas N. 2003. Horseshoe bats of the aeorld (Chiroptera: Rhinolophidae). Shropshire: Alana books.","Monadjem A, Taylor PJ, Cotterill FPD, Schoeman MC. 2010. Bats of Southern and South-central Africa: a biogeographic and taxonomic synthesis. Johannesburg: Wits University Press.","Kock D, Csorba G, Howell KM. 2000. Rhinolophus maendeleo n. sp. from Tanzania, a horseshoe bat noteworthy for its systematics and biogeography. Senckenbergiana biologica 80: 233 - 239."]}

RHINOLOPHUS RHODESIAE ROBERTS, 1946 ROBERTS' S HORSESHOE BAT Synonyms: None. Holotype: TMSA 1325, adult female, collected by A. Roberts on 16 August 1913. Type locality: 'Southern Rhodesia' (= Zimbabwe), Bezwe River, tributary of 'Wanetsi' (= Nuanetsi) River, −21.500° S, 31.167° E. Referred specimens having molecular identifications: FMNH 228942 (SMG 19017), female, 228943 (SMG 19018), male, 228944 (SMG 19019), female, 228945 (SMG 19020), male, 228946 (SMG 19021), female, 228948 (SMG 19023), male, 228949 (SMG 19024), female, 228950 (SMG 19025), female, 228951 (SMG 19026), female, 228952 (SMG 19027), male, 228953 (SMG 19028), male, 228955 (SMG 19030), male, 228957 (SMG 19032), female, 228958 (SMG 19033), female, 228959 (SMG 19034), female, 228960 (SMG 19035), female, 228961 (SMG 19036), male, 228962 (SMG 19037), male, 228964 (SMG 19039), female, all collected on 2 May 2015 by S. M. Goodman, M. C. Schoeman and G. le Minter from Mozambique, Inhambane Province, Malashane Cave, 39.1 km Efrom Inhassoro, −21,668° S, 34,847° E, and situated Referred specimens having only morphological identifications: DM 7080, adult male, KwaZulu-Natal Province, Hlabeni Forest Reserve, −29.933° S, 29.766° E collected by D. Forbes on 29 July 2000; DM 12007 (adult male); DM 14034 (adult male), collected by M. C. Schoeman from KwaZulu-Natal Province, Pietermaritzburg, Ferncliff Nature Reserve, Ferncliff Cave, −29.550° S, 30.320° E; DM 11270 (female), 11271 (male), 11272 (female), 11273 (female), 11275 (male), all collected by S. Stoffberg from Chihalatan Cave, 38.2 km Eof Inhassoro, Inhambane Province, Mozambique −21.671° S, 34.864° E;. DM 11275 collected on 8 August 2006 while the other specimens were collected on 3 September 2007; FMNH 228956 (SMG 19031), collected 2 May 2015 by S. M. Goodman, M. C. Schoeman and G. le Minter from Mozambique, Inhambane Province, Malashane Cave, 39.1 km E from Inhassoro, −21,668° S, 34,847° E; DM 13450 (female), DM13451 (female), collected on 8 May 2012 by J. Bayliss at Mozambique, Niassa Province, Mount Mecula, −12.068° S, 37.662° E. Etymology: The name refers to the location in Southern Rhodesia (now Zimbabwe) where the type specimen was collected. Re-diagnosis and description: Roberts (1946) described this subspecies as being slightly smaller than the nominate R. s. swinnyi based on slightly smaller body size, slightly longer tail, smaller ears and its bright ochraceous colour. Since the last-mentioned character is known to be an environmentally induced effect in many cave-dwelling bat species, it does not serve as a diagnostic character. In our analysis, molecular evidence closely matched our series from Chihalatan and Malashane Caves with Genbank sequences from the extreme northern South Africa (Pafuri), Zimbabwe (Dambanzara) and Zambia (Kalenda and Shimalala Caves). Since these localities encompass the type locality of rhodesiae (Bezwe River in Zimbabwe), and Pafuri is only 100 km south of Bezwe, we are confident to use this available name for this widespread taxon. The species can be further diagnosed by having echolocation peak frequencies around 100 kHz (99–102 kHz, N = 8 from Malashane Cave; Table 2), which are quite distinct from typical swinnyi (105–107kHz, N = 6; Table 2) as well as the new Gorongosa National Park taxon R. gorongosae sp. nov. (104–108 kHz, N = 16; Table 2). Noseleaf structure is distinctive, being characterised by a hastate lancet, not as concave as true swinnyi, less erect, low, rounded connecting process and more pronounced posterior lobe. Bacular structure of rhodesiae is clearly distinct from other swinnyi -like animals (see Figs 7, 8), being characterized by a distinctly longer tapered baculum with a distinctly broader base and shallow notchalong the lower portion of the shaft that isvisible in the lateral profile. Traditional morphometrics (Table 2; Fig. 4) do not differentiate rhodesiae from swinnyi proper; however, rhodesiae is clearly distinguished with minimal overlap from the distinctly smaller gorongosae sp. nov. Although quite small, the R. rhodesiae holotype falls within the range of variation of specimens assigned to R. rhodesiae from Mozambique, Zimbabwe, Zambia, northern South Africa (Pafuri, Limpopo Province), and Zanzibar (Table 2; Fig. 4). It falls clearly outside (larger than) the range of variation of the smaller gorongosae sp. nov. taxon (Fig. 4). Geometric morphometric results result in a better separation between rhodesiae and swinnyi s.s. with only minimal overlap (Fig. 5). Once again, the rhodesiae holotype from Bezwe River clusters within the range of variation of the rhodesiae taxon and outside the gorongosae sp. nov. or swinnyi taxa, thus validating the use of this name for this taxon. Distribution and biology: Combined molecular and morphometric data suggest the widespread distribution of this taxon from central and northern South Africa through Zimbabwe, Zambia and Mozambique extending to Zanzibar (Fig. 11). Based on specimen assignments on morphological grounds, the species co-occurs with R. swinnyi in central KwaZulu-Natal at Ferncliff Cave, as well as occurring in close proximity in northern KwaZulu-Natal, recorded at Hlabisi Forest close to Ngome Forest where swinnyi was recorded (Fig. 11). The widespread extent of this taxon and its occurrence in northern South Africa is confirmed by the widespread occurrence of a hitherto unidentified 100 kHz acoustic type recorded in the Soutpansberg (Taylor et al., 2013), and Pafuri Region of northern Kruger National Park (Taylor & Parker, unpublished data)., Published as part of Taylor, Peter J., Macdonald, Angus, Goodman, Steven M., Kearney, Teresa, Cotterill, Fenton P. D., Stoffberg, Sam, Monadjem, Ara, Schoeman, M. Corrie, Guyton, Jennifer, Naskrecki, Pitor & Richards, Leigh R., 2018, Integrative taxonomy resolves three new cryptic species of small southern African horseshoe bats (Rhinolophus), pp. 1249-1276 in Zoological Journal of the Linnean Society 184 on pages 1265-1269, DOI: 10.1093/zoolinnean/zly024, http://zenodo.org/record/4253044, {"references":["Taylor PJ, Sowler S, Schoeman MC, Monadjem A. 2013. Diversity of bats in the Soutpansberg and Blouberg Mountains of northern South Africa: complimentarity of acoustic and non-acoustic survey methods. South African Journal of Wildlife Research 43: 12 - 26."]}

Taylor, Peter J., Macdonald, Angus, Goodman, Steven M., Kearney, Teresa, Cotterill, Fenton P. D., Stoffberg, Sam, Monadjem, Ara, Schoeman, M. Corrie, Guyton, Jennifer, Naskrecki, Pitor, Richards, Leigh R. (2018): Integrative taxonomy resolves three new cryptic species of small southern African horseshoe bats (Rhinolophus). Zoological Journal of the Linnean Society 184: 1249-1276, DOI: https://doi.org/10.1093/zoolinnean/zly024

RHINOLOPHUS SWINNYI GOUGH, 1908 SWINNY’ S HORSESHOE BAT Synonyms: Rhinolophus swinnyi piriensis Hewitt, 1913, South Africa, Eastern Cape Province, Pirie, near King Williams Town. Types: TMSA 1021 (holotype, male), 1022 (cotype, male), collected on 22–23 February 1908 by H. H. Swinny. Type locality: South Africa, Eastern Cape Province, Pondoland, Ngqeleni District, - 31.667° S, 29,033° E. Referred specimens having molecular identifications: None. Referred specimens having only morphological identifications: All specimens originate from South Africa. DM 7084, adult male collected by P. J. Taylor on 3 March 2001 and DM 13250, 13252 and 13254 (all of unknown sex), collected by E. J. Richardson on 2 November 2010 from the Eastern Cape Province, Insizwe Mine, (- 30.804° S, 29.281° E); DM 14036 (male), collected by M. C. Schoeman on 9 May 2012 from the KwaZulu-Natal Province, Pietermaritzburg, Ferncliff Nature Reserve, (- 29.550° S, 30.320° E); DM 14291 (female) and 14292 (female) collected by L. R. Richards on 1 April 2014 from the KwaZulu-Natal Province, Eshowe, Entumeni Nature Reserve, (- 28.886° S, 31.376° E); DM 14441 (female) and 14440 (male) collected by S. Stoffberg and M. C. Schoeman on 23 July 2004 from Eastern Cape Province, Kokstad Mine, (- 30.810° S, 29.280° E); DM 15018, adult female collected by L. R. Richards on 7 April 2016 from Eastern Cape Province, Sandile’s Rest Trout and Forest Country Estate, (- 32.661° S, 27.298° E); TMSA 39848, adult male collected by G. Bronner on 14 March 1988 from KwaZulu-Natal Province, Ngome Forest Reserve, (- 27.833° S, 31.413° E). Re-diagnosis and comparisons: Gough’s (1908) original description emphasized inter alia, the very small size of the species, the ‘mouse = grey’ colour, the position of the anterior premolar within the toothrow, the medium-sized ears, the shape of the connecting process (‘forming a marked projection, rounded terminally’), the parallel-sided edges of the sella and the moderate lancet with strongly concave edges. Rhinolopus swinny s.s., as defined here, confined to the Eastern Cape, KwaZulu-Natal and possibly Mpumalanga Provinces, South Africa, is clearly distinct on molecular grounds from R. rhodesiae and R. gorongosae sp. nov., and has as its closest relative, R. capensis with which it overlaps in range in the Eastern Cape. The echolocation peak frequency of R. swinnyi s.s. calls recorded in the Eastern Cape [mean 106.6 ± 0.4 kHz, N = 10; Schoeman & Jacobs (2008; Table S1)] and KwaZulu-Natal (mean 105.6 ± 0.76 kHz; this study) Provinces, South Africa overlap considerably, but differ clearly from the 100 kHz calls of R. rhodesiae from central and northern South Africa and Mozambique (and probably more broadly). Although R. swinnyi s.s. overlaps in external and cranial characters with R. rhodesiae, it can be distinguished from the latter taxon based on both baculum and noseleaf characters, as well as on detailed cranial shape analysis (see above). It can be differentiated from R. gorongosae sp. nov. on its larger size (e.g. condylecanine skull length 14.8–15.2 mm in R. gorongosae sp. nov., 14.3–16.0 mm in R. swinnyi; zygomatic width 8.13–8.56 mm in R. gorongosae sp. nov., 8.7–9.2 mm in R. swinnyi; Table 2), as well as on noseleaf, bacular and subtle cranial shape characters (see above). Distribution and biology: We here restrict the distribution of R. swinnyi s.s. to South Africa, including the Eastern Cape and KwaZulu-Natal Provinces and possibly Mpumalanga Province (Fig. 11). However, a recent specimen collected from the montane western region of Swaziland has tentatively been assigned to R. rhodesiae based on morphology and acoustics (maximum frequency = 102 kHz), but a molecular analysis has yet to be conducted (A. Monadjem, unpublished data); this suggests that R. rhodesiae (and not R. swinnyi) occurs to the north in Mpumalanga Province. Specimens from Limpopo Province in the north of South Africa are referred to R. rhodesiae. As pointed out above, swinnyi and rhodesiae co-occur at at least one locality, Ferncliff Cave, in central KwaZulu-Natal., Published as part of Taylor, Peter J., Macdonald, Angus, Goodman, Steven M., Kearney, Teresa, Cotterill, Fenton P. D., Stoffberg, Sam, Monadjem, Ara, Schoeman, M. Corrie, Guyton, Jennifer, Naskrecki, Pitor & Richards, Leigh R., 2018, Integrative taxonomy resolves three new cryptic species of small southern African horseshoe bats (Rhinolophus), pp. 1249-1276 in Zoological Journal of the Linnean Society 184 on page 1273, DOI: 10.1093/zoolinnean/zly024, http://zenodo.org/record/4253044, {"references":["Schoeman MC, Jacobs DS. 2008. The Relative Influence of Competition and Prey Defenses on the Phenotypic Structure of Insectivorous Bat Ensembles in Southern Africa. PLoS ONE 3: e 3715. doi: 10.1371 / journal. pone. 0003715"]}

RHINOLOPHUS GORONGOSAE SP. NOV. LEAST HORSESHOE BAT Holotype: Durban Natural Science Museum (DM) No. 14820 (field number JAG196), is an adult male, preserved in 70% ethyl alcohol, originally part of series of specimens collected by J. A. Guyton on 25 April 2015. The cranium and baculum has been extracted and examined for this study. The specimen has been included in both morphometric analyses. Type locality: Bunga Inselberg, Gorongosa National Park, Sofala Province, Mozambique − 18.599° S, 34.343° E, 212 m. Paratypes: Eight specimens collected 24–25 April 2015 (DM 14815–14819), 2 May 2015 (DM 14828), 5 November 2015 (DM 14843) and 22 July 2015 (DM 14865). Referred specimens having molecular identification: DM 14815 (JAG 188) a female specimen collected by J. A. Guyton from Mozambique, Sofala Province, Gorongosa National Park, Bunga Inselberg, - 18.599° S, 34.343° E, 212 m; DM 14843 (JAG 228) a female specimen collected by J. A. Guyton from Mozambique, Sofala Province, Gorongosa National Park, - 18.694° S, 34.208° E, 308 m. Referred specimens having only morphological identification: TMSA 49116 (JAG 31), adult male, collected on 21 April 2013 by J. A. Guyton from Mozambique, Sofala Province, Gorongosa National Park, Cheringoma Plateau, Gorge Rim, Site 1, - 18.635° S, 34.808° E, 213 m. Incertae sedis: DM 14864 adult female collected by J. A. Guyton on 21 July 2015 from Mozambique, Sofala Province, Gorongosa National Park, - 18.465° S, 34.052° E, 1150 m; DM 11482, adult female collected on 1 May 2009 by J. Bayliss from Mozambique, Nampula Province, Mount Inago Forest Camp, - 15.045° S, 37.396° E 1478 m. Etymology: The species derives its name from the Gorongosa district of Mozambique, in particular Gorongosa National Park, a biologically diverse region of southern Africa. Diagnosis: The species can be clearly distinguished from both R. swinnyi s.s. and R. rhodesiae on molecular grounds (Figs 2, 3) as well as by its smaller size (Fig. 4; Table 2), distinct cranial shape (Figs 5, 6), echolocation call peakfrequency (Table 2), baculum (Figs 7, 8) and noseleaf (Figs 9, 12) characteristics. Although some measurements overlap, there is minimal overlap in condylocanine skull length and zygomatic skull width between this species (14.8–15.2 mm; 8.13–8.56 mm) and R. swinnyi (14.3–16.0 mm; 8.7–9.2 mm) and R. rhodesiae (15.1–15.7; 8.46–8.96 mm). The small size of this form makes it even smaller than denti (regarded by Csorba et al., 2003 as the smallest species in the Ethiopian region), therefore making this new species Africa’s smallest horseshoe bat. Comparing means for R. gorongosae sp. nov. (Table 2) and denti (Monadjem et al., 2010): forearm 41.3 mm cf. 43.1 mm; mass 5.6 g cf. 7.0 g. Description: The genetically-distinct R. gorongosae sp. nov. is similar in pelage colour but distinctly smaller in external (mean total length 68 mm, mean forearm length = 41 mm) and cranial (mean condylocanine length 15.1 mm) measurements (Table 2) than both rhodesiae (mean totallength 75 mm, forearmlength 44 mm, condyolocanine length 15.4 mm) and swinnyi s.s. from South Africa (mean totallength 73 mm, forearm length 44 mm, condyolocanine length 15.4 mm). Based on geometric morphometric results, the diminutive R. gorongosae sp. nov. has a reduced foramen magnum, a noticeable depression along the parietal region, low set bullae, a narrow braincase and a narrow and high set nasal inflation with a sharp slope from the nasal inflation to the maxillae (Fig. 6). Lancet is subtriangular with straight to slightly concave sides, and a high, rounded connecting process; the sella has a diminutive, pointed tip (Fig. 12). Baculum short with reduced base, with a characteristic notch on one side of the shaft tip (position variable). The slightly narrower baculum shaft with narrower tip distinguishes R. gorongosae sp. nov. from R. swinnyi (Table 5; Figs 7, 8). Maxillary toothrow with minute and laterally displaced PM 1 with a distinct gap between the alveolar borders of canine and PM 2 (Fig. 10). Mean peak echolocation CF frequency is 106 kHz (104–108 kHz; Table 2). Distribution and biology: So far, it appears that this tiny species is restricted in its distribution to Gorongosa National Park in Mozambique (Fig. 11), although we provisionally refer a very small adult individual from Mount Inago in Mozambique to this taxon. Molecular sequences are required from a wider range of localities to determine the range of this taxon. Given that the two individuals sequenced from Gorongosa National Park were distinct from each other, the possibility exists that more than one cryptic species may be present., Published as part of Taylor, Peter J., Macdonald, Angus, Goodman, Steven M., Kearney, Teresa, Cotterill, Fenton P. D., Stoffberg, Sam, Monadjem, Ara, Schoeman, M. Corrie, Guyton, Jennifer, Naskrecki, Pitor & Richards, Leigh R., 2018, Integrative taxonomy resolves three new cryptic species of small southern African horseshoe bats (Rhinolophus), pp. 1249-1276 in Zoological Journal of the Linnean Society 184 on pages 1262-1265, DOI: 10.1093/zoolinnean/zly024, http://zenodo.org/record/4253044, {"references":["Csorba G, Ujhelyip P, Thomas N. 2003. Horseshoe bats of the aeorld (Chiroptera: Rhinolophidae). Shropshire: Alana books.","Monadjem A, Taylor PJ, Cotterill FPD, Schoeman MC. 2010. Bats of Southern and South-central Africa: a biogeographic and taxonomic synthesis. Johannesburg: Wits University Press."]}

Chaerephon major (Trouessart, 1897) This species was previously recorded at Grassfield in Liberian Nimba (Wolton et al., 1982), but was not recorded during this study., Published as part of Monadjem, Ara, Richards, Leigh & Denys, Christiane, 2016, An African bat hotspot: the exceptional importance of Mount Nimba for bat diversity, pp. 359-375 in Acta Chiropterologica 18 (2) on page 367, DOI: 10.3161/15081109ACC2016.18.2.005, http://zenodo.org/record/3942481

Saccolaimus peli (Temminck, 1853) This species has only been recorded at Mt Nimba by Coe (1975) in forested habitat on the Liberian side. A. Forbes-Watson also collected a specimen from Liberian Nimba and deposited it in the National Museums of Kenya (A. Monadjem, personal observation of the specimen). Despite its characteristic and conspicuous habits of foraging during the early evening, it was not observed during this study., Published as part of Monadjem, Ara, Richards, Leigh & Denys, Christiane, 2016, An African bat hotspot: the exceptional importance of Mount Nimba for bat diversity, pp. 359-375 in Acta Chiropterologica 18 (2) on page 366, DOI: 10.3161/15081109ACC2016.18.2.005, http://zenodo.org/record/3942481, {"references":["COE, M. 1975. Mammalian ecological studies at Mount Nimba, Liberia. Mammalia, 39: 523 - 588."]}

Rhinolophus landeri Martin, 1838 During this study, this species was recorded from forested habitat in Liberian Nimba at altitudes ranging from 550 m to 1,000 m. Despite its wide distribution (Monadjem et al., 2010 a), this species had not been recorded from Mt Nimba prior to this survey. This species closely resembles R. guineensis but was distinguished from it by its smaller size (forearm and greatest skull lengths were 42.0– 43.9 mm and 18.8–18.9 mm, n = 3). Rhinolophus alcyone is typically far larger in size., Published as part of Monadjem, Ara, Richards, Leigh & Denys, Christiane, 2016, An African bat hotspot: the exceptional importance of Mount Nimba for bat diversity, pp. 359-375 in Acta Chiropterologica 18 (2) on page 366, DOI: 10.3161/15081109ACC2016.18.2.005, http://zenodo.org/record/3942481

Chaerephon bemmeleni (Jentink, 1879) The type specimen of this species was originally collected from an unspecified location in Liberia (Fahr, 2013 e). The occurrence of this species at Mt Nimba is mentioned only by Fahr (2013 e), who stated that it had been recorded in montane grassland at 1,600 m (specimen in the Senckenberg Museum, Frankfurt). It was not recorded during this survey., Published as part of Monadjem, Ara, Richards, Leigh & Denys, Christiane, 2016, An African bat hotspot: the exceptional importance of Mount Nimba for bat diversity, pp. 359-375 in Acta Chiropterologica 18 (2) on page 367, DOI: 10.3161/15081109ACC2016.18.2.005, http://zenodo.org/record/3942481, {"references":["FAHR, J. 2013 e. Tadarida bemmeleni Gland-tailed free-tailed bat. Pp. 497 - 499, in The mammals of Africa. Volume IV: Hedgehogs, shrews and bats (M. HAPPOLD and D. C. D. HAPPOLD, eds.). Bloomsbury Publishing, London, 800 pp."]}

Kerivoula lanosa (A. Smith, 1847) This species was previously recorded from Ziéla, Guinean Nimba (Aellen, 1963 [as K. harrisoni bellula), but was not recorded during this study. It has also been collected from the Simandou Range in north-eastern Guinea (Fahr and Ebigbo, 2003)., Published as part of Monadjem, Ara, Richards, Leigh & Denys, Christiane, 2016, An African bat hotspot: the exceptional importance of Mount Nimba for bat diversity, pp. 359-375 in Acta Chiropterologica 18 (2) on page 368, DOI: 10.3161/15081109ACC2016.18.2.005, http://zenodo.org/record/3942481, {"references":["AELLEN, V. 1963. La Reserve Naturelle Integrale du Mont Nimba. XXIX. Chiropteres. Memoires de l'Institut Francais d'Afrique Noire, 66: 629 - 638.","FAHR, J., and N. M. EBIGBO. 2003. A conservation assessment of the bats of the Simandou Range, Guinea, with the first record of Myotis welwitschii (Gray, 1866) from West Africa. Acta Chiropterologica, 5: 125 - 141."]}

Monadjem, Ara, Richards, Leigh, Denys, Christiane (2016): An African bat hotspot: the exceptional importance of Mount Nimba for bat diversity. Acta Chiropterologica 18 (2): 359-375, DOI: 10.3161/15081109ACC2016.18.2.005

Nycteris hispida (Schreber, 1775) This species was previously recorded from Zouguépo, Guinean Nimba (Aellen, 1963). During this survey, it was recorded sparsely in open savanna on the Liberian side of the mountain. This species was easily identifiable by its trifid upper incisors and short ears (21 mm in DM 12611)., Published as part of Monadjem, Ara, Richards, Leigh & Denys, Christiane, 2016, An African bat hotspot: the exceptional importance of Mount Nimba for bat diversity, pp. 359-375 in Acta Chiropterologica 18 (2) on page 367, DOI: 10.3161/15081109ACC2016.18.2.005, http://zenodo.org/record/3942481, {"references":["AELLEN, V. 1963. La Reserve Naturelle Integrale du Mont Nimba. XXIX. Chiropteres. Memoires de l'Institut Francais d'Afrique Noire, 66: 629 - 638."]}

Nanonycteris veldkampii (Jentink, 1888) This species was captured in forested habitats on both Liberian and Guinean sides of the mountain from 500 m to 1,200 m a.s.l. It may be a migrant to Mt Nimba as it has not been recorded during the wet season (Coe, 1975; Wolton et al., 1982)., Published as part of Monadjem, Ara, Richards, Leigh & Denys, Christiane, 2016, An African bat hotspot: the exceptional importance of Mount Nimba for bat diversity, pp. 359-375 in Acta Chiropterologica 18 (2) on page 364, DOI: 10.3161/15081109ACC2016.18.2.005, http://zenodo.org/record/3942481, {"references":["COE, M. 1975. Mammalian ecological studies at Mount Nimba, Liberia. Mammalia, 39: 523 - 588."]}

Pipistrellus nanulus Thomas, 1904 This species was previously recorded from Liberian Nimba (Hill, 1982; Wolton et al., 1982). During this survey, six specimens were recorded in lowland forest from 420 m to 480 m a.s.l. on the Liberian side of the mountain., Published as part of Monadjem, Ara, Richards, Leigh & Denys, Christiane, 2016, An African bat hotspot: the exceptional importance of Mount Nimba for bat diversity, pp. 359-375 in Acta Chiropterologica 18 (2) on page 371, DOI: 10.3161/15081109ACC2016.18.2.005, http://zenodo.org/record/3942481, {"references":["HILL, J. E. 1982. Records of bats from Mount Nimba, Liberia. Mammalia, 46: 116 - 120."]}

Myotis bocagii (Peters, 1870) During this survey, a single specimen was collected over water in old growth forest on the Liberian side, at 420 m a.s.l. This species, which has a wide African distribution, had not been previously recorded from Mt Nimba., Published as part of Monadjem, Ara, Richards, Leigh & Denys, Christiane, 2016, An African bat hotspot: the exceptional importance of Mount Nimba for bat diversity, pp. 359-375 in Acta Chiropterologica 18 (2) on page 369, DOI: 10.3161/15081109ACC2016.18.2.005, http://zenodo.org/record/3942481

Nycteris arge Thomas, 1903 This species was previously recorded from Liberian Nimba (Verschuren, 1977; Wolton et al., 1982). During this survey, it was recorded widely in forested habitats between 400 and 1,000 m a.s.l. on the Liberian side. This species was distinguished from the similar N. intermedia by its larger size (forearm length 39.8–48.8 mm, and greatest skull length 19.7–20.1 mm, n = 5)., Published as part of Monadjem, Ara, Richards, Leigh & Denys, Christiane, 2016, An African bat hotspot: the exceptional importance of Mount Nimba for bat diversity, pp. 359-375 in Acta Chiropterologica 18 (2) on page 367, DOI: 10.3161/15081109ACC2016.18.2.005, http://zenodo.org/record/3942481

Hipposideros cf. caffer (Sundevall, 1846) The occurrence of this species at Mt Nimba is mentioned by Fahr et al. (2006), who re-examined specimens, originally identified as Hipposideros beatus, collected from Zouguépo and deposited in MNHN. Hipposideros caffer sensu stricto, however, is restricted to southern Africa (Vallo et al., 2008), and it is not clear what species these specimens refer to. It was not recorded during this survey., Published as part of Monadjem, Ara, Richards, Leigh & Denys, Christiane, 2016, An African bat hotspot: the exceptional importance of Mount Nimba for bat diversity, pp. 359-375 in Acta Chiropterologica 18 (2) on page 365, DOI: 10.3161/15081109ACC2016.18.2.005, http://zenodo.org/record/3942481, {"references":["FAHR, J., B. A. DJOSSA, and H. VIERHAUS. 2006. Rapid assessment of bats (Chiroptera) in Dere, Diecke and Mt. Bero classified forests, southeastern Guinea; including a review of the distribution of bats in Guinee Forestiere. Pp. 168 - 180, 245 - 247, in A rapid biological assessment of three classified forests in Southeastern Guinea (H. E. WRIGHT, J. MCCULLOUGH, L. E. ALONSO, and M. S. DIALLO, eds.). Conservation International, RAP Bulletin of Biological Assessment, Washington, D. C., 40, 248 pp."]}