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IMPLICATIONS FOR PARATRIAENOPS The strong genetic differentiation of Paratriaenops and Triaenops evident in our concatenated intron and species tree analyses (Fig. 4; Supporting Information, Fig. S2) reinforces earlier distinctions of these taxa based on mitochondrial evidence (Russell et al., 2007, 2008) and on morphology (Benda & Vallo, 2009). Foley et al. (2015) dated the divergence of these taxa at 22 Mya; that analysis recovered both Rhinonycteris and Cloeotis in successive splits off the lineage leading to Triaenops. The paraphyly of Malagasy rhinonycterids clearly supports the conclusion of Russell et al. (2008) that Madagascar was colonized at least twice in the history of this group. However, this interpretation hinges on the phylogenetic positions of Rhinonycteris and Cloeotis, which were not included in our analysis. A sister relationship of Paratriaenops + Triaenops, given the well-supported position of T. menamena as sister to African plus Arabian Triaenops, could indicate that only one colonization of Madagascar was involved. In this scenario, a descendent of T. menamena could have colonized the African mainland and given rise to the clade of T. afer, T. persicus and T. parvus. Additional genetic sampling of Cloeotis, Rhinonicteris and the missing Triaenops species might help to distinguish these alternatives, but it seems likely that extinction has strongly shaped the extant diversity of the group. Genetic and distributional data provide mixed support for the validity of P. auritus and P. furcula as separate species. Previous morphological analyses of specimens assigned to these two taxa found consistent differences (Ranivo & Goodman, 2006). The Cytb genetic distance between these species (4.5%; Table 1) and the well-supported monophyly of P. auritus and moderately supported monophyly of P. furcula (Fig. 2) could be argued to support their current taxonomic status (also see Russell et al., 2008). In stark opposition, gene tree analyses of four independent nuclear loci under both ML and BI models did not recover any genetic structure within or between the two species (Fig. 4). Instead, the relationships inferred between P. auritus and P. furcula are consistent with extensive ongoing or recent hybridization. The allopatric distributions of these species do not provide any support for their reproductive isolation. Additionally, mitochondrial isolation by distance cannot be ruled out as the mechanism responsible for the genetic distance and topological relationship between populations assigned to P. auritus and P. furcula in our genetic analyses. Before the step is taken to combine these species, for which P. auritus would be the junior synonym, further genetic sampling is needed. In the material used in the present study, the northernmost locality for P. furcula (Namoroka, FMNH 175783) is ~ 330 km south of the southernmost locality for P. auritus (Betsiaka, FMNH 179370���179373); additional data are needed from the intervening zone to determine the nature and level of genetic separation between these forms., Published as part of Rossoni, Daniela M, Demos, Terrence C, Goodman, Steven M, Yego, Richard K, Mohlman, Jessica L, Webala, Paul W & Patterson, Bruce D, 2021, Genetic, morphological and acoustic differentiation of African trident bats (Rhinonycteridae: Triaenops), pp. 236 in Zoological Journal of the Linnean Society 192 (1) on page 236, DOI: 10.1093/zoolinnean/zlaa098, http://zenodo.org/record/5301043, {"references":["Russell AL, Ranivo J, Palkovacs EP, Goodman SM, Yoder AD. 2007. Working at the interface of phylogenetics and population genetics: a biogeographical analysis of Triaenops spp. (Chiroptera: Hipposideridae). Molecular Ecology 16: 839 - 851.","Russell AL, Goodman SM, Cox MP. 2008. Coalescent analyses support multiple mainland-to-island dispersals in the evolution of Malagasy Triaenops bats (Chiroptera: Hipposideridae). Journal of Biogeography 35: 995 - 1003.","Benda P, Vallo P. 2009. Taxonomic revision of the genus Triaenops (Chiroptera: Hipposideridae) with description of a new species from southern Arabia and definitions of a new genus and tribe. Folia Zoologica 58: 1 - 45.","Foley NM, Thong VD, Soisook P, Goodman SM, Armstrong KN, Jacobs DS, Puechmaille SJ, Teeling EC. 2015. How and why overcome the impediments to resolution: lessons from rhinolophid and hipposiderid bats. Molecular Biology and Evolution 32: 313 - 333.","Ranivo J, Goodman SM. 2006. Revision taxinomique des Triaenops malgaches (Mammalia, Chiroptera, Hipposideridae). Zoosystema 28: 963 - 985."]}

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."]}

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 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."]}