Bat diversity in the Simandou Mountain Range of Guinea, with the description of a new white-winged vespertilionid Author Decher, Jan Zoologisches Forschungsmuseum Alexander Koenig, Leibniz Institute for Animal Biodiversity, Adenauerallee 160, 53113 Bonn, Germany & Corresponding author: E-mail: J. Decher @ zfmk. de echer@zfmk.de Author Hoffmann, Anke Museum für Naturkunde, Leibniz Institute for Evolution and Biodiversity Science, Invalidenstr. 43, 10115 Berlin, Germany Author Schaer, Juliane Museum für Naturkunde, Leibniz Institute for Evolution and Biodiversity Science, Invalidenstr. 43, 10115 Berlin, Germany & Max Planck Institute for Infection Biology, Chariteplatz 1, 10117 Berlin, Germany Author N Orris, Ryan W. Department of Evolution, Ecology and Organismal Biology, Ohio State University, 4240 Campus Dr., Lima, OH 45804, USA Author Kadjo, Blaise Université Félix Houphouët-Boigny, 22 BP 582 Abidjan 22, Côte d’Ivoire Author Astrin, Jonas Zoologisches Forschungsmuseum Alexander Koenig, Leibniz Institute for Animal Biodiversity, Adenauerallee 160, 53113 Bonn, Germany Author Monadjem, Ara All Out Africa Research Unit, Department of Biological Sciences, University of Swaziland, Private Bag 4, Kwaluseni, Swaziland & Mammal Research Institute, Department of Zoology & Entomology, University of Pretoria, Private Bag 20, Hatfield 0028, Pretoria, South Africa Author Hutterer, Rainer Zoologisches Forschungsmuseum Alexander Koenig, Leibniz Institute for Animal Biodiversity, Adenauerallee 160, 53113 Bonn, Germany text Acta Chiropterologica 2015 2015-12-01 17 2 255 282 journal article 10.3161/15081109ACC2015.17.2.003 470b137b-05b5-468a-b37e-6a315108ecc1 1733-5329 3943621 Neoromicia isabella sp. nov. Decher, Hutterer and Monadjem Isabelline White-winged Serotine Neoromicia cf. rendalli ; Monadjem et al. , 2013 b Holotype ZFMK 2008.0292 , field number JD 650, collected by Jan Decher on 7 March 2008 . Adult male in reproductive condition (testes 5 × 4 mm ), preserved in 70% ethanol, skull extracted. Tissue preserved (COI sequence GenBank no. KT598187 ). Type locality Guinea , Province Macenta, Simandou Range, Foko Confluence, 8°29’48.62’’N , 8°54’48.22’’W , ca. 765 m a.s.l., captured in net set across creek in rainforest. Paratype ZFMK 2008.0291 , field number JD 647 ; adult female (pregnant with 1 embryo), preserved in ethanol. Netted same day and place as holotype . Referred specimen DM 12619 , Mount Nimba , Liberia ; reported as Neoromicia cf. rendalli by Monadjem et al. (2013 b ) ; COI sequence deposited under GenBank no. JX508832 . Diagnosis A medium-sized pipistrelloid bat of the genus Neoromicia , based on the presence of a single upper premolar (Hill and Harrison, 1987) with white wings; only likely to be confused with N. tenuipinnis (slightly smaller and different pelage colour, see below) and N. rendalli (larger). Dorsal and ventral surface of wings pure white (as in N. tenuipinnis ), tips of dorsal pelage light rusty orange-brown (‘isabelline’), not dark brown (as in N. tenuipinnis ). Baculum similar in shape to N. rendalli but smaller. COI sequence different from that of both N. tenuipinnis and N. rendalli ( Fig. 3 ). Etymology The species epithet refers (as a noun in apposition) to the ‘Isabella’ colouration (sensu Ridgeway, 1912 ) of the dorsal fur of this bat, and also recalls the name of Jan Decher’s daughter Isabelle. Description Neoromicia isabella sp. nov. is similar in size and general colour pattern to N. tenuipinnis ( Table 3 , Figs. 4 and 5 ) but slightly larger in total length, forearm length, and body mass ( Table 3 ). Tail length is 34% of the total length in the holotype , and 37% on average. This value in the holotype of N. rendalli ( Thomas, 1889 ) is 49%, and in the holotype of N. tenuipinnis 39% ( Peters, 1872 ). The tips of the dorsal hairs are of a light isabelline colour ( Ridgeway, 1912 ), much lighter than in N. tenuipinnis (compare Figs. 4 and 5 ). The dorsal hairs are 4.5 mm long. Ears, lips, wings, uropatagium, and hindfeet are almost white and covered by very short hairs; only the ear tips and dorsal surfaces of forearm, hindfeet and tail have a pale brownish hue. Ear length is 10 mm in the holotype of N. isabella ( 10–13 mm , n = 3), and its tip is rounded ( Fig. 4 ). The tragus of the holotype ( Fig. 6 ) is 4.2 mm long, and represents 42% of the ear length. A small notch is present at the basis of the anterior surface. Compared to illustrations of N. tenuipinnis and N. rendalli in Rosevear (1965) and specimens in ZFMK and DM, the tragus of N. isabella is similar to that of N. tenuipinnis (tragus length estimated from figure: 3.9 mm ) but the shape of the notch is different ( Fig. 6 ). The penis of the holotype specimen is 6.4 mm long and covered by short whitish hairs. A digital x -ray of its baculum is shown in Fig. 7A in a semilateral view. It is slightly longer ( 3.5 mm ) than in the N. isabella specimen DM12619 ( 2.8 mm ; Fig. 7B ) reported by Monadjem et al. (2013 b : Fig. 4C ), but very similar in shape, and both are still longer than that of N. tenuipinnis (in Monadjem et al. , 2013 b : Fig. 4D ), which is 1.8 mm long and strongly bent ( Fig. 7C ). A baculum figured as N. rendalli by Kearney et al. (2002 : Fig. 1F ) is about 4 mm long. FIG. 4. Views of the head and body of N. isabella sp. nov. (holotype) after capture. Note the isabelline-colored dorsal fur and the white wings (Photographs by J. Decher) The cranium of N. isabella is illustrated in Fig 8 . Neoromicia isabella is distinguished from N. tenuipinnis (skull figured in Monadjem et al. , 2010: 479 ) by slightly larger cranial and dental measurements ( Table 4 ), particularly in the greatest skull length and the upper and lower molar rows. Greatest skull length in N. rendalli (skull figured in Monadjem et al. , 2010: 475 ) is similar to that of N. isabella , but its cranium is wider, as expressed by zygomatic width, greatest width of braincase, and mastoid width ( Table 4 ). Within the genus Neoromicia the greatest skull length of N. isabella (GSKL 12.5–13.0 mm) is of medium size and less robust than in N. rendalli , with a narrow maxillary and braincase and a fragile zygomatic arch. Skulls of 15 other species assignable to the genus Neoromicia (see accounts in Monadjem et al. , 2010 , 2013 b ; Goodman et al. , 2012 ; Happold and Happold, 2013 ) are either larger or smaller in length. The dorsal profile of the skull is slightly concave at the rostrum and slopes gradually upwards to the occiput. There is no sagittal crest, but a faint lambdoid crest. The dental formula is I2/3, C1/1, P1/2, M3/3 = 32, as in the other Neoromicia . Several observers have reported the presence of a tiny anterior upper premolar present on either or both sides of the upper jaw in N. tenuipinnis ( Hayman, 1954 ; Schlitter et al. , 1982 ; Thorn and Kerbis Peterhans, 2009). The occasional presence of this tiny anterior upper premolar is not unusual in Neoromicia species (e.g. see Rosevear, 1965 ; Happold and Happold, 2013 ); however, not one of the specimens that we examined displayed such a tooth. The anterior upper incisor is more than twice as long as the posterior (about twice in N. rendalli and N. tenuipinnis — Monadjem et al. , 2010 ) and unicuspid. The upper canine is long; 1.6-times as long as the anterior upper incisor, and 2.5-times as long as the subsequent premolar. The upper molar rows (C–M 3 ) are almost parallel; the molars are shorter than wide, and all teeth from the canine to the third molar are in contact. The mandible length is intermediate between that of N. tenuipinnis and N. rendalli ( Table 4 ). The lower incisors are all trifid. FIG. 5. Dorsal and ventral view of a live-captured N. tenuipinnis from Liberia. Note the dark dorsal fur (Photographs by A. Monadjem) FIG. 6. Outlines of the left tragus in N. rendalli (left), N. tenuipinnis (middle; both redrawn after Rosevear 1965 ), and N. isabella sp. nov. ( holotype ) Comparisons and Designation of a Neotype for Neoromicia tenuipinnis Our comparisons are hampered because the holotype of N. tenuipinnis is lost (Turni and Kock, 2008), and the original description of Peters (1872) does not include skull measurements. Likewise, the description of N. rendalli by Thomas (1889) does not contain skull measurements; however, the holotype skull (BMNH 1889.3.2.3) is figured in Monadjem et al. (2010) and in the ACR (2015 : Appendix), and was examined by AM. External measurements of the types of both taxa, however, are congruent with the current treatment of the species ( Table 3 ). Externally the new species is quite distinct. Our images ( Fig. 4 ) may be compared with those of N. tenuipinnis from Liberia ( Fig. 5 ) and another one from Comoe National Park, Côte d’Ivoire (http:// www.inaturalist.org/photos/189398). The latter species has much darker body coloration. Neoromica rendalli is also darker, and the dark-brown coloration extends to the ventral surface of the body (specimen ZFMK 1977.1015 from South Sudan ). Neoromicia isabella is immediately distinguishable from its sister species N. roseveari , itself only recently described, by its white wings and the rich colour of its fur. In N. roseveari , the wings are dark and the fur is dark brown ( Monadjem et al ., 2013 b ). Furthermore, N. roseveari is far larger (for the holotype , FA is 37.1 mm and GSKL is 14.36 mm ). The whole group, however, may include further unrecognized taxa if one looks at the genetic and morphological variation throughout Africa, which is beyond the scope of our paper. Specimens from Nigeria to Central Africa mentioned by Bergmans (1977) , Viellard (1974), Thorn and Kerbis Peterhans (2009), Bates et al. (2013) , and other authors should be re-examined in the context of a revision of the genus. It should be also noted that the most current range map for N. tenuipinnis sensu lato ( Fahr, 2013 e ) may contain records of both N. tenuipinnis sensu stricto (see below) and the newly described N. isabella . FIG. 7. Comparison of bacula. A, N. rendalli , dorsal and lateral views (adopted from Kearney et al ., 2002 ); B, C, N. isabella sp. nov. , digital x -ray in semi-lateral view (ZFMK 2008.0292, holotype) and dorsal and lateral views of DM12619; D, N. tenuipinnis (DM13235, Neotype). Scale bar = 1 mm (C, D modified from Monadjem et al ., 2013 a ) FIG. 8. Dorsal, ventral and lateral views of the cranium and lateral view of the mandible of N. isabella sp. nov. (ZFMK 2008.0292, JD 650, the holotype) from the Foko Confluence, Simandou Range, Guinea. Greatest length of skull is 12.98 mm. (Photographs by R. Hutterer) Under these circumstances, however, we consider it essential to fix the morphological and genetic characters of N. tenuipinnis and to designate a neotype for Vesperus tenuipinnis Peters, 1872 (now Neoromicia tenuipinnis ), because the former holotype , an adult female from ‘Guinea’ ( Peters, 1872: 264 ), in the Berlin Museum (ZMB) is lost (Turni and Kock, 2008). Monadjem et al. (2013 b ) listed a specimen in the Natural History Museum, London (BMNH 1889.5.1.3) as a ‘syntype’ of N. tenuipinnis , which however is incorrect, as Peters (1872: 264) only mentioned a single specimen in the Berlin Museum at hand; the one, which is now lost. Here we designate specimen DM 13235 in the Durban Natural Science Museum as the neotype . It consists of a male specimen collected by A. Monadjem at Bonlah village , near Yekepa at the base of Mount Nimba , Liberia on 7 January 2012 ; external and cranial measurements are included in Tables 3 and 4 . The body is preserved in formalin with the skull extracted. The tragus and the baculum (also Fig. 7C ) are documented in Monadjem et al. (2013 b : Figs. 4D and 6D ). A COI sequence is deposited in GenBank ( JX 508831 ). The external measurements of the neotype are very similar to those of Peters’ (1872) lost holotype ( Table 3 ). TABLE 3. External measurements (mm) and body mass (g) of specimens of the new species N. isabella sp. nov. from Guinea and Liberia and of other white-winged Neoromicia species from West Africa. Values taken from published resources marked by an asterisk

Specimen or taxon	Total length	Tail length	Forearm length	Hindfoot length	Ear length	Body mass
Neoromicia isabella sp. nov.	80	27	31.7	7.3	10	5.5
Holotype ZFMK 2008.0292
Neoromicia isabella sp. nov.	78	24	29.9	6.0	12	4.0
Paratype ZFMK 2008.0291
Neoromicia isabella sp. nov. DM 12619	74	36	28.0	5	13	4.8
Nimba, Liberia ( Monadjem et al ., 2013 b )*
Neoromicia tenuipinnis lost holotype,	76	30	29	6	11.5	–
Guinea ( Peters, 1872 )*
Neoromicia tenuipinnis Neotype DM 13235	72	30	29.3	5	12	3.5
Nimba, Liberia
Neoromicia tenuipinnis	–	–	29.0	6.2	13.3	–
Ivory Coast ( De Vree, 1971 )*
Neoromicia cf. tenuipinnis	–	31	28	5.5	11.5	–
Guinea, Nimba ( Aellen, 1963 )*
Neoromicia cf. tenuipinnis	83	30	27	–	12	–
Guinea ( Konstantinov et al ., 2000 )*
Neoromicia rendalli Holotype BMNH	89	39	36	–	13	–
1889.3.2.3, Gambia ( Thomas, 1889 )*
Neoromicia rendalli DM 13712	92	36	33.7	8.0	12	6.0
Sierra Leone (A. Monadjem, unpublished data)
Neoromicia rendalli	89.5	41.5	34.5	7.1	13.3	–
Gambia, SMF 91086 ( Kock et al ., 2002 )*
Neoromicia rendalli	86.8	37.1	37.0	7.5	12.5	–
Gambia, SMF 91087 ( Kock et al ., 2002 )*

In light of our new genetic analysis ( Fig. 3 ), the genus Neoromicia , at least in West Africa, no longer appears to be paraphyletic (as suggested by Monadjem et al., 2013 b ) but consists of a monophyletic clade. The type species of the genus, N. zuluensis (Roberts, 1924) , was studied genetically by Goodman et al. (2012) showing that it is sister to N. somalica . We are therefore confident that our new species has been assigned to the correct genus. In our COI tree N. isabella is sister to N. roseveari , and both taxa sister to N. brunnea , while N. tenuipinnis and N. rendalli group more at the root of this clade ( Fig. 3 ). Hence, whereas morphological characters suggest a closer relationship between N. tenuipinnis and N. isabella , this is not supported by the genetic analysis. Remarks This is the first record of a white-winged bat from the Simandou Range, initially identified as N. tenuipinnis . Eleven individuals were captured, all between 7 and 9 March 2008 in the canopy net over the creek at FC, two of which were kept as vouchers. They group with specimen DM12619 from Mount Nimba , Liberia , that was labelled Neoromicia cf. rendalli by Monadjem et al. (2013 b ) . Neoromica isabella was not recorded during the 2002 RAP, nor the 2003 RAP. The individuals captured in the context of this study over the forest creek at FC suggests that N. isabella uses drainages lines as travel routes probably emerging from hollow trees or rock crevices upstream. The specimen from the Liberian portion of Mount Nimba was captured emerging from the roof of a hut in a small village close to the larger settlement of Yekepa ( Monadjem et al. , 2013 b ). Records from adjacent regions ( Aellen, 1963 ; De Vree, 1971 ; Bützler, 1994; Fahr et al. , 2006 ; Monadjem and Fahr, 2007) may refer either to N. tenuipinnis or to the new species. Monadjem et al. (2013 b ) collected both species on Mount Nimba . A specimen mentioned by Roche (1971) as N. tenuipinnis deviates by very small external and cranial measurements, and until a re-examination of the voucher specimen we regard this record with caution. TABLE 4. Skull measurements (mm) of specimens of the new species N. isabella sp. nov. from Guinea and Liberia and of other white-winged Neoromicia species from West Africa, partly quoted from De Vree (1971) , Kock et al. (2002) and Monadjem et al. (2013 a ) . Measurement abreviations: GSKL — greatest skull length; CIL — condylo-incisive length; CCL — condylo-canine length; ZYGO — greatest zygomatic breadth; GBW — greatest braincase width; GSH — greatest skull height; POB — postorbital width; MAST — greatest mastoid breadth; MAND — greatest mandible length; M 3 –M 3 — width across the third molars; C–M 3 — complete upper canine-molar tooth row length; C–C — width across upper canines; C–M 3 — complete mandibular canine-molar tooth row length (see Monadjem et al. , 2013 a for more details). Values taken from published resources marked by an asterisk

Taxon and/or specimen	GSKL	CIL	CCL	ZYGO	GBW	GSH	POB	MAST	MAND	M3–M3	C–M3	C–C	C–M3
N. isabella sp. nov.	12.98	12.47	11.88	8.48	6.59	6.30	3.42	7.26	9.47	5.40	4.64	4.57	5.04
Holotype ZFMK 2008.0292
N. isabella sp. nov. DM 12619,	12.99	12.55	11.85	7.58	6.53	5.97	3.77	6.96	9.32	5.33	4.46	4.15	5.35
Nimba, Liberia ( Monadjem et al ., 2013 a )
N. tenuipinnis Neotype DM 13235,	12.50	11.35	11.06	7.32	6.53	5.41	3.80	6.75	8.62	5.03	4.31	3.82	4.48
Nimba, Liberia ( Monadjem et al ., 2013 a )
N. tenuipinnis 12.0, 12.4		11.6, 11.7	–	–, 7.5	–	–	–	–	–	5.0, 5.1	–	3.7, 3.7	4.7, 4.7
Ivory Coast ( De Vree, 1971 )*
N. rendalli	13.60	13.10	12.80	–	7.50	5.80	4.00	7.80	10.00	6.00	4.80	4.80	5.10
Holotype BMNH 1889.3.2.3
N. rendalli	12.98	12.57	–	9.20	7.18	–	3.90	7.98	10.19	–	4.65	4.35	5.03
Gambia, SMF 91086 ( Kock et al ., 2002 )*
N. rendalli	13.61	12.57	–	9.90	6.93	–	3.94	8.05	10.17	–	4.79	4.28	5.26
Gambia, SMF 91087 ( Kock et al ., 2002 )*
N. rendalli Sierra Leone, DM 13712	13.36	12.90	12.58	–	7.27	5.57	4.09	7.54	9.55	5.76	4.44	4.01	4.98
N. rendalli S. Sudan, ZFMK 77.1015	13.36	12.70	12.29	8.92	7.14	6.16	3.72	7.71	9.45	5.83	4.60	4.23	5.17

Conservation status The conservation status of this species has not yet been evaluated ( IUCN, 2015 ). Neoromicia roseveari Monadjem, Richards, Taylor and Stoffberg, 2013 New material ZFMK 2008.0293, ♂ , FC, 7 March 2008 ; ZFMK 2008.0294, ♀ , FC, 7 March 2008 . In the molecular analysis ( Fig. 3 ), the two individuals of this species captured at FC group with specimens that Monadjem et al. (2013 b ) described as a new species, N. roseveari , from Mount Nimba . The dark fur colour dorsally and ventrally and measurements fit well with those published in the description. Similar to the Mount Nimba specimens, our individuals were caught in lower lying areas of the study area in dense rainforest over the creek at FC. The female carried an embryo of 20 mm crownrump length. Conservation status The conservation status of this species has not yet evaluated ( IUCN, 2015 ).