taxonID	type	description	language	source
7E05356DFFC6FFACD484FB1C94A4F9FC.taxon	discussion	REMARKS This species was described by Geoffroy (1803) from the “ île de Bourbon ” (La Réunion) based on two individuals sent to Paris by a M. Macé. Hill (1980) discusses the historical details of this material. The two specimens were apparently divided between the MNHN and the Musée des Pays-Bas in Leiden, which would be later called the Rijksmuseum van Natuurlijke Historie. Neither of these specimens has been located in the MNHN, nor are they referred to by Rode (1941) in his catalogue of type specimens in that collection. However, one of the specimens was registered in the museum’s catalog (Moutou 1982). In Jentink’s (1888: 184, entry c) catalogue of bats in the Leiden Museum it is mentioned under the heading Scotophilus borbonicus Geoffroy, “ Individu adulte monté, figuré dans la Mammalogie de Temminck, T. II, Pl. XLVII, fig. 7. Bourbon. Du voyage de M. Macé ”. It is almost without question that this specimen (RMNH 28508) is one of the two animals É. Geoffroy used in the original description of this species, and, based on this logic, Hill (1980) designated it as the lectotype of S. borbonicus. After the inspection of collections and queries at numerous natural history museums, including the MNHN and Muséum d’Histoire naturelle of Saint-Denis (La Réunion), as well as literature citations (e. g., Hill 1980; Moutou 1982; Robbins et al. 1985), the lectotype in the RMNH appears to be the only known extant specimen in the world of S. borbonicus from La Réunion. It is in very poor condition. The mounted skin has partially extended wings and associated membranes are brittle, partially broken, and the distal portions frayed. An incision made in the lower abdomen, presumably when it was prepared as a mounted skin, is only partially closed. Further, the pelage is foxed, but certain aspects of the original coloration can still be clearly discerned. The dorsum of the specimen is a reddish-brown and the ventrum a dull whitish (Table 1). In Geoffroy’s (1803: 55) original description it is noted, “ pelage marron en dessus, blanchâtre en dessous ”, which closely fits with our judgment of this specimen’s current coloration. The associat- ed skull is partially broken, with the cranium largely shattered – rendering it impossible to make numerous measurements. As mentioned earlier, Dorst (1947) noted the occurrence of S. borbonicus on Madagascar, presumably based on the Grandidier material in the MNHN. Of the three small Malagasy Scotophilus specimens in that collection, one (MNHN 1976.420) from Sarodrano has a distinctly reddish-brown dorsum and light-colored ventrum. This specimen was collected in 1868 and has been stored in alcohol over the intervening years, and thus it is not unexpected that the natural pelage coloration has become washed-out. Even given the poor condition of the lectotype, these two specimens (RMNH 28508 and MNHN 1976.420) showed considerable resemblance in basic pelage coloration and comparable cranial and dental structures (Tables 1 - 5), and we are inclined to identify MNHN 1976.420 as S. borbonicus. However, until new material from La Réunion of S. borbonicus is uncovered in museum collections or collected in the wild, final determination of MNHN 1976.420 will not be possible. At the present time we refer this specimen to S. cf. borbonicus. The tragus of the MNHN specimen is rounded towards the apex and has a simple peduncle attachment (Fig. 2). In Figure 3 we have illustrated the skull and mandible of MNHN 1976.420 for future comparative purposes. One striking aspect of the history of moderate to small Scotophilus species on Madagascar is that two different sea caves near the village of Sarodrano, presumably exactly at or close to where Grandidier collected his specimens, and the nearby village of Saint-Augustin have been extensively surveyed over the past few years by our field teams and no example of this genus has been found. While it is true that much of our capture work has been conducted at cave entrances, a site type Scotophilus rarely use for their day-roosts, we have surveyed synanthropically-living bat species in the village of Saint- Augustin. Further, the late R. L. Peterson also captured bats in the sea caves of Sarodrano in 1967 and no Scotophilus was collected. Even more exceptional is that amongst the two specimens of Scotophilus obtained by Grandidier in the Sarodrano region are two different species – S. cf. borbonicus and a second species that is new to science and described below. Although the specimen labels indicate that these individuals were captured in caves, it is possible that they were obtained in the nearby village of Sarodrano, which would have been a series of thatched houses during the period of Grandidier’s visit, a construction style Scotophilus frequently inhabit (Kingdon 1974). Considerable work has been conducted over the past century of the bats of the Mascarene Islands, particularly La Réunion. The reputed presence of S. borbonicus on Mauritius Island is erroneous (Cheke & Dahl 1981), and there is no evidence of a Scotophilus on the Comoro Islands (Louette et al. 2004). Given that members of this genus are often associated with man-made shelters and are relatively easy to catch, the absence of any s u b s e q u e n t r e c o r d s o f S. b o r b o n i c u s o n L a Réunion is rather notable (Cheke & Dahl 1981; Moutou 1982). This would imply that this species is exceptionally rare, difficult to capture, part of an extralimital migratory population, or even extinct. Given that currently available material is insufficient to properly diagnose S. borbonicus, the possibility cannot be completely eliminated that the provenance of the two Macé specimens is incorrect or that these captured individuals were vagrants to La Réunion. It has been suggested that borbonicus might be conspecific with African leucogaster (Hill in Cheke & Dahl 1981). The same point might hold for MNHN 1976.420 in that it may not be a representative of a resident population on Madagascar. In a recent action plan for microchiropteran bats, Hutson et al. (2001) list- ed S. borbonicus as occurring on La Réunion and Madagascar and considered it to be critically endangered.	en	Goodman, Steven M., Jenkins, Richard K. B., Ratrimomanarivo, Fanja H. (2005): A review of the genus Scotophilus (Mammalia, Chiroptera, Vespertilionidae) on Madagascar, with the description of a new species. Zoosystema 27 (4): 867-882, DOI: 10.5281/zenodo.4525389
7E05356DFFCAFFA4D4CBFCFC9026F9FC.taxon	description	(Figs 3 - 5; Tables 1 - 5)	en	Goodman, Steven M., Jenkins, Richard K. B., Ratrimomanarivo, Fanja H. (2005): A review of the genus Scotophilus (Mammalia, Chiroptera, Vespertilionidae) on Madagascar, with the description of a new species. Zoosystema 27 (4): 867-882, DOI: 10.5281/zenodo.4525389
7E05356DFFCAFFA4D4CBFCFC9026F9FC.taxon	materials_examined	HOLOTYPE. — Adult *, 27. VII. 2003, R. K. B. Jenkins and F. H. Ratrimomanarivo coll., field number RBJ 161 (UADBA 46923). The specimen was preserved in formalin and the skull extracted and subsequently cleaned. The specimen is in a good state of preservation, except that the skull has a cut mark across ventral posterior portion. Muscle tissue samples taken from the upper breast were preserved in EDTA, resulting in small cuts across both sides of the chest. PARATYPES. — Mahabo, 20 ° 23 ’ S, 44 ° 38 ’ E, 29. IV. 1869, A. Grandidier, 1 + (MNHN 1984. 433); Grotte de Sarodrano, 1868, A. Grandidier, 1 + (MNHN 1912.48). Both specimens were preserved in fluid and the skulls have been extracted and cleaned. TYPE LOCALITY. — Madagascar, Province de Mahajanga, just outside the limit of the Parc national de Bemaraha, 1.8 km SE from Bekopaka and 0.6 km NE from Andadoany, 19 ° 08.454 ’ S, 44 ° 48.732 ’ E, about 50 m above sea-level (Fig. 1).	en	Goodman, Steven M., Jenkins, Richard K. B., Ratrimomanarivo, Fanja H. (2005): A review of the genus Scotophilus (Mammalia, Chiroptera, Vespertilionidae) on Madagascar, with the description of a new species. Zoosystema 27 (4): 867-882, DOI: 10.5281/zenodo.4525389
7E05356DFFCAFFA4D4CBFCFC9026F9FC.taxon	etymology	ETYMOLOGY. — The name tandrefana is derived from the Malagasy meaning “ from the west ”.	en	Goodman, Steven M., Jenkins, Richard K. B., Ratrimomanarivo, Fanja H. (2005): A review of the genus Scotophilus (Mammalia, Chiroptera, Vespertilionidae) on Madagascar, with the description of a new species. Zoosystema 27 (4): 867-882, DOI: 10.5281/zenodo.4525389
7E05356DFFCAFFA4D4CBFCFC9026F9FC.taxon	description	MEASUREMENTS. — Measurements taken directly from the fluid preserved specimen are noted with an asterisk. External: total length 111 mm, tail length * 46 mm, hind foot * 7 mm, tragus length 7 mm, ear length * 13 mm, forearm length * 47 mm. Weight: 14.2 g. Skull and teeth: GSKL 17.9 mm, CBL 16.7 mm, ZYGO 12.3 mm, IOW 4.2 mm, BCW 10.5 mm, PAL 8.2 mm, C 1 - C 1 5.9 mm, M 3 - M 3 7.8 mm, C-M 3 5.8 mm, I-M 3 7.1 mm, and MAND 11.6 mm (Tables 2 - 4).	en	Goodman, Steven M., Jenkins, Richard K. B., Ratrimomanarivo, Fanja H. (2005): A review of the genus Scotophilus (Mammalia, Chiroptera, Vespertilionidae) on Madagascar, with the description of a new species. Zoosystema 27 (4): 867-882, DOI: 10.5281/zenodo.4525389
7E05356DFFCAFFA4D4CBFCFC9026F9FC.taxon	distribution	DISTRIBUTION. — Scotophilus tandrefana n. sp. is currently known from three localities in western Madagascar: Bemaraha, Mahabo, and Sarodrano (Fig. 1) – all below 100 m elevation. On the basis of current information there is no evidence that this species is strictly forest dwelling or synanthropic. In January 1993 Martin Göpfert and colleagues captured a small Scotophilus in the Kirindy (CFPF) Forest to the northeast of Morondava (20 ° 04.6 ’ S, 44 ° 40.5 ’ E, 30 m; Fig. 1). The pelage coloration of this individual was noted as “ ventrum being uniformly brown and hair on upper side light brown with dark brown tips ” (M. Göpfert pers. comm., 25 May 2004). The forearm was measured as 45.5 mm. On the basis of these characters this individual is probably referable to S. tandrefana n. sp. HABITAT. — The type specimen was captured in a 9 m mist-net placed in an open grassy clearing adjacent to rice fields and disturbed deciduous forest and within 100 m of a limestone outcrop. The original natural habitat of this region is dry semi-deciduous forest and the capture site is about 200 m from the relatively intact natural forest formations of the Parc national de Bemaraha.	en	Goodman, Steven M., Jenkins, Richard K. B., Ratrimomanarivo, Fanja H. (2005): A review of the genus Scotophilus (Mammalia, Chiroptera, Vespertilionidae) on Madagascar, with the description of a new species. Zoosystema 27 (4): 867-882, DOI: 10.5281/zenodo.4525389
7E05356DFFCAFFA4D4CBFCFC9026F9FC.taxon	diagnosis	DIAGNOSIS. — A member of the genus Scotophilus of small size with average forearm length of 44 - 47 mm. Muzzle is pronounced, relatively short, and pug-like. Slightly elongated crescent-shaped nostrils opening slightly antero-laterally (Fig. 4). Distinctly long forward projecting tragus with a slightly complex peduncle (Fig. 2). Dorsal fur is relatively long, soft, and a uniform dark brown, while the throat and ventral pelage is shorter and finer, and a lighter mediumbrown (Fig. 4). Wing membranes and uropatagium dark. Relatively well developed lamboidal and sagittal crests. Dental formula 1 / 3 - 1 / 1 - 1 / 2 - 3 / 3.	en	Goodman, Steven M., Jenkins, Richard K. B., Ratrimomanarivo, Fanja H. (2005): A review of the genus Scotophilus (Mammalia, Chiroptera, Vespertilionidae) on Madagascar, with the description of a new species. Zoosystema 27 (4): 867-882, DOI: 10.5281/zenodo.4525389
7E05356DFFCAFFA4D4CBFCFC9026F9FC.taxon	description	DESCRIPTION Amongst the three specimens available for study (holotype and two paratypes) there is not a marked difference in fur coloration between the dorsal and ventral surfaces. In the recently collected holotype specimen the dorsal pelage is a dense and rich dark chocolate brown and the basal portions are distinctly lighter brown. The ventrum pelage coloration, including the throat and upper breast, is a medium-brown, that appears to become lighter posteriorly, and basally Malagasy Scotophilus (Mammalia, Chiroptera, Vespertilionidae) a grayish-brown. The paratypes have been stored in fluid for well over a century and the pelage color is washed-out. Nonetheless, the contrasting dark dorsum and slightly lighter ventrum is clearly discernable in these specimens. The wing membrane and uropatagium are dark brownishblack in the holotype. The muzzle is relatively short and rounded (Fig. 4). Slightly elongated almost tubular nostrils open in a slightly lateral position. The upper lips have a regular, but not dense, covering of hairs. Scotophilus tandrefana n. sp. is a small species in external measurements, particularly when compared to African and Asian members of this genus (Table 2). The black ears are short (13 mm) and fall outside the range of most African species of Scotophilus. There is some variation in the sickle-shaped tragus of S. tandrefana n. sp., but the attachment peduncle of the tragus is a slightly complex structure, rather than a simple attachment stalk as in S. borbonicus (Fig. 2). The apex of the tragus terminates as a slightly roundpointed shape in S. tandrefana n. sp. The skull of S. tandrefana n. sp. is relatively diminutive in size, particularly when compared to other species of small Scotophilus such as S. leucogaster and S. viridis. S. tandrefana n. sp. has a slightly short and broad rostrum (without expanded lacrimal processes), expanded braincase, and tapered postorbital constriction (Fig. 5). The lambdoidal and sagittal crests are well developed, forming the typical “ helmet ” of members of this genus, but less prominent than in adults of most other species. Posterior palatal extension terminates as acute spine. Zygomatic arches slightly flared. Anterior emargination of palate is deep and broad. Pterygoids expanded posteriorly and wing-shaped. The dental configuration is typical of other Scotophilus species (Koopman 1994: 128). The single upper pair of incisors is trifid and upper and lower canines well developed and powerful. M 1 and M 2 have a reduced mesostyle, with a dis- torted W-shaped cusp pattern, and M 3 is greatly reduced in size. PM 1 and PM 2 have the trigonid distinctly larger than the talanoid.	en	Goodman, Steven M., Jenkins, Richard K. B., Ratrimomanarivo, Fanja H. (2005): A review of the genus Scotophilus (Mammalia, Chiroptera, Vespertilionidae) on Madagascar, with the description of a new species. Zoosystema 27 (4): 867-882, DOI: 10.5281/zenodo.4525389
7E05356DFFCAFFA4D4CBFCFC9026F9FC.taxon	discussion	REMARKS Of the 12 species of Scotophilus recognized worldwide by Simmons (in press), the following species (generalized distribution in parentheses) fall within the approximate size range of S. tandrefana n. sp. based on forearm length (Table 1): S. borbonicus (La Réunion and possibly Madagascar [see above]), S. dinganii (broad range in sub-Saharan Africa), S. kuhlii Leach, 1821 (Indonesia to Pakistan), S. leucogaster (broad range in sub-Saharan Africa), S. nucella Robbins, 1984 (montane zones of western to eastern Africa), S. nux Thomas, 1904 (montane zones of western to eastern Africa), and S. viridis (broad range in sub-Saharan Africa). Other species not included in these analyses because they are larger than S. tandrefana n. sp. include S. celebensis Sody, 1928, S. heathii Horsfield, 1831, S. nigrita (Schreber, 1774), and S. robustus (Koopman 1994; Taylor 2000). One species was excluded because its insular southeastern Asian range is a considerable distance from Madagascar (S. collinus Sody, 1936). The other Malagasy members of the genus Scotophilus can be easily distinguished from S. tandrefana n. sp. These include S. robustus based on its notably larger measurements (Tables 2 - 5) and S. cf. borbonicus using pelage coloration and certain cranial measurements (Tables 2; 3). S. tandrefana n. sp. can be easily distinguished from several extralimital African and Asiatic members of this genus with approximately the same forearm length (Table 2) by pelage characters. S. leucogaster, S. dinganii, and S. viridis have notably lighter ventrums as compared to their dorsums (Table 1). The remaining three species, S. nux, S. nucella, and S. kuhlii can be differentiated from S. tandrefana n. sp. by a variety of external, cranial, and dental measurements (Tables 2 - 5). Further, the ventral fur coloration of kuhlii is notably lighter than the backside. Of the Scotophilus species that fall in the same general size-range of S. tandrefana n. sp. based on forearm length, S. nux and S. nucella are the only two that match its basic pelage col- oration pattern (Table 1), although in both cases these two species are notably larger in numerous cranial measurements (Table 3). The phylogenetic relationships of S. tandrefana n. sp. are not addressed here, as R. Trujillo is currently conducting a taxonomic revision of African and Malagasy members of this genus based on morphological and molecular characters. Further, research on the acoustics of members of this genus might provide insight into their phylogenetic relationships.	en	Goodman, Steven M., Jenkins, Richard K. B., Ratrimomanarivo, Fanja H. (2005): A review of the genus Scotophilus (Mammalia, Chiroptera, Vespertilionidae) on Madagascar, with the description of a new species. Zoosystema 27 (4): 867-882, DOI: 10.5281/zenodo.4525389
