Scotophilus tandrefana, Goodman & Jenkins & Ratrimomanarivo, 2005

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), pp. 867-882 : 875-881

Scotophilus tandrefana n. sp.

( Figs 3-5 ; Tables 1-5)

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 ).

ETYMOLOGY. — The name tandrefana is derived from the Malagasy meaning “from the west”.

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).

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.

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.

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.

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.

Figures

FIG. 3. — Dorsal, ventral and lateral views of adult crania and mandible of Scotophilus cf. borbonicus (MNHN 1976.420; greatest skull length = 19.0 mm) from Sarodrano.

FIG. 4. — Color photograph of anterior portion holotype of Scotophilus tandrefana n. sp. (UADBA 46923, field number RBJ 161) obtained near the Parc National de Bemaraha (photograph by R. K. B. Jenkins).

FIG. 5. — Dorsal, ventral, and lateral views of adult crania and mandible of holotype of Scotophilus tandrefana n. sp. (UADBA 46923; greatest skull length = 17.9 mm) from near the Parc national de Bemaraha.

FIG. 1. — Map of Madagascar with localities mentioned in the text. Symbols: [, collection site; *, other localities.

FIG. 2. — Right ear and tragus of Scotophilus from Madagascar; A, holotype of S. tandrefana n. sp. (UADBA 46923); B, S. cf. borbonicus (MNHN 1976.420); C, paratype of S. tandrefana n. sp. (MNHN 1984.433). 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 (É. Geoffroy, 1803).