Murina feae ( Thomas 1891 )

Francis, Charles M. & Eger, Judith L., 2012, A review of tube-nosed bats (Murina) from Laos with a description of two new species, Acta Chiropterologica 14 (1), pp. 15-38 : 24-27

Murina feae ( Thomas 1891)

( Figs. 3h , 4h , 5h , 6d ; Tables 1 , 2; Map Fig. 1b )

Harpiocephalus feae: Thomas 1891: 884 ; type locality Biapo , Burma .

Murina tubinaris : authors as summarized in Csorba et al. (2011), including Francis et al. (1999, 2010), Francis (2008), but not Scully (1881).

Murina cineracea: Csorba and Furey 2011: 896 ; type locality Seima Biodiversity Conservation Centre , Mondulkiri, Cambodia .

Specimens examined from Laos

ROM: 9 ♂♂, 7 ♀♀ ; EBD 3 ♂♂, 2 ♀♀ ; SMF: 3 ♂♂, 1 ♀ (see Appendix for details, including information from adjacent countries and the literature) .

Description

The overall colour is greyish, but with multiple bands of colour on the hairs ( Fig. 6d ). Most of the hairs of the dorsum have dark grey-brown to blackish bases for about 60–65% of their length, then a buffy gray band, then dark gray brown tips. Some specimens have a more brownish cast than others. The longer, guard hairs have pale tips, giving a frost- ed appearance, and an overall impression of 4 bands of colour in the back fur. The ventral fur has dark grey or black bases for up to 75% of the length, with silvery white tips. The legs and all of the interfemoral membrane are sparsely covered, above and below, with grayish brown hairs. The hairs are denser and longer on the legs than on the interfemoral membrane. The ear is rounded with a long narrow tragus about half the length of the ear.

In the skull, the braincase is somewhat inflated, with a well developed rostral depression, and the rostrum relatively narrow ( Fig. 3h ). The anterior upper premolar (P 2) is very small, less than half the height and surface area of the posterior premolar (P 4), while the canine is only slightly longer than P 4.

Discussion

These specimens match the description of bats referred by Hill (1963) to M. tubinaris in both skull morphology and the fur coloration, including the banding pattern of the hairs.

Csorba et al. (2011) suggested that M. tubinaris is restricted to Pakistan and northwest India (the type locality is Gilgit, Kashmir), and they proposed the name M. cineracea for the species that occurs from east India through southeast Asia.

We have subsequently examined the holotype of Murina feae ( Thomas 1891) which was collected at Biapo in the Karin Hills, ≈ 70 km northeast of Toungoo, Myanmar (estimated coordinates 19°20’N, 96°40’E) and preserved in the Museo Civico di Storia Naturale in Genova, Italy (MSNG 44307; field catalogue number 306). We suggest that this represents the same species as M. cineracea and thus is the prior name for the species.

The type description by Thomas (1891: 884) was very brief, indicating that the specimen was similar to M. aurata but distinguished by “being brown instead of golden yellow, by the smaller nasal tubes and by having the forearms, hind limbs, and posterior edge of the interfemoral membrane almost naked.” Thomas (1893: 926) provided a few more details, including “general colour smoky brown, the tips of the hairs above brown, below white” and indicated that the wings were connected to the base of the ungual phalanx. He also wrote “the arms being only very thickly clothed with scattered brown hairs, the hind limbs but little more hairy, and the interfemoral membrane although thickly covered on the greater part of its surface, yet with its edge not fringed.” [The term “only” suggests he may have meant to write “thinly” rather thickly with respect to the arm hairs.] Murina feae was tentatively consider- ed a synonym of M. aurata by Hill (1992) and Maeda (1980), presumably on the basis of this description because neither author had examined the specimen.

Murina feae differs from the most similar species currently known from the region, M. eleryi (which was formerly confused with the similar looking M. aurata ), in several features of fur colour and skull shape. The dorsal fur of M. eleryi also has dark and light bands, but the banding pattern is less distinct with shorter dark bases, and the fur is more brownish with golden tips (see pictures in Furey et al., 2009 and description below). Although it is possible golden tips could fade over time, Thomas (1893) explicitly noted the lack of golden tips when he described the specimen. Furthermore, the skull of M. feae can be differentiated from M. eleryi by its slightly larger and more robust skull, thickened rostrum, slightly larger canines and thickened zygoma.

Ruedi et al. (2012) recently described M. jaintiana from Megalaya, India as a species distinct from M. tubinaris and M. cineracea based, in part, on genetic differences. Genetically, analysis of cytochrome b sequences suggest M. jaintiana is more closely allied to Lao and Vietnamese M. feae than to any other species so far sequenced, but nevertheless diverges by nearly 10% ( Ruedi et al., 2012; J. L. Eger, unpublished data). Ruedi et al. (2012) noted that M. jaintiana differed from M. cineracea in several morphological features including reduced mesostyles on M 1 and M 2, stronger zygomata, lack of a sagittal crest, more extensive dark bases to the ventral fur, and insertion of the wing membrane at the base of the claw. In most of these characters, the holotype of M. feae resembles our specimens from Laos and Vietnam. The upper molars have small but distinct mesostyles and the zygoma are weak. The sagittal crest is not apparent, but this character is variable among our specimens from Laos, Vietnam and southern China. The dark bases on the ventral fur of several of our Lao and Vietnamese specimens extend up to 75% of the length of the hairs, while others are less extensive, indicating this is not a useful diagnostic character. The wing membrane on all of our specimens insert- ed on the side of the toe slightly below the base of the claw.

Thus, we conclude that M. feae is the prior name for the species occurring in southeast Asia that was described in Csorba et al. (2011) as M. cineracea . We acknowledge the possibility that future genetic studies may demonstrate multiple cryptic species in the region resembling M. feae , in which case this conclusion would need to be reexamined. In the meantime, in the absence of evidence to the contrary, it seems appropriate to treat M. cineracea as a junior synonym of M. feae .

Distribution and ecology

Murina feae was first reported from Laos by Osgood (1932) (as M. tubinaris ) based on specimens from Phongsali in northern Laos. Although not quite as widespread as M. cyclotis , our surveys indicate the species was nevertheless relatively common, with 42 individuals captured from most major localities where we surveyed. It has also been reported from many localities in adjacent countries ( Fig. 1b , Appendix). It was caught in a range of habitats from lowland mixed forest to montane evergreen rainforest.

Figures

FIG. 3. Skull profiles of specimens of eight species of Murina from Laos: (a) M. harrisoni, EBD 24974; (b) M. fionae sp. nov. ROM 106383; (c) M. huttoni ROM 106419; (d) M. cyclotis, ROM 106431; (e) M. annamitica sp. nov. ROM 106467; (f) M. eleryi, ROM 106411; (g) M. walstoni ROM 110450; (h) M. feae, ROM 106477. Scale bar = 5 mm

FIG. 4. Upper left toothrows of specimens of eight species of Murina from Laos: (a) M. harrisoni, EBD 24974; (b) M. fionae sp. nov. ROM 106383; (c) M. huttoni ROM 106419; (d) M. cyclotis, ROM 106431; (e) M. annamitica sp. nov., ROM 106467; (f) M. eleryi, ROM 106411; (g) M. walstoni ROM 110450; (h) M. feae, ROM 106477. Scale bar = 5 mm

FIG. 5. Lower left toothrows of specimens of eight species of Murina from Laos: (a) M. harrisoni, EBD 24974; (b) M. fionae sp. nov. ROM 106383; (c) M. huttoni ROM 106419; (d) M. cyclotis, ROM 106431; (e) M. annamitica sp. nov., ROM 106467; (f) M. eleryi, ROM 106411; (g) M. walstoni ROM 110450; (h) M. feae, ROM 106477. Scale bar = 5 mm

FIG. 6. Photographs of four species of Murina in life from Laos: (a) M. walstoni ROM 110450; (b) M. fionae sp. nov. ROM 106382 or 106383; (c): M. cyclotis ROM 110647; (d) M. feae ROM 110476

FIG. 1. Distribution of known specimen records of Murina from Laos and surrounding countries for (a) M. cyclotis, (b) M. feae, (c) M. huttoni, and (d) M. eleryi. Solid circles (●) mark locations for specimens examined by the authors from Laos, Vietnam or southern China or extracted from Genbank or the Barcodes of Life databases, while open circles (○) mark locations extracted from the published literature (see Appendix for specimen details). On map (b), the solid star represents the type locality of M. feae, while the open star represents the type locality for M. cineracea. On map (d), the open star represents the type locality for M. eleryi

FIG. 7. Photographs of the dorsal and ventral side of the holotype of M. feae MSNG 44307. The specimen had been preserved in alcohol since 1888 and some of the colour has been partly bleached; the distinctive banding pattern is still apparent, but the dark colours were likely darker brown, dark grey or black

FIG. 8. Skull of the holotype of M. feae MSNG 44307. Scale bar = 5 mm