Murina jaintiana, Ruedi & Biswas & Csorba & Mr, 2012

Ruedi, Manuel, Biswas, Jayant & Csorba, Gábor, 2012, Bats from the wet: two new species of Tube-nosed bats (Chiroptera: Vespertilionidae) from Meghalaya, India, Revue suisse de Zoologie 119 (1), pp. 111-135 : 120-131

Murina jaintiana sp. nov.

in part Murina tubinaris Bates & Harrison 1997: 207 .

in part Murina cineracea Csorba et al. 2011: 896 .

HOLOTYPE: MHNG 1976 .072 (field number M1619), adult male, in spirit, skull removed, collected by MR and JB on 12 February 2011.

TYPE LOCALITY: India, Meghalaya, Jaintia Hills, 2.3 km east of the village of

Kseh, 720 m a.s.l.; geographic coordinates: N 25°26’, E 92°36’.

FIG. 3

Dorsal view of hindfoot of M. pluvialis (holotype, MHNG 1976 .071) showing the attachment point of plagiopatagium .

REFERRED SPECIMENS: Myanmar, Chin State, Chin Hills BM (NH) 16.3.26.5 .,

16.3.26.7-8, 16.3.26.85-88, HNHM 2000.20.1.

ETYMOLOGY: The name refers to the local tribe, the Jaintias and the mountains named after them where the type of the new species was collected. The proposed English name is Jaintia tube-nosed bat.

DIAGNOSIS: The plagiopatagium is attached to the base of claw of the outer toe. General impression of the dorsal aspect is medium-grey; ventrally the hairs black at their basal two-third and white at the tip. Basal area of C 1 less than that of P 4, mesostyles of M 1 and M 2 rudimentary or completely missing. Forearm 29.1-31.1 mm, maxillary toothrow length 4.81-5.09 mm.

DESCRIPTION: A small-sized species of Murina (Table 2). External measurements of the holotype male are: head and body 40 mm, tail 33 mm, ear 13.9 mm, tragus 6.3 mm, hindfoot 6.8 mm, tibia 17.1 mm and forearm 29.1 mm. On the dorsal surface, the hairs have three distinct bands: basal half dark grey, almost black, middle part dirty white, distal end brownish-grey; there are no shiny guard hairs. The upper surface of uropatagium – especially along the tibia and near the body – is well haired with long FIG. 4

Lateral, dorsal and occlusal views of skull and mandibule of M. pluvialis (holotype, MHNG 1976 .071). Scale = 10 mm .

FIG. 5

Lateral views of skulls of (a) M. pluvialis (holotype, MHNG 1976.071); (b) M. sp. A ( Laos, MHNG 1926.034); (c) M. rozendaali (holotype, BM(NH) 83.360). Scale = 5 mm.

FIG. 6

Occlusal view of the upper molar row of a) M. pluvialis (holotype, MHNG 1976.071); (b) M. cyclotis (holotype, BM(NH) 9.4.4.4); (c) M. jaintiana (holotype, MHNG 1976.072). Scale = 1 mm.

grey-brown hairs, some of which reach beyond the free edge of the tail membrane. Ventrally, hairs are black for the proximal two-thirds, while the upper portion is a well demarcated pure white. The ventral aspect of the uropatagium has sparse whitish hairs (Figs 1-2). The ear is evenly rounded and without an emargination and the plagiopatagium is attached to the base of the claw on the outer toe.

The skull is small-sized (Table 3). The braincase is domed, and the rostrum not inflated. There is no sagittal crest and the lambdoid crests are only moderately developed. The narial emargination are as wide as long and the zygoma are strong with a low dorsal prominence (Figs 7-8).

The maxillary toothrows are convergent anteriorly (C 1 C 1 W/M 3 M 3 W = 0.69- 0.71). I 2 is only partly obscured in lateral view, slightly exceeded in height by I 3 and comprises approximately one third of the latter’s basal area. The basal area of C 1 is less than that of P 4, and the tooth is slenderly built and exceeds P 4 in height. The basal area of P 2 is approximately half that of P 4 and its height hardly reaches two-thirds that of P 4. The mesostyles of M 1 and M 2 are rudimentary or missing.

The lower canine (C 1) exceeds P 4 in height and is greater in basal area; P 2 basal dimensions and height are markedly less than those of P 4. M 1 and M 2 talonids equal their corresponding trigonids in area and the entoconids of these teeth exceed their hypoconids in height. The postcristid have a deep indentation and runs straight to the tip of entoconid.

TABLE 3. Selected external and craniodental measurements (in mm) of some species within the Murina “ suilla ” group. Values are given as min–max, (n). Acronyms and definitions for measurements are given in the text.

Character M. jaintiana sp. nov. M. beelzebub M. cineracea M. tubinaris

FA 29.1-31.1 (5) 33.7-36.3 (4) 27.5-33.8 (21) 31.0-32.9 (4) STOTL 14.75-15.25 (6) 16.54-16.77 (4) 14.78-16.35 (22) 14.92-15.74 (6) CCL 13.36-13.61 (2) 14.53-14.99 (4) 12.95-14.30 (23) 13.08-13.89 (5) C 1 C 1 W 3.52 -3.74 (6) 3.82-3.95 (4) 3.4-3.96 (23) 3.59-3.78 (5) M 3 M 3 W 5.04 -5.2 (6) 5.25-5.75 (4) 4.9-5.6 (23) 4.97-5.32 (5) ZYW 8.26-8.68 (4) 8.98-9.36 (4) 8.22-9.23 (21) 8.07-8.66 (4) MAW 7.23-7.43 (5) 7.65-8.08 (4) 7.19-7.82 (23) 7.27-7.51 (5) IOW 4.02-4.36 (6) 4.46-4.74 (4) 4.09-4.62 (23) 4.26-4.51 (6) BCH 5.96-6.17 (2) 6.28-6.44 (4) 5.71-6.34 (22) 5.55-5.86 (4) CM 3 L 4.81-5.09 (6) 5.41-5.54 (4) 4.84-5.36 (23) 4.88-5.19 (6) ML 9.85-10.28 (6) 10.90-11.34 (4) 9.75-10.92 (23) 10.07-10.62 (6) CM 3 L 5.18-5.46 (6) 5.81-6.00 (4) 5.15-5.78 (23) 5.37-5.69 (6) CPH 3.2-3.4 (6) 3.72-3.77 (4) 3.04-4.02 (23) 3.04-3.39 (4)

COMPARISONS WITH OTHER TAXA: On the basis of its dentition (I 2 only partly obscured by I 3 in lateral view, C 1 basal area decidedly less than that of P 4, P 2 basal area equals half that of P 4) M. jaintiana is a member of the “ suilla -group” and is therefore readily separable from all species of the “ cyclotis -group”. Within the “ suilla -group”, only M. beelzebub , M. cineracea and M. tubinaris exhibit predominantly greyish dorsal fur; all other species in the group have reddish or brownish dorsal pelage without signs of any greyish-blackish tint. M. jaintiana is, however, easily separable from the latter three greyish species by the lack of mesostyles on M 1 and M 2 (on the other species these cusps are reduced in bulk but still clearly defined). M. jaintiana is further distinguished from these species by the following features.

M. beelzebub is larger in all respects with no overlap in forearm and craniodental measurements. The closely related and similar-sized M. cineracea differs in the ventral pelage where only the proximal half is dark brown (in M. jaintiana the very dark colouration extends to two-third of the length of individual hairs). In addition, the rostrum of M. cineracea is deep, the sagittal crest always present, albeit weak, the zygoma are stronger, C 1 and P 2 are wider at the base and more robust. In M. jaintiana , the rostrum is less massive in lateral view, there is no sagittal crest, the zygoma are weaker, and it has more slender C 1 and P 2 (Fig. 8).

In M. tubinaris the plagiopatagium is attached to the proximal phalanx of the outer toe and the zygoma are characteristically weak (whereas in M. jaintiana the attachment point is the base of the claw and the zygomatic arch is relatively strong) (Fig. 8).

Although it exhibits distinctive differences in colour and in general dental structure, the sympatrically occurring M. cyclotis is also characterised by the lack of mesostyles on M 1 and M 2. However, in the case of this species the position of paracones and metacones in relation to the protocone shows a different cusps pattern (Fig. 6).

DISTRIBUTION AND ECOLOGY: The holotype specimen of M. jaintiana was caught in a harp-trap set in the understory of a bamboo grove growing along a small

FIG. 7

Lateral, dorsal and occlusal views of skull and mandibule of M. jaintiana (holotype, MHNG 1976 .072). Scale = 10 mm .

FIG. 8

Lateral views of skulls of (a) M. jaintiana (holotype, MHNG 1976.072); (b) M. beelzebub (holotype, HNHM 2007.50.24.); (c) M. cineracea (holotype, HNHM 2005.81.35.); (d) M. tubinaris ( Pakistan, HNHM 99.14.7.). Scale = 5 mm.

tributary of the Kopili River. This narrow stretch of bamboos is surrounded by secondary, semi-deciduous forest located close to the village of Kseh. This forested area is heavily exploited for firewood and regularly burnt for the needs of shifting cultivation practice. Other species of bats captured in the same area include typical forest species ( Murina cyclotis ), and species more linked to cave roosts (Rhinolophus affinis, R. pusillus, R. macrotis, R. pearsoni, R. luctus, Hipposideros cineraceus , H. larvatus , H. lankadiva , Myotis cf. longipes, Miniopterus magnater and Ia io). Several karstic caves are indeed found in the same area and bats roosting there were observed using the bamboo grove as an alley to reach feeding areas along the Kopili River and surrounding forests. It is thus unclear if M. jaintiana was roosting, hunting or commuting through this bamboo grove. The holotype male had enlarged testis, suggesting that it was sexually active at that time of the year (February). Specimens in the Chin Hills were collected at an elevation of ca. 1500 m; according to its label, one of them was caught in a hollow in a ficus tree. The Chin Hills are covered by semi-deciduous forests similar to those found in the Jaintia Hills.

Molecular comparisons

Ten of the 12 essayed specimens yielded cyt-b and rag 2 sequences (Table 1); all sequences were in frame for coding proteins, with no stop codon or insertion that could suggest the presence of non-functional copies of these genes (paralogs). The new sequences were blasted against all available sequences in the GenBank (as of 4 November 2011), but none matched those of M. jaintiana or M. pluvialis . For the cytb gene, the divergence within species of Murina (mean K2P distance = 1.9%) was about ten times smaller than interspecific divergence (mean 17.0%), which correspond with the general pattern of divergence measured among bat taxa ( Bradley & Baker, 2001, Anrawali Khan et al., 2010). Likewise, the cyt-b sequence of M. jaintiana differed from any other Murina by at least 9.6%, whereas a minimum distance of 16.9% distinguished M. pluvialis from other taxa (Table 4).

For the more conserved nuclear rag 2 gene, these figures of divergence are lower, but still M. jaintiana and M. pluvialis had unique sequences, differing by at least 2 to 10 mutations from any other related taxa (data not shown). The mean intraspecific divergence for that gene was 0% (all conspecific sequences being identical), while they differed by a mean of 2.0% in interspecific comparisons.

Phylogenetic reconstructions were largely congruent between the two methods used (BA and NJ). The more rapidly evolving and longer (1140 bp) mitochondrial cyt-b gene expectedly yielded more resolved nodes (Fig. 9), compared to the smaller (749 bp) and more conserved fragment of nuclear rag 2 gene (Fig. 10), but the different taxon sampling in both data sets impaired further comparisons of phylogenetic performance. In all reconstructions, M. jaintiana appeared closely related but distinct from Southeast Asian M. cineracea , and also differed significantly from other sympatric species sequenced from Meghalaya (i.e. M. pluvialis and M. cyclotis ). The phylogenetic position of M. pluvialis using the cyt-b data set and BA was unresolved within the Murina radiation (Fig. 9), but NJ reconstructions placed it sister to the Southeast Asian M. sp. A, albeit with low support (34% bootstrap). The nuclear rag 2 data set was more decisive in this situation, and clearly placed pluvialis sister to M. sp. A with 100% posterior probability (BA) or 83% bootstrap support (Fig. 10).

TABLE 4: Mean K2P genetic distance within (diagonal, underlined italics) and between (below diagonal) species of Murina calculated for the mitochondrial cytochrome b gene. Values are given as percentages.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16

1 M. aena -

2 M. sp. A 18.3 1.7

3 M. bicolor 19.0 17.9 0.9

4 M. cineracea 15.8 16.0 16.9 2.3

5 M. cf. cyclotis 16.8 17.8 18.4 16.4 5.3

6 M. cyclotis 16.7 16.8 19.1 16.6 12.8 -

7 M. florium 15.7 18.4 17.9 14.9 19.1 18.0 -

8 M. hilgendorfi 16.9 18.0 14.5 17.3 18.7 17.0 18.6 -

9 M. cf. huttoni 19.4 18.1 18.6 18.4 17.5 16.3 19.2 18.8 -

10 M. jaintiana sp. nov. 18.8 17.1 17.5 9.6 17.4 18.7 15.8 18.8 18.8 -

11 M. leucogaster 18.0 18.9 9.0 16.8 16.6 18.0 16.8 14.8 19.3 17.1 0.8

12 M. peninsularis 17.4 18.3 18.3 17.3 14.8 15.8 20.1 18.5 19.8 19.1 18.9 -

13 M. pluvialis sp. nov. 17.7 16.9 19.2 17.9 18.0 17.4 18.6 19.1 17.9 18.0 20.0 17.8 -

14 M. puta 20.0 18.7 17.6 18.2 19.6 18.4 19.8 18.4 8.6 20.3 18.3 18.7 20.5 -

15 M. suilla 14.5 18.7 21.2 16.9 18.1 18.7 8.3 18.8 19.3 17.8 18.6 19.1 17.6 21.1 -

16 M. tiensa 18.0 17.5 18.7 17.4 17.8 18.3 18.4 17.9 14.8 18.2 19.6 17.3 17.3 14.3 19.3 0.7

FIG. 9

Bayesian consensus tree representing the phylogenetic relationships of Murina (M.), Kerivoula (K.) and Hapiocephalus (H.) based on sequences (1140 bp) of the mitochondrial cyt-b gene. Some outgroups (genus Myotis and Cistugo) were omitted. An asterisk (*) associated to a node denotes that it is supported by at least 95% posterior probability (BA reconstructions) and /or 95% bootstrap (NJ reconstructions): other values are given as percentages. The scale bar represents 0.1 changes.

FIG. 10

Bayesian consensus tree representing the phylogenetic relationships of Murina (M.), Kerivoula (K.) and Hapiocephalus (H.) based on sequences (749 bp) of the nuclear rag 2 gene. The legend is otherwise the same as for Fig. 9.