Rhinolophus perniger, Hodgson, 1843

Burgin, Connor, 2019, Rhinolophidae, Handbook of the Mammals of the World – Volume 9 Bats, Barcelona: Lynx Edicions, pp. 280-332 : 328-329

99 . Northern Woolly Horseshoe Bat

Rhinolophus perniger

French: Rhinolophe noir / German: NördlicheWollhaar-Hufeisennase /Spanish: Herradura lanoso norteno

Other common names: Chinese Woolly Horseshoe Bat

Taxonomy. Rhinolophus perniger Hodgson, 1843 ,

“ central region of the Sub-Himalayas ,” Nepal.

Rhinolophus perniger is in the trifoliatus species group. It was included under R luctus but is now recognized as a distinct species based on karyological and morphological data presented by M. Volleth and colleagues in 2017. They compared the karyotype of a specimen from China (considered to be lanosus ) to the recently described R. luctoides, proving their distinctiveness. This specimen from China is morphologically similar to specimens assigned to perniger from India, which demonstrates that they are the same species. Volleth and colleagues considered this taxon to be R lanosus, but the name perniger has priority. They did not investigate taxonomic status of other named forms that are included here as subspecies of R.perniger (i.e. perniger , lanosus , and spurcus), so taxonomic status of these three names is still up for debate. See R. luctus , R. luctoides , and R. morio . Specimens in Vietnam presented by Volleth and colleagues in 2017 had an identical karyotype to R. luctoides from Malaysia and were tentatively named R. cf. luctoides . In early study by Volleth and colleagues in 2015, limited genetic data indicated that specimens now identified as R.perniger from Myanmar and China clustered with specimens of R. luctoides , and R. morio was sister to this clade. This suggests that R. luctoides might be not be distinct from. perniger or that some parts of the distribution of. perniger might actually represent. luctoides . Relationships among currently recognized species in the R. luctus complex (R. pemiger, R. morio , R. luctoides , and R. luctus ) remain uncertain. There might be undescribed species in what is currently recognized as R.perniger because some specimens throughout its distribution do not match its typical description according to Volleth and colleagues in 2015. Extensive genetic, morphological, and karyological studies are needed to clarify this and relationships among species in the R. luctus complex. Distribution of. perniger relative to R. morio and R. luctoides is uncertain, and distributions given here are tentative. Three subspecies recognized.

Subspecies and Distribution.

R. p. perniger Hodgson, 1843 - N, C & SW India (Uttarakhand, Sikkim, West Bengal, Assam, Meghalaya, Nagaland, Madhya Pradesh, Karnataka, and Kerala), C & E Nepal, Bhutan, Bangladesh, Myanmar, S China (Yunnan), N Thailand, Laos, Vietnam, and Cambodia.

. p. lanosus K. Andersen, 1905 — C & SE China (Sichuan, Guizhou, Guangxi, Anhui Zhejiang, Jiangxi, Fujian, and Guangdong).

R.p. spurcus G. M. Allen, 1928 - Hainan I, China.

Possibly also peninsular Thailand, according to recorded calls.

Descriptive notes. Head—body 75—99 mm, tail 36-61 mm, ear 28—44 mm, hindfoot 16- 20 mm, forearm 68- 5-81 mm; weight 27-45 g. (Note that some of these ranges, with the exception of the forearm values, probably include data from taxa currendy regarded as separate species.) The Northern Woolly Horseshoe Bat is the largest species of horseshoe bat. The basic craniodental features are surprisingly similar in all species in the complex; for general description, see the Great Woolly Horseshoe Bat (. luctus ). This species has not yet been compared morphologically to typical Great Woolly Horseshoe Bats from Indonesia. However, it seems that currently available craniodental morphometric data clearly—albeit based on limited dataset—differentiate this species from the Selangor Woolly Horseshoe Bat (. luctoides ). Baculum, as described from Indian specimens, is longer (6-7 mm long) than in the Selangor Woolly Horseshoe Bat and the Malaysian Woolly Horseshoe Bat (. morio ), has wide base with shallow medial bifurcation in dorsal view, and has a somewhat flattened tip that is slightly bulged. A specimen from Vietnam that showed karyotypic similarities to the Selangor Woolly Horseshoe Bat had similar baculum to this species, 6- 1 mm long. Chromosomal complement has 2n = 32 and FNa = 60.

Habitat. The Northern Woolly Horseshoe Bat is a forest-dweller which can also be found in degraded forest (and is apparently somewhat tolerant of human disturbance). In China, most records have been collected in forested areas. Recorded from sea level up to elevations of 923 m.

Food and Feeding. Northern Woolly Horseshoe Bats feed on coleopterans, termites, and other insects. Appears to forage by aerial-hawking and has also been observed perch-hunting.

Breeding. No information.

Activity patterns. Northern Woolly Horseshoe Bats roost by day in caves, and occasionally tree hollows and man-made structures. They leave their roosts early in the evening. The species hibernates in parts of the Himalayas and in northern Vietnam. Call shape is FM/CF/FM with a peak F of 33 kHz (the 110 kHz value given in a publication for Vietnam is definitely an error), 32—34 kHz in Laos, and apparently 40 kHz in Thailand (probably from the first harmonic rather than the second).

Movements, Home range and Social organization. This species roosts solitarily or in pairs, rarely in groups of more than two.

Status and Conservation. Not assessed on The IUCNed List, (included under R luctus , which is listed as Least Concern). The Northern Woolly Horseshoe Bat is widespread as currently defined but until its taxonomy is settled, its conservation status cannot be properly assessed.

Bibliography. Bates & Harrison (1997), Bates eta/. (2004), Csorba eta /. (2003), Francis (2008a), Harada, Yenbutra.Yosida &Takada (1985), Hendrichsen, Bates & Hayes (2001), Kruskop (2013a), Matveev (2005), Molur et a /. (2002), Sinha (1973), Smith & XieYan (2008), Srinivasulu & Srinivasulu (2012), Thong Vu Dinh (2014 b), Volleth, Loidl eta/. (2015), Volleth, Nguyen Truong Son eta/. (2017).

Figures

On following pages: 94. Thomas's Horseshoe Bat (Rhinolophus thomasi); 95. Lesser Horseshoe Bat (Rhinolophus hipposideros); 96. rancis's Woolly Horseshoe Bat (Rhinolophus trancisi); 97. Lesser Woolly Horseshoe Bat (Rhinolophus sedulus); 98. Trefoil Horseshoe Bat (Rhinolophus trifoliatus); 99. Northern Woolly Horseshoe Bat (Rhinolophus perniger); 100. Selangor Woolly Horseshoe Bat (Rhinolophus luctoides); 101. Malaysian Woolly Horseshoe Bat (Rhinolophus mono); 102. Great Woolly Horseshoe Bat (Rhinolophus luctus); 1.03. Beddome's Woolly Horseshoe Bat (Rhinolophus beddomei); 104. Formosan Woolly Horseshoe Bat (Rhinolophus formosaë); 105. Thailand Horseshoe Bat (Rhinolophus thailandensis); 106. Dobson's Horseshoe Bat (Rhinolophus yunanensis); 107. Chiew Kwee's Horseshoe Bat (Rhinolophus chiewkweeae); 108. Pearson's Horseshoe Bat (Rhinolophus pearsonii); 109. Mitred Horseshoe Bat (Rhinolophus mitratus).

Subspecies and Distribution. R. p. perniger Hodgson, 1843 - N, C & SW India (Uttarakhand, Sikkim, West Bengal, Assam, Meghalaya, Nagaland, Madhya Pradesh, Karnataka, and Kerala), C & E Nepal, Bhutan, Bangladesh , Myanmar , S China (Yunnan), N Thailand, Laos , Vietnam , and Cambodia . . p. lanosus K. Andersen, 1905 — C & SE China (Sichuan, Guizhou, Guangxi, Anhui Zhejiang, Jiangxi, Fujian, and Guangdong). R.p. spurcus G. M. Allen, 1928 - Hainan I, China . Possibly also peninsular Thailand , according to recorded calls.