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line:xlsx:hash://sha256/181a039844a33e66a35a457b7ece741051086608e425a040051b79581d606b97!/Sheet1!/L1421	application/vnd.openxmlformats-officedocument.spreadsheetml.sheet	N/A	N/A	N/A	Rhinolophus luctus [synonym of]	Rhinolophus luctus perniger	Rhinolophus luctus perniger	Rhinolophus perniger	Rhinolophus luctus perniger	Rhinolophus perniger	Rhinolophus luctus [synonym of]	Rhinolophus luctus [synonym of]	Rhinolophus perniger	Rhinolophus perniger	Rhinolophus perniger	Rhinolophus perniger		[HMW] 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.; [MDD2022] split from R. luctus; previously known as lanosus, but perniger has nomenclatural priority; [batnames2023]  trifoliatus species group. Does not include lanosus ; see Volleth et al. (2017, 2021), but see also Ellerman and Morrison-Scott (1951). Does not include foetidus ; see Volleth et al. (2021). Does not include beddomei; see Topál and Csorba (1992) and Bates and Harrison (1997). Does not include formosae; see Yoshiyuki and Harada (1995) and Csorba et al. (2003). Includes perniger pending additional study (see Volleth et al., 2017, 2021; Burgin, 2019; our account for lanosus ). May include cryptic diversity; see Chornelia et al. (2022).; [MDD2023] split from R. luctus; previously known as lanosus, but perniger has nomenclatural priority; [MDD2025_2.0] split from R. luctus; previously known as lanosus, but perniger has nomenclatural priority; [batnames2025_1.7] Previously considered a subspecies of luctus, but distinct; see Volleth et al. (2017, 2021). We follow Burgin (2019) in considering lanosus a synonym of perniger, which is an older name and has priority over lanosus. Although Volleth et al. (2017, 2021) make clear that neither specimen that they have examined - from southern China (their lanosus) or south Vietnam (their R. cf. luctoides) - appears to be perniger, additional research, specifically comparisons with the type of perniger, are required to resolve this issue.; [MDD2025_2.2] split from R. luctus; previously known as lanosus, but perniger has nomenclatural priority										perniger, lanosus, spurcus				perniger, lanosus, spurcus		perniger, lanosus, spurcus		perniger, lanosus, spurcus	perniger, lanosus, spurcus, pernigei	lanosus, perniger, spurcus 		perniger B. H. Hodgson, 1843|lanosus Andersen, 1905|spurcus G. M. Allen, 1928|pernigei Cao Zhong, Wang Kunhui, Wen Jiawei, Li Chuxian, Wu Yi, Wang Xiaoyun, & Yu Wenhua, 2024 [incorrect subsequent spelling]						N/A																																								_R. p. lanosus_ Andersen, 1905; _R. p. perniger_ Hodgson, 1843; _R. p. spurcus_ Allen, 1928																											885887A2FFD58A33FF63F31AF6DED7F2	Handbook of the Mammals of the World – Volume 9 Bats, Barcelona: Lynx Edicions	978-84-16728-19-0	hbmw_9_Rhinolophidae.pdf.imf	hash://md5/7461ffdaffcf8a29ffccffa1ff85d963	328	zip:hash://sha256/ec5fd314a06aba1a7b0b72f23e54ac625ae272bd98f82f1d01f4c09627d9e8e0!/treatments-xml-main/data/88/58/87/885887A2FFFE8A1BF8B4EFE6FC7FC876.xml	Rhinolophus perniger	Rhinolophidae	Rhinolophus	perniger	Hodgson	1843	Rhinolophe noir @fr | Nördliche Wollhaar-Hufeisennase @de | Herradura lanosonorteno @es | Chinese Woolly Horseshoe Bat @en	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.	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.	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.	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 .	Northern Woolly Horseshoe Bats feed on coleopterans, termites, and other insects. Appears to forage by aerial-hawking and has also been observed perch-hunting.	No information.	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 ).	This species roosts solitarily or in pairs, rarely in groups of more than two.	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.	Bates & Harrison (1997) | Bates et al. (2004) | Csorba et al. (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 et al. (2015) | Volleth, Nguyen Truong Son et al. (2017)	https://zenodo.org/record/3750110/files/figure.png	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).	Simmons, N.B. and A.L. Cirranello. 2022B. Bat Species of the World: A taxonomic and geographic database. Accessed on 10/11/2022.	Rhinolophidae	Rhinolophus luctus perniger	Rhinolophus		lanosus													Mammal Diversity Database. (2023). Mammal Diversity Database (Version 1.11) [Data set]. Zenodo. https://doi.org/10.5281/zenodo.7830771 released 15 April 2023	Rhinolophus perniger	23	Northern Woolly Horseshoe Bat	Chinese Woolly Horseshoe Bat	Theria	Placentalia	Boreoeutheria	Laurasiatheria	CHIROPTERA	PTEROPODIFORMES	NA	NA	RHINOLOPHOIDEA	RHINOLOPHIDAE	NA	NA	Rhinolophus	NA	perniger	Hodgson	1843	0						"central region of the Sub-Himalayas," Nepal.			perniger Hodgson, 1843|lanosus K. Andersen, 1905|spurcus G. M. Allen, 1928	split from R. luctus; previously known as lanosus, but perniger has nomenclatural priority	Volleth, M., Son, N. T., Wu, Y., Li, Y., Yu, W., Lin, L. K., ... & Harada, M. (2017). Comparative Chromosomal Studies in Rhinolophus formosae and R. luctus from China and Vietnam: Elevation of R. l. lanosus to Species Rank. Acta Chiropterologica, 19(1), 41-50.	India|Nepal|Bhutan|Bangladesh|Myanmar|China|Thailand|Laos|Vietnam|Cambodia	Asia	Palearctic|Indomalaya	NA	0	0	0	Rhinolophus_perniger	0	unmatched	NA	1																																			Simmons, N. B., & Cirranello, A. L. (2023). Batnames.org Species List Version 1.4 (1.4). Zenodo. https://doi.org/10.5281/zenodo.8136157 	Rhinolophidae	Rhinolophus		luctus	Temminck	1834	0	Tijdschrift Natuurl. Gesch. Physiol.	1:23	Greater Woolly Horseshoe Bat	 geminus K. Andersen, 1905; <b> foetidus </b> K. Andersen, 1918; <b> perniger </b> Hodgson, 1843; <b>spurcus</b> Allen, 1928.	Indonesia, Java, Tapos.	India, Nepal, Burma, Sri Lanka, S China, Vietnam, Cambodia, Laos, Thailand, Peninsular Malaysia; Borneo, Sumatra, Java, and Bali (Indonesia).	Not listed.	Least Concern	 trifoliatus species group. Does not include lanosus ; see Volleth et al. (2017, 2021), but see also Ellerman and Morrison-Scott (1951). Does not include foetidus ; see Volleth et al. (2021). Does not include beddomei; see Topál and Csorba (1992) and Bates and Harrison (1997). Does not include formosae; see Yoshiyuki and Harada (1995) and Csorba et al. (2003). Includes perniger pending additional study (see Volleth et al., 2017, 2021; Burgin, 2019; our account for lanosus ). May include cryptic diversity; see Chornelia et al. (2022).	Rhinolophus perniger	1004726	23	Northern Woolly Horseshoe Bat	Chinese Woolly Horseshoe Bat	Theria	Placentalia	Boreoeutheria	Laurasiatheria	CHIROPTERA	PTEROPODIFORMES	NA	NA	RHINOLOPHOIDEA	Rhinolophidae	NA	NA	Rhinolophus	NA	perniger	Hodgson	1843	0						"central region of the Sub-Himalayas," Nepal.			perniger Hodgson, 1843|lanosus K. Andersen, 1905|spurcus G. M. Allen, 1928	split from R. luctus; previously known as lanosus, but perniger has nomenclatural priority	Volleth, M., Son, N. T., Wu, Y., Li, Y., Yu, W., Lin, L. K., ... & Harada, M. (2017). Comparative Chromosomal Studies in Rhinolophus formosae and R. luctus from China and Vietnam: Elevation of R. l. lanosus to Species Rank. Acta Chiropterologica, 19(1), 41-50.				India|Nepal|Bhutan|Bangladesh|Myanmar|China|Thailand|Laos|Vietnam|Cambodia	Asia	Palearctic|Indomalaya	NA	0	0	0	Rhinolophus_perniger	0	unmatched	NA	1	Burgin, C. J., Zijlstra, J. S., Becker, M. A., Handika, H., Alston, J. M., Widness, J., Liphardt, S., Huckaby, D. G., and Upham, N. S. (2025). How many mammal species are there now? Updates and trends in taxonomic, nomenclatural, and geographic knowledge. Journal of Mammalogy in revision: TBD. https://doi.org/10.1101/2025.02.27.640393	Rhinolophus_perniger	1004726	23	Northern Woolly Horseshoe Bat	Chinese Woolly Horseshoe Bat	Theria	Placentalia	Boreoeutheria	Laurasiatheria	Chiroptera	Yinpterochiroptera	NA	NA	Rhinolophoidea	Rhinolophidae	NA	NA	Rhinolophus	NA	perniger	B. H. Hodgson	0	Rhinolophus perniger	Hodgson, B.H. 1843. Notice of two Marmots inhabiting respectively the plains of Tibet and the Himalayan Slopes near to the Snows, and also of a _Rhinolophus_ of the central region of Nepal. Journal of the Asiatic Society of Bengal 12(137):409-414.	https://www.biodiversitylibrary.org/page/40057799				"central region of the Sub-Himalayas," Nepal.			split from R. luctus; previously known as lanosus, but perniger has nomenclatural priority	Volleth, M., Son, N. T., Wu, Y., Li, Y., Yu, W., Lin, L. K., ... & Harada, M. (2017). Comparative Chromosomal Studies in Rhinolophus formosae and R. luctus from China and Vietnam: Elevation of R. l. lanosus to Species Rank. Acta Chiropterologica, 19(1), 41-50.|Srinivasulu, A., Srinivasulu, B., Kusuminda, T., Amarasinghe, C., Ukuwela, K. D., Karunarathna, M., ... & Srinivasulu, C. (2023). Taxonomic status of the Lesser Woolly Horseshoe bats (Chiroptera, Rhinolophidae, Rhinolophus beddomei) in peninsular India and Sri Lanka. Zootaxa, 5301(2), 199-218.				India|Nepal|Bhutan|Bangladesh|Myanmar|China|Thailand|Laos|Vietnam|Cambodia	Asia	Palearctic|Indomalaya	NE	0	0	0	Rhinolophus_perniger	0	unmatched	NA	1	Simmons, N. B., & Cirranello, A. L. (2025). Batnames.org Species List Version 1.7 (1.7). Zenodo. https://doi.org/10.5281/zenodo.14796586	Rhinolophidae	Rhinolophus		perniger	Hodgson	1843	0	J. Asiat. Soc. Bengal	12(1): 414	Northern Woolly Horseshoe Bat	lanosus	China, NW Fokien, Kuatun	China	<a href='https://cites.org/eng/app/appendices.php' target='_blank'>Not Listed</a>	Not Evaluated	Previously considered a subspecies of luctus, but distinct; see Volleth et al. (2017, 2021). We follow Burgin (2019) in considering lanosus a synonym of perniger, which is an older name and has priority over lanosus. Although Volleth et al. (2017, 2021) make clear that neither specimen that they have examined - from southern China (their lanosus) or south Vietnam (their R. cf. luctoides) - appears to be perniger, additional research, specifically comparisons with the type of perniger, are required to resolve this issue.		Mammal Diversity Database. (2025). Mammal Diversity Database (Version 2.2) [Data set]. Zenodo. https://doi.org/10.5281/zenodo.15007505	NA	Rhinolophus perniger; Rhinolophus luctus perniger; Rhinolophus perniger; Rhinolophus luctus; perniger; lanosus; spurcus; perniger; lanosus; spurcus; Rhinolophe noir; Nördliche Wollhaar-Hufeisennase; Herradura lanosonorteno; Chinese Woolly Horseshoe Bat; Northern Woolly Horseshoe Bat; Chinese Woolly Horseshoe Bat; Greater Woolly Horseshoe Bat; R. perniger