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line:xlsx:hash://sha256/181a039844a33e66a35a457b7ece741051086608e425a040051b79581d606b97!/Sheet1!/L947	application/vnd.openxmlformats-officedocument.spreadsheetml.sheet	Myotis pruinosus	Myotis pruinosus	Myotis pruinosus	Myotis pruinosus	Myotis pruinosus	Myotis pruinosus	Myotis pruinosus	Myotis pruinosus	Myotis pruinosus	Myotis pruinosus	Myotis pruinosus	Myotis pruinosus	Myotis pruinosus	Myotis pruinosus	Myotis pruinosus		[MSW2] Subgenus Leuconoe.; [MSW3] Reviewed by Yoshiyuki (1989), also see Horácek et al. (2000) and Kawai et al. (2003).; [HMW] Myotis pruinosus Yoshiyuki, 1971 , Geto Hot Spring, Waga-Machi, Waga-Gun, Iwate prefecture , north-eastern Honshu, Japan . Subgenus Myotis ; montivagus species group. See M. annectans . Most recent phylogenetic studies show that M. pruinosus as sister to M. yanbarensis and M. secundus and more distantly related to montivagus species group. Ancestral M. pruinosus and M. yanbarensis might have adapted to warm temperatures, with probable origin in South-east Asia. Monotypic.; [batnames2022] Reviewed by Yoshiyuki (1989), also see Horácek et al. (2000) and Kawai et al. (2003).; [IUCN] Woodman (1993) notes that many mammalian generic names ending in -otis use the wrong gender for specific names. If this advice is to be followed, the species name should be M. pruinosa . The Chiroptera SG advise keeping the names as they are for now while a decision is still to be made on these.; [batnames2023] Reviewed by Yoshiyuki (1989), also see Horácek et al. (2000) and Kawai et al. (2003).; [batnames2025_1.7] Reviewed by Yoshiyuki (1989), also see Horácek et al. (2000) and Kawai et al. (2003).														pruinosus	Woodman (1993) notes that many mammalian generic names ending in -otis use the wrong gender for specific names. If this advice is to be followed, the species name should be M. pruinosa . The Chiroptera SG advise keeping the names as they are for now while a decision is still to be made on these.			pruinosus 	pruinosus 			pruinosus Yoshiyuki, 1971		Corbet, G.B. and Hill, J.E. 1980. A World List of Mammalian Species. British Museum (Natural History), London, 226 pp.		N Honshu, Japan	Honacki, J.H., Kinman, K.E. and Koeppl, J.W. 1982. Mammal Species of the World: A Taxonomic and Geographic Reference. Allen Press, Lawrence, 694 pp.	Myotis pruinosus	Japan, Honshu, Iwate Pref., Waga-Gun, Waga-Machi.	Yoshiyuki	1971	Bull. Nat. Sci. Mus. Tokyo, 14:305.	Distribution: Known only from Honshu in Japan.		Corbet, G.B. and Hill, J.E. 1991. A World List of Mammalian Species. Third edition. Oxford University Press, London, 243 pp. ISBN 0-19-854017-5		N Honshu, Japan	Koopman, K.F. 1993. Order Chiroptera. Pp. 137–242 in Wilson, D.E. and Reeder, D.M. (eds.). Mammal Species of the World: A Taxonomic and Geographic Reference. Second edition. Smithsonian Institution Press, Washington, 1206 pp.	Yoshiyuki	1971	Bull. Natl. Sci. Mus. Tokyo, 14:305.	Subgenus Leuconoe.	Honshu and Shikoku (Japan).	Japan, Honshu, Iwate Pref., Waga-Gun, Waga-Machi.		YOSHIYUKI	1971	Size fairly small (forearm length, 30-33 mm; condylobasal length, 12-13 mm). Margin of uropatagium attached to side of foot, well removed from the ankle.	Distribution: Known only from Honshu in Japan.	No subspecies.		108	species	M. pruinosus	YOSHIYUKI	1971	Leuconoe	subgenus	Myotis pruinosus				Size fairly small (forearm length, 30-33 mm; condylobasal length, 12-13 mm). Margin of uropatagium attached to side of foot, well removed from the ankle.	No subspecies.		72. M. pruinosus YOSHIYUKI 1971 [daubentoni group].	72	NA			Don E. Wilson & DeeAnn M. Reeder (editors). 2005. Mammal Species of the World. A Taxonomic and Geographic Reference (3rd ed), Johns Hopkins University Press, 2,142 pp. (Available from Johns Hopkins University Press, 1-800-537-5487 or (410) 516-6900, or at http://www.press.jhu.edu).	CHIROPTERA	Vespertilionidae	Myotinae		Myotis pruinosus	Myotis		pruinosus	Yoshiyuki		1971		Bull. Natl. Sci. Mus. Tokyo	14		305		Frosted Myotis	Japan, NE Honshu, Iwate Pref., Waga-Gun, Waga-Machi, Geto Hot Spring.	Honshu and Shikoku (Japan).	IUCN 2003 and IUCN/SSC Action Plan (2001) – Endangered.		Reviewed by Yoshiyuki (1989), also see Horácek et al. (2000) and Kawai et al. (2003).	4C3D87E8FF3C6A83FA769446198EB086	Handbook of the Mammals of the World – Volume 9 Bats, Barcelona: Lynx Edicions	978-84-16728-19-0	hbmw_9_Vespertilionidae_716.pdf.imf	hash://md5/b004ff90fffb6a44fffc96591e00bb32	961	zip:hash://sha256/ec5fd314a06aba1a7b0b72f23e54ac625ae272bd98f82f1d01f4c09627d9e8e0!/treatments-xml-main/data/4C/3D/87/4C3D87E8FF3C6A83FA769446198EB086.xml	Myotis pruinosus	Vespertilionidae	Myotis	pruinosus	Yoshiyuki	1971	Murin givré @fr | Honshu-Bartfledermaus @de | Ratonero de Japén @es | Blackish \Whiskered Bat @en	Myotis pruinosus Yoshiyuki, 1971 , Geto Hot Spring, Waga-Machi, Waga-Gun, Iwate prefecture , north-eastern Honshu, Japan . Subgenus Myotis ; montivagus species group. See M. annectans . Most recent phylogenetic studies show that M. pruinosus as sister to M. yanbarensis and M. secundus and more distantly related to montivagus species group. Ancestral M. pruinosus and M. yanbarensis might have adapted to warm temperatures, with probable origin in South-east Asia. Monotypic.	Endemic to Japan , recorded from Honshu, Shikoku, and Kyushu Is.	Head-body 37-44 mm ,tail 33-40 mm , ear 10-3-13- 3 mm , hindfoot 7-5—8- 6 mm , forearm 30-34 mm ; weight 4-7 g . Fur dense, velvety, and short (c.5- 5 mm in length on middle of back). Dorsum is dark grayish brown, conspicuously frosted with ashy or whitish tips. Venteris slightly lighter than dorsum,frosted with buffy hair tips in adults and whitish in young. Ears are narrow and short; tragusis relatively short (5- 5-7 mm ). Hindfeet with claws are large (c.64% oftibia); lower leg is slender. Plaglopatagium is attached at base offirst toe. Tail and calcar are long, and calcar lacks keel. Skull has broad zygomatic breadth (zygomatic width ¢.65% of condylo-basal length) and narrow interorbital constriction (width ¢.50% of braincase). In dorsal view, braincase is laterally swollen; sagittal and temporal crests are poorly developed; and lambdoid crestis laterally strong. Dorsal profile from anterior nasal part to posterior nasal region is almoststraight and then rises abruptly to frontal region. I? and I? have secondary cusps and are subequal in height and crown area, distance between I* and canine narrower than transverse diameter of I’; P? and P*® are similar in shape, the latter c.33% the former in crown area and height and positioned in tooth row. Upper molars have distinct protoconules. There are three pairs of lower incisors; I, is larger than I, and I, forming V-shaped row between canines. P, and P, are subequal in size and in tooth row. Condylo-basal lengths are 12-12- 8 mm ; maxillary tooth row lengths are 4-9-5- 3 mm . Chromosomal complement has 2n = 44 and FN = 52. The Frosted Myotis has pericentric inversion on chromosome 1. Y-chromosome is submetacentric, and X-chromosomeis subtelocentric.	Intact mature primary mountain forests down to elevations of 200-300 m . The Frosted Myotis was assumed to be more widely distributed in warm temperature evergreen lowland broadleaf forests on south-western Japanese plains, but it is presently absent due to lack of large trees containing crevices and cavities and survives only in low-elevation mountains that contain nearly mature forests with tree holes for roosting.	No information.	Parturition occurs in mid-July. Litter size is one young.	The Frosted Myotis has been observed flying along forest edges, trails, and streams and above grasslands at night. There are no records from caves except for one hibernating individual from Shikoku, which was found in a crack in the ceiling. Tree hollows are primary diurnal refuges. Echolocation call is FM type, with average end frequency of 38-7 kHz (33—41-8 kHz), peak frequency of 52-4 kHz (49-7-54-7 kHz), and duration of 2:8 milliseconds (2-6-3 milliseconds).	No information.	Classified as Endangered on The IUCN Red List. Extent of occurrence of the Frosted Myotisis less than 5000 km ?, whichis severely fragmented and faces ongoing decline of mature forests. Deforestation of old-growth evergreen forests for agriculture, conifer plantations, logging, and infrastructure development for expanding human settlements is a major threat. It occurs in some protected areas.	Abe et al. (2005) | Amadoretal. (2018) | Corbet (1978) | Funakoshi (2010) | Harada & Uchida (1982) | Horacek et al. (2000) | Kawai et al. (2003) | Koopman (1994) | Maeda (2008f) | Ohdachi et al. (2009) | Ruedi, Csorba et al. (2015) | Ruedi, Stadelmann etal. (2013) | Simmons (2005) | Tsytsulina (2004) | Volleth & Heller (2012) | Yoshiyuki (1971, 1989) | Zhang Zhenzhen et al. (2009)	https://zenodo.org/record/6398890/files/figure.png	450. Frosted Myotis Myotis pruinosus French: Murin givré / German: Honshu-Bartfledermaus / Spanish: Ratonero de Japén Other common names: Blackish \Whiskered Bat Taxonomy. Myotis pruinosus Yoshiyuki, 1971 , Geto Hot Spring, Waga-Machi, Waga-Gun, Iwate prefecture , north-eastern Honshu, Japan . Subgenus Myotis ; montivagus species group. See M. annectans . Most recent phylogenetic studies show that M. pruinosus as sister to M. yanbarensis and M. secundus and more distantly related to montivagus species group. Ancestral M. pruinosus and M. yanbarensis might have adapted to warm temperatures, with probable origin in South-east Asia. Monotypic. Distribution. Endemic to Japan , recorded from Honshu, Shikoku, and Kyushu Is. Descriptive notes. Head-body 37-44 mm ,tail 33-40 mm , ear 10-3-13- 3 mm , hindfoot 7-5—8- 6 mm , forearm 30-34 mm ; weight 4-7 g . Fur dense, velvety, and short (c.5- 5 mm in length on middle of back). Dorsum is dark grayish brown, conspicuously frosted with ashy or whitish tips. Venteris slightly lighter than dorsum,frosted with buffy hair tips in adults and whitish in young. Ears are narrow and short; tragusis relatively short (5- 5-7 mm ). Hindfeet with claws are large (c.64% oftibia); lower leg is slender. Plaglopatagium is attached at base offirst toe. Tail and calcar are long, and calcar lacks keel. Skull has broad zygomatic breadth (zygomatic width ¢.65% of condylo-basal length) and narrow interorbital constriction (width ¢.50% of braincase). In dorsal view, braincase is laterally swollen; sagittal and temporal crests are poorly developed; and lambdoid crestis laterally strong. Dorsal profile from anterior nasal part to posterior nasal region is almoststraight and then rises abruptly to frontal region. I? and I? have secondary cusps and are subequal in height and crown area, distance between I* and canine narrower than transverse diameter of I’; P? and P*® are similar in shape, the latter c.33% the former in crown area and height and positioned in tooth row. Upper molars have distinct protoconules. There are three pairs of lower incisors; I, is larger than I, and I, forming V-shaped row between canines. P, and P, are subequal in size and in tooth row. Condylo-basal lengths are 12-12- 8 mm ; maxillary tooth row lengths are 4-9-5- 3 mm . Chromosomal complement has 2n = 44 and FN = 52. The Frosted Myotis has pericentric inversion on chromosome 1. Y-chromosome is submetacentric, and X-chromosomeis subtelocentric. Habitat. Intact mature primary mountain forests down to elevations of 200-300 m . The Frosted Myotis was assumed to be more widely distributed in warm temperature evergreen lowland broadleaf forests on south-western Japanese plains, but it is presently absent due to lack of large trees containing crevices and cavities and survives only in low-elevation mountains that contain nearly mature forests with tree holes for roosting. Food and Feeding. No information. Breeding. Parturition occurs in mid-July. Litter size is one young. Activity patterns. The Frosted Myotis has been observed flying along forest edges, trails, and streams and above grasslands at night. There are no records from caves except for one hibernating individual from Shikoku, which was found in a crack in the ceiling. Tree hollows are primary diurnal refuges. Echolocation call is FM type, with average end frequency of 38-7 kHz (33—41-8 kHz), peak frequency of 52-4 kHz (49-7-54-7 kHz), and duration of 2:8 milliseconds (2-6-3 milliseconds). Movements, Home range and Social organization. No information. Status and Conservation. Classified as Endangered on The IUCN Red List. Extent of occurrence of the Frosted Myotisis less than 5000 km ?, whichis severely fragmented and faces ongoing decline of mature forests. Deforestation of old-growth evergreen forests for agriculture, conifer plantations, logging, and infrastructure development for expanding human settlements is a major threat. It occurs in some protected areas. Bibliography. Abe et al. (2005), Amadoretal. (2018), Corbet (1978), Funakoshi (2010), Harada & Uchida (1982), Horacek et al. (2000), Kawai et al. (2003), Koopman (1994), Maeda (2008f), Ohdachi et al. (2009), Ruedi, Csorba et al. (2015), Ruedi, Stadelmann etal. (2013), Simmons (2005), Tsytsulina (2004), Volleth & Heller (2012), Yoshiyuki (1971, 1989), Zhang Zhenzhen et al. (2009).	Simmons, N.B. and A.L. Cirranello. 2022B. Bat Species of the World: A taxonomic and geographic database. Accessed on 10/11/2022.	Vespertilionidae	Myotis pruinosus	Myotis	Unassigned-Myotis	pruinosus	Yoshiyuki	1971	0	Bull. Natl. Sci. Mus. Tokyo	0.7951	Frosted Myotis	None.	Japan, NE Honshu, Iwate Pref., Waga-Gun, Waga-Machi, Geto Hot Spring.	Honshu and Shikoku (Japan).	Not listed.	Endangered	Reviewed by Yoshiyuki (1989), also see Horácek et al. (2000) and Kawai et al. (2003).	Mammal Diversity Database. (2023). Mammal Diversity Database (Version 1.11) [Data set]. Zenodo. https://doi.org/10.5281/zenodo.7830771 released 15 April 2023	Myotis pruinosus	23	Frosted Myotis	Blackish Whiskered Bat	Theria	Placentalia	Boreoeutheria	Laurasiatheria	CHIROPTERA	VESPERTILIONIFORMES	NA	NA	VESPERTILIONOIDEA	VESPERTILIONIDAE	MYOTINAE	NA	Myotis	Myotis	pruinosus	Yoshiyuki	1971	0	Myotis_pruinosus	Yoshiyuki, M. (1971). A new bat of the Leuconoe group in the genus Myotis from Honshu, Japan. Bulletin of the National Science Museum, Tokyo, 14(3), 305.		NSMT-M14842		Geto Hot Spring, Waga-Machi, Waga-Gun, Iwate prefecture, north-eastern Honshu, Japan.			pruinosus Yoshiyuki, 1971	NA	NA	Japan	Asia	Palearctic	EN	0	0	0	Myotis_pruinosus	0	sciname match	Myotis_pruinosus	0	IUCN. 2022. The IUCN Red List of Threatened Species. Version 2022-1. https://www.iucnredlist.org. Accessed on [28 September, 2022].	14192	Myotis pruinosus	ANIMALIA	CHORDATA	MAMMALIA	CHIROPTERA	VESPERTILIONIDAE	Myotis	pruinosus	Yoshiyuki, 1971	Woodman (1993) notes that many mammalian generic names ending in -otis use the wrong gender for specific names. If this advice is to be followed, the species name should be M. pruinosa . The Chiroptera SG advise keeping the names as they are for now while a decision is still to be made on these.	200000000	Myotis pruinosus	Endangered	B2ab(iii)	2019	2018-07-23 00:00:00 UTC	3.1	English	This species is undergoing a continuing decline in quality of habitat, its population is severely fragmented, and its area of occupancy (AOO) is estimated at 120 km², distributed in nine (9) locations. Its forest habitat is declining due to deforestation primarily for agriculture, conversion to conifer plantations, logging and infrastructure development for expanding human settlements. This species therefore is listed as Endangered under criterion B2ab(iii).	This species has been recorded widely within Japan, but most of the localities are from natural forest. Although it has been considered as a forest-dwelling species, several individuals were recorded from caves (Maeda et al. 2013, Yoshiyuki 1974). It has been observed foraging along forest edges, trails and above streams and grasslands (Endo 1976). One hibernating bat was recorded in a crevice in the roof of a cave (Yoshiyuki 1974).	Main threats are represented by deforestation of the evergreen old-growth forests in warm temperate regions, primarily for agriculture, conversion to conifer plantations, logging and infrastructure development for expanding human settlements.	There is no information on population size at present.	Unknown	<p><span lang="EN-US">This species is endemic to Japan, and has been recorded from Honshu, Shikoku and Kyushu (Kawai 2015). The species range is severely fragmented and it is currently known from nine (9) locations.</p>		Terrestrial	Some of the recorded localities are from protected areas. It is listed as Endangered VU) in the Japanese Red List (2014).	Palearctic		FALSE	FALSE	Global	Simmons, N. B., & Cirranello, A. L. (2023). Batnames.org Species List Version 1.4 (1.4). Zenodo. https://doi.org/10.5281/zenodo.8136157 	Vespertilionidae	Myotis	Unassigned - Myotis	pruinosus	Yoshiyuki	1971	0	Bull. Natl. Sci. Mus. Tokyo	0.795139	Frosted Myotis	None.	Japan, NE Honshu, Iwate Pref., Waga-Gun, Waga-Machi, Geto Hot Spring.	Honshu and Shikoku (Japan).	Not listed.	Endangered	Reviewed by Yoshiyuki (1989), also see Horácek et al. (2000) and Kawai et al. (2003).	Myotis pruinosus	1005459	23	Frosted Myotis	Blackish Whiskered Bat	Theria	Placentalia	Boreoeutheria	Laurasiatheria	CHIROPTERA	VESPERTILIONIFORMES	NA	NA	VESPERTILIONOIDEA	Vespertilionidae	MYOTINAE	NA	Myotis	Myotis	pruinosus	Yoshiyuki	1971	0	Myotis_pruinosus	Yoshiyuki, M. (1971). A new bat of the Leuconoe group in the genus Myotis from Honshu, Japan. Bulletin of the National Science Museum, Tokyo, 14(3), 305.		NSMT-M14842		Geto Hot Spring, Waga-Machi, Waga-Gun, Iwate prefecture, north-eastern Honshu, Japan.			pruinosus Yoshiyuki, 1971	NA	NA				Japan	Asia	Palearctic	EN	0	0	0	Myotis_pruinosus	0	sciname match	Myotis_pruinosus	0	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	Myotis_pruinosus	1005459	23	Frosted Myotis	Blackish Whiskered Bat	Theria	Placentalia	Boreoeutheria	Laurasiatheria	Chiroptera	Yangochiroptera	NA	NA	Vespertilionoidea	Vespertilionidae	Myotinae	NA	Myotis	Myotis	pruinosus	Yoshiyuki	0	Myotis pruinosus	Yoshiyuki, M. 1971-09-30. A new bat of the _Leuconoe_ group in the genus _Myotis_ from Honshu, Japan. Bulletin of the National Science Museum, Tokyo (n.s.)14(3):305-310.		NSMT-M 14842	holotype		Geto Hot Spring, Waga-Machi, Waga-Gun, Iwate prefecture, north-eastern Honshu, Japan.			NA	NA				Japan	Asia	Palearctic	EN	0	0	0	Myotis_pruinosus	0	sciname match	Myotis_pruinosus	0	Simmons, N. B., & Cirranello, A. L. (2025). Batnames.org Species List Version 1.7 (1.7). Zenodo. https://doi.org/10.5281/zenodo.14796586	Vespertilionidae	Myotis	Myotis	pruinosus	Yoshiyuki	1971	0	Bull. Natl. Sci. Mus. Tokyo	0.795139	Frosted Myotis	None.	Japan, NE Honshu, Iwate Pref., Waga-Gun, Waga-Machi, Geto Hot Spring.	Honshu and Shikoku (Japan).	<a href='https://cites.org/eng/app/appendices.php' target='_blank'>Not Listed</a>	<a href='https://www.iucnredlist.org/species/14192/209551299/' target='_blank'>Endangered</a>	Reviewed by Yoshiyuki (1989), also see Horácek et al. (2000) and Kawai et al. (2003).		Mammal Diversity Database. (2025). Mammal Diversity Database (Version 2.2) [Data set]. Zenodo. https://doi.org/10.5281/zenodo.15007505	NA	Myotis pruinosus; Myotis pruinosus; Myotis pruinosus; Myotis pruinosus; Myotis pruinosus; Myotis pruinosus; pruinosus; Murin givré; Honshu-Bartfledermaus; Ratonero de Japén; Blackish \Whiskered Bat; Frosted Myotis; Blackish Whiskered Bat; Frosted Myotis; Frosted Myotis; M. pruinosus