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(1=author & date in parentheses)	Citation	Pages	Common Name	Synonyms	Type Locality	Distribution	CITES	IUCN	Comments	column3781	column3791	subtribe	CONCAT_ALTNAMES
line:xlsx:hash://sha256/181a039844a33e66a35a457b7ece741051086608e425a040051b79581d606b97!/Sheet1!/L1425	application/vnd.openxmlformats-officedocument.spreadsheetml.sheet	N/A	N/A	N/A	Rhinolophus cornutus [synonym of]	N/A	Rhinolophus cornutus pumilus	Rhinolophus pumilus	Rhinolophus pumilus	Rhinolophus pumilus	N/A	Rhinolophus pumilus	Rhinolophus pumilus	Rhinolophus pumilus	Rhinolophus pumilus	Rhinolophus pumilus		[MSW3] Formerly included in hirundo, but apparently distinct; see Happold et al. (1987), Happold and Happold (1989), Taylor and Van Der Merwe (1998), and Cotterill (2001d).; [HMW] Rhinolophus comutus pumilus K. Andersen, 1905 , “ Loo-choo Islands ,” Okinawa Prefecture , Japan . Rhinolophus pumilus is included in the pusillus species group and appears to be sister species to. comutus. Echolocation frequency of. pumilus is higher than in. cornutus , but this increase follows the generally increasing cline in the echolocation frequency within R comutus from north to south. Nevertheless, based on unpublished genetic data collected by K. Armstrong, A pumilus is tentatively treated as a species here, but an in-depth study is urgently needed; the probably extinct taxon miyakonis from Miyako Island, occasionally treated as subspecies of A pumilus , is here treated as a synonym. Monotypic.; [batnames2022]  pusillus species group. Previously considered a subspecies of cornutus . We follow Burgin (2019) who provisionally considers pumilus distinct from cornutus based on unpublished genetic data. Note also that based on Li et al. (2006) and our decision to treat pusillus , monoceros , and cornutus as separate species, we further consider the position of pumilus in this study as potential evidence of its specific distinctiveness. Provisionally includes miyakonis (Burgin, 2019). See Corbet and Hill (1992), Csorba et al. (2003), and Wu et al. (2012) for an alternate view.; [MDD2022] split from R. cornutus; includes the name miyakonis; [batnames2023]  pusillus species group. Previously considered a subspecies of cornutus . We follow Burgin (2019) who provisionally considers pumilus distinct from cornutus based on unpublished genetic data. Note also that based on Li et al. (2006) and our decision to treat pusillus , monoceros , and cornutus as separate species, we further consider the position of pumilus in this study as potential evidence of its specific distinctiveness. Provisionally includes miyakonis (Burgin, 2019). See Corbet and Hill (1992), Csorba et al. (2003), and Wu et al. (2012) for an alternate view.; [MDD2023] split from R. cornutus; includes the name miyakonis; [MDD2025_2.0] split from R. cornutus; includes the name miyakonis; [batnames2025_1.7] pusillus species group. Previously considered a subspecies of cornutus. We follow Burgin (2019) who provisionally considers pumilus distinct from cornutus based on unpublished genetic data. Note also that based on Li et al. (2006) and our decision to treat pusillus, monoceros, and cornutus as separate species, we further consider the position of pumilus in this study as potential evidence of its specific distinctiveness. Provisionally includes miyakonis (Burgin, 2019). See Corbet and Hill (1992), Csorba et al. (2003), and Wu et al. (2012) for an alternate view.; [MDD2025_2.2] split from R. cornutus; includes the name miyakonis											miyakonis	pumilus, miyakonis, orii		pumilus, miyakonis		pumilus	pumilus - miyakonis	pumilus, miyakonis 	pumilus, miyakonis 	pumilus 	pumilus - miyakonis	pumilus Andersen, 1905|miyakonis Kuroda, 1924						N/A																																								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	Vespertilioninae	Nycticeiini	Scotoecus hindei	Scotoecus		hindei	Thomas		1901		Ann. Mag. Nat. Hist., ser. 7	7		264		Hinde's Lesser House Bat	Kenya, Kitui.	Nigeria and Cameroon to S Sudan and Somalia; south to SE Dem. Rep. Congo, Kenya, Tanzania, Zambia, Mozambique, Malawi.	IUCN 2003 – Not evaluated; not considered in IUCN/SSC Action Plan (2001).	falabae Thomas, 1915.	Formerly included in hirundo, but apparently distinct; see Happold et al. (1987), Happold and Happold (1989), Taylor and Van Der Merwe (1998), and Cotterill (2001d).	885887A2FFED8A0BFF69F855F71AD1EF	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	311	zip:hash://sha256/ec5fd314a06aba1a7b0b72f23e54ac625ae272bd98f82f1d01f4c09627d9e8e0!/treatments-xml-main/data/88/58/87/885887A2FFEF8A09FF7EF311F69DD03C.xml	Rhinolophus pumilus	Rhinolophidae	Rhinolophus	pumilus	K. Andersen	1905	Okinawa Horseshoe Bat @en | Rhinolophe d'Okinawa @fr | Okinawa-Hufeisennase @de | Herradura de Okinawa @es | Spanish @en | Okinawa Little Horseshoe Bat @en	Rhinolophus comutus pumilus K. Andersen, 1905 , “ Loo-choo Islands ,” Okinawa Prefecture , Japan . Rhinolophus pumilus is included in the pusillus species group and appears to be sister species to. comutus. Echolocation frequency of. pumilus is higher than in. cornutus , but this increase follows the generally increasing cline in the echolocation frequency within R comutus from north to south. Nevertheless, based on unpublished genetic data collected by K. Armstrong, A pumilus is tentatively treated as a species here, but an in-depth study is urgently needed; the probably extinct taxon miyakonis from Miyako Island, occasionally treated as subspecies of A pumilus , is here treated as a synonym. Monotypic.	S Japan , Okinawa Is, Iheyajima (one record in 1992), Tokashikijima (one record in 1978), Kumejima, Miyakojima (probably extinct after 1971), and Irabu Is.	Head—body 36-43- 3 mm , ear 19- 1 mm (average), hindfoot 8- 6 mm (average), forearm 38-40- 5 mm . The Okinawa Horseshoe Bat is similar to the Little Japanese Horseshoe Bat (. cornutus ). Pelage is woolly and glossy and the dorsal pelage is medium brown (bases of hair being lighter) whereas the ventral pelage is similar but paler (darker than the Littlejapanese Horseshoe Bat ). Ears are moderate-sized. Noseleaf has a concave-sided lancet; connecting process is subtriangular and curved with a slightly concave or straight anterior margin; sella is pandurate, being constricted above the middle, and has a broadly rounded tip; horseshoe is not very broad but broader than in the Littlejapanese Horseshoe Bat (7- 2 mm ), with no lateral leaflets, and short, narrow median emargination. Lower lip has three mental grooves. Baculum averages 4- 2 mm long and has a very deep dorsal notch on basal cone with small ventral notch; shaft is nearly cylindrical and is bent on distal one-half. Skull is long (longer than in the Littlejapanese Horseshoe Bat ) and narrow, and zygomatic width is subequal to mastoid width; anterior median swellings are small and subcircular; posterior compartments are moderate in size; rostral profile slopes gently posteriorly or is rarely almost straight; sagittal crest is weakly to moderately developed; frontal depression varies from moderate to fairly deep; supraorbital crests are conspicuous but not generally sharp. P2 is minute and completely external to tooth row ( occasionally absent); P3 is medium-sized to small and is external to tooth row. Dental formula is the usual of 32 teeth for the genus or only 30 teeth when an upper premolar is missing. Chromosomal complement has 2n = 62 and FNa = 60.	Woodland habitats.	The Okinawa Horseshoe Bat is insectivorous.	The species appears to exhibit restricted seasonal monoestry and has been observed breeding in November to January. Females give birth to a single young from May to June.	The Okinawa Horseshoe Bat forages during the night and roosts by day in natural caves, abandoned mines, and bomb shelters. Search call shape is FM/ CF/ FM . On northern Okinawajima, males had peak frequencies of 105-6—110-9 kHz (mean of 108-2 kHz) whereas females had 107-7—113 kHz (mean of 110-6 kHz). Peak frequencies on southern Okinawajima were even higher, with males at 112-8— 119-3 kHz (mean of 116 kHz) and females at 104-6—120-1 kHz (mean of 117-4 kHz). Females had significantly higher average frequencies than males on Okinawajima.	The Okinawa Horseshoe Bat roosts to colonies.	Not assessed as a separate species on The IUCN Red List, where it is included under the Least Horseshoe Bat ( pusillus } as Least Concern. Listed as endangered in theJapanese Red List . The Okinawa Horseshoe Bat probably became extinct on Miyako c.40 years ago and on Irabu c .15 years ago, although more intensive surveys are needed to confirm this. Population on Kumejima has experienced declines. The species appears to be threatened by habitat destruction and by disturbances related to tourism in the Ryukyu Islands .	Ando et al. (1980c) | Chen Shiangfan etal. (2006) | Fukui etal. (2009) | Hutson, Kingston & Walston (2008) | Li Gang etal. (2006) | Ohdachi et al. (2015) | WuYi, Motokawa et al. (2012) | Yoshino etal. (2006) | Yoshiyuki (1989) | Zhang Lin et al. (2018)	https://zenodo.org/record/3750034/files/figure.png	64 . Okinawa Horseshoe Bat Rhinolophus pumilus French : Rhinolophe d'Okinawa / German: Okinawa-Hufeisennase I Spanish: Herradura de Okinawa Other common names: Okinawa Little Horseshoe Bat Taxonomy. Rhinolophus comutus pumilus K. Andersen, 1905 , “ Loo-choo Islands ,” Okinawa Prefecture , Japan . Rhinolophus pumilus is included in the pusillus species group and appears to be sister species to. comutus. Echolocation frequency of. pumilus is higher than in. cornutus , but this increase follows the generally increasing cline in the echolocation frequency within R comutus from north to south. Nevertheless, based on unpublished genetic data collected by K. Armstrong, A pumilus is tentatively treated as a species here, but an in-depth study is urgently needed; the probably extinct taxon miyakonis from Miyako Island, occasionally treated as subspecies of A pumilus , is here treated as a synonym. Monotypic. Distribution. S Japan , Okinawa Is, Iheyajima (one record in 1992), Tokashikijima (one record in 1978), Kumejima, Miyakojima (probably extinct after 1971), and Irabu Is. Descriptive notes. Head—body 36-43- 3 mm , ear 19- 1 mm (average), hindfoot 8- 6 mm (average), forearm 38-40- 5 mm . The Okinawa Horseshoe Bat is similar to the Little Japanese Horseshoe Bat (. cornutus ). Pelage is woolly and glossy and the dorsal pelage is medium brown (bases of hair being lighter) whereas the ventral pelage is similar but paler (darker than the Littlejapanese Horseshoe Bat ). Ears are moderate-sized. Noseleaf has a concave-sided lancet; connecting process is subtriangular and curved with a slightly concave or straight anterior margin; sella is pandurate, being constricted above the middle, and has a broadly rounded tip; horseshoe is not very broad but broader than in the Littlejapanese Horseshoe Bat (7- 2 mm ), with no lateral leaflets, and short, narrow median emargination. Lower lip has three mental grooves. Baculum averages 4- 2 mm long and has a very deep dorsal notch on basal cone with small ventral notch; shaft is nearly cylindrical and is bent on distal one-half. Skull is long (longer than in the Littlejapanese Horseshoe Bat ) and narrow, and zygomatic width is subequal to mastoid width; anterior median swellings are small and subcircular; posterior compartments are moderate in size; rostral profile slopes gently posteriorly or is rarely almost straight; sagittal crest is weakly to moderately developed; frontal depression varies from moderate to fairly deep; supraorbital crests are conspicuous but not generally sharp. P2 is minute and completely external to tooth row ( occasionally absent); P3 is medium-sized to small and is external to tooth row. Dental formula is the usual of 32 teeth for the genus or only 30 teeth when an upper premolar is missing. Chromosomal complement has 2n = 62 and FNa = 60. Habitat. Woodland habitats. Food and Feeding. The Okinawa Horseshoe Bat is insectivorous. Breeding. The species appears to exhibit restricted seasonal monoestry and has been observed breeding in November to January. Females give birth to a single young from May to June. Activity patterns. The Okinawa Horseshoe Bat forages during the night and roosts by day in natural caves, abandoned mines, and bomb shelters. Search call shape is FM/ CF/ FM . On northern Okinawajima, males had peak frequencies of 105-6—110-9 kHz (mean of 108-2 kHz) whereas females had 107-7—113 kHz (mean of 110-6 kHz). Peak frequencies on southern Okinawajima were even higher, with males at 112-8— 119-3 kHz (mean of 116 kHz) and females at 104-6—120-1 kHz (mean of 117-4 kHz). Females had significantly higher average frequencies than males on Okinawajima. Movements, Home range and Social organization. The Okinawa Horseshoe Bat roosts to colonies. Status and Conservation. Not assessed as a separate species on The IUCN Red List, where it is included under the Least Horseshoe Bat ( pusillus } as Least Concern. Listed as endangered in theJapanese Red List . The Okinawa Horseshoe Bat probably became extinct on Miyako c.40 years ago and on Irabu c .15 years ago, although more intensive surveys are needed to confirm this. Population on Kumejima has experienced declines. The species appears to be threatened by habitat destruction and by disturbances related to tourism in the Ryukyu Islands . Bibliography. Ando et al. (1980c), Chen Shiangfan etal. (2006), Fukui etal. (2009), Hutson, Kingston & Walston (2008), Li Gang etal. (2006), Ohdachi et al. (2015), WuYi, Motokawa et al. (2012),Yoshino etal. (2006), Yoshiyuki (1989), Zhang Lin et al. (2018).	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 pumilus	Rhinolophus		pumilus	K. Andersen	1905	0	Proc. Zool. Soc. London	1905(2): 127	Little Okinawan Horseshoe Bat	<b> miyakonis</b> Kuroda, 1924; <b> orii</b> Kuroda, 1924; <b> pumilus</b> K. Andersen, 1905.	Japan, Okinawa Prefecture ("Loo-Choo Islands")	Okinawa, Kerama, and Miyako Isls. (S Japan)	Not listed.	Least Concern under Rhinlophus pusillus cornutus 	 pusillus species group. Previously considered a subspecies of cornutus . We follow Burgin (2019) who provisionally considers pumilus distinct from cornutus based on unpublished genetic data. Note also that based on Li et al. (2006) and our decision to treat pusillus , monoceros , and cornutus as separate species, we further consider the position of pumilus in this study as potential evidence of its specific distinctiveness. Provisionally includes miyakonis (Burgin, 2019). See Corbet and Hill (1992), Csorba et al. (2003), and Wu et al. (2012) for an alternate view.	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 pumilus	23	Okinawa Horseshoe Bat	Okinawa Little Horseshoe Bat	Theria	Placentalia	Boreoeutheria	Laurasiatheria	CHIROPTERA	PTEROPODIFORMES	NA	NA	RHINOLOPHOIDEA	RHINOLOPHIDAE	NA	NA	Rhinolophus	NA	pumilus	K. Andersen	1905	0	Rhinolophus_cornutus_pumilus	Andersen, K. (1905). On Some Bats of the Genus Rhinolophus, with Remarks on Their Natural Affinities, and Descriptions of Twenty-six New Forms. Proceedings of the Zoological Society of London, 1905(2), 127.	https://www.biodiversitylibrary.org/page/31207536#page/169/mode/1up	BM 1902.10.7.18		"Loo-choo Islands," Okinawa Prefecture, Japan.			pumilus K. Andersen, 1905|miyakonis Kuroda, 1924	split from R. cornutus; includes the name miyakonis	Wu, Y., Motokawa, M., Harada, M., Thong, V.D., Lin, L.-K., Li, Y.-C. 2012. Morphometric variation in the pusillus group of teh genus Rhinolophys (Mammalia: Chiroptera: Rhinolophidae) in East Asia. Zoological Science 29: 396-402.	Japan	Asia	Palearctic	NA	0	0	0	Rhinolophus_pumilus	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		pumilus	K. Andersen	1905	0	Proc. Zool. Soc. London	1905(2): 127	Little Okinawan Horseshoe Bat	<b> miyakonis</b> Kuroda, 1924; <b> orii</b> Kuroda, 1924; <b> pumilus</b> K. Andersen, 1905.	Japan, Okinawa Prefecture ("Loo-Choo Islands")	Okinawa, Kerama, and Miyako Isls. (S Japan)	Not listed.	Least Concern under Rhinlophus pusillus cornutus 	 pusillus species group. Previously considered a subspecies of cornutus . We follow Burgin (2019) who provisionally considers pumilus distinct from cornutus based on unpublished genetic data. Note also that based on Li et al. (2006) and our decision to treat pusillus , monoceros , and cornutus as separate species, we further consider the position of pumilus in this study as potential evidence of its specific distinctiveness. Provisionally includes miyakonis (Burgin, 2019). See Corbet and Hill (1992), Csorba et al. (2003), and Wu et al. (2012) for an alternate view.	Rhinolophus pumilus	1004729	23	Okinawa Horseshoe Bat	Okinawa Little Horseshoe Bat	Theria	Placentalia	Boreoeutheria	Laurasiatheria	CHIROPTERA	PTEROPODIFORMES	NA	NA	RHINOLOPHOIDEA	Rhinolophidae	NA	NA	Rhinolophus	NA	pumilus	K. Andersen	1905	0	Rhinolophus_cornutus_pumilus	Andersen, K. (1905). On Some Bats of the Genus Rhinolophus, with Remarks on Their Natural Affinities, and Descriptions of Twenty-six New Forms. Proceedings of the Zoological Society of London, 1905(2), 127.	https://www.biodiversitylibrary.org/page/31207536#page/169/mode/1up	BM 1902.10.7.18		"Loo-choo Islands," Okinawa Prefecture, Japan.			pumilus K. Andersen, 1905|miyakonis Kuroda, 1924	split from R. cornutus; includes the name miyakonis	Wu, Y., Motokawa, M., Harada, M., Thong, V.D., Lin, L.-K., Li, Y.-C. 2012. Morphometric variation in the pusillus group of teh genus Rhinolophys (Mammalia: Chiroptera: Rhinolophidae) in East Asia. Zoological Science 29: 396-402.				Japan	Asia	Palearctic	NA	0	0	0	Rhinolophus_pumilus	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_pumilus	1004729	23	Okinawa Horseshoe Bat	Okinawa Little Horseshoe Bat	Theria	Placentalia	Boreoeutheria	Laurasiatheria	Chiroptera	Yinpterochiroptera	NA	NA	Rhinolophoidea	Rhinolophidae	NA	NA	Rhinolophus	NA	pumilus	Andersen	0	Rhinolophus cornutus pumilus	Andersen, K.C. 1905-10-07. On some bats of the genus _Rhinolophus_, with remarks on their mutual affinities, and descriptions of twenty-six new forms. Proceedings of the Zoological Society of London 1905-II(1):75-145.	https://www.biodiversitylibrary.org/page/31207588	BMNH:Mamm:1902.10.7.18	holotype	https://data.nhm.ac.uk/object/2500559f-3fad-467f-8618-94cd7164d47e	"Loo-choo Islands," Okinawa Prefecture, Japan.			split from R. cornutus; includes the name miyakonis	Wu, Y., Motokawa, M., Harada, M., Thong, V.D., Lin, L.-K., Li, Y.-C. 2012. Morphometric variation in the pusillus group of teh genus Rhinolophys (Mammalia: Chiroptera: Rhinolophidae) in East Asia. Zoological Science 29: 396-402.				Japan	Asia	Palearctic	NE	0	0	0	Rhinolophus_pumilus	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		pumilus	K. Andersen	1905	0	Proc. Zool. Soc. London	1905(2): 127	Little Okinawan Horseshoe Bat	miyakonis Kuroda, 1924; orii Kuroda, 1924; pumilus K. Andersen, 1905.	Japan, Okinawa Prefecture ("Loo-Choo Islands")	Okinawa, Kerama, and Miyako Isls. (S Japan)	<a href='https://cites.org/eng/app/appendices.php' target='_blank'>Not Listed</a>	Not Evaluated	pusillus species group. Previously considered a subspecies of cornutus. We follow Burgin (2019) who provisionally considers pumilus distinct from cornutus based on unpublished genetic data. Note also that based on Li et al. (2006) and our decision to treat pusillus, monoceros, and cornutus as separate species, we further consider the position of pumilus in this study as potential evidence of its specific distinctiveness. Provisionally includes miyakonis (Burgin, 2019). See Corbet and Hill (1992), Csorba et al. (2003), and Wu et al. (2012) for an alternate view.		Mammal Diversity Database. (2025). Mammal Diversity Database (Version 2.2) [Data set]. Zenodo. https://doi.org/10.5281/zenodo.15007505	NA	Rhinolophus pumilus; Rhinolophus pumilus; Rhinolophus pumilus; Rhinolophus pumilus; Rhinolophus pumilus; miyakonis; miyakonis; orii; pumilus; pumilus; miyakonis; Okinawa Horseshoe Bat; Rhinolophe d'Okinawa; Okinawa-Hufeisennase; Herradura de Okinawa; Spanish; Okinawa Little Horseshoe Bat; Okinawa Horseshoe Bat; Okinawa Little Horseshoe Bat; Hinde's Lesser House Bat; Little Okinawan Horseshoe Bat; R. pumilus