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line:xlsx:hash://sha256/181a039844a33e66a35a457b7ece741051086608e425a040051b79581d606b97!/Sheet1!/L1621	application/vnd.openxmlformats-officedocument.spreadsheetml.sheet	Thoopterus nigrescens	Thoopterus nigrescens	Thoopterus nigrescens	Thoopterus nigrescens	Thoopterus nigrescens	Thoopterus nigrescens	Thoopterus nigrescens	Thoopterus nigrescens	Thoopterus nigrescens	Thoopterus nigrescens	Thoopterus nigrescens	Thoopterus nigrescens	Thoopterus nigrescens	Thoopterus nigrescens	Thoopterus nigrescens		[MSW3] See Flannery (1995b).; [HMW] Cynopterus marginatus var. nigrescens J. E. Gray, 1870 , “Morty [= Morotai] Island ,” Molucca Islands, Indonesia . This species is monotypic.; [batnames2022] See Flannery (1995 b ).; [IUCN] A second, undescribed species of Thoopterus occurs at higher altitudes in north, central and southwest Sulawesi. A third, undescribed species occupies the island of Mangole in the Sula Islands. The Morotai record may be based on a mislabeled specimen from northern Sulawesi. According to L. Ruedas (pers. comm.) molecular data suggest a north-south species level split in Sulawesi rather than a high elevation-low elevation dichotomy as described above.; [batnames2023] See Flannery (1995 b ).; [batnames2025_1.7] See Flannery (1995b).						latidens.			latidens					nigrescens, latidens	A second, undescribed species of Thoopterus occurs at higher altitudes in north, central and southwest Sulawesi. A third, undescribed species occupies the island of Mangole in the Sula Islands. The Morotai record may be based on a mislabeled specimen from northern Sulawesi. According to L. Ruedas (pers. comm.) molecular data suggest a north-south species level split in Sulawesi rather than a high elevation-low elevation dichotomy as described above.	nigrescens 	nigrescens - tailiniensis 	nigrescens, latidens	nigrescens, latidens	nigrescens 	nigrescens - tailiniensis 	nigrescens (J. E. Gray, 1871)|latidens (Dobson, 1878)		Corbet, G.B. and Hill, J.E. 1980. A World List of Mammalian Species. British Museum (Natural History), London, 226 pp.	Swift fruit bat	N Celebes, Morotai I, N Moluccas; ? Luzon	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.	Thoopterus nigrescens	Indonesia, Molucca Isis., Morotai.	Gray	1870	Cat. Monkeys, Lemurs, and Fruit-eating Bats Br. Mus., p. 123.	Distribution: Same as for genus.		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	Swift fruit bat	N Sulawesi, Sanghir Is, Morotai I, N Moluccas; ? Luzon	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.	Gray	1870	Cat. Monkeys, Lemurs, Fruit-eating Bats Brit. Mus., p. 123.		Molucca Isis, Sulawesi and Sangihe Isis (Indonesia).	Indonesia, Molucca Isis, Morotai.		GRAY	1870	Size relatively large (forearm length, 73-7 9 mm).	Distribution: Same as for genus.	No subspecies.		34	species	T. nigrescens	GRAY	1870	Thoopterus	genus	Thoopterus nigrescens				Size relatively large (forearm length, 73-7 9 mm).	No subspecies.		1. T.nigrescens (GRAY 1870).	1	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	Pteropodidae			Thoopterus nigrescens	Thoopterus		nigrescens	Gray	y	1870		Cat. Monkeys, Lemurs, Fruit-eating Bats Brit. Mus.			123		Swift Fruit Bat	Indonesia, Molucca Isls, Morotai.	Sulawesi, Sula Isls, Sangihe Isls, Karakelang (Talaud Isls), and Morotai (Indonesia).	IUCN/SSC Action Plan (1992) – No Data. IUCN 2003 – Lower Risk (nt).	latidens Dobson, 1878.	See Flannery (1995b).	03AD87FAFFD5F63B8996310CF87CFE41	Handbook of the Mammals of the World – Volume 9 Bats, Barcelona: Lynx Edicions	978-84-16728-19-0	hbmw_9_Pteropodidae_16.pdf.imf	hash://md5/ff94ff82ffc4f62a891e341cffa5ff9b	74	zip:hash://sha256/ec5fd314a06aba1a7b0b72f23e54ac625ae272bd98f82f1d01f4c09627d9e8e0!/treatments-xml-main/data/03/AD/87/03AD87FAFFD4F63B8CB5330DFBE9FB5E.xml	Thoopterus nigrescens	Pteropodidae	Thoopterus	nigrescens		1871	Cynoptere sombre @fr | Schwarzflligelflughund @de | Toptérido cenizo @es | Swift Fruit Bat @en	Cynopterus marginatus var. nigrescens J. E. Gray, 1870 , “Morty [= Morotai] Island ,” Molucca Islands, Indonesia . This species is monotypic.	Talaud Is (Karakelong), Sangihe Is (Sangihe), N Moluccas (Morotai), Sulawesi (including nearby Wowoni and Buton Is), and Sula Is (Taliabu, Mangole and Sanana).	Head-body 94- 111 mm , tail 6 mm (single individual), ear 13-17 mm , hindfoot 18-20 mm , forearm 70-82 mm ; weight 67-99 g . Head of the Common Swift Fruit Bat has relatively long muzzle, nostrils are short and barely divergent, philtrum reaches upper lip, and lowerlip has two large and flat triangular pads. Eyes are moderately large; iris is warm brown to chocolate-brown. Ears are short and rounded. Head pelage is darker than body, moderately long, dense, and dark grayish brown, with short guard hairs on nape; dorsal pelage is long (¢. 9 mm ), soft, silky, and dark grayish brown, somewhat irregularly washed in light brown especially on dorsal forearm. Uropatagium is narrow at center, with dorsal hairs extending over tibia; underside of uropatagium is naked; tail is minute, infrequently absent; and calcar is short ( 7-8 mm ). Chest and belly fur is dark gray, occasionally silvery, and dense. Wing membranes are black from sides of body and attach to second toe; index claw is present. Skull lacks basicranial deflection, rostrum is moderately long, orbit is large with marked rim, zygomatic root is from above alveolar line, zygoma is moderately thick and strongly arched posteriorly, and braincase is domed. Dorsally, rostrum is long and slightly tapering; paranasal recesses are inflated, passing short postorbital processes posteriorly; there is no postorbital foramen; postorbital constriction is obvious; braincase is oval; sagittal crest is present but low; and nuchal crest is obvious. Ventrally, palate is flat, with diverging tooth rows; post-dental is moderately short, and sides are slightly convergent; end of post-dental palate is straight; and ectotympanic is small and wide anteriorly, narrowly edged internally by entotympanic. Mandible is moderately thick; coronoid is tall, almost vertical with wide tip; condyle is slightly above lower alveolar line; and angle is distinct and rounded. There are nine arched interdental palatal ridges, and posterior one-half is divided at middle, followed by three post-dental divided ridges, the last one is denticulate, plus a last, ill-defined ridge, denticulate, near end of post-dental palate. Dental formula for all species of Thoopterusis12/2,C1/1,P 3/3, M 1/2 (x2) = 30. Upper incisors are conical, with I longer than I*; C' is long and decurved, with anterointernal groove, longer and bulkier in males; P' is small; and posterior cheekteeth are large, decreasing in size, with squarish outline, and lingual cusps reduced. Lower incisors are small and crowded; C, 1s rather small and straight, with strong cingulum and cusp-like lobule; P, is well developed, with triangular crown; posterior cheekteeth are tall, very wide, and squarish, with reduced lingual cusp and strong surface cusp, and M, very low and peg-like. Diploid number is 2n = 38.	[Lowland and montane primary rainforests and lightly disturbed forests from sea level up to elevations of 2400 m .	The Common Swift Fruit Bat is primarily frugivorous. There are miscellaneous records of use offig trees ( Ficus spp. , Moraceae ). Strong dentition suggests consumption of hard fruits.	Young Common Swift Fruit Bats are present in an area year-round; two pregnant females with single embryos were recorded in January; and females with large nipples were found in January and March.	No information.	Infestation with parasites on Common Swift Fruit Bats suggests communal roosting in permanent locations, perhaps cavities or caves.	Classified as Least Concern on The IUCN Red List. The Common Swift Fruit Bat is common, even dominant, and able to adapt to disturbed forest habitats. Population trend is unknown, but most of the population not fragmented.	Andersen (1912b) | Bergmans & Rozendaal (1988) | Maryanto (2004) | Maryanto et al. (2011) | Mubarok et al. (2018) | Ruedas, Kingston, Helgen & Sinaga (2008) | Wiantoro et al. (2017)		23. Common Swift Fruit Bat Thoopterus nigrescens French: Cynoptere sombre / German: Schwarzflligelflughund / Spanish: Toptérido cenizo Other common names: Swift Fruit Bat Taxonomy. Cynopterus marginatus var. nigrescens J. E. Gray, 1870 , “Morty [= Morotai] Island ,” Molucca Islands, Indonesia . This species is monotypic. Distribution. Talaud Is (Karakelong), Sangihe Is (Sangihe), N Moluccas (Morotai), Sulawesi (including nearby Wowoni and Buton Is), and Sula Is (Taliabu, Mangole and Sanana). Descriptive notes. Head-body 94- 111 mm , tail 6 mm (single individual), ear 13-17 mm , hindfoot 18-20 mm , forearm 70-82 mm ; weight 67-99 g . Head of the Common Swift Fruit Bat has relatively long muzzle, nostrils are short and barely divergent, philtrum reaches upper lip, and lowerlip has two large and flat triangular pads. Eyes are moderately large; iris is warm brown to chocolate-brown. Ears are short and rounded. Head pelage is darker than body, moderately long, dense, and dark grayish brown, with short guard hairs on nape; dorsal pelage is long (¢. 9 mm ), soft, silky, and dark grayish brown, somewhat irregularly washed in light brown especially on dorsal forearm. Uropatagium is narrow at center, with dorsal hairs extending over tibia; underside of uropatagium is naked; tail is minute, infrequently absent; and calcar is short ( 7-8 mm ). Chest and belly fur is dark gray, occasionally silvery, and dense. Wing membranes are black from sides of body and attach to second toe; index claw is present. Skull lacks basicranial deflection, rostrum is moderately long, orbit is large with marked rim, zygomatic root is from above alveolar line, zygoma is moderately thick and strongly arched posteriorly, and braincase is domed. Dorsally, rostrum is long and slightly tapering; paranasal recesses are inflated, passing short postorbital processes posteriorly; there is no postorbital foramen; postorbital constriction is obvious; braincase is oval; sagittal crest is present but low; and nuchal crest is obvious. Ventrally, palate is flat, with diverging tooth rows; post-dental is moderately short, and sides are slightly convergent; end of post-dental palate is straight; and ectotympanic is small and wide anteriorly, narrowly edged internally by entotympanic. Mandible is moderately thick; coronoid is tall, almost vertical with wide tip; condyle is slightly above lower alveolar line; and angle is distinct and rounded. There are nine arched interdental palatal ridges, and posterior one-half is divided at middle, followed by three post-dental divided ridges, the last one is denticulate, plus a last, ill-defined ridge, denticulate, near end of post-dental palate. Dental formula for all species of Thoopterusis12/2,C1/1,P 3/3, M 1/2 (x2) = 30. Upper incisors are conical, with I longer than I*; C' is long and decurved, with anterointernal groove, longer and bulkier in males; P' is small; and posterior cheekteeth are large, decreasing in size, with squarish outline, and lingual cusps reduced. Lower incisors are small and crowded; C, 1s rather small and straight, with strong cingulum and cusp-like lobule; P, is well developed, with triangular crown; posterior cheekteeth are tall, very wide, and squarish, with reduced lingual cusp and strong surface cusp, and M, very low and peg-like. Diploid number is 2n = 38. Habitat. [Lowland and montane primary rainforests and lightly disturbed forests from sea level up to elevations of 2400 m . Food and Feeding. The Common Swift Fruit Bat is primarily frugivorous. There are miscellaneous records of use offig trees ( Ficus spp. , Moraceae ). Strong dentition suggests consumption of hard fruits. Breeding. Young Common Swift Fruit Bats are present in an area year-round; two pregnant females with single embryos were recorded in January; and females with large nipples were found in January and March. Activity patterns. No information. Movements, Home range and Social organization. Infestation with parasites on Common Swift Fruit Bats suggests communal roosting in permanent locations, perhaps cavities or caves. Status and Conservation. Classified as Least Concern on The IUCN Red List. The Common Swift Fruit Bat is common, even dominant, and able to adapt to disturbed forest habitats. Population trend is unknown, but most of the population not fragmented. Bibliography. Andersen (1912b), Bergmans & Rozendaal (1988), Maryanto (2004), Maryanto et al. (2011), Mubarok et al. (2018), Ruedas, Kingston, Helgen & Sinaga (2008), Wiantoro et al. (2017).	Simmons, N.B. and A.L. Cirranello. 2022B. Bat Species of the World: A taxonomic and geographic database. Accessed on 10/11/2022.	Pteropodidae	Thoopterus nigrescens	Thoopterus		nigrescens	Gray	1870	1	Cat. Monkeys, Lemurs, Fruit-eating Bats Brit. Mus.	p. 123	Swift Fruit Bat	Yes.	Indonesia, Molucca Isls, Morotai.	Sulawesi, Sula Isls, Sangihe Isls, Karakelang (Talaud Isls), and Morotai (Indonesia).	Not listed.	Least Concern	See Flannery (1995 b ).	Mammal Diversity Database. (2023). Mammal Diversity Database (Version 1.11) [Data set]. Zenodo. https://doi.org/10.5281/zenodo.7830771 released 15 April 2023	Thoopterus nigrescens	23	Common Swift Fruit Bat	Swift Fruit Bat	Theria	Placentalia	Boreoeutheria	Laurasiatheria	CHIROPTERA	PTEROPODIFORMES	NA	NA	PTEROPODOIDEA	PTEROPODIDAE	CYNOPTERINAE	BALIONYCTERINI	Thoopterus	NA	nigrescens	J. E. Gray	1870	1						"Morty [= Morotai] Island," Molucca Islands, Indonesia.			nigrescens (J. E. Gray, 1870)|latidens (Dobson, 1878)	NA	NA	Indonesia	Asia|Oceania	Australasia/Oceania	LC	0	0	0	Thoopterus_nigrescens	0	sciname match	Thoopterus_nigrescens	0	IUCN. 2022. The IUCN Red List of Threatened Species. Version 2022-1. https://www.iucnredlist.org. Accessed on [28 September, 2022].	21815	Thoopterus nigrescens	ANIMALIA	CHORDATA	MAMMALIA	CHIROPTERA	PTEROPODIDAE	Thoopterus	nigrescens	(Gray, 1870)	A second, undescribed species of Thoopterus occurs at higher altitudes in north, central and southwest Sulawesi. A third, undescribed species occupies the island of Mangole in the Sula Islands. The Morotai record may be based on a mislabeled specimen from northern Sulawesi. According to L. Ruedas (pers. comm.) molecular data suggest a north-south species level split in Sulawesi rather than a high elevation-low elevation dichotomy as described above.	20000000	Thoopterus nigrescens	Least Concern		2020	2019-05-14 00:00:00 UTC	3.1	English	Listed as Least Concern because its extent of occurrence is greater than 20,000 km², it is very common, and its populations are not declining fast enough for listing in a more threatened category.	It prefers intact forest but has been collected from lowland to 2,400 m asl in many different habitats including disturbed habitats.	This species is hunted for bushmeat, but not currently at levels that are considered a major threat to the species.	This species is common and widespread in suitable habitat.	Unknown	This species occurs throughout Sulawesi and on the adjacent islands of Buton, Mangole, the Talaud and Sangihe island groups, and probably Morotai, all in Indonesia. Specimens have also been collected from Wowoni (U. Sinaga pers. comm.).		Terrestrial	This species occurs in several protected areas throughout its range.	Australasian		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 	Pteropodidae	Thoopterus		nigrescens	Gray	1870	1	Cat. Monkeys, Lemurs, Fruit-eating Bats Brit. Mus.	p. 123	Swift Fruit Bat	Yes.	Indonesia, Molucca Isls, Morotai.	Sulawesi, Sula Isls, Sangihe Isls, Karakelang (Talaud Isls), and Morotai (Indonesia).	Not listed.	Least Concern	See Flannery (1995 b ).	Thoopterus nigrescens	1004372	23	Common Swift Fruit Bat	Swift Fruit Bat	Theria	Placentalia	Boreoeutheria	Laurasiatheria	CHIROPTERA	PTEROPODIFORMES	NA	NA	PTEROPODOIDEA	Pteropodidae	CYNOPTERINAE	BALIONYCTERINI	Thoopterus	NA	nigrescens	J. E. Gray	1870	1						"Morty [= Morotai] Island," Molucca Islands, Indonesia.			nigrescens (J. E. Gray, 1870)|latidens (Dobson, 1878)	NA	NA				Indonesia	Asia|Oceania	Australasia/Oceania	LC	0	0	0	Thoopterus_nigrescens	0	sciname match	Thoopterus_nigrescens	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	Thoopterus_nigrescens	1004372	23	Common Swift Fruit Bat	Swift Fruit Bat	Theria	Placentalia	Boreoeutheria	Laurasiatheria	Chiroptera	Yinpterochiroptera	NA	NA	Pteropodoidea	Pteropodidae	Cynopterinae	Balionycterini	Thoopterus	NA	nigrescens	J. E. Gray	1	Cynopterus marginatus var. nigrescens	Gray, J.E. 1871-01-14. Catalogue of monkeys, lemurs, and fruit-eating bats in the collection of the British Museum. British Museum, London, 137 pp.	https://www.biodiversitylibrary.org/page/9994640	BMNH:Mamm:1862.10.21.6	holotype		"Morty [= Morotai] Island," Molucca Islands, Indonesia.			NA	NA				Indonesia	Asia	Australasia	LC	0	0	0	Thoopterus_nigrescens	0	sciname match	Thoopterus_nigrescens	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	Pteropodidae	Thoopterus		nigrescens	Gray	1870	1	Cat. Monkeys, Lemurs, Fruit-eating Bats Brit. Mus.	p. 123	Swift Fruit Bat	Yes.	Indonesia, Molucca Isls, Morotai.	Sulawesi, Sula Isls, Sangihe Isls, Karakelang (Talaud Isls), and Morotai (Indonesia).	<a href='https://cites.org/eng/app/appendices.php' target='_blank'>Not Listed</a>	<a href='https://www.iucnredlist.org/species/21815/21989441/' target='_blank'>Least Concern</a>	See Flannery (1995b).		Mammal Diversity Database. (2025). Mammal Diversity Database (Version 2.2) [Data set]. Zenodo. https://doi.org/10.5281/zenodo.15007505	NA	Thoopterus nigrescens; Thoopterus nigrescens; Thoopterus nigrescens; Thoopterus nigrescens; Thoopterus nigrescens; Thoopterus nigrescens; latidens; nigrescens; latidens; Cynoptere sombre; Schwarzflligelflughund; Toptérido cenizo; Swift Fruit Bat; Common Swift Fruit Bat; Swift Fruit Bat; Swift Fruit Bat; Swift Fruit Bat; T. nigrescens