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line:xlsx:hash://sha256/181a039844a33e66a35a457b7ece741051086608e425a040051b79581d606b97!/Sheet1!/L105	application/vnd.openxmlformats-officedocument.spreadsheetml.sheet	Boneia bidens	Boneia bidens	Rousettus bidens	Boneia bidens	Boneia bidens	Rousettus bidens	Boneia bidens	Boneia bidens	Boneia bidens	Boneia bidens	Boneia bidens	Boneia bidens	Boneia bidens	Boneia bidens	Boneia bidens		[MSW3] Subgenus Boneia. Placed in its own genus (Boneia) by some authors (e.g., Andersen, 1912; Koopman, 1993) but see Bergmans and Rozendaal (1988) and Bergmans (1994). Corbet and Hill (1992) referred this species to the subgenus Boneia, the arrangement followed here.; [HMW] Boneia bidens Jentink, 1879 , “ Celebes [= Sulawesi , Indonesia ], Boné [= near Gerontalo].” Boneia has long been considered a subgenus of Rousettus , but recent molecular phylogenies indicate that itis a valid genus in the Harpyionycterinae clade, related to Harpyionycterss Monotypic.; [batnames2022] Previously considered a member of Rousettus ; see Bergmanns (1997).; [MDD2022] moved to Boneia from Rousettus in many recent taxonomies, summarized in the given publication; [IUCN] Simmons (2005) follows Corbet and Hill (1992) by referring this species to the subgenus Boneia . Some authors (Andersen 1912, Koopman 1993) place it in its own genus (Boneia ) but see Bergmans and Rozendaal (1988) and Bergmans (1994).; [batnames2023] Previously considered a member of Rousettus ; see Bergmanns (1997).; [MDD2023] moved to Boneia from Rousettus in many recent taxonomies, summarized in the given publication; [MDD2025_2.0] moved to Boneia from Rousettus in many recent taxonomies, summarized in the given publication; [batnames2025_1.7] Previously considered a member of Rousettus; see Bergmanns (1997).; [MDD2025_2.2] moved to Boneia from Rousettus in many recent taxonomies, summarized in the given publication						menadensis.			menadensis			bidens	bidens - menadensis	bidens, menadensis	Simmons (2005) follows Corbet and Hill (1992) by referring this species to the subgenus Boneia . Some authors (Andersen 1912, Koopman 1993) place it in its own genus (Boneia ) but see Bergmans and Rozendaal (1988) and Bergmans (1994).	bidens	bidens - menadensis	bidens, menadensis	bidens, menadensis	bidens 	bidens - menadensis	bidens Jentink, 1879|menadensis O. Thomas, 1896		Corbet, G.B. and Hill, J.E. 1980. A World List of Mammalian Species. British Museum (Natural History), London, 226 pp.		N Celebes	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.	Boneia bidens	Indonesia, N. Sulawesi, Bone (near Gerontalo).	Jentink	1879	Notes Leyden Mus., 1:117.	This species is only known from a small area, the eastern part of the northern peninsula of Celebes.		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 Sulawesi; ref. 4.110	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.	Jentink	1879	Notes Leyden Mus., 1:117.		N Sulawesi (Indonesia).	Indonesia, N Sulawesi, Bone (near Gerontalo).		JENTINCK	1879	Medium sized (forearm length, 92-102 mm).	This species is only known from a small area, the eastern part of the northern peninsula of Celebes.	No sub species.		21	species	B. bidens	JENTINCK	1879	Boneia	genus	Boneia bidens				Medium sized (forearm length, 92-102 mm).	No sub species.		1. B. bidens JENTINCK 1879.	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			Rousettus bidens	Rousettus	Boneia	bidens	Jentink	y	1879		Notes Leyden Mus.	1		117		Manado Rousette	Indonesia, N Sulawesi, Boné (near Gerontalo).	N Sulawesi (Indonesia).	IUCN/SSC Action Plan (1992) – No Data: Limited Distribution. IUCN 2003 – Lower Risk (nt).	menadensis Thomas, 1896.	Subgenus Boneia. Placed in its own genus (Boneia) by some authors (e.g., Andersen, 1912; Koopman, 1993) but see Bergmans and Rozendaal (1988) and Bergmans (1994). Corbet and Hill (1992) referred this species to the subgenus Boneia, the arrangement followed here.	03AD87FAFFF4F61A89B53602FD30F212	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	106	zip:hash://sha256/ec5fd314a06aba1a7b0b72f23e54ac625ae272bd98f82f1d01f4c09627d9e8e0!/treatments-xml-main/data/03/AD/87/03AD87FAFFF4F61A89B53602FD30F212.xml	Boneia bidens	Pteropodidae	Boneia	bidens	Jentink	1879	Roussette de Jentink @fr | Manado-Flughund @de | Boneia de Manado @es | Manado Rousette @en	Boneia bidens Jentink, 1879 , “ Celebes [= Sulawesi , Indonesia ], Boné [= near Gerontalo].” Boneia has long been considered a subgenus of Rousettus , but recent molecular phylogenies indicate that itis a valid genus in the Harpyionycterinae clade, related to Harpyionycterss Monotypic.	Sulawesi and adjacent Manado, Lembeh, and Buton Is.	Head-body 185 mm , tail 22-24 mm , ear 23-26 mm , hindfoot 30-31 mm , forearm 94-103 mm ; weight 142-194 g . The Manado Fruit Bat is Rousette-like, with long muzzle, sparse hair, and short nostrils. Eyes are moderately large, with brown irises. Ears are relatively long, with narrowly rounded-off tips; antitragus is small. Head pelage is brown, darker on crown; hairs on nape are spread, golden buffy in males with tuft ofrigid, unctuous hairs on neck side, ocherous at bases; dorsal pelage is relatively long and adpressed and reaches knees; and rump is tinged with russet-brown. Uropatagium is half furred, wide, joined to dorsal side of relatively long tail; calcar is short. Chin and throat are sparsely haired; chest, belly, and flanks are drab, lightly washed in brown. Wing membranes are grayish brown, from sides of back and inserted onto first toes; index claw is present. Skull has pronounced basicranial deflection; rostrum is relatively thin and long; premaxillae are thin and concave anteriorly, with single incisor in middle and projecting edentulous tips; forehead is gently sloping; orbit is large; zygomatic root is level with upper alveolar line; zygoma is thin anteriorly, arched, and thicker behind orbit; braincase is domed; and occiput projects slightly backward. Dorsally, rostrum is long and straight; nasals protrude slightly; paranasal recesses are inflated and reach level of small postorbital foramina; postorbital process has triangular base and is long and decurved pointing posterolaterally and slightly ventrally; there is no postorbital constriction in females and slight constriction in males; temporal lines are low and parallel; braincase is oval; and nuchal crest is inconspicuous. Ventrally, rostrum is broad anteriorly; premaxillae are separated in front, without palatine processes; palate is flat and rectangular; tooth rows are subparallel; post-dental palate is moderately short; end of palate is slightly concave; small spine is joined to thin sphenoidal crest; ectotympanic is small and wide; and entotympanic is not ossified. Mandible is straight and long, with widened symphysis; coronoid is very low and sloping; condyle is below level of lower alveolar line; and angle is weak and indistinct. There are six interdental palatal ridges; anterior four are arched, smooth, and undivided; next two are divided at midline and serrated; and two or three post-dental ridges are divided and serrated. Dental formula isl 1/2, C 1/1, P 3/3, M 2/3 (x2) = 32. I is tiny (1' absent); C' is long, slender, grooved anteriorly, and larger in males; P' is small, almost peg-like; posterior cheekteeth are generally small and very low to flat; occlusal outline is oval to rectangular; and cusps are poorly differentiated. I, is much smaller than I; C, is small but heavy, widely separated, and strongly slanted laterally; P, is moderately large, with low cusps; posterior cheekteeth are small, very low, and narrow; occlusal outline is rectangular; M, is minute, occasionally missing on one or both sides. Chromosomal complement has 2n = 30 and FN = 53 (FNa = 50), with four pairs of metacentric, seven pairs submetacentric, and three pairs telocentric autosomes and one additional unpaired autosome. X-chromosome is submetacentric, and Y-chromosome is telocentric.	Lowland to upper montane rainforests and marsh forests at elevations of 200-2100 m .	Long rostrum, widened anteriorly, broad symphysis, weak dentition, and deflected skull suggest the Manado Fruit Bat has a diet based on flower products and soft fruit.	Pregnant Manado Fruit Bats with small, single embryos were caught in June-July.	Manado Fruit Bats roost in caves.	The Manado Fruit Bat is gregarious.	Classified as Vulnerable on The IUCN Red List (as Rousettus bidens ). Populations of the Manado Fruit Bat are not fragmented but are estimated to be decreasing (30% over the next ten years) due to hunting for bushmeat in Sulawesi and changes in land use (logging and conversion to agriculture). It occurs in several protected areas such as Bogani Nani Wartabone (Dumoga Bone) National Park.	Almeida et al. (2011) | Amador et al. (2018) | Andersen (1912b) | Bergmans (1994) | Bergmans & Rozendaal (1988) | Giannini & Simmons (2007a) | Giannini, Almeida et al. (2006) | Helgen, Kingston et al. (2008) | Maryanto et al. (2011) | Mubarok et al. (2018)		78. Manado Fruit Bat Boneia bidens French: Roussette de Jentink / German: Manado-Flughund / Spanish: Boneia de Manado Other common names: Manado Rousette Taxonomy. Boneia bidens Jentink, 1879 , “ Celebes [= Sulawesi , Indonesia ], Boné [= near Gerontalo].” Boneia has long been considered a subgenus of Rousettus , but recent molecular phylogenies indicate that itis a valid genus in the Harpyionycterinae clade, related to Harpyionycterss Monotypic. Distribution. Sulawesi and adjacent Manado, Lembeh, and Buton Is. Descriptive notes. Head-body 185 mm , tail 22-24 mm , ear 23-26 mm , hindfoot 30-31 mm , forearm 94-103 mm ; weight 142-194 g . The Manado Fruit Bat is Rousette-like, with long muzzle, sparse hair, and short nostrils. Eyes are moderately large, with brown irises. Ears are relatively long, with narrowly rounded-off tips; antitragus is small. Head pelage is brown, darker on crown; hairs on nape are spread, golden buffy in males with tuft ofrigid, unctuous hairs on neck side, ocherous at bases; dorsal pelage is relatively long and adpressed and reaches knees; and rump is tinged with russet-brown. Uropatagium is half furred, wide, joined to dorsal side of relatively long tail; calcar is short. Chin and throat are sparsely haired; chest, belly, and flanks are drab, lightly washed in brown. Wing membranes are grayish brown, from sides of back and inserted onto first toes; index claw is present. Skull has pronounced basicranial deflection; rostrum is relatively thin and long; premaxillae are thin and concave anteriorly, with single incisor in middle and projecting edentulous tips; forehead is gently sloping; orbit is large; zygomatic root is level with upper alveolar line; zygoma is thin anteriorly, arched, and thicker behind orbit; braincase is domed; and occiput projects slightly backward. Dorsally, rostrum is long and straight; nasals protrude slightly; paranasal recesses are inflated and reach level of small postorbital foramina; postorbital process has triangular base and is long and decurved pointing posterolaterally and slightly ventrally; there is no postorbital constriction in females and slight constriction in males; temporal lines are low and parallel; braincase is oval; and nuchal crest is inconspicuous. Ventrally, rostrum is broad anteriorly; premaxillae are separated in front, without palatine processes; palate is flat and rectangular; tooth rows are subparallel; post-dental palate is moderately short; end of palate is slightly concave; small spine is joined to thin sphenoidal crest; ectotympanic is small and wide; and entotympanic is not ossified. Mandible is straight and long, with widened symphysis; coronoid is very low and sloping; condyle is below level of lower alveolar line; and angle is weak and indistinct. There are six interdental palatal ridges; anterior four are arched, smooth, and undivided; next two are divided at midline and serrated; and two or three post-dental ridges are divided and serrated. Dental formula isl 1/2, C 1/1, P 3/3, M 2/3 (x2) = 32. I is tiny (1' absent); C' is long, slender, grooved anteriorly, and larger in males; P' is small, almost peg-like; posterior cheekteeth are generally small and very low to flat; occlusal outline is oval to rectangular; and cusps are poorly differentiated. I, is much smaller than I; C, is small but heavy, widely separated, and strongly slanted laterally; P, is moderately large, with low cusps; posterior cheekteeth are small, very low, and narrow; occlusal outline is rectangular; M, is minute, occasionally missing on one or both sides. Chromosomal complement has 2n = 30 and FN = 53 (FNa = 50), with four pairs of metacentric, seven pairs submetacentric, and three pairs telocentric autosomes and one additional unpaired autosome. X-chromosome is submetacentric, and Y-chromosome is telocentric. Habitat. Lowland to upper montane rainforests and marsh forests at elevations of 200-2100 m . Food and Feeding. Long rostrum, widened anteriorly, broad symphysis, weak dentition, and deflected skull suggest the Manado Fruit Bat has a diet based on flower products and soft fruit. Breeding. Pregnant Manado Fruit Bats with small, single embryos were caught in June-July. Activity patterns. Manado Fruit Bats roost in caves. Movements, Home range and Social organization. The Manado Fruit Bat is gregarious. Status and Conservation. Classified as Vulnerable on The IUCN Red List (as Rousettus bidens ). Populations of the Manado Fruit Bat are not fragmented but are estimated to be decreasing (30% over the next ten years) due to hunting for bushmeat in Sulawesi and changes in land use (logging and conversion to agriculture). It occurs in several protected areas such as Bogani Nani Wartabone (Dumoga Bone) National Park. Bibliography. Almeida et al. (2011), Amador et al. (2018), Andersen (1912b), Bergmans (1994), Bergmans & Rozendaal (1988), Giannini & Simmons (2007a), Giannini, Almeida et al. (2006), Helgen, Kingston et al. (2008), Maryanto et al. (2011), Mubarok 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.	Pteropodidae	Boneia bidens	Boneia	Boneia	bidens	Jentink	1879	1	Notes Leyden Mus.	0.1229	Manado Rousette	 menadensis Thomas, 1896.	Indonesia, N Sulawesi, Boné (near Gerontalo).	N Sulawesi (Indonesia).	Not listed.	Vulnerable	Previously considered a member of Rousettus ; see Bergmanns (1997).	Mammal Diversity Database. (2023). Mammal Diversity Database (Version 1.11) [Data set]. Zenodo. https://doi.org/10.5281/zenodo.7830771 released 15 April 2023	Boneia bidens	23	Manado Fruit Bat	Manado Rousette	Theria	Placentalia	Boreoeutheria	Laurasiatheria	CHIROPTERA	PTEROPODIFORMES	NA	NA	PTEROPODOIDEA	PTEROPODIDAE	HARPYIONYCTERINAE	HARPYIONYCTERINI	Boneia	NA	bidens	Jentink	1879	0						"Celebes [= Sulawesi, Indonesia], Boné [= near Gerontalo]."			bidens Jentink, 1879|menadensis O. Thomas, 1896	moved to Boneia from Rousettus in many recent taxonomies, summarized in the given publication	Almeida, F. C., Giannini, N. P., & Simmons, N. B. (2016). The evolutionary history of the African fruit bats (Chiroptera: Pteropodidae). Acta Chiropterologica, 18(1), 73-108.	Indonesia	Asia	Australasia/Oceania	VU	0	0	0	Boneia_bidens	0	oldname match	Rousettus_bidens	0	IUCN. 2022. The IUCN Red List of Threatened Species. Version 2022-1. https://www.iucnredlist.org. Accessed on [28 September, 2022].	19749	Boneia bidens	ANIMALIA	CHORDATA	MAMMALIA	CHIROPTERA	PTEROPODIDAE	Boneia	bidens	Jentink, 1879	Simmons (2005) follows Corbet and Hill (1992) by referring this species to the subgenus Boneia . Some authors (Andersen 1912, Koopman 1993) place it in its own genus (Boneia ) but see Bergmans and Rozendaal (1988) and Bergmans (1994).	20000000	Boneia bidens	Vulnerable	A3cd	2020	2019-05-14 00:00:00 UTC	3.1	English	This species is assessed as Vulnerable under A3cd as its global population is suspected to decline by more than 30% over the next 14.9 years (3 generations; generation length of 4.9 years; Pacifici et al. 2013) due to an ongoing decline in the extent and quality of its forest habitat as well as actual levels of hunting.	This is a lowland forest bat that is also able to forage in partly cultivated habitats. It roosts in caves.	This species is threatened by hunting, including the national-level bushmeat trade, and habitat loss through logging and conversion to agriculture.	This species is rare across its range.	Decreasing	This species is found on Sulawesi and the adjacent islands of Lembeh, Buton and Manado in Indonesia (Bergmans and Rozendaal 1988). The type locality is Boné (near Gorontalo), north Sulawesi, Indonesia. The known elevation range for this species is from 200 to 1,060 m asl (Bergmans and Rozendaal 1988).		Terrestrial	This species is known to occur in the Bogani Nani Wartabone National Park, Kakenauwe Nature Reserve on the island of Buton. Its range also includes three smaller reserves along the eastern tip of north Sulawesi where it is likely to occur (Bergmans and Rozendaal 1988). Currently, this species was found in two forest reserve in South Sulawesi, in 2013 and 2016 (S. Wiantoro pers. comm).	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	Boneia	Boneia	bidens	Jentink	1879	1	Notes Leyden Mus.	0.122917	Manado Rousette	 menadensis Thomas, 1896.	Indonesia, N Sulawesi, Boné (near Gerontalo).	N Sulawesi (Indonesia).	Not listed.	Vulnerable	Previously considered a member of Rousettus ; see Bergmanns (1997).	Boneia bidens	1004404	23	Manado Fruit Bat	Manado Rousette	Theria	Placentalia	Boreoeutheria	Laurasiatheria	CHIROPTERA	PTEROPODIFORMES	NA	NA	PTEROPODOIDEA	Pteropodidae	HARPYIONYCTERINAE	HARPYIONYCTERINI	Boneia	NA	bidens	Jentink	1879	0						"Celebes [= Sulawesi, Indonesia], Boné [= near Gerontalo]."			bidens Jentink, 1879|menadensis O. Thomas, 1896	moved to Boneia from Rousettus in many recent taxonomies, summarized in the given publication	Almeida, F. C., Giannini, N. P., & Simmons, N. B. (2016). The evolutionary history of the African fruit bats (Chiroptera: Pteropodidae). Acta Chiropterologica, 18(1), 73-108.				Indonesia	Asia	Australasia/Oceania	VU	0	0	0	Boneia_bidens	0	oldname match	Rousettus_bidens	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	Boneia_bidens	1004404	23	Manado Fruit Bat	Manado Rousette	Theria	Placentalia	Boreoeutheria	Laurasiatheria	Chiroptera	Yinpterochiroptera	NA	NA	Pteropodoidea	Pteropodidae	Harpyionycterinae	Harpyionycterini	Boneia	NA	bidens	Jentink	0	Boneia bidens	Jentink, F.A. 1879. On a new genus and species of bat from Celebes. Notes from the Royal Zoological Museum of the Netherlands at Leyden 1:117-119.	https://www.biodiversitylibrary.org/page/9630996	RMNH.MAM.38015	holotype	https://data.biodiversitydata.nl/naturalis/specimen/RMNH.MAM.38015.a | https://data.biodiversitydata.nl/naturalis/specimen/RMNH.MAM.38015.b	"Celebes [= Sulawesi, Indonesia], Boné [= near Gerontalo]."			moved to Boneia from Rousettus in many recent taxonomies, summarized in the given publication	Almeida, F. C., Giannini, N. P., & Simmons, N. B. (2016). The evolutionary history of the African fruit bats (Chiroptera: Pteropodidae). Acta Chiropterologica, 18(1), 73-108.				Indonesia	Asia	Australasia	VU	0	0	0	Boneia_bidens	0	oldname match	Rousettus_bidens	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	Boneia	Boneia	bidens	Jentink	1879	1	Notes Leyden Mus.	0.122917	Manado Rousette	menadensis Thomas, 1896.	Indonesia, N Sulawesi, Boné (near Gerontalo).	N Sulawesi (Indonesia).	<a href='https://cites.org/eng/app/appendices.php' target='_blank'>Not Listed</a>	<a href='https://www.iucnredlist.org/species/19749/22002714/' target='_blank'>Vulnerable</a>	Previously considered a member of Rousettus; see Bergmanns (1997).		Mammal Diversity Database. (2025). Mammal Diversity Database (Version 2.2) [Data set]. Zenodo. https://doi.org/10.5281/zenodo.15007505	NA	Rousettus bidens; Boneia bidens; Boneia bidens; Boneia bidens; Boneia bidens; Boneia bidens; menadensis; menadensis; bidens; menadensis; Roussette de Jentink; Manado-Flughund; Boneia de Manado; Manado Rousette; Manado Fruit Bat; Manado Rousette; Manado Rousette; Manado Rousette; B. bidens