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line:xlsx:hash://sha256/181a039844a33e66a35a457b7ece741051086608e425a040051b79581d606b97!/Sheet1!/L1064	application/vnd.openxmlformats-officedocument.spreadsheetml.sheet	N/A	N/A	N/A	N/A	N/A	Nyctimene keasti	Nyctimene keasti	Nyctimene keasti	Nyctimene keasti	Nyctimene keasti	Nyctimene keasti	Nyctimene keasti	Nyctimene keasti	Nyctimene keasti	Nyctimene keasti		[MSW3] cephalotes species group. Originally described as a subspecies of albiventer, but recognized as a distinct species by Kitchener et al. (1995c) and Bergmans (2001). Revised by Bergmans (2001).; [HMW] Nyctimene albiventer keasti Kitchener in Kitchener et al., 1993, “Near Taman Anggrek, 12 km N.Tual, Pulau Dullah, closely associated with Pulan Kai Reeil, ( 5°38’S, 132°44'E ) ... at sea level,” Moluccas , Indonesia . Nyctimene keasti is currently in the cephalotes species group. It was originally described as a subspecies of N. albiventer but subsequently elevated to a species and is considered more closely related to N. cephalotes on morphological grounds. Limited genetic data showed that N. keasti from the Kai Islands, Bandaneira Island, and Yamdena Island were not genetically distinct from N. cephalotes , indicating that N. keasti might be best recognized as a subspecies or synonym of N. cephalotes with which it is morphologically very similar. Nevertheless, ongoing revision of Nyctimene is currently underway, and more thorough sampling is needed before any conclusion can be made regarding these species. Three subspecies recognized.; [batnames2022]  cephalotes species group. Originally described as a subspecies of albiventer, but recognized as a distinct species by Kitchener et al. (1995c)and Bergmans (2001). Revised by Bergmans (2001).; [IUCN] This species was originally described as a subspecies of Nyctimene albiventer , but has been elevated to species status by Kitchener et al . (1995) and Bergmans (2001).; [batnames2023]  cephalotes species group. Originally described as a subspecies of albiventer, but recognized as a distinct species by Kitchener et al. (1995c)and Bergmans (2001). Revised by Bergmans (2001).; [batnames2025_1.7] cephalotesspecies group. Originally described as a subspecies of albiventer, but recognized as a distinct species by Kitchener et al. (1995c)and Bergmans (2001). Revised by Bergmans (2001).								keasti, babari, tozeri		keasti, babari, tozeri	keasti?	keasti, babari, tozeri		keasti, tozeri, babari	This species was originally described as a subspecies of Nyctimene albiventer , but has been elevated to species status by Kitchener et al . (1995) and Bergmans (2001).	keasti, babari, tozeri		keasti, tozeri, babari 	keasti, tozeri, babari 	babari, tozeri, tozeri 		keasti D. J. Kitchener in D. J. Kitchener, Packer, & Maryanto, 1993|tozeri D. J. Kitchener in D. J. Kitchener, Packer, & Suyanto, 1995|babari Bergmans, 2001						N/A																																								_N. k. babari_ Bergmans, 2001; _N. k. keasti_ Kitchener, 1993; _N. k. tozeri_ Kitchener, 1995			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			Nyctimene keasti	Nyctimene		keasti	Kitchener in Kitchener, Packer, and Maryanto		1993		Rec. West. Aust. Mus.	16		408		Keast's Tube-nosed Fruit Bat	Indonesia, Maluku, Pulau Dullah (closely associated with Pulau Kai Kecil), 12 km N Tual, near Taman Anggrek, 5°38'S, 132°44'E, sea level.	Babar, Tanimbar, and Kai Isls (Molucca Isls, Indonesia); probably Timor and Flores (K. Helgen, pers. comm.).	Described after completion of IUCN/SSC Action Plan (1992); IUCN 2003 – not evaluated.	babari Bergmans, 2001; tozeri Kitchener, 1995 [in Kitchener et al., 1995c].	cephalotes species group. Originally described as a subspecies of albiventer, but recognized as a distinct species by Kitchener et al. (1995c) and Bergmans (2001). Revised by Bergmans (2001).	03AD87FAFF84F66A89B23516FD6EF53B	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	122	zip:hash://sha256/ec5fd314a06aba1a7b0b72f23e54ac625ae272bd98f82f1d01f4c09627d9e8e0!/treatments-xml-main/data/03/AD/87/03AD87FAFF84F66A89B23516FD6EF53B.xml	Nyctimene keasti	Pteropodidae	Nyctimene	keasti		1993	Nyctimene de Keast @fr | Keast-Rohrennasenflughund @de | Nyctimeno de Keast @es	Nyctimene albiventer keasti Kitchener in Kitchener et al., 1993, “Near Taman Anggrek, 12 km N.Tual, Pulau Dullah, closely associated with Pulan Kai Reeil, ( 5°38’S, 132°44'E ) ... at sea level,” Moluccas , Indonesia . Nyctimene keasti is currently in the cephalotes species group. It was originally described as a subspecies of N. albiventer but subsequently elevated to a species and is considered more closely related to N. cephalotes on morphological grounds. Limited genetic data showed that N. keasti from the Kai Islands, Bandaneira Island, and Yamdena Island were not genetically distinct from N. cephalotes , indicating that N. keasti might be best recognized as a subspecies or synonym of N. cephalotes with which it is morphologically very similar. Nevertheless, ongoing revision of Nyctimene is currently underway, and more thorough sampling is needed before any conclusion can be made regarding these species. Three subspecies recognized.	N.k.keastiKitchener,1993—KaiIs(Kur,KaiBesar,KaiKecil,andDullah). N.k.babariBergmans,2001—ELesserSundaIs(Babar). N. k. tozeri Kitchener, 1995 — Tanimbar Is (Larat, Yamdena, and Salaru). The species also probably exists on Flores and Timor Is, although this has yet to be confirmed and there is also apparently a record from Gag I (Raja Ampat Is).	Head-body 69-3-89- 3 mm , tail 18-25- 4 mm , ear 14-2-16- 9 mm , hindfoot 11:3-13- 9 mm , forearm 40-7-45- 7 mm ; weight 29- 5-30 g . Females generally have considerably paler dorsum than males. Head of Keast’s Tube-nosed Fruit Bat is broad, with deep face, broad, bluntly pointed ears, and tubular divergent nostrils. Eyes are large, with amber irises. Pelage is thick and woolly. Dorsal pelage is yellowish brown (males) to buffy yellowish brown (females), with distinctive thin dark blackish brown gradually widening mid-dorsal stripe (0-5- 1-5 mm thick at top and 2-5- 3-5 mm at lower end) stretching from neck to rump. Ventral pelage is buffy yellowish or yellowish brown (in some males). Wings are dark brown, with irregular darker and yellow spots throughout (more yellow spotting is on digits). Ears are light yellowish brown, with some yellow spotting. Second digit of wing has a claw, and wing attaches at second digit of foot. Tail is short, black, and wrinkled, and narrow uropatagium connects at base and stretches to calcar at ankles. Claws are brown. Skull and mandible are robust; rostrum is short; zygomatic breadth is wide; sagittal and lambdoidal crests are moderately developed; posterior palate extends well beyond M' and margin varies from U-shaped to sharply rectangular; and condylo-basal length is generally shorter relative to both rostrum height, interorbital breadth, and cranial height compared with Pallas’s Tube-nosed Fruit Bat (N. cephalotes ). Single lower incisor is completely deciduous, falling out before adulthood; lower molars are broad and rounded in dorsal view; C, replaces incisors and is long and powerful; P, is elongated and longer than P, and P,; and C' has moderately developed secondary labial cusp.	Primary tropical moist forests, dominated by Ficus ( Moraceae ), Erythrina ( Fabaceae ), Podocarpus ( Podocarpaceae ), and Albizia ( Fabaceae ), and vine forest from sea level up to elevations ¢. 1000 m .	No information.	No information.	No information.	No information.	Classified as Vulnerable on The IUCN Red List. Keast’s Tubenosed Fruit Bat has a restricted distribution and is threatened by deforestation. Additional research is needed to validate its taxonomic status and to understand its ecology and threats.	Bergmans (2001) | Helgen & Hutson (2008) | Kitchener, Packer & Maryanto (1993) | Kitchener, Packer & Suyanto (1995) | Newbound et al. (2008)		112. Keast’s Tube-nosed Fruit Bat Nyctimene keasti French: Nyctimene de Keast / German: Keast-Rohrennasenflughund / Spanish: Nyctimeno de Keast Taxonomy. Nyctimene albiventer keasti Kitchener in Kitchener et al., 1993, “Near Taman Anggrek, 12 km N.Tual, Pulau Dullah, closely associated with Pulan Kai Reeil, ( 5°38’S, 132°44'E ) ... at sea level,” Moluccas , Indonesia . Nyctimene keasti is currently in the cephalotes species group. It was originally described as a subspecies of N. albiventer but subsequently elevated to a species and is considered more closely related to N. cephalotes on morphological grounds. Limited genetic data showed that N. keasti from the Kai Islands, Bandaneira Island, and Yamdena Island were not genetically distinct from N. cephalotes , indicating that N. keasti might be best recognized as a subspecies or synonym of N. cephalotes with which it is morphologically very similar. Nevertheless, ongoing revision of Nyctimene is currently underway, and more thorough sampling is needed before any conclusion can be made regarding these species. Three subspecies recognized. Subspecies and Distribution. N.k.keastiKitchener,1993—KaiIs(Kur,KaiBesar,KaiKecil,andDullah). N.k.babariBergmans,2001—ELesserSundaIs(Babar). N. k. tozeri Kitchener, 1995 — Tanimbar Is (Larat, Yamdena, and Salaru). The species also probably exists on Flores and Timor Is, although this has yet to be confirmed and there is also apparently a record from Gag I (Raja Ampat Is). Descriptive notes. Head-body 69-3-89- 3 mm , tail 18-25- 4 mm , ear 14-2-16- 9 mm , hindfoot 11:3-13- 9 mm , forearm 40-7-45- 7 mm ; weight 29- 5-30 g . Females generally have considerably paler dorsum than males. Head of Keast’s Tube-nosed Fruit Bat is broad, with deep face, broad, bluntly pointed ears, and tubular divergent nostrils. Eyes are large, with amber irises. Pelage is thick and woolly. Dorsal pelage is yellowish brown (males) to buffy yellowish brown (females), with distinctive thin dark blackish brown gradually widening mid-dorsal stripe (0-5- 1-5 mm thick at top and 2-5- 3-5 mm at lower end) stretching from neck to rump. Ventral pelage is buffy yellowish or yellowish brown (in some males). Wings are dark brown, with irregular darker and yellow spots throughout (more yellow spotting is on digits). Ears are light yellowish brown, with some yellow spotting. Second digit of wing has a claw, and wing attaches at second digit of foot. Tail is short, black, and wrinkled, and narrow uropatagium connects at base and stretches to calcar at ankles. Claws are brown. Skull and mandible are robust; rostrum is short; zygomatic breadth is wide; sagittal and lambdoidal crests are moderately developed; posterior palate extends well beyond M' and margin varies from U-shaped to sharply rectangular; and condylo-basal length is generally shorter relative to both rostrum height, interorbital breadth, and cranial height compared with Pallas’s Tube-nosed Fruit Bat (N. cephalotes ). Single lower incisor is completely deciduous, falling out before adulthood; lower molars are broad and rounded in dorsal view; C, replaces incisors and is long and powerful; P, is elongated and longer than P, and P,; and C' has moderately developed secondary labial cusp. Habitat. Primary tropical moist forests, dominated by Ficus ( Moraceae ), Erythrina ( Fabaceae ), Podocarpus ( Podocarpaceae ), and Albizia ( Fabaceae ), and vine forest from sea level up to elevations ¢. 1000 m . Food and Feeding. No information. Breeding. No information. Activity patterns. No information. Movements, Home range and Social organization. No information. Status and Conservation. Classified as Vulnerable on The IUCN Red List. Keast’s Tubenosed Fruit Bat has a restricted distribution and is threatened by deforestation. Additional research is needed to validate its taxonomic status and to understand its ecology and threats. Bibliography. Bergmans (2001), Helgen & Hutson (2008), Kitchener, Packer & Maryanto (1993), Kitchener, Packer & Suyanto (1995), Newbound et al. (2008).	Simmons, N.B. and A.L. Cirranello. 2022B. Bat Species of the World: A taxonomic and geographic database. Accessed on 10/11/2022.	Pteropodidae	Nyctimene keasti	Nyctimene		keasti	Kitchener in Kitchener, Packer, and Maryanto	1993	0	Rec. West. Aust. Mus.	0.95	Keast's Tube-nosed Fruit Bat	<b> babari </b>Bergmans, 2001; <b> tozeri </b>Kitchener, 1995 [in Kitchener et al., 1995].	Indonesia, Maluku, Pulau Dullah (closely associated with Pulau Kai Kecil), 12 km N Tual, near Taman Anggrek, 5°38'S, 132°44'E, sea level.	Babar, Tanimbar, and Kai Isls (Molucca Isls, Indonesia); probably Timor and Flores (K. Helgen, pers. comm.).	Not listed.	Near Threatened	 cephalotes species group. Originally described as a subspecies of albiventer, but recognized as a distinct species by Kitchener et al. (1995c)and Bergmans (2001). Revised by Bergmans (2001).	Mammal Diversity Database. (2023). Mammal Diversity Database (Version 1.11) [Data set]. Zenodo. https://doi.org/10.5281/zenodo.7830771 released 15 April 2023	Nyctimene keasti	23	Keast's Tube-nosed Fruit Bat		Theria	Placentalia	Boreoeutheria	Laurasiatheria	CHIROPTERA	PTEROPODIFORMES	NA	NA	PTEROPODOIDEA	PTEROPODIDAE	NYCTIMENINAE	NA	Nyctimene	NA	keasti	Kitchener in Kitchener, Packer, & Maryanto	1993	0						"Near Taman Anggrek, 12 km N.Tual, Pulau Dullah, closely associated with Pulan Kai Kecil, (5°38'S, 132°44'E) … at sea level," Moluccas, Indonesia.	-5.633	132.73	keasti Kitchener in Kitchener, Packer, & Maryanto, 1993|tozeri Kitchener in Kitchener, Packer, & Suyanto, 1995|babari Bergmans, 2001	NA	NA	Indonesia|East Timor?	Asia|Oceania	Indomalaya|Australasia/Oceania	NT	0	0	0	Nyctimene_keasti	0	sciname match	Nyctimene_keasti	0	IUCN. 2022. The IUCN Red List of Threatened Species. Version 2022-1. https://www.iucnredlist.org. Accessed on [28 September, 2022].	136441	Nyctimene keasti	ANIMALIA	CHORDATA	MAMMALIA	CHIROPTERA	PTEROPODIDAE	Nyctimene	keasti	Kitchener in Kitchener, Packer &; Maryanto, 1993	This species was originally described as a subspecies of Nyctimene albiventer , but has been elevated to species status by Kitchener et al . (1995) and Bergmans (2001).	20000000	Nyctimene keasti	Near Threatened	A2c	2020	2020-03-01 00:00:00 UTC	3.1	English	Nyctimene keasti is assessed as Near Threatened under criterion A2c as the species ;range is severely fragmented (3 islands) and it is ;suspected to have experienced a decline of 25-30% over the past 12.9 years (3 generations; GL = 4.3 years; Pacifici et al. 2013) ;due to extensive native forest loss due to conversion to from agriculture and increased mining. The threats have not stopped and are projected to continue.	It has been caught by mist-netting in disturbed vine forest, with an understorey of lantana (Ficus, Erythrina, Podocoarpus, Albizia ). It has also been found in mixed gardens (I. Maryanto pers. comm.).	There has been and continues to be extensive loss of native forest habitat due to conversion to agriculture and mining; the loss of habitat is continuing. This species is not hunted.	Historically, this species was relatively common on the islands where it occurs. The species global populations is suspected to be decreasing by 25-29% over the past 12.9 years (3 generations; GL = 4.3 years; Pacifici et al. 2013) due to extensive forest loss and degradation due to conversion of native forest to agriculture and mining.	Decreasing	This species has been recorded from Maluku, the Kai Islands, Tanimbar (Timorlaut), and Babar island groups in Indonesia (Simmons 2005), and is probably also on Timor and Flores (K. Helgen pers. comm.). The type locality for this species is 12 km north of Tual, near Taman Anggrek (5°38'S, 132°44'E), Pulau Dullah (closely associated with Pulau Kai Kecil), Maluku, Indonesia. There is a record of the species also from Gag Island, Indonesia (I. Maryanto pers. comm.). Its elevation range is thought to be below 1,000 m asl.	The species is not hunted.	Terrestrial	Pulau Nustaram Nature Reserve is within the range of this species. This species needs further taxonomic studies as it may be conspecific with N. albifenta which is more widespread (M. Sinaga 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	Nyctimene		keasti	Kitchener in Kitchener, Packer, and Maryanto	1993	0	Rec. West. Aust. Mus.	0.95	Keast's Tube-nosed Fruit Bat	<b> babari </b>Bergmans, 2001; <b> tozeri </b>Kitchener, 1995 [in Kitchener et al., 1995].	Indonesia, Maluku, Pulau Dullah (closely associated with Pulau Kai Kecil), 12 km N Tual, near Taman Anggrek, 5°38'S, 132°44'E, sea level.	Babar, Tanimbar, and Kai Isls (Molucca Isls, Indonesia); probably Timor and Flores (K. Helgen, pers. comm.).	Not listed.	Near Threatened	 cephalotes species group. Originally described as a subspecies of albiventer, but recognized as a distinct species by Kitchener et al. (1995c)and Bergmans (2001). Revised by Bergmans (2001).	Nyctimene keasti	1004420	23	Keast's Tube-nosed Fruit Bat		Theria	Placentalia	Boreoeutheria	Laurasiatheria	CHIROPTERA	PTEROPODIFORMES	NA	NA	PTEROPODOIDEA	Pteropodidae	NYCTIMENINAE	NA	Nyctimene	NA	keasti	Kitchener in Kitchener, Packer, & Maryanto	1993	0						"Near Taman Anggrek, 12 km N.Tual, Pulau Dullah, closely associated with Pulan Kai Kecil, (5°38'S, 132°44'E) … at sea level," Moluccas, Indonesia.	-5.63333	132.7333	keasti Kitchener in Kitchener, Packer, & Maryanto, 1993|tozeri Kitchener in Kitchener, Packer, & Suyanto, 1995|babari Bergmans, 2001	NA	NA				Indonesia|East Timor?	Asia|Oceania	Indomalaya|Australasia/Oceania	NT	0	0	0	Nyctimene_keasti	0	sciname match	Nyctimene_keasti	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	Nyctimene_keasti	1004420	23	Keast's Tube-nosed Fruit Bat		Theria	Placentalia	Boreoeutheria	Laurasiatheria	Chiroptera	Yinpterochiroptera	NA	NA	Pteropodoidea	Pteropodidae	Nyctimeninae	NA	Nyctimene	NA	keasti	D. J. Kitchener in D. J. Kitchener, Packer, & Maryanto	0	Nyctimene albiventer keasti	Kitchener, D.J., Packer, W.C. and Maryanto, I. 1993. Taxonomic status of _Nyctimene_ (Chiroptera: Pteropodidae) from the Banda, Kai and Aru Islands, Maluku, Indonesia – implications for biogeography. Records of the Western Australian Museum 16(3):399-417.	https://www.biodiversitylibrary.org/page/54768845	MZB 15300	holotype		"Near Taman Anggrek, 12 km N.Tual, Pulau Dullah, closely associated with Pulan Kai Kecil, (5°38'S, 132°44'E) … at sea level," Moluccas, Indonesia.	-5.63333	132.7333	NA	NA				Indonesia|East Timor?	Asia|Oceania (Continent)	Indomalaya|Australasia	NT	0	0	0	Nyctimene_keasti	0	sciname match	Nyctimene_keasti	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	Nyctimene		keasti	Kitchener in Kitchener, Packer, and Maryanto	1993	0	Rec. West. Aust. Mus.	0.95	Keast's Tube-nosed Fruit Bat	babari Bergmans, 2001; tozeri Kitchener, 1995 [in Kitchener et al., 1995].	Indonesia, Maluku, Pulau Dullah (closely associated with Pulau Kai Kecil), 12 km N Tual, near Taman Anggrek, 5°38'S, 132°44'E, sea level.	Babar, Tanimbar, and Kai Isls (Molucca Isls, Indonesia); probably Timor and Flores (K. Helgen, pers. comm.).	<a href='https://cites.org/eng/app/appendices.php' target='_blank'>Not Listed</a>	<a href='https://www.iucnredlist.org/species/136441/21983677/' target='_blank'>Near Threatened</a>	cephalotesspecies group. Originally described as a subspecies of albiventer, but recognized as a distinct species by Kitchener et al. (1995c)and Bergmans (2001). Revised by Bergmans (2001).		Mammal Diversity Database. (2025). Mammal Diversity Database (Version 2.2) [Data set]. Zenodo. https://doi.org/10.5281/zenodo.15007505	NA	Nyctimene keasti; Nyctimene keasti; Nyctimene keasti; Nyctimene keasti; Nyctimene keasti; Nyctimene keasti; keasti; babari; tozeri; keasti; babari; tozeri; keasti?; babari; tozeri; keasti; tozeri; babari; Nyctimene de Keast; Keast-Rohrennasenflughund; Nyctimeno de Keast; Keast's Tube-nosed Fruit Bat; Keast's Tube-nosed Fruit Bat; Keast's Tube-nosed Fruit Bat; N. keasti