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line:xlsx:hash://sha256/181a039844a33e66a35a457b7ece741051086608e425a040051b79581d606b97!/Sheet1!/L1130	application/vnd.openxmlformats-officedocument.spreadsheetml.sheet	Penthetor lucasi	Penthetor lucasi	Penthetor lucasi	Penthetor lucasi	Penthetor lucasi	Penthetor lucasi	Penthetor lucasii	Penthetor lucasii	Penthetor lucasii	Penthetor lucasi	Penthetor lucasii	Penthetor lucasii	Penthetor lucasii	Penthetor lucasii	Penthetor lucasii		[HMW] Cynopterus (Ptenochirus) lucasii Dobson, 1880 , “ Sarawak , Borneo,” Malaysia . Specimens of P. lucasii from Peninsular Malaysia are referred here to subspecies suyantoi with caution. Two subspecies recognized.; [batnames2022] The misspelling lucasi is in widespread use and is attributed to Dobson, 1880, the original describer. However, the correct form remains lucasii , following ICZN article 33.4. 'Use of -i for -ii and vice versa, and other alternative spellings, in subsequent spellings of species-group names.'; [MDD2022] often spelt with only one 'i', but the original description has two, which is what is used here; [batnames2023] The misspelling lucasi is in widespread use and is attributed to Dobson, 1880, the original describer. However, the correct form remains lucasii , following ICZN article 33.4. 'Use of -i for -ii and vice versa, and other alternative spellings, in subsequent spellings of species-group names.'; [MDD2023] often spelt with only one 'i', but the original description has two, which is what is used here; [MDD2025_2.0] often spelt with only one 'i', but the original description has two, which is what is used here; [batnames2025_1.7] The misspelling lucasi is in widespread use and is attributed to Dobson, 1880, the original describer. However, the correct form remains lucasii, following ICZN article 33.4. 'Use of -i for -ii and vice versa, and other alternative spellings, in subsequent spellings of species-group names.'; [MDD2025_2.2] often spelt with only one 'i', but the original description has two, which is what is used here										lucasii, suyantoi				lucasii, suyantoi				lucasii, suyantoi	lucasii, lucasi, suyantoi			lucasii (Dobson, 1880)|lucasi (Bonhote, 1901) [incorrect subsequent spelling]|suyantoi Maryanto, 2003		Corbet, G.B. and Hill, J.E. 1980. A World List of Mammalian Species. British Museum (Natural History), London, 226 pp.	Lucas' short-nosed fruit	Malaya, Borneo bat	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.	Penthetor lucasi	Malaysia, Sarawak.	Dobson	1880	Ann. Mag. Nat. Hist., ser. 5, 6:163.	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	Lucas's short-nosed fruit	Malaya, Borneo bat	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.	Dobson	1880	Ann. Mag. Nat. Hist., ser. 5, 6:163.		W Malaysia, Borneo, Riau Arch. (Indonesia).	Malaysia, Sarawak.		DOBSON	1880	Size medium (fore arm length, 58-63 mm).	Distribution: Same as for genus.	No subspecies.		35	species	P. lucasi	DOBSON	1880	Penthetor	genus	Penthetor lucasi				Size medium (fore arm length, 58-63 mm).	No subspecies.		1. P. lucasi (DOBSON 1880).	1	_P. l. lucasii_ (Dobson, 1880); _P. l. suyantoi_ Maryanto, 2003			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			Penthetor lucasi	Penthetor		lucasi	Dobson	y	1880		Ann. Mag. Nat. Hist., ser. 5	6		163		Lucas's Short-nosed Fruit Bat	Malaysia, N Borneo, Sarawak.	W Malaysia, Borneo, Sumatra, Riau Arch. (Indonesia).	IUCN/SSC Action Plan (1992) – Not Threatened. IUCN 2003 – Lower Risk (lc).			03AD87FAFFD0F63E89B93C3BF6D1F5CB	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	78	zip:hash://sha256/ec5fd314a06aba1a7b0b72f23e54ac625ae272bd98f82f1d01f4c09627d9e8e0!/treatments-xml-main/data/03/AD/87/03AD87FAFFD0F63E89B93C3BF6D1F5CB.xml	Penthetor lucasii	Pteropodidae	Penthetor	lucasii		1880	Cynoptére de Lucas @fr | Lucas-Kurznasenflughund @de | Penthetor de Lucas @es | Dusky Fruit Bat @en	Cynopterus (Ptenochirus) lucasii Dobson, 1880 , “ Sarawak , Borneo,” Malaysia . Specimens of P. lucasii from Peninsular Malaysia are referred here to subspecies suyantoi with caution. Two subspecies recognized.	PL lucasii Dobson, 1880 — Borneo. P. l. suyantoi Maryanto, 2004 — Peninsular Malaysia and Sumatra, including Riau Archipelago.	Head—body 78-103 mm , tail 8-13 mm , ear 13-17 mm , hindfoot 10- 13 mm , forearm 54-66 mm (males) and 57-63 mm (females); weight 34-48 g (males) and 42-55 g (females). Head is stout; muzzle is broad at base, tapering, and moderately deep; nostrils are shortly tubular; philtrum is divided into two parallel grooves. Eyes are large and bulging; iris is warm brown. Fars are moderately long and rounded, pale brown skin is thin, and antitragus is very small. Head is very wide; pelage is sparse on muzzle and cheek; pelage is very short, dense, and darker on cap, with some individuals presenting a distinctly darker, wide central line on the cap; and very long guard hairs occur in occiput. Neck and back are sparsely haired, especially in scapular area; dorsum is brown to grayish brown. Uropatagium is naked, tail is moderately long and very thin, calcar is short, and tibia is long. Throatis thinly furred but has isolated, very long hairs, and sides of neck lack tufts or distinctly colored ruff. Chest and belly are light grayish brown, and flanks are distinctly warmer and browner. Wing membranes are dark brown from sides of body and attach on first toe, pollex and index finger are distinctly short, and index claw is present. Skull lacks basicranial deflection, rostrum is short and moderately deep, orbit is large, zygomatic rootis slightly above upper alveolar line, and zygomais rather thin and little arched posteriorly. Dorsally, rostrum is wide at base and slightly tapering; paranasal recesses are inflated, passing root of small, triangular postorbital processes; there is no postorbital foramen or postorbital constriction; barely obvious temporal lines do notjoin in a sagittal crest; braincase is oval and long; and nuchal crest is moderately developed. Ventrally, palate is moderately long (wider in nominate lucasit) and flat; tooth rows are almost straight; post-dental is short and converging; end of post-dental palate is concave; there is no palatine spine; sphenoidalcrest is thin and sharp; and ectotympanic is small and wide anteriorly, and edged internally by ribbon-like entotympanic. Mandible is thin (less so in subspecies suyantoi); coronoid slopes butis rather tall, with rounded tip; condyle is level with lower alveolar line; and angle is distinctly marked. Dental formula is12/1,C1/1,P3/3 M 1/2 (x2) = 28. Upper incisors are very thin; I! is longer; Cl is short with marked anterointernal groove; P' is a spicule; posterior cheekteeth decrease in height, without additional cusps, and have rectangular outlines; and P23 is broad. Lower incisors are thin (possibly representing L,); C, is relatively wide at base and short; P, is peg-like; posterior cheekteeth decrease in height and are rather wide and somewhat squarish in occlusal outline; and M, is peg-like. Diploid numberis 2n = 48.	[Lowland to hill rainforest from sea level up to elevations of ¢. 600 m .	L.ucas’s Short-nosed Fruit Bat is primarily frugivorous.	In Brunei , Lucas’s Short-nosed Fruit Bat is seasonally monoestrous, with synchronized births centered in October and estrus and mating within 2-3 weeks postpartum, followed by prolonged delayed embryonic development; rapid development resumes in the next year’s July-September, and births occur again around October. This birth peak does not correlate with rainfall or fruiting/flowering peaks and might be related to specific resources. Neonates weigh c. 15 g at birth and are able to fly in 2-3 weeks old when they weigh more than 20 g . Lactation lasts 2-4 months. Females achieve sexual maturity at weights of 33-34 g and males at c.12 months old when they weigh 27-30 g . Other patterns might exist elsewhere based on near-term pregnant females recorded in January and lactating female in May in Peninsular Malaysia .	Lucas’s Short-nosed Fruit Bat roosts in caves and rock shelters.	Colonies of Lucas’s Short-nosed Fruit Bats can have more than 100 individuals that cluster together in crevices and cave ceilings in the twilight zones. Nursing females roost together in clusters in caves. Lucas’s Short-nosed Fruit Bat likely commutes between a fixed cave roost and shifting feeding areas.	Classified as Least Concern on The IUCN Red List. Lucas’s Short-nosed Fruit Bats can be locally very common, generally in the proximity of occupied caves, but are generally rare. Limestone extraction and to a lesser extent deforestation are likely conservation threats.	Andersen (1912b) | Bates, Bumrungsri, Suyanto, Francis, Kingston & Maryanto (2008) | Giannini & Simmons (2007a) | Kingston et al. (2006) | Kofron (2007b) | Maryanto (2004) | Mohd-Ridwan & Abdullah (2012) | Payne et al. (1985) | Rahman & Abdullah (2010) | Rosli et al. (2018) | Yong & Dhaliwal (1976)		31. Lucas’s Short-nosed Fruit Bat Penthetor lucasii French: Cynoptére de Lucas / German: Lucas-Kurznasenflughund / Spanish: Penthetor de Lucas Other common names: Dusky Fruit Bat Taxonomy. Cynopterus (Ptenochirus) lucasii Dobson, 1880 , “ Sarawak , Borneo,” Malaysia . Specimens of P. lucasii from Peninsular Malaysia are referred here to subspecies suyantoi with caution. Two subspecies recognized. Subspecies and Distribution. PL lucasii Dobson, 1880 — Borneo. P. l. suyantoi Maryanto, 2004 — Peninsular Malaysia and Sumatra, including Riau Archipelago. Descriptive notes. Head—body 78-103 mm , tail 8-13 mm , ear 13-17 mm , hindfoot 10- 13 mm , forearm 54-66 mm (males) and 57-63 mm (females); weight 34-48 g (males) and 42-55 g (females). Head is stout; muzzle is broad at base, tapering, and moderately deep; nostrils are shortly tubular; philtrum is divided into two parallel grooves. Eyes are large and bulging; iris is warm brown. Fars are moderately long and rounded, pale brown skin is thin, and antitragus is very small. Head is very wide; pelage is sparse on muzzle and cheek; pelage is very short, dense, and darker on cap, with some individuals presenting a distinctly darker, wide central line on the cap; and very long guard hairs occur in occiput. Neck and back are sparsely haired, especially in scapular area; dorsum is brown to grayish brown. Uropatagium is naked, tail is moderately long and very thin, calcar is short, and tibia is long. Throatis thinly furred but has isolated, very long hairs, and sides of neck lack tufts or distinctly colored ruff. Chest and belly are light grayish brown, and flanks are distinctly warmer and browner. Wing membranes are dark brown from sides of body and attach on first toe, pollex and index finger are distinctly short, and index claw is present. Skull lacks basicranial deflection, rostrum is short and moderately deep, orbit is large, zygomatic rootis slightly above upper alveolar line, and zygomais rather thin and little arched posteriorly. Dorsally, rostrum is wide at base and slightly tapering; paranasal recesses are inflated, passing root of small, triangular postorbital processes; there is no postorbital foramen or postorbital constriction; barely obvious temporal lines do notjoin in a sagittal crest; braincase is oval and long; and nuchal crest is moderately developed. Ventrally, palate is moderately long (wider in nominate lucasit) and flat; tooth rows are almost straight; post-dental is short and converging; end of post-dental palate is concave; there is no palatine spine; sphenoidalcrest is thin and sharp; and ectotympanic is small and wide anteriorly, and edged internally by ribbon-like entotympanic. Mandible is thin (less so in subspecies suyantoi); coronoid slopes butis rather tall, with rounded tip; condyle is level with lower alveolar line; and angle is distinctly marked. Dental formula is12/1,C1/1,P3/3 M 1/2 (x2) = 28. Upper incisors are very thin; I! is longer; Cl is short with marked anterointernal groove; P' is a spicule; posterior cheekteeth decrease in height, without additional cusps, and have rectangular outlines; and P23 is broad. Lower incisors are thin (possibly representing L,); C, is relatively wide at base and short; P, is peg-like; posterior cheekteeth decrease in height and are rather wide and somewhat squarish in occlusal outline; and M, is peg-like. Diploid numberis 2n = 48. Habitat. [Lowland to hill rainforest from sea level up to elevations of ¢. 600 m . Food and Feeding. L.ucas’s Short-nosed Fruit Bat is primarily frugivorous. Breeding. In Brunei , Lucas’s Short-nosed Fruit Bat is seasonally monoestrous, with synchronized births centered in October and estrus and mating within 2-3 weeks postpartum, followed by prolonged delayed embryonic development; rapid development resumes in the next year’s July-September, and births occur again around October. This birth peak does not correlate with rainfall or fruiting/flowering peaks and might be related to specific resources. Neonates weigh c. 15 g at birth and are able to fly in 2-3 weeks old when they weigh more than 20 g . Lactation lasts 2-4 months. Females achieve sexual maturity at weights of 33-34 g and males at c.12 months old when they weigh 27-30 g . Other patterns might exist elsewhere based on near-term pregnant females recorded in January and lactating female in May in Peninsular Malaysia . Activity patterns. Lucas’s Short-nosed Fruit Bat roosts in caves and rock shelters. Movements, Home range and Social organization. Colonies of Lucas’s Short-nosed Fruit Bats can have more than 100 individuals that cluster together in crevices and cave ceilings in the twilight zones. Nursing females roost together in clusters in caves. Lucas’s Short-nosed Fruit Bat likely commutes between a fixed cave roost and shifting feeding areas. Status and Conservation. Classified as Least Concern on The IUCN Red List. Lucas’s Short-nosed Fruit Bats can be locally very common, generally in the proximity of occupied caves, but are generally rare. Limestone extraction and to a lesser extent deforestation are likely conservation threats. Bibliography. Andersen (1912b), Bates, Bumrungsri, Suyanto, Francis, Kingston & Maryanto (2008), Giannini & Simmons (2007a), Kingston et al. (2006), Kofron (2007b), Maryanto (2004), Mohd-Ridwan & Abdullah (2012), Payne et al. (1985), Rahman & Abdullah (2010), Rosli et al. (2018), Yong & Dhaliwal (1976).	Simmons, N.B. and A.L. Cirranello. 2022B. Bat Species of the World: A taxonomic and geographic database. Accessed on 10/11/2022.	Pteropodidae	Penthetor lucasii	Penthetor		lucasii	Dobson	1880	1	Ann. Mag. Nat. Hist.	ser. 5, 6: 163	Lucas' Short-nosed Fruit Bat	None.	Malaysia, N Borneo, Sarawak.	W Malaysia, Borneo, Sumatra, Riau Arch. (Indonesia).	Not listed.	Least Concern	The misspelling lucasi is in widespread use and is attributed to Dobson, 1880, the original describer. However, the correct form remains lucasii , following ICZN article 33.4. 'Use of -i for -ii and vice versa, and other alternative spellings, in subsequent spellings of species-group names.'	Mammal Diversity Database. (2023). Mammal Diversity Database (Version 1.11) [Data set]. Zenodo. https://doi.org/10.5281/zenodo.7830771 released 15 April 2023	Penthetor lucasii	23	Lucas's Short-nosed Fruit Bat	Dusky Fruit Bat	Theria	Placentalia	Boreoeutheria	Laurasiatheria	CHIROPTERA	PTEROPODIFORMES	NA	NA	PTEROPODOIDEA	PTEROPODIDAE	CYNOPTERINAE	BALIONYCTERINI	Penthetor	NA	lucasii	Dobson	1880	1						"Saráwak, Borneo," Malaysia.			lucasii (Dobson, 1880)|suyantoi Maryanto, 2004	often spelt with only one 'i', but the original description has two, which is what is used here	NA	Thailand|Malaysia|Singapore|Indonesia|Brunei	Asia	Indomalaya	LC	0	0	0	Penthetor_lucasii	0	sciname match	Penthetor_lucasi	0	IUCN. 2022. The IUCN Red List of Threatened Species. Version 2022-1. https://www.iucnredlist.org. Accessed on [28 September, 2022].	16563	Penthetor lucasi	ANIMALIA	CHORDATA	MAMMALIA	CHIROPTERA	PTEROPODIDAE	Penthetor	lucasi	(Dobson, 1880)		20000000	Penthetor lucasi	Least Concern		2020	2019-07-16 00:00:00 UTC	3.1	English	This species is listed as Least Concern in view of its wide distribution, it occurs in a number of protected areas, it is tolerant of some disturbance of its habitat, and because it is unlikely to be declining at nearly the rate required to qualify for listing in a threatened category.	The species roosts in caves and can be found in colonies of more than 100 individuals. It inhabits secondary and primary lowland to hill forest (Medway 1983). Following mating, there appears to be a relatively long period of delayed embryonic development with a relatively rapid embryonic development from July to September (Kofon 2007). There is generally a synchronized birth season around October followed by 2-4 months of lactation (Kofon 2007). Reproduction appears to be linked to the flowering/fruiting and availability of specific plant species (Kofon 2007).	This species is most likely affected by limestone extraction and cave disturbance. The significance of deforestation for agriculture, plantations, logging, and fire through a lot of the species' range is unknown and should be evaluated.	The species is uncommon throughout its range. The global population is thought to be declining, but not at a rate that would qualify it as threatened.	Decreasing	This species occurs in Peninsular Malaysia, and throughout Borneo, and Sumatra.		Terrestrial	It is found in protected areas throughout its range. Cave roosts warrant protection and research is needed on the ecology, life history, population and threats of the species.	Indomalayan		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	Penthetor		lucasii	Dobson	1880	1	Ann. Mag. Nat. Hist.	ser. 5, 6: 163	Lucas' Short-nosed Fruit Bat	None.	Malaysia, N Borneo, Sarawak.	W Malaysia, Borneo, Sumatra, Riau Arch. (Indonesia).	Not listed.	Least Concern	The misspelling lucasi is in widespread use and is attributed to Dobson, 1880, the original describer. However, the correct form remains lucasii , following ICZN article 33.4. 'Use of -i for -ii and vice versa, and other alternative spellings, in subsequent spellings of species-group names.'	Penthetor lucasii	1004370	23	Lucas's Short-nosed Fruit Bat	Dusky Fruit Bat	Theria	Placentalia	Boreoeutheria	Laurasiatheria	CHIROPTERA	PTEROPODIFORMES	NA	NA	PTEROPODOIDEA	Pteropodidae	CYNOPTERINAE	BALIONYCTERINI	Penthetor	NA	lucasii	Dobson	1880	1						"Saráwak, Borneo," Malaysia.			lucasii (Dobson, 1880)|suyantoi Maryanto, 2004	often spelt with only one 'i', but the original description has two, which is what is used here	NA				Thailand|Malaysia|Singapore|Indonesia|Brunei	Asia	Indomalaya	LC	0	0	0	Penthetor_lucasii	0	sciname match	Penthetor_lucasi	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	Penthetor_lucasii	1004370	23	Lucas's Short-nosed Fruit Bat	Dusky Fruit Bat	Theria	Placentalia	Boreoeutheria	Laurasiatheria	Chiroptera	Yinpterochiroptera	NA	NA	Pteropodoidea	Pteropodidae	Cynopterinae	Balionycterini	Penthetor	NA	lucasii	Dobson	1	Cynopterus (Ptenochirus) Lucasii	Dobson, G.E. 1880-08-01. Description of a new species of _Cynopterus_ (Ptenochirus) from Saráwak. Annals and Magazine of Natural History (5)6(32):163-164.	https://www.biodiversitylibrary.org/page/25170388	BMNH:Mamm:1880.8.13.1	holotype		"Saráwak, Borneo," Malaysia.			often spelt with only one 'i', but the original description has two, which is what is used here	NA				Thailand|Malaysia|Singapore|Indonesia|Brunei	Asia	Indomalaya	LC (as Penthetor lucasi)	0	0	0	Penthetor_lucasii	0	sciname match	Penthetor_lucasi	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	Penthetor		lucasii	Dobson	1880	1	Ann. Mag. Nat. Hist.	ser. 5, 6: 163	Lucas' Short-nosed Fruit Bat	None.	Malaysia, N Borneo, Sarawak.	W Malaysia, Borneo, Sumatra, Riau Arch. (Indonesia).	<a href='https://cites.org/eng/app/appendices.php' target='_blank'>Not Listed</a>	<a href='https://www.iucnredlist.org/species/16563/22055450/' target='_blank'>Least Concern as Penthetor lucasi</a>	The misspelling lucasi is in widespread use and is attributed to Dobson, 1880, the original describer. However, the correct form remains lucasii, following ICZN article 33.4. 'Use of -i for -ii and vice versa, and other alternative spellings, in subsequent spellings of species-group names.'		Mammal Diversity Database. (2025). Mammal Diversity Database (Version 2.2) [Data set]. Zenodo. https://doi.org/10.5281/zenodo.15007505	NA	Penthetor lucasi; Penthetor lucasii; Penthetor lucasii; Penthetor lucasii; Penthetor lucasi; Penthetor lucasii; lucasii; suyantoi; lucasii; suyantoi; Cynoptére de Lucas; Lucas-Kurznasenflughund; Penthetor de Lucas; Dusky Fruit Bat; Lucas's Short-nosed Fruit Bat; Dusky Fruit Bat; Lucas's Short-nosed Fruit Bat; Lucas' Short-nosed Fruit Bat; P. lucasii