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line:xlsx:hash://sha256/181a039844a33e66a35a457b7ece741051086608e425a040051b79581d606b97!/Sheet1!/L1123	application/vnd.openxmlformats-officedocument.spreadsheetml.sheet	N/A	N/A	N/A	N/A	N/A	Paranyctimene tenax	Paranyctimene tenax	Paranyctimene tenax	Paranyctimene tenax	Paranyctimene tenax	Paranyctimene tenax	Paranyctimene tenax	Paranyctimene tenax	Paranyctimene tenax	Paranyctimene tenax		[MSW3] Originally placed in Nyctimene (subgenus Paranyctimene) by Bergmans (2001); see comments under those genera. Many published records referred to raptor may actually represent tenax, see Bergmans (2001).; [HMW] Nyctimene (Paranyctimene) tenax Bergmans, 2001 , “upstream of Anadea (about 07°36’S , 146° 37 °E), 32 km SSW of Wau, Morobe Province [ Papua New Guinea ], alt. 850 m .” Paranyctimene tenax is very similar to P. raptor , and they are often confused; thus, distributional limits between the two species have not been fully differentiated. Two subspecies recognized.; [batnames2022] Originally placed in Nyctimene (subgenus Paranyctimene ) by Bergmans (2001); see comments under those genera. Many publishedrecords referred to raptor may actually represent tenax , see Bergmans (2001).; [IUCN] Very similar in appearance and overall size but distinct from Paranyctimene raptor (see Bergmans 2001), with findings from a recent genetic study supporting the recognition of two Paranyctimene taxa (K.N. Armstrong unpublished data).; [batnames2023] Originally placed in Nyctimene (subgenus Paranyctimene ) by Bergmans (2001); see comments under those genera. Many publishedrecords referred to raptor may actually represent tenax , see Bergmans (2001).; [batnames2025_1.7] Originally placed in Nyctimene (subgenus Paranyctimene) by Bergmans (2001); see comments under those genera. Many published records referred to raptor may actually represent tenax, see Bergmans (2001).								tenax, marculus		tenax, marculus		tenax, marculus		tenax, marculus	Very similar in appearance and overall size but distinct from Paranyctimene raptor (see Bergmans 2001), with findings from a recent genetic study supporting the recognition of two Paranyctimene taxa (K.N. Armstrong unpublished data).	tenax, marculus		tenax, marculus 	marculus, tenax	marculus, tenax		marculus (Bergmans, 2001)|tenax (Bergmans, 2001)						N/A																																								_P. t. marculus_ (Bergmans, 2001); _P. t. tenax_ (Bergmans, 2001)			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			Paranyctimene tenax	Paranyctimene		tenax	Bergmans	y	2001		Beaufortia	51		146		Steadfast Tube-nosed Fruit Bat	Papua New Guinea, Morobe Province, 32 km SSW of Wau, upstream of Anadea, about 07°36'S, 146°37'E, 850 m.	Mainland New Guinea; Waigeo Isl (Indonesia, Prov. of Papua).	IUCN 2003 – not evaluated (new species).	marculus Bergmans, 2001.	Originally placed in Nyctimene (subgenus Paranyctimene) by Bergmans (2001); see comments under those genera. Many published records referred to raptor may actually represent tenax, see Bergmans (2001).	03AD87FAFFFEF6118CAF38BDFE93F642	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	116	zip:hash://sha256/ec5fd314a06aba1a7b0b72f23e54ac625ae272bd98f82f1d01f4c09627d9e8e0!/treatments-xml-main/data/03/AD/87/03AD87FAFFFEF6118CAF38BDFE93F642.xml	Paranyctimene tenax	Pteropodidae	Paranyctimene	tenax		2001	Nyctiméne de Nijhoff @fr | Standhafter Rohrennasenflughund @de | Paranyctimeno de Nijhoff @es	Nyctimene (Paranyctimene) tenax Bergmans, 2001 , “upstream of Anadea (about 07°36’S , 146° 37 °E), 32 km SSW of Wau, Morobe Province [ Papua New Guinea ], alt. 850 m .” Paranyctimene tenax is very similar to P. raptor , and they are often confused; thus, distributional limits between the two species have not been fully differentiated. Two subspecies recognized.	P.t.tenaxBergmans,2001—presumablymuchofNewGuinea(possiblyfoundpri-marilyinSlowlands);distributionallimitsbetweenthisspeciesandtheUnstripedTube-noseFruitBat(FP.raptor)areuncertainandthusdistributiongivenhereistentative. P. t. marculus Bergmans, 2001 — Waigeo I.	Tail 15-3-21- 5 mm , ear 11:6-12- 7 mm , hindfoot 11:5-13- 5 mm , forearm 51-54- 9 mm ; weight 25- 5 g . The Steadfast Tube-nosed Fruit Bat is very similar to the Unstriped Tube-nosed Fruit Bat ( P. raptor ) but can be distinguished by its larger average body and skull measurements and its more robust skull, along with some skull and dental differences. Rostrum is short, with long tubular divergent nostrils. Ears are somewhat elongated, with bluntly pointed tips and thickened along top edges. Eyes are large, with amberirises, and there is often greenish ring around eyes and darker fur behind nostrils. Dorsal pelage is dark brown, being grayish brown on head and fading to medium brown on back of head and darker medium to dark brown on back. Females are typically lighter overall than males, with slightly orangish hue to back of head. Unlike most species of Nyctimene , there is no dorsal stripe down middle of back. Ventral pelage is yellowish white, being more yellowish brown on flanks, genital region, upper arms, and legs. Ears are dark blackish brown, occasionally with a few yellow spots on edges. Wings and uropatagium are uniquely tinged dark lime-green, being browner near dark brown digits. Wings occasionally have some randomly dispersed small white dots. Second digit of wing has a claw, and wing attaches at second digit of foot. Tail is black and wrinkled, and narrow uropatagium connects at base and stretches to calcar at ankles. Claws are black. Skull and mandible are comparatively robust, rostrum is short and wide, post-dental palate is longer than in species of Nyctimene and the Unstriped Tube-nosed Fruit Bat, sagittal crest is well developed, and occipital crest 1s well developed. Single lower incisor is completely deciduous, falling out before adulthood; canines and premolars are considerably longer and more slender than in Nyctimene , lower molars are broad and rounded in dorsal view; lower canines replace incisors and are long and powerful; P| is some distance from C, and next premolar (P,) and is within tooth row; P, is narrow; and P, and M, have close and basally touching inner and outer cusps.	Primary and secondary tropical moist forests, rural gardens, and swamps at an elevation of 850 m (type locality). Specimens attributed to the Steadfast Tube-nosed Fruit Bat and the Unstriped Tube-nosed Fruit Bat have been recorded from sea level up to elevations of ¢. 1350 m .	Diets of the Steadfast Tube-nosed Fruit Bat are probably similar to those of the Unstriped Tube-nosed Fruit Bat.	Records of pregnant females listed under the Unstriped Tube-nosed Fruit Bat might apply to the Steadfast Tube-nosed Fruit Bat.	Steadfast Tube-nosed Fruit Bats are nocturnal and roost in vegetation.	Steadfast Tube-nosed Fruit Bats are largely solitary.	Classified as Least Concern on The IUCN Red List. Distributional limits and threats to the Steadfast Tube-nosed Fruit Bat are confused by the lack of taxonomic clarity between it and the Unstriped Tube-nosed Fruit Bat. Additional studies are needed to clarify conservation status of both species.	Bergmans (2001) | Bonaccorso (1998) | Flannery (1995a, 1995b) | Helgen (2007a) | Helgen, Opiang & Thomas (2009)		98. Steadfast Tube-nosed Fruit Bat Paranyctimene tenax French: Nyctiméne de Nijhoff / German: Standhafter Rohrennasenflughund / Spanish: Paranyctimeno de Nijhoff Taxonomy. Nyctimene (Paranyctimene) tenax Bergmans, 2001 , “upstream of Anadea (about 07°36’S , 146° 37 °E), 32 km SSW of Wau, Morobe Province [ Papua New Guinea ], alt. 850 m .” Paranyctimene tenax is very similar to P. raptor , and they are often confused; thus, distributional limits between the two species have not been fully differentiated. Two subspecies recognized. Subspecies and Distribution. P.t.tenaxBergmans,2001—presumablymuchofNewGuinea(possiblyfoundpri-marilyinSlowlands);distributionallimitsbetweenthisspeciesandtheUnstripedTube-noseFruitBat(FP.raptor)areuncertainandthusdistributiongivenhereistentative. P. t. marculus Bergmans, 2001 — Waigeo I. Descriptive notes. Tail 15-3-21- 5 mm , ear 11:6-12- 7 mm , hindfoot 11:5-13- 5 mm , forearm 51-54- 9 mm ; weight 25- 5 g . The Steadfast Tube-nosed Fruit Bat is very similar to the Unstriped Tube-nosed Fruit Bat ( P. raptor ) but can be distinguished by its larger average body and skull measurements and its more robust skull, along with some skull and dental differences. Rostrum is short, with long tubular divergent nostrils. Ears are somewhat elongated, with bluntly pointed tips and thickened along top edges. Eyes are large, with amberirises, and there is often greenish ring around eyes and darker fur behind nostrils. Dorsal pelage is dark brown, being grayish brown on head and fading to medium brown on back of head and darker medium to dark brown on back. Females are typically lighter overall than males, with slightly orangish hue to back of head. Unlike most species of Nyctimene , there is no dorsal stripe down middle of back. Ventral pelage is yellowish white, being more yellowish brown on flanks, genital region, upper arms, and legs. Ears are dark blackish brown, occasionally with a few yellow spots on edges. Wings and uropatagium are uniquely tinged dark lime-green, being browner near dark brown digits. Wings occasionally have some randomly dispersed small white dots. Second digit of wing has a claw, and wing attaches at second digit of foot. Tail is black and wrinkled, and narrow uropatagium connects at base and stretches to calcar at ankles. Claws are black. Skull and mandible are comparatively robust, rostrum is short and wide, post-dental palate is longer than in species of Nyctimene and the Unstriped Tube-nosed Fruit Bat, sagittal crest is well developed, and occipital crest 1s well developed. Single lower incisor is completely deciduous, falling out before adulthood; canines and premolars are considerably longer and more slender than in Nyctimene , lower molars are broad and rounded in dorsal view; lower canines replace incisors and are long and powerful; P| is some distance from C, and next premolar (P,) and is within tooth row; P, is narrow; and P, and M, have close and basally touching inner and outer cusps. Habitat. Primary and secondary tropical moist forests, rural gardens, and swamps at an elevation of 850 m (type locality). Specimens attributed to the Steadfast Tube-nosed Fruit Bat and the Unstriped Tube-nosed Fruit Bat have been recorded from sea level up to elevations of ¢. 1350 m . Food and Feeding. Diets of the Steadfast Tube-nosed Fruit Bat are probably similar to those of the Unstriped Tube-nosed Fruit Bat. Breeding. Records of pregnant females listed under the Unstriped Tube-nosed Fruit Bat might apply to the Steadfast Tube-nosed Fruit Bat. Activity patterns. Steadfast Tube-nosed Fruit Bats are nocturnal and roost in vegetation. Movements, Home range and Social organization. Steadfast Tube-nosed Fruit Bats are largely solitary. Status and Conservation. Classified as Least Concern on The IUCN Red List. Distributional limits and threats to the Steadfast Tube-nosed Fruit Bat are confused by the lack of taxonomic clarity between it and the Unstriped Tube-nosed Fruit Bat. Additional studies are needed to clarify conservation status of both species. Bibliography. Bergmans (2001), Bonaccorso (1998), Flannery (1995a, 1995b), Helgen (2007a), Helgen, Opiang & Thomas (2009).	Simmons, N.B. and A.L. Cirranello. 2022B. Bat Species of the World: A taxonomic and geographic database. Accessed on 10/11/2022.	Pteropodidae	Paranyctimene tenax	Paranyctimene		tenax	Bergmans	2001	1	Beaufortia	53:26:00	Steadfast Tube-nosed Fruit Bat	<b> marculus </b> Bergmans, 2001.	Papua New Guinea, Morobe Province, 32 km SSW of Wau, upstream of Anadea, about 07°36'S, 146°37'E, 850 m.	Mainland New Guinea; Waigeo Isl (Indonesia, Prov. of Papua).	Not listed.	Least Concern	Originally placed in Nyctimene (subgenus Paranyctimene ) by Bergmans (2001); see comments under those genera. Many publishedrecords referred to raptor may actually represent tenax , see 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	Paranyctimene tenax	23	Steadfast Tube-nosed Fruit Bat		Theria	Placentalia	Boreoeutheria	Laurasiatheria	CHIROPTERA	PTEROPODIFORMES	NA	NA	PTEROPODOIDEA	PTEROPODIDAE	NYCTIMENINAE	NA	Paranyctimene	NA	tenax	Bergmans	2001	1						"upstream of Anadea (about 07°36'S, 146°37'E), 32 km SSW of Wau, Morobe Province [Papua New Guinea], alt. 850 m."	-7.6	146.62	tenax (Bergmans, 2001)|marculus (Bergmans, 2001)	NA	NA	Indonesia|Papua New Guinea	Oceania	Australasia/Oceania	LC	0	0	0	Paranyctimene_tenax	0	sciname match	Paranyctimene_tenax	0	IUCN. 2022. The IUCN Red List of Threatened Species. Version 2022-1. https://www.iucnredlist.org. Accessed on [28 September, 2022].	136836	Paranyctimene tenax	ANIMALIA	CHORDATA	MAMMALIA	CHIROPTERA	PTEROPODIDAE	Paranyctimene	tenax	Bergmans, 2001	Very similar in appearance and overall size but distinct from Paranyctimene raptor (see Bergmans 2001), with findings from a recent genetic study supporting the recognition of two Paranyctimene taxa (K.N. Armstrong unpublished data).	20000000	Paranyctimene tenax	Least Concern		2021	2020-11-21 00:00:00 UTC	3.1	English	This species is assessed as Least Concern given its wide distribution, use of a broad range of forested habitats, presumed large population size, occurrence in protected areas, and the absence of evidence for a decline.	This morphologically cryptic species is largely solitary. It can be found in primary and secondary tropical moist forest, rural gardens, and swamps. Roosts occur in vegetation. Females give birth to single young.	There appear to be no major threats to this species that could cause widespread or imminent decline, mainly because of the species’ small size, solitary roosting behaviour and broad distribution.	It can be relatively common and population size is assumed to be large and stable.	Unknown	Until recently, the assumed distribution of both Paranyctimene raptor and P. tenax  was considered to span the island of New Guinea, plus the islands of Salawati and Waigeo. Unpublished genetic results (K.N. Armstrong unpublished data) support information in Helgen (2007) that: “Paranyctimene tenax may be widely distributed in the northern lowlands of New Guinea and P. raptor in the south (N. Irwin pers. comm.).” The type locality is near Wau in Morobe Province, Papua New Guinea, which is also consistent with a northern distribution of P. tenax in New Guinea. It ranges from sea level to 1,350 m asl.	The species is not known to hunted, used, or traded.	Terrestrial	It is presumed to be present in most protected areas throughout its range. Broadscale protection of forests will help maintain population size in this species.	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	Paranyctimene		tenax	Bergmans	2001	1	Beaufortia	53:26:00	Steadfast Tube-nosed Fruit Bat	<b> marculus </b> Bergmans, 2001.	Papua New Guinea, Morobe Province, 32 km SSW of Wau, upstream of Anadea, about 07°36'S, 146°37'E, 850 m.	Mainland New Guinea; Waigeo Isl (Indonesia, Prov. of Papua).	Not listed.	Least Concern	Originally placed in Nyctimene (subgenus Paranyctimene ) by Bergmans (2001); see comments under those genera. Many publishedrecords referred to raptor may actually represent tenax , see Bergmans (2001).	Paranyctimene tenax	1004430	23	Steadfast Tube-nosed Fruit Bat		Theria	Placentalia	Boreoeutheria	Laurasiatheria	CHIROPTERA	PTEROPODIFORMES	NA	NA	PTEROPODOIDEA	Pteropodidae	NYCTIMENINAE	NA	Paranyctimene	NA	tenax	Bergmans	2001	1						"upstream of Anadea (about 07°36'S, 146°37'E), 32 km SSW of Wau, Morobe Province [Papua New Guinea], alt. 850 m."	-7.6	146.6167	tenax (Bergmans, 2001)|marculus (Bergmans, 2001)	NA	NA				Indonesia|Papua New Guinea	Oceania	Australasia/Oceania	LC	0	0	0	Paranyctimene_tenax	0	sciname match	Paranyctimene_tenax	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	Paranyctimene_tenax	1004430	23	Steadfast Tube-nosed Fruit Bat		Theria	Placentalia	Boreoeutheria	Laurasiatheria	Chiroptera	Yinpterochiroptera	NA	NA	Pteropodoidea	Pteropodidae	Nyctimeninae	NA	Paranyctimene	NA	tenax	Bergmans	1	Nyctimene (Paranyctimene) tenax	Bergmans, W. 2001. Notes on distribution and taxonomy of Australasian bats. I. Pteropodinae and Nyctimeninae (Mammalia, Megachiroptera, Pteropodidae). Beaufortia 51(8):119-152.		ZMA.MAM.25360.a, ZMA.MAM.25360.b	holotype	https://data.biodiversitydata.nl/naturalis/specimen/ZMA.MAM.25360.a | https://data.biodiversitydata.nl/naturalis/specimen/ZMA.MAM.25360.b	"upstream of Anadea (about 07°36'S, 146°37'E), 32 km SSW of Wau, Morobe Province [Papua New Guinea], alt. 850 m."	-7.6	146.6167	NA	NA				Indonesia|Papua New Guinea	Oceania (Continent)	Australasia	LC	0	0	0	Paranyctimene_tenax	0	sciname match	Paranyctimene_tenax	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	Paranyctimene		tenax	Bergmans	2001	1	Beaufortia	53:26:00	Steadfast Tube-nosed Fruit Bat	marculus  Bergmans, 2001.	Papua New Guinea, Morobe Province, 32 km SSW of Wau, upstream of Anadea	Mainland New Guinea; Waigeo Isl (Indonesia, Prov. of Papua)	<a href='https://cites.org/eng/app/appendices.php' target='_blank'>Not Listed</a>	<a href='https://www.iucnredlist.org/species/136836/22041117/' target='_blank'>Least Concern</a>	Originally placed in Nyctimene (subgenus Paranyctimene) by Bergmans (2001); see comments under those genera. Many published records referred to raptor may actually represent tenax, see Bergmans (2001).		Mammal Diversity Database. (2025). Mammal Diversity Database (Version 2.2) [Data set]. Zenodo. https://doi.org/10.5281/zenodo.15007505	NA	Paranyctimene tenax; Paranyctimene tenax; Paranyctimene tenax; Paranyctimene tenax; Paranyctimene tenax; Paranyctimene tenax; tenax; marculus; tenax; marculus; marculus; tenax; marculus; Nyctiméne de Nijhoff; Standhafter Rohrennasenflughund; Paranyctimeno de Nijhoff; Steadfast Tube-nosed Fruit Bat; Steadfast Tube-nosed Fruit Bat; Steadfast Tube-nosed Fruit Bat; P. tenax