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(1=author & date in parentheses)	Citation	Pages	Common Name	Synonyms	Type Locality	Distribution	CITES	IUCN	Comments	column3781	column3791	subtribe	CONCAT_ALTNAMES
line:xlsx:hash://sha256/181a039844a33e66a35a457b7ece741051086608e425a040051b79581d606b97!/Sheet1!/L1279	application/vnd.openxmlformats-officedocument.spreadsheetml.sheet	Pteropus livingstonei	Pteropus livingstonei	Pteropus livingstonii	Pteropus livingstonei	Pteropus livingstonei	Pteropus livingstonii	Pteropus livingstonii	Pteropus livingstonii	Pteropus livingstonii	Pteropus livingstonii	Pteropus livingstonii	Pteropus livingstonii	Pteropus livingstonii	Pteropus livingstonii	Pteropus livingstonii		[MSW2] See Bergmans (1990).; [MSW3] livingstonii species group. Reviewed by Bergmans (1990). Misspelled livingstonei by Koopman (1993, 1994).; [HMW] Pteropus livingstonii J. E. Gray, 1866 , “Island of Johanna [= Anjouan Island ], Comoro group [= Comoros ].” Pteropus livingstonii is in the livingstonii species group. Monotypic.; [batnames2022]  livingstonii species group; see Almeida et al. (2014). Reviewed by Bergmans (1990). Misspelled livingstonei by Koopman (1993, 1994).; [IUCN] P. livingstonei was used on the 2000 Red List. CITES also uses P. livingstonei , and livingstonei should be the grammatically correct spelling. T. Hutson (Chiroptera Red List Authority) notes that the correct spelling should be livingstonii (also by Simmons in Wilson and Reeder 2005) . ; Andersen’s 1912 emendation of the species name is now considered unjustified (Smith and Leslie 2006).; [batnames2023]  livingstonii species group; see Almeida et al. (2014). Reviewed by Bergmans (1990). Misspelled livingstonei by Koopman (1993, 1994).; [batnames2025_1.7] livingstoniispecies group; see Almeida et al. (2014). Reviewed by Bergmans (1990). Misspelled livingstonei by Koopman (1993, 1994).														livingstonii	P. livingstonei was used on the 2000 Red List. CITES also uses P. livingstonei , and livingstonei should be the grammatically correct spelling. T. Hutson (Chiroptera Red List Authority) notes that the correct spelling should be livingstonii (also by Simmons in Wilson and Reeder 2005) . ; Andersen’s 1912 emendation of the species name is now considered unjustified (Smith and Leslie 2006).			livingstonii 	livingstonii, livingstonei			livingstonii J. E. Gray, 1866|livingstonei Andersen, 1912 [unjustified emendation]		Corbet, G.B. and Hill, J.E. 1980. A World List of Mammalian Species. British Museum (Natural History), London, 226 pp.	Comoro black flying fox	Johanna I, Comoro Is	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.	Pteropus livingstonei	Comoro Isis., Anjouan Isl.	Gray	1866	Proc. Zool. Soc. Lond., 1866:66.	Distribution: Confined to the Comoros (north west of Madagascar).		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	Comoro black flying fox	Anjouan I, Moheli I, Comoro Is; ref. 4.17; E	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	1866	Proc. Zool. Soc. Lond., 1866:66.	See Bergmans (1990).	Comoro Isis.	Comoro Isis, Anjouan Isl.		GRAY	1866	Ears semicircularly rounded. Smaller than P. melanopogon (forearm length, 162-172 mm).	Distribution: Confined to the Comoros (north west of Madagascar).	No subspecies.		24	species	P. livingstonei	GRAY	1866	Pteropus	genus	Pteropus livingstonei				Ears semicircularly rounded. Smaller than P. melanopogon (forearm length, 162-172 mm).	No subspecies.		24. P. livingstonei GRAY 1866 [livingstonei group],	24	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			Pteropus livingstonii	Pteropus		livingstonii	Gray		1866		Proc. Zool. Soc. Lond.	1866		66		Comoro Flying Fox	Comoro Isls, Anjouan Isl.	Comoro Isls.	CITES – Appendix II. IUCN/SSC Action Plan (1992) – Endangered: Limited Distribution. IUCN 2003 – Critically Endangered.		livingstonii species group. Reviewed by Bergmans (1990). Misspelled livingstonei by Koopman (1993, 1994).	03AD87FAFF8AF6648CBB351DF67AF489	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	155	zip:hash://sha256/ec5fd314a06aba1a7b0b72f23e54ac625ae272bd98f82f1d01f4c09627d9e8e0!/treatments-xml-main/data/03/AD/87/03AD87FAFFA5F64B8964351EFD05F301.xml	Pteropus livingstonii	Pteropodidae	Pteropus	livingstonii	J. E. Gray	1866	Roussette de Livingstone @fr | Livingstone-Flughund @de | Zorro volador de Livingstone @es | Comoro Flying Fox @en | Livingstone's Fruit Bat @en	Pteropus livingstonii J. E. Gray, 1866 , “Island of Johanna [= Anjouan Island ], Comoro group [= Comoros ].” Pteropus livingstonii is in the livingstonii species group. Monotypic.	Comoro Is ( Anjouan and Mohéli ).	Head-body 340 mm (tailless), ear 30-31 mm , hindfoot 5H 1- 53 mm , forearm 161-172 mm ; weight 576- 3 g (males) and 601- 5 g (females). Livingstone’s Flying Fox is the largest species of western Indian Ocean Pteropus . Head 1s fox-like; muzzle is long and black. Eyes are relatively large, with orange to scarlet-red irises; ears are large, protruding, and semicircular above; inner margin is convex; outer margin is straight or slightly convex; and exposed concha is smooth, without ribs or grooves. Males and females are alike, generally blackish brown on head and body. Back is dark to blackish brown, with dark russet tinge, and speckled with golden or tawny hairs; tawny hairs are also variably found in mantle, lower back, and flanks; tawny-tipped hairs in mantle form a band c. 6 cm wide and 10-15 mm long anteroposteriorly, with tufts of similarly colored fur on each shoulder. Dense pelage is present on upper inside of femur and uropatagium,; upper side of tibia is naked. Belly and flanks are similarly colored as back. Genital area can be surrounded by ring of deep reddish brown hair, extending laterally as a short stripe. Wings are dark, often jet-black, shiny, and hairless; index claw is present; tail membrane is poorly developed. Skull is typical pteropine, with very long rostrum; orbit is often closed by postorbital process reaching zygomatic arch; and crests and mandible are strong. Dentition is typical pteropine, with relatively small canines grooved in their outer surface; P' is lost in most adults; and cheekteeth are relatively simple but robust.	Undisturbed moist forests and, to a lesser extent, degraded forests at elevations above 300 m on Anjouan and 40 m on Mohéli . Roosts of Livingstone’s Flying Foxes are predominantly found in Ficus exasperata and FE lutea ( Moraceae ); Gyrostipula comorensis ( Rubiaceae ); and especially Gambeya spp. ( Sapotaceae ) and Nuxia pseudodentata ( Stilbaceae ). Roosts are in steep-sided (30-50°) valleys with south-east facing slopes, suggesting that they are sensitive to wind and sun.	Livingstone’s Flying Fox is primarily frugivorous, eating fruits of native trees but also nectar and leaves. Ficus antandronarumis preferred during wet season and F lutea during dry season; other foods come from at least 14 genera in 13 families. Flowers of Gyrostipula and Neonauclea ( Rubiaceae ) and flowers, nectar, and to a lesser extent unripe fruit of Ceiba ( Malvaceae ) are eaten. Captive bats caught and ate insects.	Captive female Livingstone’s Flying Foxesstart to reproduce at 3-4 years old. After gestation of c.152 days, young are generally born in first part of September and weigh c. 137 g . Females with young were also observed in July-August, October, and January. Litter size is one.	Livingstone’s Flying Foxes forage at night. During the day, they remain in roosts and bask and stretch their wings in the sun to warm up, or urinate on their wings while flapping them to cool down. They take advantage of afternoon thermals to soar long distances. Peak activity is between 22:00 h and 02:00 h, and it declines to ¢.04:00 h. They start urinating immediately after taking flight.	Livingstone’s Flying Foxes form colonies of 15-150 individuals, exceptionally 250-349 during rainy seasons. Intra-island movements between roosts are common; inter-island movements are unknown. A dominant male has a harem of adult females and their female offspring.	CITES Appendix II. Classified as Critically Endangered on The IUCN Red List. Livingstone’s Flying Fox has a small distribution ( 1856 km ? overall but probably 65-188 km *) and suspected significant population decline (current population is ¢.1300 animals: 940 on Anjouan and 360 on Mohéli ) from habitat loss due to logging for firewood, construction, and conversion to agricultural use. Recent disappearance of lowland forests has forced individuals higher on slopes up to 400 m on Anjouan and 200 m on Mohéli . It is not threatened by hunting. “The Conservation Action Plan for Livingstone’s Flying Fox” was set up and adopted by the government of Comoros , and a captive breeding program was established that led to a captive population of 59 individuals spread over four institutions as of 2014. Based on high levels of inter-island genetic differentiation, island populations probably need to be considered separate management units. Livingstone’s Flying Foxes do not occur in any protected area, but plans for appropriate reserves are being developed.	Almeida et al. (2014) | Andersen (1912b) | Bergmans (1991) | Cheke & Dahl (1981) | Courts (1998) | Daniel et al. (2017) | Granek (2000) | Ibouroi, Cheha, Arnal et al. (2018) | Ibouroi, Cheha, Astruc et al. (2018) | O'Brien et al. (2009) | Sewall (2004) | Sewall, Granek et al. (2007) | Sewall, Young et al. (2016) | Smith & Leslie (2006) | Szekely et al. (2015) | Trewhella et al. (2001)		174. Livingstone’s Flying Fox Pteropus livingstonii French: Roussette de Livingstone / German: Livingstone-Flughund / Spanish: Zorro volador de Livingstone Other common names: Comoro Flying Fox , Livingstone's Fruit Bat Taxonomy. Pteropus livingstonii J. E. Gray, 1866 , “Island of Johanna [= Anjouan Island ], Comoro group [= Comoros ].” Pteropus livingstonii is in the livingstonii species group. Monotypic. Distribution. Comoro Is ( Anjouan and Mohéli ). Descriptive notes. Head-body 340 mm (tailless), ear 30-31 mm , hindfoot 5H 1- 53 mm , forearm 161-172 mm ; weight 576- 3 g (males) and 601- 5 g (females). Livingstone’s Flying Fox is the largest species of western Indian Ocean Pteropus . Head 1s fox-like; muzzle is long and black. Eyes are relatively large, with orange to scarlet-red irises; ears are large, protruding, and semicircular above; inner margin is convex; outer margin is straight or slightly convex; and exposed concha is smooth, without ribs or grooves. Males and females are alike, generally blackish brown on head and body. Back is dark to blackish brown, with dark russet tinge, and speckled with golden or tawny hairs; tawny hairs are also variably found in mantle, lower back, and flanks; tawny-tipped hairs in mantle form a band c. 6 cm wide and 10-15 mm long anteroposteriorly, with tufts of similarly colored fur on each shoulder. Dense pelage is present on upper inside of femur and uropatagium,; upper side of tibia is naked. Belly and flanks are similarly colored as back. Genital area can be surrounded by ring of deep reddish brown hair, extending laterally as a short stripe. Wings are dark, often jet-black, shiny, and hairless; index claw is present; tail membrane is poorly developed. Skull is typical pteropine, with very long rostrum; orbit is often closed by postorbital process reaching zygomatic arch; and crests and mandible are strong. Dentition is typical pteropine, with relatively small canines grooved in their outer surface; P' is lost in most adults; and cheekteeth are relatively simple but robust. Habitat. Undisturbed moist forests and, to a lesser extent, degraded forests at elevations above 300 m on Anjouan and 40 m on Mohéli . Roosts of Livingstone’s Flying Foxes are predominantly found in Ficus exasperata and FE lutea ( Moraceae ); Gyrostipula comorensis ( Rubiaceae ); and especially Gambeya spp. ( Sapotaceae ) and Nuxia pseudodentata ( Stilbaceae ). Roosts are in steep-sided (30-50°) valleys with south-east facing slopes, suggesting that they are sensitive to wind and sun. Food and Feeding. Livingstone’s Flying Fox is primarily frugivorous, eating fruits of native trees but also nectar and leaves. Ficus antandronarumis preferred during wet season and F lutea during dry season; other foods come from at least 14 genera in 13 families. Flowers of Gyrostipula and Neonauclea ( Rubiaceae ) and flowers, nectar, and to a lesser extent unripe fruit of Ceiba ( Malvaceae ) are eaten. Captive bats caught and ate insects. Breeding. Captive female Livingstone’s Flying Foxesstart to reproduce at 3-4 years old. After gestation of c.152 days, young are generally born in first part of September and weigh c. 137 g . Females with young were also observed in July-August, October, and January. Litter size is one. Activity patterns. Livingstone’s Flying Foxes forage at night. During the day, they remain in roosts and bask and stretch their wings in the sun to warm up, or urinate on their wings while flapping them to cool down. They take advantage of afternoon thermals to soar long distances. Peak activity is between 22:00 h and 02:00 h, and it declines to ¢.04:00 h. They start urinating immediately after taking flight. Movements, Home range and Social organization. Livingstone’s Flying Foxes form colonies of 15-150 individuals, exceptionally 250-349 during rainy seasons. Intra-island movements between roosts are common; inter-island movements are unknown. A dominant male has a harem of adult females and their female offspring. Status and Conservation. CITES Appendix II. Classified as Critically Endangered on The IUCN Red List. Livingstone’s Flying Fox has a small distribution ( 1856 km ? overall but probably 65-188 km *) and suspected significant population decline (current population is ¢.1300 animals: 940 on Anjouan and 360 on Mohéli ) from habitat loss due to logging for firewood, construction, and conversion to agricultural use. Recent disappearance of lowland forests has forced individuals higher on slopes up to 400 m on Anjouan and 200 m on Mohéli . It is not threatened by hunting. “The Conservation Action Plan for Livingstone’s Flying Fox” was set up and adopted by the government of Comoros , and a captive breeding program was established that led to a captive population of 59 individuals spread over four institutions as of 2014. Based on high levels of inter-island genetic differentiation, island populations probably need to be considered separate management units. Livingstone’s Flying Foxes do not occur in any protected area, but plans for appropriate reserves are being developed. Bibliography. Almeida et al. (2014), Andersen (1912b), Bergmans (1991), Cheke & Dahl (1981), Courts (1998), Daniel et al. (2017), Granek (2000), Ibouroi, Cheha, Arnal et al. (2018), Ibouroi, Cheha, Astruc et al. (2018), O'Brien et al. (2009), Sewall (2004), Sewall, Granek et al. (2007), Sewall, Young et al. (2016), Smith & Leslie (2006), Szekely et al. (2015), Trewhella et al. (2001).	Simmons, N.B. and A.L. Cirranello. 2022B. Bat Species of the World: A taxonomic and geographic database. Accessed on 10/11/2022.	Pteropodidae	Pteropus livingstonii	Pteropus		livingstonii	Gray	1866	0	Proc. Zool. Soc. Lond.	1867:06:00	Comoro Flying Fox	None.	Comoro Isls, Anjouan Isl.	Comoro Isls.	Appendix II	Critically Endangered	 livingstonii species group; see Almeida et al. (2014). Reviewed by Bergmans (1990). Misspelled livingstonei by Koopman (1993, 1994).	Mammal Diversity Database. (2023). Mammal Diversity Database (Version 1.11) [Data set]. Zenodo. https://doi.org/10.5281/zenodo.7830771 released 15 April 2023	Pteropus livingstonii	23	Livingstone's Flying Fox	Comoro Flying Fox|Livingstone's Fruit Bat	Theria	Placentalia	Boreoeutheria	Laurasiatheria	CHIROPTERA	PTEROPODIFORMES	NA	NA	PTEROPODOIDEA	PTEROPODIDAE	PTEROPODINAE	PTEROPODINI	Pteropus	NA	livingstonii	J. E. Gray	1866	0	Pteropus_livingstonii	Gray, J. E. (1866). A revision of the genera of Pteropine bats (Pteropidae), and the descriptions of some apparently undescribed species. Proceedings of the Zoological Society of London, 1866, 66.	https://www.biodiversitylibrary.org/item/90954#page/100/mode/1up	BM 1863.12.11.2		"Island of Johanna [= Anjouan Island], Comoro group [= Comoros]."			livingstonii J. E. Gray, 1866	NA	NA	Comoros	Africa	Afrotropic	CR	0	0	0	Pteropus_livingstonii	0	sciname match	Pteropus_livingstonii	0	IUCN. 2022. The IUCN Red List of Threatened Species. Version 2022-1. https://www.iucnredlist.org. Accessed on [28 September, 2022].	18732	Pteropus livingstonii	ANIMALIA	CHORDATA	MAMMALIA	CHIROPTERA	PTEROPODIDAE	Pteropus	livingstonii	Gray, 1866	P. livingstonei was used on the 2000 Red List. CITES also uses P. livingstonei , and livingstonei should be the grammatically correct spelling. T. Hutson (Chiroptera Red List Authority) notes that the correct spelling should be livingstonii (also by Simmons in Wilson and Reeder 2005) . ; Andersen’s 1912 emendation of the species name is now considered unjustified (Smith and Leslie 2006).	20000000	Pteropus livingstonii	Critically Endangered	A2c	2016	2016-05-31 00:00:00 UTC	3.1	English	Listed as Critically Endangered because of a serious population decline suspected from catastrophic habitat decline caused by cutting of trees for fuelwood and construction, and by conversion of all but the steepest upland areas to agricultural use; extensive declines in area of occupancy, extent of occurrence and quality of habitat. True population change in P. livingstonii is unknown and data on habitat change over time is limited. However, the species qualifies for CR under the A2c criterion because best estimates indicate habitat loss has exceeded 80% over the past three generations (estimated generation length: 8.1 years/generation), because remaining habitat is increasingly degraded and fragmented, and because declines in the extent and quality of habitat are continuing. P. livingstonii also meets the Endangered threshold under a second criterion, B1. The geographic range is small, such that the current extent of occurrence is estimated to be 1,856 km², less than the EN threshold of 5,000 km² under criterion B1. In addition, the habitat is severely fragmented, and there have been clear, continuing, observed declines in extent of occurrence, area of occupancy, habitat quality, and number of locations. Declines in number of mature individuals are also inferred from these declines in habitat. Therefore, this species would also be listed as EN under criterion B1ab(i, ii, iii, iv, v).	This species roosts in emergent trees, primarily on steep-sided valleys with south-east facing slopes, near ridge tops and in areas generally associated with natural vegetation (Granek 2002). Ficus esperata , Girostpula comoriensis , Gambeya spp., Ficus lutea and Nuxia pseudodentata are the five most commonly used species for roosting (Daniel et al.  in press.). Populations of this bat are largely confined to primary tropical moist forest (Mickleburgh et al.  1992), especially montane areas. Specifically, on Anjouan, all the major roosts are restricted to a narrow mid-altitudinal range (600-960 m asl) and are strongly associated with the presence of native and endemic trees, with most of the largest roosts located in dense canopy, old growth forest (Granek 2002, Sewall et al.  2007, Sewall et al.  2011, Daniel et al.  in press.). The bats feed primarily on fruit, but also to a lesser degree on nectar and leaves (Trewhella et al.  2001, Sewall 2002). Feeding is principally on native species but also includes non-native kapok (Ceiba pentandra ) (Trewhella et al.  2001, Sewall 2002); particularly important food sources are native Ficus lutea and Ficus antandronarum (Sewall 2002).	Pteropus livingstonii is threatened principally by the continuing degradation, fragmentation, and loss of its forest habitat (Sewall et al.  2007). Over the last few decades, the Comoros have undergone a sustained, rapid deforestation, resulting in the loss of nearly all its native forests. Data quality on rates of habitat loss is poor (FAO 2010), but best estimates are that during the past 20 years alone, the country has lost 75% of its remaining forests, the fastest rate of any country in the world (UNDP 2013). This habitat change is caused by cutting of trees for fuelwood and construction, and by conversion of all but the steepest upland areas to agricultural use (for subsistence agriculture and for export crops such as cloves) (Weightman 1987, Sewall et al.  2007, Sewall et al.  2011b, Doulton et al . 2015). With continuing habitat change, this species has retreated to higher elevations (above 450 m asl on Anjouan and above 250 m asl on Mohéli). Even above these elevations, forest habitat on both islands, and especially Anjouan, has undergone extensive and continuing habitat degradation, fragmentation, and loss (Sewall et al.  2007, 2011a; R. Young unpublished), leaving large areas increasingly unsuitable for this species. Human pressures on native forests are expected to continue, as the growth rate of the human population remains high, with an annual increase of 2.5% (UNDP 2013). Roost sites represent critical habitat for Pteropus livingstonii (Granek 2002, Sewall et al.  2007, Sewall et al.  2011b), yet tree felling has destroyed a number of roosts, leading to displacement of bats to less impacted areas, often at higher elevations (Trewhella et al.  2005, Sewall et al.  2011b). Of 23 roosts occupied in 2007 (Sewall et al.  2007), three roosts have been abandoned due to clear felling of forest in the past five years, and only one new but very small roost (15 bats) has been uncovered despite extensive searching effort (Daniel et al.  in prep.). This new roost was found in a patch of degraded forest at 1050 m, the first to be found at such a high elevation, and is believed to have been established very recently. On Anjouan, forest clearance, underplanting or significant soil erosion following deforestation upslope of the roost was found within 50 m of all but one of the 16 occupied roosts (Daniel et al.  in prep.). Further, the replacement of native forests with agricultural lands on Anjouan and Mohéli may render formerly inaccessible roost sites on Anjouan and Mohéli more accessible, possibly exposing roosts to increased levels of human disturbance. Finally, the loss of native forests has resulted in impermanence or complete drying of nearly all rivers on Anjouan and many on Mohéli (Louette et al.  2004, Fernandez Astudillo 2012). As P. livingstonii roosts are often associated with rivers and other humid environments, a factor which may be related to their thermal sensitivity (Granek 2002), this loss of rivers may additionally affect the quality of roosting habitat. There is no evidence that the species is hunted for food (Trewhella et al.  2005), a key factor affecting other fruit bat species (Mickleburgh et al.  1992). This may be due in part to cultural taboos among the majority of the population that surround consumption of fruit bats. However, the apparent lack of hunting may also be due in part to P. livingstonii ’s habit of roosting in inaccessible areas remote from towns; some limited hunting of the other two fruit bat species in the Comoros has been occasionally recorded (Sewall et al.  2003, B.J. Sewall, pers. obs.). It is unclear if the increased access to formerly inaccessible roost areas could expose the bats to any hunting pressure in the future, but any such hunting would likely first affect P. livingstonii ’s congener, P. seychellensis comorensis , which sometimes roosts and forages in visible locations in or near towns. The mean surface temperature of the Comoros is thought to have increased about 1° C since 1900 and is expected to increase by 1-3° C more by 2100 due to global climate change (IPCC 2013). It is unclear whether or how such changes will affect P. livingstonii , but since thermal characteristics are important components of roost suitability for this and other Pteropus species (Pierson and Rainey 1992, Granek 2002) predicted temperature increases could affect availability of suitable roosting habitat. P. livingstonii ’s small population is found within a restricted range solely on two small, adjacent islands. The species is therefore susceptible to single threatening processes, like cyclones, that could simultaneously or rapidly affect its entire range (Sewall et al.  2007). Further, the species may have naturally taken advantage of elevational differences in tree fruiting phenology to compensate for seasonal variation in fruit availability (Sewall 2002). Thus, the loss of large areas of foraging habitat including all the lower parts of its elevational range, could result in seasonally restricted food availability.	Repeated simultaneous surveys of all 23 known roosts during 1998-2006 typically recorded about 1,200 bats (Sewall et al.    2007); many of these roosts were located as a result of the implementation of a national monitoring program (Trewhella et al.    2005). Surveys have been conducted only sporadically since 2007, so it is unclear whether populations have changed recently. However, a survey conducted with sequential visits to all previously-known roosts in 2011 and 2012 estimated 1,300 bats across 22 roosts (Anjouan: 940 bats at 16 roosts, Mohéli: 360 bats at 6 roosts) (Daniel et al.    in press.). Colony size typically ranges from 15 to 150 individuals (Sewall et al.    2007). The largest known roosts have sometimes reached about 250 individuals during the rainy season, but not all of these are mature individuals (Sewall et al.    2011 a,b). Generation length for the wild population of P. livingstonii is unknown, but several demographic parameters can be derived from the studbook for captive bats of this species (Glendewar 2014). Bats of this species can be long-lived in captivity (e.g., one founder captured as an adult in 1992 is still alive and has now lived 22 years in captivity). Females do not reproduce during the first few years of life in captivity (minimum dam age at first reproduction = 3.4 years, mean dam age at first reproduction = 5.9 years), and the average age of females giving birth is elevated (of all dams born in captivity that have since died or are &gt;10 years of age, mean age when giving birth = 8.1 years). Thus, based on studbook data (Glendewar 2014), this last figure of 8.1 years represents our best estimate of the generation length of the captive population. This estimate may be conservative as most (75%) of these females are still alive and at least some presumably will reproduce again at a later age. It is unclear to what extent the generation length of the captive population correlates with the generation length of the wild population of P. livingstonii . However, the generation length of captive P. livingstonii correlates well with the few published generation length values for wild populations of other Pteropus species. In a wild population of P. poliocephalus , generation length was estimated conservatively as 7.4 years (Tidemann and Nelson 2011), though the true generation length could be much higher. Further, a generation length of 5.0 years was estimated for a rapidly declining wild population of P. conspicillatus (Fox et al.    2008); however, our calculations using these same data suggest that in a stable population, generation length for P. conspicillatus would be about 7.7 years. Thus, generation length values for the captive population of P. livingstonii correlate well with the only published generation length values for wild Pteropus . Further, an estimate derived from taxonomically-adjusted allometric equations for mammals suggests generation length for P. livingstonii is 8.1 years (Pacifici et al.    2013). Thus, the captive estimate appears in line with estimates for wild populations, and we conservatively estimate the generation length in the wild population of P. livingstonii to be 8.1 years. True changes in the population size of P. livingstonii are currently unknown, but are likely to be linked to changes in forest and underplanted forest habitat. Our estimates, derived from the little available data on forest loss rates (FAO 2010, UNDP 2013), suggest that habitat decline over the most recent three-generation (or 24.3-year) period was 83%. Thus, our best estimate of loss of P. livingstonii habitat exceeds the threshold under the A2 criterion for CR status (≥80% loss of population, as suspected by habitat change, over a three-generation period, where habitat loss is ongoing).	Decreasing	P. livingstonii is endemic to the Union of the Comoros, where it is only found on the islands of Anjouan and Mohéli. The species roosts and forages primarily in native forest and underplanted forest, and avoids areas heavily affected by human disturbance. On Anjouan the species avoids all lower elevations on the island (lowest record of feeding is 300 m asl), but on Mohéli some limited forest patches extend to near sea level (lowest record of feeding is 40 m asl). However, nearly all lower-elevation native forests have been lost, and in the past five years, there have been no records of this species below approximately 450 m on Anjouan or below 200 m on Mohéli (Daniel et al.     in press.). The Extent of Occurrence (calculated as the minimum convex polygon around area above minimum observed altitude above sea level on each of the two islands) is estimated as 1,856 km². ; This includes unsuitable areas of land and ocean located between patches of suitable habitat. The total land area of Anjouan and Mohéli is a combined 635 km² (Louette et al.     2004). However, not all this area is occupied by P. livingstonii . The species is found within a combined area on the two islands ranging from 99.1 km² (minimum convex polygons around all known currently occupied long-term roost sites: Anjouan = 90.2 km² Mohéli = 8.9 km²) to 462.5 km² (minimum convex polygons around all potential but unconfirmed foraging and roosting areas in native forest, degraded forest and agroforestry habitats above the lowest elevation of recent sightings on each island [450m-1050m asl on Anjouan, and above 200m asl on Moheli]: Anjouan = 326.9 km², Mohéli = 135.6 km²) (Daniel et al.     in press.). These two figures represent the possible range of this species on the two islands it inhabits. This represents a substantial decline over the past five years. A land cover classification conducted in 2012 using high resolution satellite imagery and exenstive ground-truthing (ECDD, BCSF and Durrell, 2013) found the combined area of native forest (i.e. forest with little trace of human impact and a closed canopy) on both islands was 54.7 km² (29.6 km² in Anjouan; 25.1 km² in Moheli). The combined area of native forest and degraded forest (i.e. forest either underplanted with crops, or with localised logging or clearance for fuelwood) on both islands was 143.1 km².  Within the smallest total range on both islands (the area around all known currently occupied long-term roost sites), the Area of Occupancy or surface area of potentially occupied habitat (i.e. native and degraded forest and agroforestry habitats that could be used for either roosting and foraging) is estimated to be 65.0 km² (Anjouan = 56.8 km², Moheli = 8.2 km²). Within the largest total range on both islands, the equivalent estimate for Area of Occupancy is 188.4 km² (Anojuan = 114.07 km², Moheli = 74.34 km²). Thus, the estimated total Area of Occupancy for this species is 65.0 - 188.4 km².		Terrestrial	A detailed conservation plan for the species, the Conservation Action Plan for Livingstone’s Flying Fox (Sewall et al.  2007) was developed by the non-governmental organizations Action Comores Anjouan and Action Comores International, and a consortium of other conservation groups, through a participatory process involving a broad range of stakeholders. The plan identifies a conservation strategy with several key elements, including habitat protection, forest management, environmental education, population monitoring, ecological research, ex-situ breeding, and conservation partnerships. This conservation plan was adopted by the government of the Union of the Comoros as the national conservation strategy for this species and has served as a guide for conservation action. A long-term comprehensive citizen science program involving Comorian villagers in population monitoring for this species was led by Action Comores Anjouan and Action Comores International between 1992 and 2006. Population surveys and habitat evaluation around roost sites was conducted by Engagement Communautaire pour le Développement Durable of the Bristol Zoological Society and Durrell Wildlife Conservation Trust between 2009 and 2012, and by Dahari more recently. A national environmental education programme was implemented by Action Comores Anjouan and Action Comores International, and it was successful in raising local and international awareness of this species and the threats it faces, in improving training of local personnel, and in increasing local participation in conservation and science programs (Trewhella et al.  2005). A small community-based ecotourism program for this species has been implemented on Mohéli by Projet Conservation de la Biodiversité et Développement Durable and local communities and is maintained by the local community in Ouallah 2 (Sewall et al.  2011b, Doulton et al.  2015). Dahari is implementing integrated landscape management around the southern forest block of Anjouan; this involves sustainable land management at the roosting and foraging habitat of this species and a pilot program of payment for ecosystem services both within the Moya Forest area (Doulton et al.  2015). ; Research by Dahari and partners into feeding ecology and the genetics of roost-site populations is also planned.  There is an active captive-breeding programme underway for this species, initiated by Durrell Wildlife Conservation Trust in 1992. The program began with 17 founder individuals (7 females and 10 males) captured from the wild on Anjouan during 1992-1995 (Clark et al.  1997). The captive population of P. livingstonii subsequently expanded at a slower rate than other fruit bat species in captive breeding programs (e.g., P. rodricensis ), at least initially (O’Brien 2011). However, by 2014, the P. livingstonii captive population had reached 59 individuals (22 females and 37 males) housed at four institutions (Durrell Wildlife Conservation Trust [Jersey, U.K.]; Bristol, Clifton, and West of England Zoological Society [Bristol, U.K.]; North of England Zoological Society [Chester, U.K.]; and Lisieux Cerza [Lisieux, France])(Glenewar 2014). This species receives the highest level of legal protection available within the Union of the Comoros. It is listed as an ‘integrally-protected species’ (list 1 of RFIC 2001), which prohibits the capture or detention of P. livingstonii individuals without a permit. This law also expressly prohibits the killing of flying fox individuals; transport, purchase, sale, export or re-export of live or dead flying fox individuals or body parts; all disruption during the period of reproduction and raising of young; and the destruction of roosts (RFIC 2001). The Union of the Comoros also ratified the Convention on Biological Diversity in 1994, and in response has developed a National Biodiversity Conservation Strategy (Roby and Dossar 2000). This strategy highlights the importance of, threats to, and conservation recommendations for fruit bats of the Union of the Comoros (Sewall and Granek 2000). P. livingstonii is also listed on Appendix II of the Convention on International Trade in Endangered Species of Wild Fauna and Flora, or CITES (UN 1973), which prevents international trade in specimens of this species without a permit. In practice, however, enforcement activities within the Comoros for these laws and treaties have been very limited in scope (Sewall et al.  2007). P. livingstonii also does not occur in any protected areas. However, critical roosting habitat at seven key roost sites that together harboured more than half the population was identified during the development of the conservation action plan (Sewall et al.  2007). Habitat conservation of these roost sites was identified as a key goal in this plan. As part of its implementation, Action Comores Anjouan and the Comoros Forest Reserves Project conducted a conservation assessment to identify the two highest-priority sites (roost sites Yiméré on Anjouan and Hassera-Ndrengé on Mohéli), where the establishment of protected areas for roost habitat would be both highly beneficial to biodiversity conservation and highly feasible (Sewall et al.  2011a). These two groups also completed conservation planning for protection of critical roosting habitat at all seven of these critical roost sites (Sewall et al.  2011b). Implementation efforts by Action Comores Anjouan and partners to conserve habitat around critical roosting habitat on Anjouan and Mohéli are under development. A project to establish larger reserves to protect rainforest and cloud forest on both Anjouan and Mohéli, which would include roosting and foraging habitat for this species, has been proposed by the United Nations Development Programme, the Global Environmental Facility, and the Comorian government. The success of habitat conservation and restoration efforts will be particularly critical to the long-term prospects for this species.	Afrotropical		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	Pteropus		livingstonii	Gray	1866	0	Proc. Zool. Soc. Lond.	1867:06:00	Comoro Flying Fox	None.	Comoro Isls, Anjouan Isl.	Comoro Isls.	Appendix II	Critically Endangered	 livingstonii species group; see Almeida et al. (2014). Reviewed by Bergmans (1990). Misspelled livingstonei by Koopman (1993, 1994).	Pteropus livingstonii	1004470	23	Livingstone's Flying Fox	Comoro Flying Fox|Livingstone's Fruit Bat	Theria	Placentalia	Boreoeutheria	Laurasiatheria	CHIROPTERA	PTEROPODIFORMES	NA	NA	PTEROPODOIDEA	Pteropodidae	PTEROPODINAE	PTEROPODINI	Pteropus	NA	livingstonii	J. E. Gray	1866	0	Pteropus_livingstonii	Gray, J. E. (1866). A revision of the genera of Pteropine bats (Pteropidae), and the descriptions of some apparently undescribed species. Proceedings of the Zoological Society of London, 1866, 66.	https://www.biodiversitylibrary.org/item/90954#page/100/mode/1up	BM 1863.12.11.2		"Island of Johanna [= Anjouan Island], Comoro group [= Comoros]."			livingstonii J. E. Gray, 1866	NA	NA				Comoros	Africa	Afrotropic	CR	0	0	0	Pteropus_livingstonii	0	sciname match	Pteropus_livingstonii	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	Pteropus_livingstonii	1004470	23	Livingstone's Flying Fox	Comoro Flying Fox|Livingstone's Fruit Bat	Theria	Placentalia	Boreoeutheria	Laurasiatheria	Chiroptera	Yinpterochiroptera	NA	NA	Pteropodoidea	Pteropodidae	Pteropodinae	Pteropodini	Pteropus	NA	livingstonii	J. E. Gray	0	Pteropus livingstonii	Gray, J.E. 1866-05. A revision of the genera of pteropine bats (Pteropidee), and the descriptions of some apparently undescribed species. Proceedings of the Zoological Society of London 1866(1):62-67.	https://www.biodiversitylibrary.org/page/28627502	BMNH:Mamm:1863.12.11.2	holotype	https://data.nhm.ac.uk/object/9ef4994a-cd65-4ad9-a996-368dd0e6a8b7	"Island of Johanna [= Anjouan Island], Comoro group [= Comoros]."			NA	NA				Comoros	Africa	Afrotropic	CR	0	0	0	Pteropus_livingstonii	0	sciname match	Pteropus_livingstonii	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	Pteropus		livingstonii	Gray	1866	0	Proc. Zool. Soc. Lond.	1867:06:00	Comoro Flying Fox	None.	Comoro Isls, Anjouan Isl.	Comoro Isls.	<a href='https://cites.org/eng/app/appendices.php' target='_blank'>Appendix II</a>	<a href='https://www.iucnredlist.org/species/18732/22081502/' target='_blank'>Critically Endangered</a>	livingstoniispecies group; see Almeida et al. (2014). Reviewed by Bergmans (1990). Misspelled livingstonei by Koopman (1993, 1994).		Mammal Diversity Database. (2025). Mammal Diversity Database (Version 2.2) [Data set]. Zenodo. https://doi.org/10.5281/zenodo.15007505	NA	Pteropus livingstonii; Pteropus livingstonii; Pteropus livingstonii; Pteropus livingstonii; Pteropus livingstonii; Pteropus livingstonii; livingstonii; Roussette de Livingstone; Livingstone-Flughund; Zorro volador de Livingstone; Comoro Flying Fox; Livingstone's Fruit Bat; Livingstone's Flying Fox; Comoro Flying Fox; Livingstone's Fruit Bat; Comoro Flying Fox; Comoro Flying Fox; P. livingstonii