Pteropus vampyrus, Linnaeus, 1758

Don E. Wilson & Russell A. Mittermeier, 2019, Pteropodidae, Handbook of the Mammals of the World – Volume 9 Bats, Barcelona: Lynx Edicions, pp. 16-162 : 158-159

182.

Large Flying Fox

Pteropus vampyrus

French: Roussette de Malaisie / German: Kalong-Flughund / Spanish: Zorro volador de cuello rojizo

Other common names: Andersen's Flying Fox (intermedius)

Taxonomy. Vespertilio vampyrus Linnaeus, 1758 ,

“Asia.” Restricted by K. Andersen in 1912 to “ Java,” Indonesia.

Pteropus vampyrus is in the vampyrus species group. Pteropus intermedius is considered conspecific but requires full taxonomic review. Six subspecies recognized.

Subspecies and Distribution.

P. v. vampyrus Linnaeus, 1758 — Java.

P.v. edulis E. Geoffroy Saint-Hilaire, 1810 — Lesser Sunda Is (Timor).

P. v. lanensis Mearns, 1905 — Philippines.

P.v. natunae K. Andersen, 1908 — Borneo and Natuna Is.

P. v. pluton Temminck, 1853 — W Lesser Sunda Is.

P. v. sumatrensis Ludeking, 1862 — S mainland SE Asia and Malay Peninsula, Sumatra, Lingga Archipelago, Bangka, and S Mentawai Is (Sipora, North Pagai, and South Pagai).

Chinese record from Shaanxi Province likely alien.

Descriptive notes. Head-body 225-290 mm (tailless), ear 35-45 mm, hindfoot 52- 68 mm, forearm 175-230 mm; weight 0.7-1.4 kg. Greatest lengths of skulls are 78- 90 mm and tibias 84-105 mm. Forearm lengths and body mass varies among subspecies and geographical regions: smallest subspecies is natunae (forearm 180-196 mm) and largest is pluton (200-230 mm). There are minor variations in color of subspecies. The Large Flying Fox generally has dark brown to blackish body, with distinct mantle varying from tawny brown, reddish brown, and orange yellow to blond. Head can be reddish brown to brown to blackish. Dorsal side of body is generally dark brown to blackish, with some flecks of gray. Males are generally larger than females. Ears are long, with pointed tips. Wing membranes are dark brown to blackish, with mildly rounded tips and fur only near body. Baculum is similar to that in other pteropodids, saddle-shaped in lateral view but without an apical prominence. Proximal prongs on baculum are separated instead of fused as in other pteropodids. Baculum is large (4-5-8- 2 mm long). Skull is typical pteropine and generally large and robust, with large orbits and wide zygomatic arches. Upper incisors are separated from one another, and I' is slightly larger than I°. C! is long and pointed, with distinct cingulum but no secondary cusp, grooved labially and with smaller groove lingually. C, is smaller and lack grooves. Premolars and molars are broad and short, with no prominent cusps. P' is very small and lost in adults. Chromosomal complement has 2n=38 and EN = 72,

Habitat. Primary and secondary forests, mangrove forests, and swamps in coastal areas and on offshore islands from sea level and generally below elevations of ¢. 500 m (but up to 1370 m). In the Philippines, Indonesia, and Malaysia, the Large Flying Fox prefers to roost in areas with relatively pristine mangroves. It can be found in some mixed landscapes of natural, modified, and intensely farmed plots. In highly modified landscapes, it can be found in relatively undisturbed areas, such as a temple forests, botanical gardens, atop steep cliffs, or at other culturally significant sites such as burial grounds. In the Philippines, it prefers natural forest to human-modified habitats. It roosts in a variety of trees, often in riparian zones; trees become defoliated, suggesting high roost fidelity.

Food and Feeding. The Large Flying Fox has a broad diet that includes fruits, flowers, leaves (rarely), pollen, and nectar from natural and agroforestry habitats. Various species of Ficus (Moraceae) are the most significant parts of diets throughoutits distribution. It eats pollen, nectar, and flowers of coconut ( Cocos nucifera, Arecaceae ), petai ( Parkia speciosa and P. javanica, Fabaceae ), and durian ( Durio zibethinus, Malvaceae ). It defends flower resources from conspecifics and feeds on flowers of Parkia , Erythrina (Fabaceae) , Octomeles (Tetramelaceae) , Pterocymbium (Malvaceae) , and Syzygium (Myrtaceae) . It has been recorded more frequently in orchards during fruiting in recent decades.

Breeding. Female Large Flying Foxes typically give birth synchronously to one young/ year. Mating and courtship occur in November in Cambodia and May in Indonesia. Birthing peaks vary geographically: March-April ( Thailand and presumably Malaysia), April ( Cambodia), and April-June ( Philippines). Births usually coincide with fruiting peaks. In captivity at Lubee Bat Conservancy (Gainesville, Florida), births peak in May-June. Young are carried by their mothers for the first few days after birth and then are left at roosts while mothers forage. Young nurse for 2-3 months.

Activity patterns. The Large Flying Fox is nocturnal. It leaves the roost around dusk (typically ¢.18:30-19:30 h) in a scattered stream high above the canopy (above 100 m) for nightly foraging. At foraging sites, the colony may form smaller feeding groups (fewer than 50 individuals). In Malaysia, two feeding periods have been recorded (19:30-21:30 h and 22:30-23:30 h), with individuals resting in trees between feeding periods. Individuals return to roosts around dawn, exhibiting territorial vocalizations and fighting that promotes some level of spacing among individuals. During the day, individuals rest and exhibit typical pteropodine activity, such as wing flapping and occasional conspecific territorial interactions. Some individuals take short flights around the roost during the day.

Movements, Home range and Social organization. The Large Flying Fox is generally gregarious and roosts colonially in tall trees. It is tolerant of some habitat disturbance but prefers undisturbed habitat. Currently, the largest known colony is in Subic Bay on Luzon, Philippines (c.20,000 individuals). It might roost with otherflying foxes in large colonies (up to 52,000 individuals). In Malaysia, severe declines in abundance have been recorded, and no roosts are known to have more than 1500 individuals. Foraging range of up to 50 km /night occurs in some parts of its distribution, but in others, they can be as short as 0-4- 12 km. Home range with 90% occupancy of the Large Flying Fox can be as large as 128,000 km * (i.e. almost 13 million ha) on the Malay Peninsula. Inter-island movement likely occurs at higher altitudes. It switches roosts according to food availability. In Sarawak, it has a continuous movement phase (roost sites appear to move daily) and a restricted movement phase (roost sites confined to narrow area).

Status and Conservation. CITES Appendix II. Classified as Near Threatened on The IUCN Red List. The Large Flying Fox has experienced significant declines throughout its distribution due to overhunting and ongoing degradation of habitat. Nevertheless, rate of decline is less than 30%, but additional decline will likely qualify it as Vulnerable. Previously, the Large Flying Fox occurred in many large colonies throughout its distribution; colonies are now much rarer due to overhunting and habitat loss. Hunted individuals are normally sold only at a local level, making detection of illegal take difficult. Resident populations occur in urban parks or forests where they receive informal protection. Hunting of flying foxes in the Philippines is illegal, except for use by a few indigenous groups, but hunting is unregulated and therefore continues even now.

Bibliography. Andersen (1912b), Bates, Francis, Gumal, Bumrungsri, Walston et al. (2008), Corbet & Hill (1992), Corlett (1992), Epstein et al. (2009), Gumal (2004), Heaney et al. (2016), Ingle & Heaney (1992), Kunz & Jones (2000), Mildenstein et al. (2005), Mohd-Azlan et al. (2001), Patagwang et al. (2014), Ravon et al. (2014), Simmons (2005), Stier & Mildenstein (2005), Tsang et al. (2018), Yap (2016), Zhang Jinshuo et al. (2010).

Figures

174. Livingstone’s Flying Fox (Pteropus livingstonii), 175. Pemba Flying Fox (Pleropus voeltzkowi), 176. Ryukyu Flying Fox (Pleropus dasymallus), 177. Bonin Flying Fox (Pteropus pselaphon), 178. Little Golden-mantled Flying Fox (Pleropus pumilus), 179. Lyle’s Flying Fox (Pteropus lylei), 180. Indian Flying Fox (Pleropus medius), 181. Rodrigues Flying Fox (Pteropus rodricensus), 182. Large Flying Fox (Pteropus vampyrus), 183. Aldabra Flying Fox (Pleropus aldabrensis), 184. Malagasy Flying Fox (Pteropus rufus), 185. Seychelles Flying Fox (Pleropus seychellensis), 186. Greater Mascarene Flying Fox (Pteropus niger), 187. Aru Flying Fox (Pteropus aruensis), 188. Kai Flying Fox (Pleropus keyensis), 189. Black-bearded Flying Fox (Pleropus melanopogon), 190. South Moluccan Flying Fox (Pteropus chrysoproctus), 191. North Moluccan Flying Fox (Pteropus caniceps)

Subspecies and Distribution. P. v. vampyrus Linnaeus, 1758 — Java. P.v. edulis E. Geoffroy Saint-Hilaire, 1810 — Lesser Sunda Is (Timor). P. v. lanensis Mearns, 1905 — Philippines . P.v. natunae K. Andersen, 1908 — Borneo and Natuna Is. P. v. pluton Temminck, 1853 — W Lesser Sunda Is. P. v. sumatrensis Ludeking, 1862 — S mainland SE Asia and Malay Peninsula, Sumatra, Lingga Archipelago, Bangka, and S Mentawai Is (Sipora, North Pagai, and South Pagai). Chinese record from Shaanxi Province likely alien.