Pteropus vampyrus (*)
Superregnum: Eukaryota
Cladus: Unikonta
Cladus: Opisthokonta
Cladus: Holozoa
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Superphylum: Deuterostomia
Phylum: Chordata
Subphylum: Vertebrata
Infraphylum: Gnathostomata
Megaclassis: Osteichthyes
Cladus: Sarcopterygii
Cladus: Rhipidistia
Cladus: Tetrapodomorpha
Cladus: Eotetrapodiformes
Cladus: Elpistostegalia
Superclassis: Tetrapoda
Cladus: Reptiliomorpha
Cladus: Amniota
Cladus: Synapsida
Cladus: Eupelycosauria
Cladus: Sphenacodontia
Cladus: Sphenacodontoidea
Cladus: Therapsida
Cladus: Theriodontia
Cladus: Cynodontia
Cladus: Eucynodontia
Cladus: Probainognathia
Cladus: Prozostrodontia
Cladus: Mammaliaformes
Classis: Mammalia
Subclassis: Trechnotheria
Infraclassis: Zatheria
Supercohors: Theria
Cohors: Eutheria
Infraclassis: Placentalia
Cladus: Boreoeutheria
Superordo: Laurasiatheria
Cladus: Scrotifera
Ordo: Chiroptera
Subordo: Yinpterochiroptera
Familia: Pteropodidae
Subfamilia: Pteropodinae
Tribus: Pteropodini
Genus: Pteropus
Species: P. vampyrus
Subspecies (6): P. v. edulis – P. v. lanensis – P. v. natunae – P. v. pluton – P. v. sumatrensis – P. v. vampyrus
Name
Pteropus vampyrus (Linnaeus, 1758)
Type material: unknown.
Type locality: “Asia”, restricted to “Java”, Indonesia, by Andersen (1912).
Combinations
Vespertilio vampyrus Linnaeus, 1758: 31 [original combination]
Pteropus vampyrus: Thomas, 1892: 316 [subsequent combination]
Native distribution areas
Vietnam, Burma, Malay Peninsula, Borneo, Philippines, Sumatra, Java, and Lesser Sunda Islands, adjacent small islands including Anak Krakatau.
References
Primary references
Linnaeus, C. 1758. Systema Naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Editio Decima, Reformata. Tomus I. Holmiæ (Stockholm): impensis direct. Laurentii Salvii. 824 pp. DOI: 10.5962/bhl.title.542 BHL Reference page.
Links
Bates, P., Francis, C., Gumal, M., Bumrungsri, S., Walston, J., Heaney, L. & Mildenstein, T. 2008. IUCN: Pteropus vampyrus (Near Threatened). The IUCN Red List of Threatened Species 2008: e.T18766A8593657. DOI: 10.2305/IUCN.UK.2008.RLTS.T18766A8593657.en.
Vernacular names
Boarisch: Kalong-Fliaghund
čeština: Kaloň malajský
Deutsch: Kalong, Kalong-Flughund
English: Large Flying Fox
español: Zorro volador de cuello rojo
eesti: Kalong
suomi: Isolentäväkoira
français: Roussette de Malaisie
magyar: Óriás repülőkutya
Bahasa Melayu: Keluang Besar
Nederlands: Kalong
polski: Rudawka malajska
русский: Гигантская летучая лисица
svenska: Kalong
ไทย: ค้างคาวแม่ไก่ป่าฝน
українська: Калонг
Tiếng Việt: Dơi quạ lớn
The large flying fox (Pteropus vampyrus, formerly Pteropus giganteus), also known as the greater flying fox, Malayan flying fox, Malaysian flying fox, large fruit bat, kalang, or kalong, is a southeast Asian species of megabat in the family Pteropodidae.[3] Despite its scientific name, it feeds exclusively on fruits, nectar, and flowers, like the other flying foxes of the genus Pteropus. It is noted for being one of the largest bats.[4] As with nearly all other Old World fruit bats, it lacks the ability to echolocate but compensates for it with well-developed eyesight.[5]
Taxonomy
P. livingstonii
P. voeltzkowi
"vampyrus" group
P. dasymallus
P. pumilus
P. rodricensis
P. vampyrus
P. lylei
P. medius
P. aldabrensis
P. rufus
P. seychellensis*
P. niger*
P. seychellensis*
P. niger*
P. pselaphon
Location of the large flying fox (Pteropus vampyrus) relative to some other members of its genus[6]
The large flying fox was one of the many mammal species originally described by Carl Linnaeus in the landmark 1758 10th edition of his Systema Naturae, receiving the name Vespertilio vampyrus.[2] The holotype was collected on Java.[7]: 70 Its species name "vampyrus" is derived from Slavic "wampir" meaning "blood-sucking ghost or demon: vampire".[4] This name was chosen in reference to its "alleged blood-sucking habits",[7]: 87 although it is entirely vegetarian and largely frugivorous.[4]
Based on phylogenetic analysis using mitochondrial DNA, the closest relative of the large flying fox is the Rodrigues flying fox (Pteropus rodricensis).[6] Because the genus Pteropus is so speciose, it is further subdivided into species groups. The large flying fox is the namesake of the "vampyrus" group, which also includes the following species:[6]
Aldabra flying fox (Pteropus aldabrensis)
Ryukyu flying fox (Pteropus dasymallus)
Lyle's flying fox (Pteropus lylei)
Indian flying fox (Pteropus medius)
Mauritian flying fox (Pteropus niger)
Bonin flying fox (Pteropus pselaphon)
Little golden-mantled flying fox (Pteropus pumilus)
Rodrigues flying fox (Pteropus rodricensis)
Madagascan flying fox (Pteropus rufus)
Seychelles fruit bat (Pteropus seychellensis)
Description
Main articles: Megabat § Description, and Pteropus § Description
The large flying fox is among the largest species of bat.[4] It weighs 0.65–1.1 kg (1.4–2.4 lb) and has a wingspan of up to 1.5 m (4 ft 11 in).[8][9] Its head-body length is 27–32 cm (11–13 in).[10] Its forearm length is 180–220 mm (7.1–8.7 in).[4] As is common with most megabats, it has a fox-like face. It lacks a tail and has pointed ears. The hairs on much of its body are long and woolly, but are shorter and more erect on the upper back.[4] The mantle hairs tend to be the longest.[11] The color and texture of the coat differ between sexes and age classes.[12] Males tend to have slightly stiffer and thicker coats than females.[4] Immature individuals are almost all dull gray-brown.[8] Young have a dark-colored mantle that becomes lighter in males when they mature.[4] The head has hairs that range in color from mahogany-red and orange-ochreous to blackish. The ventral areas are brown or blackish, tinged with chocolate, gray or silver.[12] The mantle can vary from pale dirty-buff to orange-yellow, while the chest is usually dark-golden brown or dark russet.[4] The large flying fox has a large and robust skull. The dental formula is 2.1.3.22.1.3.3. It has a total of 34 teeth.[11][13] The large flying fox's wings are short and somewhat rounded at the tips. This allows them to fly slowly, but with great maneuverability.[4] The wing membranes are only haired near the body.
Biology and ecology
This species primarily feeds on flowers, nectar and fruit. When all three food items are available, flowers and nectar are preferred.[8][11][12] The pollen, nectar, and flower of coconut and durian trees, as well as the fruits of rambutan, fig and langsat trees, are consumed. Flying foxes will also eat mangoes and bananas.[12][14][15] With fruit, the flying fox prefers the pulp, and slices open the rind to get it.[15] With durian tree flowers, the flying fox can lick up the nectar without doing apparent damage to the flower.[4] The large flying fox is a host of the Acanthocephalan intestinal parasite Moniliformis convolutus.[16]
Behavior and life history
Colonies of large flying foxes fly in a scattered stream.[11] They may fly up to 50 km (31 mi) to their feeding grounds in one night. Vocalizations are not made during flight.[11] Large flocks fuse into family or feeding groups upon arrival at feeding grounds.[4] Flying foxes may circle a fruit tree before landing, and usually land on the tips of branches in an upright position, then fall into a head-down position from which they feed.[4] Feeding aggregations tend to be very noisy.[17]
Flowering trees form the basis of territories in this species. Territorial behavior includes growling and the spreading of wings.[17] During antagonistic behavior, individuals maintain spacing with wrists/thumbs sparring, bites, and loud vocalizations.[12] When moving to a suitable resting place after landing, an individual may fight with conspecifics along the way.[12] A roosting flying fox is positioned upside down with its wings wrapped up.[18] When it gets too warm, a flying fox fans itself with its wings.[12] Roosting bats are restless until midmorning.
Female large flying fox gestations are at their highest between November and January in Peninsular Malaysia, but some births occur in other months.[19] In Thailand, gestation may take place during the same period with young being born in March or early April.[11][19] Females apparently give birth during April and May in the Philippines,[18] and usually give birth to only one young.[11] For the first days, the mothers carry their young, but leave them at the roost when they go on their foraging trips.[4] The young are weaned by two to three months.[11]
Range and habitat
The large flying fox ranges from Malay Peninsula, to the Philippines in the east and Indonesian Archipelago of Sumatra, Java, Borneo and Timor in the south.[20] In certain areas, the bat prefers coastal regions, but it can also be found at elevations up to 1,370 m (4,490 ft).[21]
Flying foxes inhabit primary forest, mangrove forest, coconut groves, mixed fruit orchards, and a number of other habitats.[19] During the day, trees in mangrove forests and coconut groves may be used as roosts.[12] In Malaysia, flying foxes prefer lowland habitats below 365 m.[14] In Borneo, they inhabit the coastal areas, but move to nearby islands to feed on fruit.[8] Flying foxes roost in the thousands (maximum). One colony was recorded numbering around 2,000 individuals in a mangrove forest in Timor[12] and colonies of 10,000–20,000 have also been reported.[4] In general, mangrove roosts have lower numbers of resting bats compared to lowland roost sites, which could mean mangrove forests are only used temporarily.[14]
Relationship to humans
The large flying fox is hunted for bushmeat. In Peninsular Malaysia, 1,756 hunting licenses were issued for the large flying fox from 2002–2006. In total, these hunting licenses permitted the hunting of 87,800 large flying foxes, or about 22,000 each year. Based on population modeling, the loss of the estimated 22,000 large flying foxes annually is unlikely to be sustainable. A 2009 study predicted extinction of the Peninsular Malaysian population within 6–81 years if 22,000 individuals are lost to hunting each year.[22]
The large flying fox is a natural reservoir of the Nipah virus. It is generally considered as the reservoir that led to the 1998 Malaysian outbreak, which was the first emergence of the disease in humans and pigs.[23] In a study of seventeen large flying foxes, Nipah virus was only isolated from one individual, which was at the time of capture. However, in maintaining the bats in quarantine for one year, researchers found that the bat was negative for antibodies against Nipah virus for the first eleven months, but was then seropositive once more. Two other bats—from which the Nipah virus was never detected—also registered as seropositive at points within the year. This suggested that the Nipah virus can recrudesce in the large flying fox, or maintain itself after periods of remission. The virus also recrudesces in humans, with humans becoming fatally ill with the disease up to four years after first exposure.[24]
Conservation
As of 2022, the large flying fox is evaluated as an endangered species by the International Union for Conservation of Nature. It meets the criteria for this designation because it is likely experiencing significant population decline. The bushmeat trade is resulting in unsustainable harvest of this species. Additionally, it is experiencing habitat loss through deforestation.[1] The large flying fox is on Appendix II of CITES, which restricts international trade.[25]
One threat to the large flying fox is habitat destruction.[19] Flying foxes are sometimes hunted for food, and the controls on hunting seem to be unenforceable.[4] In some areas, farmers consider them pests as they sometimes feed on their orchards.[14] This species is also hunted for bushmeat in Indonesia, contributing to its decline.[26]
References
Mildenstein, T.; Abdul Aziz, S.; Paguntalan, L.; Jakosalem, P.G.; Mohd-Azlan, J.; Tagtag, A.; Bansa, L.; Reintar, A.R.; Struebig, M.; Fredriksson, G.; Lee, B.; Thong, V.D.; Sheherazade. (2022). "Pteropus vampyrus". IUCN Red List of Threatened Species. 2022: e.T18766A22088824. Retrieved 9 December 2022.
Linnaeus, Carl (1758). Systema naturæ per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I (in Latin) (10th ed.). Holmiæ: Laurentius Salvius. p. 31. Retrieved 21 November 2012.
Simmons, N.B. (2005). "Order Chiroptera". In Wilson, D.E.; Reeder, D.M (eds.). Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. pp. 345–346. ISBN 978-0-8018-8221-0. OCLC 62265494.
Kunz, T.; Jones, D. (2000). "Pteropus vampyrus" (PDF). Mammalian Species. 642: 1–6. doi:10.1644/1545-1410(2000)642<0001:PV>2.0.CO;2. S2CID 31875754. Archived from the original (PDF) on 2015-02-21. Retrieved 2012-12-29.
Matti Airas. "Echolocation in bats" (PDF). HUT, Laboratory of Acoustics and Audio Signal Processing. p. 4. Archived from the original (PDF) on October 19, 2013. Retrieved July 19, 2013.
Almeida, Francisca C; Giannini, Norberto P; Simmons, Nancy B; Helgen, Kristofer M (2014). "Each flying fox on its own branch: A phylogenetic tree for Pteropus and related genera (Chiroptera: Pteropodidae)". Molecular Phylogenetics and Evolution. 77: 83–95. Bibcode:2014MolPE..77...83A. doi:10.1016/j.ympev.2014.03.009. hdl:11336/12485. PMID 24662680.
Andersen, Knud (1912). Catalogue of the Chiroptera in the collection of the British Museum. Vol. 1 (2nd ed.). London: British Museum (Natural History). Department of Zoology . [Mammals].
Payne J., Francis, C. M. and Philps, K. (1985). A field guide to the mammals of Borneo. The Sabah Society, Kota Kinabalu, Malaysia ISBN 9679994716
Francis, C. M. (2008). Mammals of Southeast Asia. pp. 195-196. ISBN 978-0-691-13551-9
Shepherd, Chris R.; Shepherd, Loretta Ann (2012). A Naturalist's Guide to the Mammals of Southeast Asia. Wiltshire, UK: John Beaufoy Publishing. p. 19. ISBN 978-1-906780-71-5.
Lekagul B., J. A. McNeely. 1977. Mammals of Thailand. Association for the Conservation of Wildlife, Bangkok, Thailand.
Goodwin R. E. (1979). "The bats of Timor". Bulletin of the American Museum of Natural History. 163: 75–122. hdl:2246/1288.
"Pteropus vampyrus (large flying fox)". Animal Diversity Web.
Lim, B. L. (1966). "Abundance and distribution of Malaysian bats in different ecological habitats". Federated Museums Journal. 11: 61–76.
Davis D. D. (1962). "Mammals of the lowland rainforest of North Borneo". Bulletin of the National Museum of Singapore. 31: 1–129.
Amin, Omar M.; Heckmann, Richard A.; Osama, Mohammed; Evans, R. Paul (2016). "Morphological and molecular descriptions of Moniliformis saudi sp. n. (Acanthocephala: Moniliformidae) from the desert hedgehog, Paraechinus aethiopicus (Ehrenberg) in Saudi Arabia, with a key to species and notes on histopathology". Folia Parasitologica. 63. doi:10.14411/fp.2016.014. ISSN 0015-5683. PMID 27189420.
Gould, E. (1978). "Foraging behavior of Malaysian nectar-feeding bats". Biotropica. 10 (3): 184–193. Bibcode:1978Biotr..10..184G. doi:10.2307/2387904. JSTOR 2387904.
Rabor D. (1977). Philippine birds and mammals, University of Philippines Press.
Heideman P. D., L. D. Heaney. (1992). "Pteropus vampyrus". pp. 140-143 in Old World fruit bats: an action plan for the family Pteropodidae (S. P. Mickleburgh, A. M. Hutson, P. A. Racy, eds). ICUN Survival Commission, Gland, Switzerland.
Corbet G. B., Hill, J. E. (1992). Mammals of the Indomalayan region: a systematic review. Oxford University Press ISBN 0198546939.
Medway L. (1965). Wild mammals of Malaya (Peninsular Malaysia) and Singapore. Oxford University Press.
Epstein, Jonathan H.; Olival, Kevin J.; Pulliam, Juliet R.C.; Smith, Craig; Westrum, Justin; Hughes, Tom; Dobson, Andrew P.; Zubaid, Akbar; Rahman, Sohayati Abdul; Basir, Misliah Mohamad; Field, Hume E.; Daszak, Peter (2009). "Pteropus vampyrus, a hunted migratory species with a multinational home-range and a need for regional management". Journal of Applied Ecology. 46 (5): 991–1002. Bibcode:2009JApEc..46..991E. doi:10.1111/j.1365-2664.2009.01699.x.
Constable, Harriet (2021-01-12). "The other virus that worries Asia". www.bbc.com. Retrieved 2022-01-02.
Rahman, Sohayati A.; Hassan, Sharifah S.; Olival, Kevin J.; Mohamed, Maizan; Chang, Li-Yen; Hassan, Latiffah; Saad, Norsharina M.; Shohaimi, Syamsiah A.; Mamat, Zaini C.; Naim, M.S.; Epstein, Jonathan H.; Suri, Arshad S.; Field, Hume E.; Daszak, Peter; the Henipavirus Ecology Research Group (2010). "Characterization of Nipah Virus from Naturally Infected Pteropus vampyrus Bats, Malaysia". Emerging Infectious Diseases. 16 (12): 1990–1993. doi:10.3201/eid1612.091790. PMC 3294568. PMID 21122240.
"Appendices I, II and III". CITES. 4 October 2017. Retrieved 28 August 2019.
Mickleburgh, S., Waylen, K., & Racey, P. (2009). Bats as bushmeat: a global review. Oryx, 43(02), 217-234.
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