View source for Large flying fox

{{Short description|Species of fruit bat}}
{{Redirect|Greater flying fox|the species from New Guinea and nearby islands|great flying fox}}
{{Speciesbox
| name = Large flying fox
| image = Wilhelma Kalong-Flughund Pteropus vampyrus 0513.jpg
| status = EN
| status_system = IUCN3.1
| status_ref = <ref name="iucn status 15 November 2021">{{cite iucn |author1=Mildenstein, T. |author2=Abdul Aziz, S. |author3=Paguntalan, L. |author4=Jakosalem, P.G. |author5=Mohd-Azlan, J. |author6=Tagtag, A. |author7=Bansa, L. |author8=Reintar, A.R. |author9=Struebig, M. |author10=Fredriksson, G. |author11=Lee, B. |author12=Thong, V.D. |author13=Sheherazade. |date=2022 |title=''Pteropus vampyrus'' |volume=2022 |page=e.T18766A22088824|url=https://www.iucnredlist.org/species/18766/22088824 |access-date=9 December 2022}}</ref>
| genus = Pteropus
| species = vampyrus
| authority = ([[Carl Linnaeus|Linnaeus]], [[10th edition of Systema Naturae|1758]])<ref name=SN>{{cite book|last=Linnaeus|first=Carl|title=Systema naturæ per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I|year=1758|publisher=Laurentius Salvius|location=Holmiæ|pages=31|url=https://www.biodiversitylibrary.org/item/80764#page/41/mode/1up|edition=10th|access-date=21 November 2012|language=la}}</ref> 
| range_map = Large Flying Fox area.png
| range_map_caption = Large flying fox range
| synonyms = {{species list
|Vespertilio vampyrus|Linnaeus, 1758
|Pteropus giganteus|(Brünnich, 1825)}}
}}

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 Asia]]n species of [[megabat]] in the family Pteropodidae.<ref name = MSW3/> 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.<ref name=Kunz2000/> As with nearly all other Old World fruit bats, it lacks the ability to [[Animal echolocation|echolocate]] but compensates for it with well-developed eyesight.<ref>{{cite web|url=http://www.acoustics.hut.fi/u/mairas/pubs/echolocation_in_bats.pdf |title=Echolocation in bats |author=Matti Airas |publisher=HUT, Laboratory of Acoustics and Audio Signal Processing |page=4 |access-date=July 19, 2013 |url-status=dead |archive-url=https://web.archive.org/web/20131019094829/http://www.acoustics.hut.fi/u/mairas/pubs/echolocation_in_bats.pdf |archive-date=October 19, 2013}}</ref>

==Taxonomy==
{{cladogram|align=left|style=width:390px;font-size:85%;line-height:100%|caption=Location of the large flying fox (''Pteropus vampyrus'') relative to some other members of its genus<ref name="Almeida 2014">{{cite journal|doi=10.1016/j.ympev.2014.03.009|pmid=24662680|title=Each flying fox on its own branch: A phylogenetic tree for Pteropus and related genera (Chiroptera: Pteropodidae)|journal=Molecular Phylogenetics and Evolution|volume=77|pages=83–95|year=2014|last1=Almeida|first1=Francisca C|last2=Giannini|first2=Norberto P|last3=Simmons|first3=Nancy B|last4=Helgen|first4=Kristofer M|doi-access=free|bibcode=2014MolPE..77...83A |hdl=11336/12485|hdl-access=free}}</ref>
|cladogram={{clade
|1={{clade
 |1={{clade
   |1=''[[Pteropus livingstonii|P. livingstonii]]''
   |2=''[[Pteropus voeltzkowi|P. voeltzkowi]]'' }}
 |label2="''vampyrus''" group
 |2={{clade
   |1={{clade
     |1=''[[Pteropus dasymallus|P. dasymallus]]''
     |2=''[[Pteropus pumilus|P. pumilus]]'' }}
   |2={{clade
     |1={{clade
       |1=''[[Pteropus rodricensis|P. rodricensis]]''
       |2='''''P. vampyrus''''' }}
     |2={{clade
       |1={{clade
         |1=''[[Pteropus lylei|P. lylei]]''
         |2=''[[Pteropus giganteus|P. medius]]''}}
       |2={{clade
         |1=''[[Pteropus aldabrensis|P. aldabrensis]]''
         |2=''[[Pteropus rufus|P. rufus]]''
         |3={{clade
           |1={{clade
             |1=''[[Pteropus seychellensis|P. seychellensis]]''*
             |2={{clade
               |1=''[[Pteropus niger|P. niger]]''*
               |2={{clade
                 |1=''[[Pteropus seychellensis|P. seychellensis]]''*
                 |2=''[[Pteropus niger|P. niger]]''* }} }} }} }} }} }}
        |3=''[[Pteropus pselaphon|P. pselaphon]]'' }} }} }} }} }}
The large flying fox was one of the many [[Mammalia in the 10th edition of Systema Naturae|mammal species originally described]] by [[Carl Linnaeus]] in the landmark 1758 [[10th edition of Systema Naturae|10th edition]] of his ''[[Systema Naturae]]'', receiving the name ''Vespertilio vampyrus''.<ref name=SN/> The [[holotype]] was collected on [[Java]].<ref name="Andersen 1912">{{cite book |last=Andersen |first=Knud |date=1912 |title=Catalogue of the Chiroptera in the collection of the British Museum |url= https://biodiversitylibrary.org/page/8409413|location=London |publisher=British Museum (Natural History). Department of Zoology . [Mammals] |edition=2nd|volume=1 |author-link=Knud Andersen (mammalogist)}}</ref>{{rp|70}} Its [[specific epithet (zoology)|species name]] "''vampyrus''" is derived from [[Slavs|Slavic]] "wampir" meaning "blood-sucking ghost or demon: [[vampire]]".<ref name=Kunz2000/> This name was chosen in reference to its "alleged blood-sucking habits",<ref name="Andersen 1912"/>{{rp|87}} although it is entirely vegetarian and largely [[frugivorous]].<ref name=Kunz2000/>

Based on phylogenetic analysis using [[mitochondrial DNA]], the closest relative of the large flying fox is the [[Rodrigues flying fox]] (''Pteropus rodricensis'').<ref name="Almeida 2014"/> 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:<ref name="Almeida 2014"/>
{{div col}}
*[[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'')
{{div col end}}

==Description==
{{Main|Megabat#Description|Pteropus#Description}}
The large flying fox is among the largest species of bat.<ref name=Kunz2000/> It weighs {{convert|0.65|-|1.1|kg|abbr=on}} and has a wingspan of up to {{convert|1.5|m|abbr=on}}.<ref name=Payne1985/><ref name=b3/> Its head-body length is {{convert|27|-|32|cm|in|abbr=on}}.<ref>{{Cite book |title = A Naturalist's Guide to the Mammals of Southeast Asia|last1 = Shepherd|first1 = Chris R.|publisher = John Beaufoy Publishing|year = 2012|isbn = 978-1-906780-71-5|location = Wiltshire, UK|pages = 19|first2 = Loretta Ann|last2 = Shepherd}}</ref> Its forearm length is {{cvt|180-220|mm|in}}.<ref name=Kunz2000/> 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.<ref name=Kunz2000/> The mantle hairs tend to be the longest.<ref name=Leagul1977/> The color and texture of the coat differ between sexes and age classes.<ref name=Goodwin1979/> Males tend to have slightly stiffer and thicker coats than females.<ref name=Kunz2000/> Immature individuals are almost all dull gray-brown.<ref name=Payne1985/> Young have a dark-colored mantle that becomes lighter in males when they mature.<ref name=Kunz2000/> 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.<ref name=Goodwin1979/> The mantle can vary from pale dirty-buff to orange-yellow, while the chest is usually dark-golden brown or dark russet.<ref name=Kunz2000/> The large flying fox has a large and robust skull. The [[dental formula]] is {{DentalFormula|upper=2.1.3.2|lower=2.1.3.3}}. It has a total of 34 teeth.<ref name=Leagul1977/><ref>{{cite web|url=http://animaldiversity.org/accounts/Pteropus_vampyrus/ |title=''Pteropus vampyrus'' (large flying fox)|website=Animal Diversity Web}}</ref> The large flying fox's wings are short and somewhat rounded at the tips. This allows them to fly slowly, but with great maneuverability.<ref name=Kunz2000/> 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.<ref name=Payne1985/><ref name=Leagul1977/><ref name=Goodwin1979/> 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.<ref name=Goodwin1979/><ref name=Lim1966/><ref name=Davis1962/> With fruit, the flying fox prefers the pulp, and slices open the rind to get it.<ref name=Davis1962/> With durian tree flowers, the flying fox can lick up the nectar without doing apparent damage to the flower.<ref name=Kunz2000/> The large flying fox is a host of the [[Acanthocephala]]n intestinal parasite ''[[Moniliformis convolutus]]''.<ref name="AminHeckmann2016">{{cite journal|last1=Amin|first1=Omar M.|last2=Heckmann|first2=Richard A.|last3=Osama|first3=Mohammed|last4=Evans|first4=R. Paul |title=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|journal=Folia Parasitologica|volume=63|year=2016|issn=0015-5683|doi=10.14411/fp.2016.014|pmid=27189420|doi-access=free}}</ref>

==Behavior and life history==
Colonies of large flying foxes [[Bat flight|fly]] in a scattered stream.<ref name=Leagul1977/> They may fly up to {{convert|50|km|abbr=on}} to their feeding grounds in one night. Vocalizations are not made during flight.<ref name=Leagul1977/> Large flocks fuse into family or feeding groups upon arrival at feeding grounds.<ref name=Kunz2000/> 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.<ref name=Kunz2000/> Feeding aggregations tend to be very noisy.<ref name=Gould1978/>

Flowering trees form the basis of territories in this species. Territorial behavior includes growling and the spreading of wings.<ref name=Gould1978/> During antagonistic behavior, individuals maintain spacing with wrists/thumbs sparring, bites, and loud vocalizations.<ref name=Goodwin1979/> When moving to a suitable resting place after landing, an individual may fight with conspecifics along the way.<ref name=Goodwin1979/> A roosting flying fox is positioned upside down with its wings wrapped up.<ref name=Rabor1977/> When it gets too warm, a flying fox fans itself with its wings.<ref name=Goodwin1979/> 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.<ref name=Heideman1992/> In Thailand, gestation may take place during the same period with young being born in March or early April.<ref name=Leagul1977/><ref name=Heideman1992/> Females apparently give birth during April and May in the Philippines,<ref name=Rabor1977/> and usually give birth to only one young.<ref name=Leagul1977/> For the first days, the mothers carry their young, but leave them at the roost when they go on their foraging trips.<ref name=Kunz2000/> The young are weaned by two to three months.<ref name=Leagul1977/>

==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.<ref name=b1/> In certain areas, the bat prefers coastal regions, but it can also be found at elevations up to {{convert|1370|m|abbr=on}}.<ref name=b2/>

Flying foxes inhabit [[primary forest]], mangrove forest, coconut groves, mixed fruit orchards, and a number of other habitats.<ref name=Heideman1992/> During the day, trees in mangrove forests and coconut groves may be used as roosts.<ref name=Goodwin1979/> In Malaysia, flying foxes prefer lowland habitats below 365 m.<ref name=Lim1966/> In Borneo, they inhabit the coastal areas, but move to nearby islands to feed on fruit.<ref name=Payne1985/> Flying foxes roost in the thousands (maximum). One colony was recorded numbering around 2,000 individuals in a mangrove forest in Timor<ref name=Goodwin1979/> and colonies of 10,000–⁠20,000 have also been reported.<ref name=Kunz2000/> In general, mangrove roosts have lower numbers of resting bats compared to lowland roost sites, which could mean mangrove forests are only used temporarily.<ref name=Lim1966/>

==Relationship to humans==
The large flying fox is hunted for [[bushmeat]]. In [[Peninsular Malaysia]], 1,756 [[hunting license]]s were issued for the large flying fox from 2002&ndash;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 [[sustainability|sustainable]]. A 2009 study predicted extinction of the Peninsular Malaysian population within 6&ndash;81 years if 22,000 individuals are lost to hunting each year.<ref>{{Cite journal| doi = 10.1111/j.1365-2664.2009.01699.x| volume = 46| issue = 5| pages = 991–1002| last1 = Epstein| first1 = Jonathan H.| last2 = Olival| first2 = Kevin J.| last3 = Pulliam| first3 = Juliet R.C.| last4 = Smith| first4 = Craig| last5 = Westrum| first5 = Justin| last6 = Hughes| first6 = Tom| last7 = Dobson| first7 = Andrew P.| last8 = Zubaid| first8 = Akbar| last9 = Rahman| first9 = Sohayati Abdul| last10 = Basir| first10 = Misliah Mohamad| last11 = Field| first11 = Hume E.| last12 = Daszak| first12 = Peter| title = ''Pteropus vampyrus'', a hunted migratory species with a multinational home-range and a need for regional management| journal = Journal of Applied Ecology| date = 2009| doi-access = | bibcode = 2009JApEc..46..991E}}</ref>

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 [[pig]]s.<ref>{{Cite web|last=Constable|first=Harriet|date=2021-01-12|title=The other virus that worries Asia|url=https://www.bbc.com/future/article/20210106-nipah-virus-how-bats-could-cause-the-next-pandemic|access-date=2022-01-02|website=www.bbc.com|language=en}}</ref> 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 [[antibody|antibodies]] against Nipah virus for the first eleven months, but was then [[seropositive]] once more. Two other bats&mdash;from which the Nipah virus was never detected&mdash;also registered as seropositive at points within the year. This suggested that the Nipah virus can [[recrudescence|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.<ref>{{Cite journal| doi = 10.3201/eid1612.091790| pmid = 21122240| pmc = 3294568| volume = 16| issue = 12| pages = 1990–1993| last1 = Rahman| first1 = Sohayati A.| last2 = Hassan| first2 = Sharifah S.| last3 = Olival| first3 = Kevin J.| last4 = Mohamed| first4 = Maizan| last5 = Chang| first5 = Li-Yen| last6 = Hassan| first6 = Latiffah| last7 = Saad| first7 = Norsharina M.| last8 = Shohaimi| first8 = Syamsiah A.| last9 = Mamat| first9 = Zaini C.| last10 = Naim| first10 = M.S.| last11 = Epstein| first11 = Jonathan H.| last12 = Suri| first12 = Arshad S.| last13 = Field| first13 = Hume E.| last14 = Daszak| first14 = Peter| last15 = the Henipavirus Ecology Research Group| title = Characterization of Nipah Virus from Naturally Infected ''Pteropus vampyrus'' Bats, Malaysia| journal = Emerging Infectious Diseases| date = 2010}}</ref>

==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]].<ref name="iucn status 15 November 2021" />
The large flying fox is on [[CITES#Appendix II|Appendix II]] of [[CITES]], which restricts international trade.<ref>{{cite web| title=Appendices I, II and III | date=4 October 2017 | url=https://cites.org/eng/app/appendices.php | publisher=CITES | access-date=28 August 2019}}</ref>

One threat to the large flying fox is [[habitat destruction]].<ref name=Heideman1992/> Flying foxes are sometimes hunted for food, and the controls on hunting seem to be unenforceable.<ref name=Kunz2000/> In some areas, farmers consider them pests as they sometimes feed on their orchards.<ref name=Lim1966/> This species is also hunted for bushmeat in [[Indonesia]], contributing to its decline.<ref name="Mickleburgh">Mickleburgh, S., Waylen, K., & Racey, P. (2009). Bats as bushmeat: a global review. Oryx, 43(02), 217-234.</ref>

==References==
{{Reflist|35em|refs=
<ref name=Davis1962>{{cite journal|author=Davis D. D. |year=1962|title=Mammals of the lowland rainforest of North Borneo|journal=Bulletin of the National Museum of Singapore |volume=31|pages=1–129}}</ref>

<ref name=Goodwin1979>{{cite journal|author=Goodwin R. E. |year=1979|title=The bats of Timor|journal= Bulletin of the American Museum of Natural History|volume= 163|pages=75–122|hdl=2246/1288}}</ref>

<ref name=Gould1978>{{cite journal|author=Gould, E. |year=1978|title=Foraging behavior of Malaysian nectar-feeding bats|journal=Biotropica|volume= 10|pages=184–193|jstor=2387904|doi=10.2307/2387904|issue=3|bibcode=1978Biotr..10..184G }}</ref>

<ref name=Heideman1992>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.</ref>

<ref name=Kunz2000>{{cite journal|author=Kunz, T.|author2=Jones, D.|year=2000|title=''Pteropus vampyrus''|journal=Mammalian Species|volume=642|pages=1–6|url=http://www.science.smith.edu/msi/pdf/642_Pteropus_vampyrus.pdf|doi=10.1644/1545-1410(2000)642<0001:PV>2.0.CO;2|s2cid=31875754 |access-date=2012-12-29|archive-url=https://web.archive.org/web/20150221051148/http://www.science.smith.edu/msi/pdf/642_Pteropus_vampyrus.pdf|archive-date=2015-02-21|url-status=dead}}</ref>

<ref name=Leagul1977>Lekagul B., J. A. McNeely. 1977. Mammals of Thailand. Association for the Conservation of Wildlife, Bangkok, Thailand.</ref>

<ref name=Lim1966>{{cite journal|author=Lim, B. L. |year=1966|title=Abundance and distribution of Malaysian bats in different ecological habitats|journal=Federated Museums Journal|volume= 11|pages=61–76}}</ref>

<ref name = MSW3>{{MSW3 Chiroptera | id = 13800382 | pages = 345–346}}</ref>

<ref name=Payne1985>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}}</ref>

<ref name=Rabor1977>Rabor D. (1977). ''Philippine birds and mammals'', University of Philippines Press.</ref>

<ref name=b1>Corbet G. B., Hill, J. E. (1992). ''Mammals of the Indomalayan region: a systematic review''. Oxford University Press {{ISBN|0198546939}}.</ref>

<ref name=b2>Medway L. (1965). ''Wild mammals of Malaya (Peninsular Malaysia) and Singapore.'' Oxford University Press.</ref>

<ref name=b3>Francis, C. M. (2008). ''Mammals of Southeast Asia.'' pp. 195-196. {{ISBN|978-0-691-13551-9}}</ref>
}}

==External links==
* {{Commons category-inline|Pteropus vampyrus}}
* {{Wikispecies-inline|Pteropus vampyrus}}

{{Pteropodidae|P.}}
{{Taxonbar |from=Q649684}}

[[Category:Pteropus]]
[[Category:Bats of Oceania]]
[[Category:Bats of Southeast Asia]]
[[Category:Bats of Borneo]]
[[Category:Bats of Indonesia]]
[[Category:Bats of Malaysia]]
[[Category:Bats of the Philippines]]
[[Category:Mammals of Thailand]]
[[Category:Bats as food]]
[[Category:Mammals described in 1758]]
[[Category:Taxa named by Carl Linnaeus]]
[[Category:Taxonomy articles created by Polbot]]