View source for Pteropus

{{Short description|Genus of large bats}}
{{Redirect|Flying fox|other bats known as flying fox|Acerodon|other uses|Flying Fox (disambiguation)}}
{{Automatic taxobox
| fossil_range = {{fossilrange|Holocene|ref=<ref>{{cite web|url=https://paleobiodb.org/classic/checkTaxonInfo?taxon_no=103145|title=Pteropus Brisson 1762 (flying fox)|website=Fossilworks|access-date=17 December 2021|archive-date=12 March 2023|archive-url=https://web.archive.org/web/20230312002730/http://www.fossilworks.org/cgi-bin/bridge.pl?a=taxonInfo&taxon_no=103145|url-status=live}}</ref>}}
| image = Polynesian flying fox (Pteropus tonganus) in flight Taveuni.jpg
| image_caption = Polynesian flying fox<br>''[[Pteropus tonganus]]'', Taveuni, Fiji
| taxon = Pteropus
| authority = [[Mathurin Jacques Brisson|Brisson]], 1762
| type_species = ''[[Pteropus niger|Vespertilio vampyrus niger]]''<ref name="ICZN 1998">{{cite journal |author1=ICZN |title=Opinion 1894. ''Regnum Animale ...'', Ed. 2 (M.J. Brisson, 1762): rejected for nomenclatural purposes, with the conservation of the mammalian generic names ''Philander'' (Marsupialia), ''Pteropus'' (Chiroptera), ''Glis'', ''Cuniculus'', and ''Hydrochoerus'' (Rodentia), ''Meles'', ''Lutra'' and ''Hyaena'' (Carnivora), ''Tapirus'' (Perissodactyla), ''Tragulus'' and ''Giraffa'' (Artiodactyla) |journal=Bulletin of Zoological Nomenclature |date=1998 |volume=55 |issue=1 |pages=64–71 |url=https://biodiversitylibrary.org/page/12441432}}</ref>
| type_species_authority = Kerr, 1792
| range_map = Pteropus range.jpg
| range_map_caption = Worldwide distribution of flying foxes
}}
[[File:Grey headed flying fox - skimming water - AndrewMercer - DSC00530.jpg|thumb|[[Gray-headed flying fox]]]]
'''''Pteropus''''' (suborder [[Yinpterochiroptera]]) is a [[genus]] of [[megabat]]s which are among the largest bats in the world. They are commonly known as '''fruit bats''' or '''flying foxes''', among other colloquial names. 
They live in [[South Asia]], [[Southeast Asia]], [[Australia]], [[East Africa]], and some oceanic islands in the Indian and Pacific Oceans.<ref name="Walker 1999">{{cite book| editor-last=Nowak| editor-first= R. M.| year=1999| title= Walker's Mammals of the World| volume= 1| edition= 6| pages=264–271| publisher= Johns Hopkins University Press| isbn=978-0-8018-5789-8}}</ref> 
There are at least 60 [[Extant taxon|extant]] species in the genus.<ref name = MSW3>{{MSW3 Chiroptera | id = 13800239 | pages = 334–346 | heading = Genus ''Pteropus''}}</ref>

Flying foxes eat fruit and other plant matter, and occasionally consume insects as well. They locate resources with their keen sense of smell.
Most, but not all, are [[nocturnality|nocturnal]]. They navigate with keen eyesight, as they cannot [[Animal echolocation|echolocate]].
They have [[R/K selection theory#K-selection|long life spans and low reproductive outputs]], with females of most species producing only one offspring per year. Their slow life history makes their populations vulnerable to threats such as [[Overexploitation|overhunting]], [[culling]], and natural disasters. Six flying fox species have been made [[extinction|extinct]] in modern times by overhunting. Flying foxes are often persecuted for their real or perceived role in [[pest (organism)|damaging crops]]. They are ecologically beneficial by assisting in the regeneration of forests via [[Seed dispersal#Animals: epi- and endozoochory|seed dispersal]]. They benefit ecosystems and human interests by [[pollinator|pollinating]] plants.

Like other bats, flying foxes are relevant to humans as a source of disease, as they are the [[Natural reservoir|reservoirs]] of rare but fatal disease agents including [[Australian bat lyssavirus]], which causes [[rabies]], and [[Henipavirus#Hendra virus|Hendra virus]]; seven known human deaths have resulted from these two diseases. [[Nipah virus]] is also transmitted by flying foxes&mdash;it affects more people, with over 100 attributed fatalities. They have cultural significance to indigenous people, with appearances in traditional art, folklore, and weaponry. Their fur and teeth were used as currency in the past. Some cultures still use their teeth as currency today.

==Taxonomy and etymology==
The genus name ''Pteropus'', from [[Ancient Greek]] 'πτερόν' (pterón), meaning "wing", and 'πούς' (poús), meaning "foot", was coined by French zoologist [[Mathurin Jacques Brisson]] in 1762.<ref name="Brisson 1762">{{cite book| last=Brisson| first= A. D.| year=1762| title= Regnum Animale in Classes IX distributum sive synopsis methodica| publisher=Lugduni Batavorum :Apud Theodorum Haak,1762| pages=153–155| url=https://biodiversitylibrary.org/page/28278666}}</ref>
Prior to 1998, genus authority was sometimes given to German naturalist [[Johann Christian Polycarp Erxleben]].<ref name="Erxleben 1777">{{cite book| last=Erxleben| first=J. C. P.| year=1777| title=Io. Christ. Polyc. Erxleben Phil. D. ... Systema regni animalis per classes, ordines, genera, species, varietates : cvm synonymia et historia animalivm : Classis I. Mammalia| publisher=Lipsiae :Impensis Weygandianis,1777| pages=130–141| url=https://biodiversitylibrary.org/page/15872927}}</ref> Although the Brisson publication (1762) predated the Erxleben publication (1777), thus giving him preference under the [[Principle of Priority]], some authors gave preference to Erxleben as genus authority because Brisson's publication did not consistently use [[binomial nomenclature]].<ref name="Jackson 2015">{{cite book| last1=Jackson| first1= S.| last2= Jackson| first2= S. M.| last3= Groves| first3= C.| year=2015| title= Taxonomy of Australian mammals| publisher= Csiro Publishing| isbn=978-1486300136| page=238}}</ref> In 1998, the [[International Commission on Zoological Nomenclature]] (ICZN) decided that Brisson's 1762 publication was a "rejected work" for nomenclatural purposes. Despite rejecting the majority of the publication, the ICZN decided to conserve a dozen generic names from the work and retain Brisson as authority, including ''Pteropus''.<ref name="ICZN 1998"/>

The [[type species]] of the genus is the [[Mauritian flying fox]], ''Pteropus niger'' (described as ''Vespertilio vampyrus niger'' by [[Robert Kerr (writer)|Robert Kerr]] in 1792).<ref name="Kerr 1792">{{cite book| last1=Kerr| first1= R.| last2= Gmelin| first2=J. F.|year=1792| title= The Animal Kingdom Or Zoological System of the Celebrated Sir Charles Linnaeus: Class I. Mammalia (class II. the Birds): Containing a Complete Systematic Description, arrangement, and nomenclature, of all the known species and varieties of the mammalia, or animals which give suck to their young| publisher=Edinburgh :Printed for A. Strahan, and T. Cadell, London, and W. Creech, Edinburgh,1792| pages=90–91| url=https://biodiversitylibrary.org/page/38664108}}</ref>
The decision to designate ''P.&nbsp;niger'' as the type species was made by the ICZN through their [[plenary power]]s over biological nomenclature.<ref name="Jackson 2015"/> The phrase "flying fox" has been used to refer to ''Pteropus'' bats since at least 1759.<ref name="merriam-webster">{{cite web|title=Flying Fox|website=Merriam-Webster.com| publisher= Merriam-Webster, n.d.| access-date= 5 June 2018| url=https://www.merriam-webster.com/dictionary/flying%20fox}}</ref>

===Species===
{{main|List of fruit bats#Subfamily Pteropodinae}}
{{cladogram|align=center|style=width:500px;font-size:85%;line-height:100%|caption=Phylogeny of ''Pteropus'' based on mitochondrial [[cytochrome b]] and rRNA12S genes.  Species without enough data are excluded.<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=''[[Acerodon jubatus]]''
        |2=''[[Acerodon celebensis]]''}}
     |2={{clade
       |1=''[[Pteropus personatus|P. personatus]]''
       |2=''[[Neopteryx frosti]]''
       |3={{clade
         |1={{clade
           |1=''[[Pteropus macrotis|P. macrotis]]''
           |2={{clade
             |1={{clade
               |1=''[[Pteropus mahaganus|P. mahaganus]]''
               |2={{clade
                 |1=''[[Pteropus gilliardi|P. gilliardi]]''
                 |2=''[[Pteropus woodfordi|P. woodfordi]]'' }} }}
             |2={{clade
               |1=''[[Pteropus molossinus|P. molossinus]]''
               |2={{clade 
                 |1=''†[[Pteropus tokudae|P. tokudae]]''
                 |2=''[[Pteropus pelagicus|P. pelagicus]]'' }} }} }} }}
         |2={{clade
           |1=''[[Pteropus scapulatus|P. scapulatus]]''
           |2={{clade
              |1=''[[Pteropus lombocensis|P. lombocensis]]''
              |2={{clade
                 |1={{clade
                    |1={{clade
                      |1=''[[Pteropus livingstonii|P. livingstonii]]''
                      |2=''[[Pteropus voeltzkowi|P. voeltzkowi]]'' }}
                    |2={{clade
                      |1={{clade
                         |1=''[[Pteropus dasymallus|P. dasymallus]]''
                         |2=''[[Pteropus pumilus|P. pumilus]]'' }}
                      |2={{clade
                         |1={{clade
                           |1=''[[Pteropus rodricensis|P. rodricensis]]''
                           |2=''[[Pteropus vampyrus|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]]'' }} }}
                 |2={{clade
                   |1={{clade
                     |1={{clade
                       |1=''[[Pteropus capistratus|P. capistratus]]''
                       |2=''[[Pteropus ennisae|P. ennisae]]'' }}
                     |2={{clade
                       |1=''[[Pteropus vetulus|P. vetulus]]''
                       |2={{clade
                         |1={{clade
                           |1=''[[Pteropus nitendiensis|P. nitendiensis]]''
                           |2=''[[Pteropus tuberculatus|P. tuberculatus]]'' }}
                         |2={{clade
                           |1=''[[Pteropus anetianus|P. anetianus]]''
                           |2={{clade
                             |1=''[[Pteropus samoensis|P. samoensis]]''
                             |2=''[[Pteropus fundatus|P. fundatus]]''
                             |3={{clade
                               |1=''[[Pteropus rayneri|P. rayneri]]''
                               |2={{clade
                                 |1=''[[Pteropus rennelli|P. rennelli]]''
                                 |2=''[[Pteropus cognatus|P. cognatus]]'' }} }} }} }} }} }} }}
                   |2={{clade
                     |1=''[[Pteropus poliocephalus|P. poliocephalus]]''
                     |2={{clade
                       |1=''[[Pteropus ornatus|P. ornatus]]''
                       |2={{clade
                         |1={{clade
                           |1={{clade
                             |1=''[[Pteropus hypomelanus|P. hypomelanus]]''*
                             |2={{clade
                               |1=''[[Pteropus griseus|P. griseus]]''
                               |2={{clade
                                 |1=''[[Pteropus speciosus|P. speciosus]]''
                                 |2=''[[Pteropus hypomelanus|P. hypomelanus]]''* }} }} }}
                           |2={{clade
                             |1=''[[Pteropus neohibernicus|P. neohibernicus]]''
                             |2={{clade
                               |1=''[[Pteropus conspicillatus|P. conspicillatus]]''
                               |2=''[[Pteropus alecto|P. alecto]]'' }} }} }}
                         |2={{clade
                           |1={{clade
                             |1=''[[Pteropus tonganus|P. tonganus]]''
                             |2={{clade
                               |1=''[[Pteropus ualanus|P. ualanus]]''
                               |2=''[[Pteropus admiralitatum|P. admiralitatum]]'' }} }} 
                           |2={{clade
                              |1=''[[Pteropus pohlei|P. pohlei]]''
                              |2={{clade
                                |1=''[[Pteropus mariannus|P. mariannus]]''
                                |2={{clade
                                  |1=''[[Pteropus pelewensis|P. pelewensis]]''*
                                  |2={{clade
                                    |1=''[[Pteropus yapensis|P. yapensis]]''
                                    |2=''[[Pteropus pelewensis|P. pelewensis]]''* }} }} }} }}

}} }} }} }} }} }} }} }} }} }} }} }} }}

==Description==
===External characteristics===
[[File:Flying-Fox-Bat.jpg|thumb|Flying fox in flight]]
Flying fox species vary in body weight, ranging from {{cvt|120-1600|g|lb}}. Across all species, males are usually larger than females.<ref name="McNab 2001">{{cite journal|doi=10.1644/1545-1542(2001)082<0709:SDASOE>2.0.CO;2|year=2001|volume=82|issue=3|pages=709|title=Sexual Dimorphism and Scaling of Energetics in Flying Foxes of the Genus Pteropus|journal=Journal of Mammalogy|last1=McNab|first1=Brian K|last2=Armstrong|first2=Monica I|doi-access=free}}</ref>
The [[large flying fox]] has the longest forearm length and reported wingspan of any bat species,<ref name="Walker 1999"/> but some bat species exceed it in weight. 
Its wingspan is up to {{convert|1.5|m|ftin|abbr=on}}, and it can weigh up to {{cvt|1.1|kg|lb|frac=4}}.<ref name="Kunz 2000">{{cite journal|doi=10.1644/1545-1410(2000)642<0001:PV>2.0.CO;2|year=2000|volume=642|pages=1|title=Pteropus vampyrus|journal=Mammalian Species|last1=Kunz|first1=Thomas H|last2=Jones|first2=Deborah P|s2cid=31875754 }}</ref> The [[Indian flying fox|Indian]] and [[great flying fox|great]] flying foxes are heavier, at {{convert|1.6|and|1.45|kg|lb|frac=8|abbr=on}}, respectively.<ref name="Walker 1999"/><ref name="Flannery 1995b">{{cite book| last=Flannery| first= T.| year=1995| title=Mammals of New Guinea| pages= 376–377| isbn=978-0801431494| publisher=Cornell University Press}}</ref> 
Outside this genus, the [[giant golden-crowned flying fox]] (genus ''[[Acerodon]]'') is the only bat with similar dimensions.<ref name="Walker 1999"/>

Most flying fox species are considerably smaller and generally weigh less than {{convert|600|g|oz|frac=2|abbr=on}}.<ref name="Flannery 1995">{{cite book| last=Flannery| first= T.| year=1995| title=Mammals of the South-West Pacific & Moluccan Islands| pages= 245–303| isbn=978-0801431500| publisher=Cornell University Press}}</ref> 
Smaller species such as the [[masked flying fox|masked]], [[Temminck's flying fox|Temminck's]], [[Guam flying fox|Guam]], and [[dwarf flying fox|dwarf]] flying foxes all weigh less than {{convert|170|g|oz|frac=2|abbr=on}}.<ref name="Flannery 1995"/>

The pelage is long and silky with a dense underfur.<ref name="Walker 1999"/> In many species, individuals have a "mantle" of contrasting fur color on the back of their head, the shoulders, and the upper back.<ref name="Nelson"/> They lack tails. As the common name "flying fox" suggests, their heads resemble that of a small fox because of their small ears and large eyes. Females have one pair of [[mammary gland|mammae]] located in the chest region. Their ears are long and pointed at the tip and lack [[tragus (ear)#other animals|tragi]], the outer margin of each ear forming an unbroken ring.<ref name="Walker 1999"/><ref name="Nelson">{{cite report|url=https://www.environment.gov.au/system/files/pages/a117ced5-9a94-4586-afdb-1f333618e1e3/files/37-ind.pdf| title=Fauna of Australia| last=Nelson| first=John E.| volume=1B|publisher=Australian Government Department of the Environment and Energy}}</ref> The toes have sharp, curved claws. While [[microbat]]s only have a claw on each thumb of their forelimbs, flying foxes additionally have a claw on each index finger.<ref name="Walker 1999"/>

===Skull and dentition===
[[File:Pteropus melanotus 04 MWNH 237.jpg|thumb|Skull of the [[black-eared flying fox]]]]
The skulls of ''Pteropus'' species are composed of 24 bones, the [[snout]] is made of 7, the [[cranium]] of 16 and the [[mandible]] is a single bone. It has a large and bulbous [[Neurocranium|braincase]]. Like all mammals, flying foxes have three [[Evolution of mammalian auditory ossicles|middle ear ossicles]] which assist in transmitting sound to the brain. Flying fox skulls continue to develop after they are born. Compared to adults, young flying foxes have very short snouts; as they reach maturity, the [[maxilla]] elongates, gaining bone between the [[Zygomatic process of maxilla|zygomatic process]]es and the [[Canine tooth|canine teeth]].<ref name="Norberto 2006">{{cite journal|doi=10.1206/0003-0090(2006)295[0001:OTCOOC]2.0.CO;2|year=2006|volume=295|pages=1–134|title=On the Cranial Osteology of Chiroptera. I. Pteropus (Megachiroptera: Pteropodidae)|journal=Bulletin of the American Museum of Natural History|last1=Giannini|first1=Norberto P|last2=Wible|first2=John R|last3=Simmons|first3=Nancy B|s2cid=86010208 }}</ref>

Based on the [[grey-headed flying fox]]'s development, pups are born with some [[Deciduous teeth|milk teeth]] already erupted: canines and [[incisor]]s. By 9 days old, all the milk teeth have emerged, with a [[dentition#Dental formula|dental formula]] of {{DentalFormula|upper=2.1.2.0|lower=2.1.2.0}} and a total of 20 teeth. By 140 days old (4.6 months), all the milk teeth have fallen and been replaced by [[permanent teeth]]. The canines are usually replaced first, followed by the [[premolar]]s, incisors, and then [[Molar (tooth)|molars]].<ref name="Divljan 2011">{{cite book|last1=Divljan| first1= A.| last2= Parry-Jones| first2= K.| last3= Wardle| first3= G. M.| editor1-last=Law| editor1-first=B.|editor2-last=Eby| editor2-first=P. |editor3-last=Lunney| editor3-first=D.| editor4-last=Lumsden| editor4-first=L.| year=2011| chapter= One hundred and forty days in the life of a flying-fox tooth-fairy: estimating the age of pups using tooth eruption and replacement|title=The Biology and Conservation of Australasian bats| publisher=Royal Zoological Society of New South Wales| pages=97–105| isbn= 978-0-9803272-4-3}}</ref> The adult dental formula is {{DentalFormula|upper=2.1.3.2|lower=2.1.3.3}} for a total of 34 teeth. The [[Glossary of dentistry#Occlusal|occlusal surface]] of the molars is generally smooth but with longitudinal furrows.<ref name="Walker 1999"/>

===Internal systems===
Flying foxes have large hearts and a relatively fast [[heart rate]]: resting individuals have a heart rate of 100–400 beats per minute.<ref name="Parish 2012"/>

Flying foxes have simple [[Gastrointestinal tract|digestive tracts]];<ref name="Walker 1999"/> the time between ingestion and excretion is as short as 12 minutes. They lack both a [[cecum]] and an [[Appendix (anatomy)|appendix]].<ref name="Tedman 1985">{{cite journal|doi=10.1071/ZO9850625|title=The Morphology of the Gastrointestinal Tract and Food Transit Time in the Fruit Bats Pteropus Alecto and P. Poliocephalus (Megachiroptera)|journal=Australian Journal of Zoology|volume=33|issue=5|pages=625|year=1985|last1=Tedman|first1=RA|last2=Hall|first2=LS}}</ref> The stomach has marked [[Gastric glands#Types of gland|cardiac and fundic regions]].<ref name="Nelson"/>

===Intelligence===
The megabats, including flying foxes, have the greatest [[encephalization quotient]] (brain size relative to body size) of any bat family at 1.20.<ref name="Eisenberg 1978">{{cite journal|doi=10.1111/j.1558-5646.1978.tb04627.x|pmid=28567922|title=Relative Brain Size and Feeding Strategies in the Chiroptera|journal=Evolution|volume=32|issue=4|pages=740–751|year=1978|last1=Eisenberg|first1=John F|last2=Wilson|first2=Don E|doi-access=free}}</ref> 
This value is equivalent to that of domestic dogs.<ref name="Deamer 2009">{{cite web| url=http://www.science20.com/stars_planets_life/calculating_animal_intelligence| last=Deamer| first=D.|date=5 November 2009| title=Calculating Animal Intelligence| website=Science2.0| publisher=ION Publications LLC| access-date=24 June 2018}}</ref> Flying foxes display behaviors that indicate a reliance on long-term information storage. Though they have wide-ranging movements and cover thousands of square kilometers annually, they are consistently able to locate the same resource patches and roosts. They will visit these resource patches consistently in a strategy known as [[trap-lining]]. They can also be [[Operant conditioning|conditioned]] to perform behaviors, such as one study where [[spectacled flying fox]]es were trained to pull a lever using juice as a [[reinforcement]]. In a follow-up to the initial study, individuals who had learned to pull the lever to receive juice still did so 3.5 years later.<ref name="Flick 2011">{{cite book| last1=Flick| first1= B.| last2= Spencer| first2= H.| last3=Van Der Zwan| first3= R.| editor1-last=Law| editor1-first=B.|editor2-last=Eby| editor2-first=P. |editor3-last=Lunney| editor3-first=D.| editor4-last=Lumsden| editor4-first=L.| year=2011| chapter= Are hand-raised flying-foxes (Pteropus conspicillatus) better learners than wild-raised ones in an operant conditioning situation?| title=The Biology and Conservation of Australasian bats| publisher=Royal Zoological Society of New South Wales| pages=86–91| doi= 10.7882/FS.2011.011| isbn= 978-0-9803272-4-3}}</ref>

===Senses===
====Smell====
[[File:CSIRO ScienceImage 3220 Spectacled flying fox.jpg|thumb|The prominent eyes of the [[spectacled flying fox]]]]
Flying foxes rely heavily on their sense of smell.
They have large [[olfactory bulb]]s to process scents. They use scent to locate food, for mothers to locate their pups, and for mates to locate each other.<ref name="Nelson"/> Males have enlarged [[androgen]]-sensitive [[sebaceous gland]]s on their shoulders that they use for [[Territory (animal)#Spraying|scent-marking]] their territories, particularly during the mating season. The secretions of these glands vary by species&mdash;of the 65 chemical compounds identified from the glands of four species, no compound was found in all species.<ref name="Wood 2005"/> Males also engage in "[[Urination#Other animals|urine washing]]", meaning that they coat themselves in their own urine.<ref name="Wood 2005">{{cite journal| last1=Wood| first1= W. F.| last2= Walsh| first2= A.| last3= Seyjagat| first3= J.| last4= Weldon| first4= P. J.| title= Volatile Compounds in Shoulder Gland Secretions of Male Flying Foxes, Genus Pteropus (Pteropodidae, Chiroptera)| year=2005| journal=Z Naturforsch C| volume=60| issue=9–10| pages= 779–84|url=https://www.researchgate.net/publication/7447695| doi= 10.1515/znc-2005-9-1019| pmid= 16320623| s2cid= 1938648| doi-access= free}}</ref><ref name="Wagner 2008">{{cite journal| last=Wagner| first= J.| year=2008| title= Glandular secretions of male Pteropus (Flying foxes): preliminary chemical comparisons among species|journal=Independent Study Project (Isp) Collection| url=http://digitalcollections.sit.edu/isp_collection/559/}}</ref>

====Sight====
Flying foxes do not [[Animal echolocation|echolocate]], and therefore rely on sight to navigate. Their eyes are relatively large and positioned on the front of their heads, giving them [[binocular vision]].<ref>{{cite journal|doi=10.1371/journal.pone.0034564|pmid=22509324|title=Multiple Episodes of Convergence in Genes of the Dim Light Vision Pathway in Bats|journal=PLOS ONE|volume=7|issue=4|pages=e34564|year=2012|last1=Shen|first1=Yong-Yi|last2=Lim|first2=Burton K|last3=Liu|first3=He-Qun|last4=Liu|first4=Jie|last5=Irwin|first5=David M|last6=Zhang|first6=Ya-Ping|bibcode=2012PLoSO...734564S|pmc=3324491|doi-access=free}}</ref>
Like most mammals, though not [[primate]]s, they are [[dichromacy|dichromatic]].<ref name="Melin 2014">{{cite journal|doi=10.1007/s00359-014-0951-x|pmid=25319538|title=Dichromatic vision in a fruit bat with diurnal proclivities: The Samoan flying fox (Pteropus samoensis)|journal=Journal of Comparative Physiology A|volume=200|issue=12|pages=1015–22|year=2014|last1=Melin|first1=Amanda D|last2=Danosi|first2=Christina F|last3=McCracken|first3=Gary F|last4=Dominy|first4=Nathaniel J|s2cid=12167082}}</ref> They have both [[Rod cell|rods]] and [[Cone cell|cones]]; they have "blue" cones that detect short-wavelength light and "green" cones that detect medium-to-long-wavelengths. The rods greatly outnumber the cones, however, as cones comprise only 0.5% of photoreceptors. Flying foxes are adapted to [[Scotopic vision|seeing in low-light conditions]].<ref name="Max 2007">{{cite web| url=https://www.sciencedaily.com/releases/2007/06/070615093131.htm| author=Max-Planck-Gesellschaft| title=Fruit Bats Are Not 'Blind As A Bat'| date=16 June 2007| website=ScienceDaily|access-date=25 June 2018}}</ref>

==Evolutionary history==
{{cladogram|align=right|style=width:250px;font-size:85%;line-height:85%|caption=Phylogeny of Pteropodidae<ref name="Giannini 2003">{{cite journal|doi=10.1111/j.1096-0031.2003.tb00385.x|title=A phylogeny of megachiropteran bats (Mammalia: Chiroptera: Pteropodidae) based on direct optimization analysis of one nuclear and four mitochondrial genes|journal=Cladistics|volume=19|issue=6|pages=496–511|year=2003|last1=Giannini|first1=Norberto P|last2=Simmons|first2=Nancy B|pmid=34905855 |s2cid=84696546}}</ref>
|cladogram={{clade
|label1='''Pteropodidae'''
   |1={{clade
     |1=''[[Melonycteris]]''
     |2={{clade
       |1=''[[Nyctimene (genus)|Nyctimene]]''
       |2={{clade
         |1={{clade
           |1=''[[Pteralopex]]''
           |2=''[[Eidolon (genus)|Eidolon]]'' }} 
         |2={{clade
           |1={{clade
              |1=''[[Acerodon]]''
              |2=<big>'''''Pteropus'''''</big>   }}
           |2={{clade
              |1={{clade
                 |1=''[[Notopteris]]''
                 |2={{clade
                    |1={{clade
                       |1=''[[Aproteles]]''
                       |2=''[[Dobsonia]]''}} 
                    |2={{clade
                       |1={{clade
                          |1=''[[Syconycteris]]''
                          |2=''[[Macroglossus]]''}}
                       |2={{clade
                          |1={{clade
                             |1=''[[Cynopterus]]''
                             |2={{clade
                                |1=''[[Megaerops]]''
                                |2=''[[Ptenochirus]]'' }} }} 
                          |2={{clade
                            |1=''[[Thoopterus]]''
                            |2=''[[Chironax]]''
                            |3=''[[Aethalops]]''
                            |4=''[[Balionycteris]]'' }} }} }} }} }}
              |2={{clade
                |1={{clade
                   |1=''[[Eonycteris]]''
                   |2={{clade
                     |1=''[[Stenonycteris]]''
                     |2=''[[Rousettus]]'' }} }} 
                |2={{clade
                   |1={{clade
                     |1=''[[Epomops]]''
                     |2={{clade
                        |1=''[[Epomophorus]]''
                        |2=''[[Micropteropus]]'' }} }} 
                   |2={{clade
                     |1=''[[Megaloglossus]]''
                     |2={{clade
                        |1=''[[Lissonycteris]]''
                        |2=''[[Myonycteris]]'' }} }} }}
 }} }} }}

}} }} }} }} }}
Flying foxes are poorly represented in the fossil record. Relative to the current number of extant species, the Pteropodidae has one of the most incomplete fossil records of any bat group.<ref name="Eiting 2009"/> As of 2014, no flying fox fossils are known from before the [[Holocene]].<ref name="Almeida 2014"/>
Many flying foxes live in the [[tropics]], where conditions for fossilization are poor.<ref name="Eiting 2009">{{cite journal|doi=10.1007/s10914-009-9118-x|title=Global Completeness of the Bat Fossil Record|journal=Journal of Mammalian Evolution|volume=16|issue=3|pages=151|year=2009|last1=Eiting|first1=Thomas P|last2=Gunnell|first2=Gregg F|s2cid=5923450}}</ref> Based on [[molecular evolution]], flying foxes diverged from a common ancestor with ''[[Rousettus]]'' 28&ndash;18 million years ago<ref name="Teeling 2005">{{cite journal|doi=10.1126/science.1105113|pmid=15681385|title=A Molecular Phylogeny for Bats Illuminates Biogeography and the Fossil Record|journal=Science|volume=307|issue=5709|pages=580–4|year=2005|last1=Teeling|first1=E. C|last2=Springer|first2=M. S|last3=Madsen|first3=O|last4=Bates|first4=P|last5=O'Brien|first5=S. J|last6=Murphy|first6=W. J|bibcode=2005Sci...307..580T|s2cid=25912333}}</ref> and from their [[Sister group|sister taxa]] ''[[Neopteryx]]'' and ''Acerodon'' 6.6&ndash;10.6 million years ago.<ref name="Almeida 2014"/> ''Neopteryx'', ''Acerodon'', ''[[Desmalopex]]'', ''[[Melonycteris]]'', ''[[Mirimiri]]'', ''[[Pteralopex]]'', and ''[[Styloctenium]]'' are all relatively closely related to the flying foxes, as they are the other members of its [[subfamily]] [[Pteropodinae]].

Phylogenetic analysis indicates that flying foxes diversified rapidly in an explosive [[evolutionary radiation]], creating many taxa in a relatively short time frame.<ref name="Almeida 2011">{{cite journal|doi=10.1186/1471-2148-11-281|pmid=21961908|pmc=3199269|title=Evolutionary relationships of the old world fruit bats (Chiroptera, Pteropodidae): Another star phylogeny?|journal=BMC Evolutionary Biology|volume=11|pages=281|year=2011|last1=Almeida|first1=Francisca C|last2=Giannini|first2=Norberto P|last3=Desalle|first3=Rob|last4=Simmons|first4=Nancy B |issue=1 |doi-access=free |bibcode=2011BMCEE..11..281A }}</ref> Most flying fox lineages emerged after the [[Zanclean]], with two major [[clade]]s created: one consisting of the [[Indian Ocean]] species and the other of the [[Melanesia]]n, [[Micronesia]]n, Australian, and insular [[Southeast Asia]]n species. Flying foxes likely originated on mainland Asia; molecular data suggests that there were at least three [[Colonisation (biology)|colonization events]] into the Indian Ocean. One event resulted in [[Livingstone's fruit bat]] and the [[Pemba flying fox]], which are the westernmost flying foxes. A second colonization event resulted in the [[Rodrigues flying fox]] to [[Rodrigues]] Island; while a third event resulted in several species diverging to [[Mauritius]], the [[Seychelles]], [[Madagascar]], and [[Aldabra]].<ref name="O'Brien 2009"/>

With one possible exception - the masked flying fox (''P.&nbsp;personatus''), flying foxes are likely [[Monophyly|monophyletic]]. There are over 60 extant species of flying fox.<ref name="Almeida 2014"/> Flying foxes are now present from the western Indian Ocean midway through the [[Pacific Ocean]] as far east as the [[Cook Islands]].<ref name="O'Brien 2009"/><ref name="Almeida 2014"/> They are found in tropical and subtropical climates.<ref name="O'Brien 2009">{{cite journal|doi=10.1016/j.ympev.2009.02.010|pmid=19249376|title=Multiple colonisations of the western Indian Ocean by Pteropus fruit bats (Megachiroptera: Pteropodidae): The furthest islands were colonised first|journal=Molecular Phylogenetics and Evolution|volume=51|issue=2|pages=294–303|year=2009|last1=O'Brien|first1=John|last2=Mariani|first2=Carol|last3=Olson|first3=Link|last4=Russell|first4=Amy L|last5=Say|first5=Ludovic|last6=Yoder|first6=Anne D|last7=Hayden|first7=Tom J|bibcode=2009MolPE..51..294O }}</ref>

==Biology and ecology==
===Reproduction and life cycle===
[[File:Pteropus conspicillatus with baby.jpg|thumb|Female spectacled flying fox with pup]]
Many species of flying fox are [[Polygynandry|polygynandrous]], meaning that each individual will mate with several other individuals.<ref name="Maruthupandian 2013">{{cite journal|doi=10.1371/journal.pone.0059743|pmid=23544092|title=Cunnilingus Apparently Increases Duration of Copulation in the Indian Flying Fox, Pteropus giganteus|journal=PLOS ONE| volume=8| issue=3| pages=e59743 |year=2013 |last1=Maruthupandian |first1=Jayabalan |last2=Marimuthu |first2=Ganapathy|bibcode=2013PLoSO...859743M|pmc=3609797|doi-access=free}}</ref> The [[Samoa flying fox]] is a notable exception because it is [[Monogamy in animals|monogamous]].<ref name="Sugita 2009">{{cite journal|doi=10.1644/07-MAMM-A-368.1|title=Roosting Pattern and Reproductive Cycle of Bonin Flying Foxes (Pteropus pselaphon)|journal=Journal of Mammalogy| volume=90|pages=195–202|year=2009|last1=Sugita|first1=Norimasa|last2=Inaba|first2=Makoto|last3=Ueda|first3=Keisuke|doi-access=free}}</ref> Flying fox sexual behaviors include oral sex in addition to intercourse, with [[fellatio]] and [[cunnilingus]] observed between opposite sexes, as well as homosexual fellatio in at least one species, the [[Bonin flying fox]].<ref name="Sugita 2016">{{cite journal|doi=10.1371/journal.pone.0166024|pmid=27824953|pmc=5100941|title=Homosexual Fellatio: Erect Penis Licking between Male Bonin Flying Foxes Pteropus pselaphon|journal=PLOS ONE|volume=11|issue=11|pages=e0166024|year=2016|last1=Sugita|first1=Norimasa|bibcode=2016PLoSO..1166024S|doi-access=free}}</ref><ref name="Tan 2009">{{cite journal|doi=10.1371/journal.pone.0007595|pmid=19862320|title=Fellatio by Fruit Bats Prolongs Copulation Time|journal=PLOS ONE|volume=4|issue=10|pages=e7595|year=2009|last1=Tan|first1=Min|last2=Jones|first2=Gareth|last3=Zhu|first3=Guangjian|last4=Ye|first4=Jianping|last5=Hong|first5=Tiyu|last6=Zhou|first6=Shanyi|last7=Zhang|first7=Shuyi|last8=Zhang|first8=Libiao|bibcode=2009PLoSO...4.7595T|pmc=2762080|doi-access=free}}</ref> Opposite-sex oral sex is associated with increased duration of intercourse, while same-sex fellatio is hypothesized to encourage colony formation of otherwise-antagonistic males in colder climates.<ref name="Sugita 2016"/><ref name="Tan 2009"/>

Flying fox [[Pregnancy (mammals)#Gestation periods|gestation length]] varies among species; gestation length is 140&ndash;190 days (4.6&ndash;6.3 months).
Females have a [[Litter (animal)|litter size]] of one young at a time, called a pup.<ref name="Pierson 1992">{{cite journal|last1=Pierson| first1= E. D.| last2= Rainey| first2= W. E.| year=1992| title= The biology of flying foxes of the genus Pteropus: a review| journal=Biological Report| volume=90| issue=23| url=https://www.researchgate.net/publication/235094381}}</ref>
Twins have been occasionally documented in some species, however. Twins can be [[Twin#Dizygotic (fraternal) twins|fraternal]], [[Twin#Monozygotic (identical) twins|identical]], or the result of [[superfetation]].<ref name="Fox 2008">{{cite journal|doi=10.3161/150811008X414845|title=Analysis of twinning in flying-foxes (Megachiroptera) reveals superfoetation and multiple-paternity|journal=Acta Chiropterologica| volume=10|issue=2 |pages=271|year=2008|last1=Fox|first1=Samantha|last2=Spencer|first2=Hugh|last3=O'Brien|first3=Gemma M|s2cid=83506361}}</ref> Pups are [[altricial]] and sparsely furred at birth, thereby dependent on their mothers for care.<ref name="Pierson 1992"/> Pups are relatively small at birth, weighing approximately 12% of the mother's weight. Bats in other genera can have pups that weigh as much as 30% of the mother's weight at birth.<ref name="Orr 2012">{{cite book| last=Orr| first=R. T.| editor-last=Wimsatt| editor-first= W.| year=2012| chapter=Development: Prenatal and Postnatal| title= Biology of Bats| publisher=Elsevier| pages=221–222| isbn=978-0323151191}}</ref><ref name="Kunz 1987">{{cite journal|last1=Kunz| first1= T. H.| last2= Kurta| first2= A.| year=1987| title= Size of bats at birth and maternal investment during pregnancy| journal= Symposia of the Zoological Society of London| volume=57| url=http://www.bu.edu/cecb/files/2009/08/Symp.pdf}}</ref> They cling to their mothers' abdomens, gripping her fur with their thumb claws and teeth; females carry the pups for the first several weeks of life. After this, the females may leave the pups behind at the roost at night while they forage.<ref name="Pierson 1992"/> As with nearly all bat species, males do not assist females in parental care.<ref name="Safi 2008">{{cite journal|doi=10.1644/08-MAMM-S-058.1|title=Social Bats: The Males' Perspective|journal=Journal of Mammalogy|volume=89|issue=6|pages=1342–1350|year=2008|last1=Safi|first1=Kamran|doi-access=free}}</ref> While male flying foxes of at least one species, the [[Bismarck masked flying fox]], can [[lactation|lactate]], it is unclear if the lactation is functional and males actually nurse pups or if it is a result of [[stress (biology)|stress]] or [[malnutrition]].<ref name="Kunz 2009">{{cite journal|doi=10.1016/j.tree.2008.09.009|pmid=19100649|title=Male lactation: Why, why not and is it care?|journal=Trends in Ecology & Evolution|volume=24|issue=2|pages=80–5|year=2009|last1=Kunz|first1=Thomas H|last2=Hosken|first2=David J|bibcode=2009TEcoE..24...80K }}</ref> Pups [[fledge]] beginning at 3 months old, but may not be [[Weaning|weaned]] until 4&ndash;6 months old. Pups may stay with their mothers until age one. Flying foxes do not reach [[sexual maturity]] until 1.5&ndash;2 years old. Females can have up to two litters annually, though one is the norm due to the long weaning period. Most flying foxes are [[seasonal breeder]]s and give birth in the [[Spring (season)|spring]], though the [[Mariana fruit bat]] seems to have aseasonal breeding with new pups documented throughout the year.<ref name="Pierson 1992"/> Females remain [[fertility|fertile]] with no decrease in reproductive capability for at least the first 12 or 13 years of life.<ref name="McIlwee 2002"/>

Flying foxes, like all bats, are long-lived relative to their size.<ref name="Ma 2015">{{cite journal|doi=10.1016/j.cmet.2015.07.005| pmid=26244935|title=Organization of the Mammalian Metabolome according to Organ Function, Lineage Specialization, and Longevity| journal=Cell Metabolism| volume=22|issue=2|pages=332–43|year=2015|last1=Ma |first1=Siming|last2=Yim|first2=Sun Hee|last3=Lee|first3=Sang-Goo |last4=Kim|first4=Eun Bae|last5=Lee|first5=Sang-Rae|last6=Chang|first6=Kyu-Tae| last7=Buffenstein|first7=Rochelle|last8=Lewis |first8=Kaitlyn N|last9=Park|first9=Thomas J|last10=Miller|first10=Richard A|last11=Clish |first11=Clary B|last12=Gladyshev|first12=Vadim N|pmc=4758382}}</ref> In the wild, average lifespans are likely 15 years.<ref name="Vincenot 2017c"/> However, individuals part of populations that face excessive [[Disturbance (ecology)|disturbance]] may have lifespans as short as 7.1 years.<ref name="Tidemann 2011">{{cite journal|doi=10.3161/150811011X624901|title=Life Expectancy, Causes of Death and Movements of the Grey-Headed Flying-Fox (Pteropus poliocephalus) Inferred from Banding|journal=Acta Chiropterologica|volume=13|issue=2|pages=419|year=2011|last1=Tidemann|first1=Christopher R|last2=Nelson|first2=John E|s2cid=84322011}}</ref> In captivity, individuals can live approximately 20&ndash;28 years.<ref name="Vincenot 2017c"/> The longest-lived flying fox was an Indian flying fox named Statler, who was a resident at Bat World Sanctuary for his last few years. He was born at a zoo in 1987, and was 34 years old at the time of his death. <ref name="Walker 1999"/>

===Social systems===
[[File:Pteropus giganteus colony.JPG|thumb|A roosting colony of Indian flying foxes]]
Most flying fox species are [[gregarious]] and form large aggregations of individuals called [[colony (biology)|colonies]] or "camps." The large flying fox forms colonies of up to 15,000 individuals,<ref name="Reeder 2006">{{cite journal|doi=10.1016/j.yhbeh.2005.11.001|pmid=16380123|title=The hormonal and behavioral response to group formation, seasonal changes, and restraint stress in the highly social Malayan Flying Fox (Pteropus vampyrus) and the less social Little Golden-mantled Flying Fox (Pteropus pumilus) (Chiroptera: Pteropodidae)|journal=Hormones and Behavior|volume=49|issue=4|pages=484–500|year=2006|last1=Reeder|first1=Deeann M|last2=Kosteczko|first2=Nicole S|last3=Kunz|first3=Thomas H|last4=Widmaier|first4=Eric P|s2cid=7443825|doi-access=free}}</ref><ref name="Epstein 2009">{{cite journal|doi=10.1111/j.1365-2664.2009.01699.x|title=Pteropus vampyrus, a hunted migratory species with a multinational home-range and a need for regional management|journal=Journal of Applied Ecology|volume=46|issue=5|pages=991|year=2009|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|doi-access=free|bibcode=2009JApEc..46..991E }}</ref> while the [[little red flying fox]] forms colonies of up to 100,000 individuals.<ref name="Pierson 1992"/> A few species and subspecies, such as [[Ryukyu flying fox|Orii's flying fox]] (''P.&nbsp;dasymallus inopinatus'') and the [[Ceram fruit bat]], are solitary.<ref name="Nakamoto 2012">{{cite journal|doi=10.3161/150811012X661701|title=Ranging Patterns and Habitat Use of a Solitary Flying Fox (Pteropus dasymallus) on Okinawa-jima Island, Japan|journal=Acta Chiropterologica|volume=14|issue=2|pages=387|year=2012|last1=Nakamoto|first1=Atsushi|last2=Kinjo|first2=Kazumitsu|last3=Izawa|first3=Masako|s2cid=84641539}}</ref><ref name="Tsang 2015">{{cite journal|doi=10.1206/3842.1|title=New Records of Flying Foxes (Chiroptera:Pteropussp.) from Seram, Indonesia, with Notes on Ecology and Conservation Status|journal=American Museum Novitates|issue=3842|pages=1–23|year=2015|last1=Tsang|first1=Susan M|last2=Wiantoro|first2=Sigit|last3=Simmons|first3=Nancy B|hdl=2246/6622|s2cid=54209937|url=https://www.biodiversitylibrary.org/item/270956}}</ref>

Colony size varies throughout the year in response to biological needs. The grey-headed flying fox forms [[harem (zoology)|harems]] during the breeding season consisting of one male and up to six females. These colonies break up after the breeding season is over.<ref name="Klose 2009">{{cite journal|doi=10.1098/rsbl.2009.0563|pmid=19689980|pmc=2828011|title=Testosterone is associated with harem maintenance ability in free-ranging grey-headed flying-foxes, Pteropus poliocephalus|journal=Biology Letters|volume=5|issue=6|pages=758–61|year=2009|last1=Klose|first1=S. M|last2=Welbergen|first2=J. A|last3=Kalko|first3=E. K. V}}</ref> In the Bonin flying fox, colony formation is based on both the sex and age of individuals, as well as the season. In the winter breeding season, adult females will form colonies that include a few adult males (likely harems). Adult males who do not roost with females will form colonies with other adult and [[juvenile (organism)|subadult]] males. Subadults will form mixed-sex "subadult groups" with each other. In the summer, however, individuals are solitary, with the exception of [[lactation|nursing]] females, who roost with their pups.<ref name="Sugita 2009"/>

===Diet and foraging===
Flying foxes consume 25–35% of their body weight daily.<ref name="sa.gov.au">{{cite report| title=Guidelines for the Captive Management of Flying Foxes in South Australia| author=Department of Environment and Natural Resources| publisher=South Australian Government|pages=18–19}}</ref> They are [[Generalist and specialist species|generalists]] that will consume a variety of items to meet their nutritional needs. Food items include fruit, flowers, nectar, and leaves.<ref name="Marshall 1985">{{cite journal|doi=10.1111/j.1096-3642.1985.tb01181.x|title=Old World phytophagous bats (Megachiroptera) and their food plants: A survey|journal=Zoological Journal of the Linnean Society|volume=83|issue=4|pages=351–369|year=1985|last1=Marshall|first1=Adrian G}}</ref><ref name="Aziz 2017">{{cite journal|doi=10.7717/peerj.3176|pmid=28413729|pmc=5391789|title=Elucidating the diet of the island flying fox (Pteropus hypomelanus) in Peninsular Malaysia through Illumina Next-Generation Sequencing|journal=PeerJ|volume=5|pages=e3176|year=2017|last1=Aziz|first1=Sheema Abdul|last2=Clements|first2=Gopalasamy Reuben|last3=Peng|first3=Lee Yin|last4=Campos-Arceiz|first4=Ahimsa|last5=McConkey|first5=Kim R|last6=Forget|first6=Pierre-Michel|last7=Gan|first7=Han Ming |doi-access=free }}</ref> They will sometimes deliberately consume insects such as [[cicada]]s as well.<ref name="Clulow 2011">{{cite journal|doi=10.3161/150811011X578750|title=Deliberate Insectivory by the Fruit Bat ''Pteropus'' poliocephalusby Aerial Hunting|journal=Acta Chiropterologica|volume=13|pages=201–205|year=2011|last1=Clulow|first1=Simon|last2=Blundell|first2=Adam T|s2cid=86754699}}</ref>
In Australia, [[eucalyptus|eucalypt]] blossoms and pollen are preferred food sources, followed by ''[[Melaleuca]]'' and ''[[Banksia]]'' flowers <ref name="NWC 2010">{{cite report|url=http://www.nwc.org.au/wp-content/uploads/2016/12/Flying_Fox_Article_June2010.pdf| title=Flying-foxes| author=NWC - NSW Wildlife Council| date=June 2010}}</ref> They feed on a wide variety of [[crop]]s as well, causing conflicts with farmers. Crops eaten by flying foxes include [[sisal]], [[cashew]], [[pineapple]], [[areca]], [[breadfruit]], [[jackfruit]], [[neem]], [[papaya]], [[citrus]], [[ficus|fig]], [[mango]], [[banana]], [[avocado]], [[guava]], [[sugar cane]], [[tamarind]], [[grape]]s, and more.<ref name="Aziz 2016"/>

In captivity, the recommended diet for flying foxes consists of two-thirds hard fruits like pears and apples and one-third soft fruits. Bananas and other high-fiber fruits should only be offered occasionally, as flying foxes are not adapted to high-fiber diets. Protein supplements are recommended for captive flying foxes; other supplements such as [[vitamin C]], [[calcium]], [[chondroitin sulfate]], and [[glucosamine]] can be recommended periodically.<ref name="sa.gov.au"/>

The majority of flying fox species are [[nocturnality|nocturnal]] and forage at night. A few island species and subspecies are [[diurnality|diurnal]], however, hypothesized as a response to a lack of predators. Diurnal taxa include ''[[Black-eared flying fox|P.&nbsp;melanotus natalis]]'', the Mauritian flying fox, the [[Caroline flying fox]], ''[[Pteropus pelagicus|P.&nbsp;p.&nbsp;insularis]]'', and the [[Seychelles fruit bat]]. Foraging resources are often far from roosts, with individuals traveling up to {{cvt|40-60|km|mi}} to reach them.<ref name="Pierson 1992"/> Flying foxes can travel at {{cvt|6|m/s|mph}} for three hours or more, and can reach top speeds of {{cvt|8.6|m/s|mph}}.<ref name="Carpenter 1985">{{cite journal|last1=Carpenter| first1= R. E.| year=1985| title= Flight physiology of flying foxes, Pteropus poliocephalus| journal= Journal of Experimental Biology| volume= 114| issue=1| pages= 619–647| doi= 10.1242/jeb.114.1.619| url=http://jeb.biologists.org/content/jexbio/114/1/619.full.pdf| doi-access= free| bibcode= 1985JExpB.114..619C}}</ref> Some colonial species will forage in groups, especially when resources are abundant. Less social species will forage alone.<ref name="Pierson 1992"/> When they land on a tree with food, they will hang onto the branch with their clawed hind feet and use their clawed thumbs to pull branches bearing flowers or fruits towards them.<ref name="Flick 2011"/> As they forage on fruit, flying foxes will compress the fruit against the [[palate]] with the tongue to squeeze out and consume the juices. The rest of the fruit is then discarded in "ejecta pellets."<ref name="Banack 1998">{{cite journal|doi=10.1890/0012-9658(1998)079[1949:DSARUB]2.0.CO;2|year=1998|volume=79|issue=6|pages=1949–1967|title=Diet Selection and Resource Use by Flying Foxes (Genus: Pteropus)|journal=Ecology|last1=Banack|first1=Sandra Anne}}</ref>

===Role in ecosystems===
Flying foxes have important roles as [[Seed dispersal#Animals: epi- and endozoochory|seed dispersers]] and [[pollinator]]s. They help spread the seeds in the fruit they eat by discarding them in ejecta pellets or through their [[guano]]. In Madagascar, fig seeds have better germination success if they have passed through the gut of a flying fox, which is important because fig trees are a vital [[pioneer species]] in regenerating lost forest. Even though flying foxes can have a gut transit time as fast as 12 minutes, seeds can be retained in the gut for as long as 20 hours. As the flying foxes travel large distances, seeds can be deposited up to {{cvt|20|km|mi}} from the parent tree. They are particularly important in fragmented forests, as many other frugivores are terrestrial and often confined to forest fragments.
Flying foxes have the capability to spread seeds beyond the forest fragments through flight.<ref name="Oleksy 2017">{{cite journal|doi=10.1371/journal.pone.0184023|pmid=28877248|pmc=5587229|title=Flying foxes create extensive seed shadows and enhance germination success of pioneer plant species in deforested Madagascan landscapes|journal=PLOS ONE|volume=12 |issue=9|pages=e0184023|year=2017|last1=Oleksy| first1=Ryszard|last2=Giuggioli| first2=Luca|last3=McKetterick |first3=Thomas J|last4=Racey|first4=Paul A|last5=Jones|first5=Gareth|bibcode=2017PLoSO..1284023O|doi-access=free}}</ref>

Flying foxes pollinate a variety of plants, including the economically valuable [[durian]]. They forage on its nectar in such a way that the flowers (and eventual fruit production) are not usually harmed. Flying fox pollination has a positive effect on durian reproductive success, suggesting that both flying foxes and durian trees [[Mutualism (biology)|benefit]] from this relationship.<ref name="Aziz 2017b">{{cite journal|doi=10.1002/ece3.3213|pmid=29152168|pmc=5677486|title=Pollination by the locally endangered island flying fox (Pteropus hypomelanus) enhances fruit production of the economically important durian (Durio zibethinus)|journal=Ecology and Evolution|volume=7|issue=21|pages=8670–8684|year=2017|last1=Aziz|first1=Sheema A|last2=Clements|first2=Gopalasamy R|last3=McConkey|first3=Kim R|last4=Sritongchuay|first4=Tuanjit|last5=Pathil|first5=Saifful|last6=Abu Yazid|first6=Muhammad Nur Hafizi|last7=Campos-Arceiz|first7=Ahimsa|last8=Forget|first8=Pierre-Michel|last9=Bumrungsri|first9=Sara|bibcode=2017EcoEv...7.8670A }}</ref>

==Conservation==
===Conservation status===
[[File:Flying fox conservation status.jpg|thumb|upright=1.1|IUCN status of ''Pteropus'' species]]
Of the 62 flying fox species evaluated by the [[IUCN]] as of 2018, 3 are considered [[critically endangered]]: the [[Aru flying fox]], Livingstone's fruit bat, and the [[Vanikoro flying fox]]. Another 7 species are listed as [[endangered species|endangered]]; 20 are listed as [[vulnerable species|vulnerable]], 6 as [[near-threatened species|near threatened]], 14 as [[least-concern species|least concern]], and 8 as [[data deficient]]. A further 4 are listed as [[extinct]]: the [[dusky flying fox]], the [[large Palau flying fox]], the [[small Mauritian flying fox]], and the Guam flying fox.<ref name="iucn search">{{cite web| title= The IUCN Red List of Threatened Species Version 2017-3| url=http://www.iucnredlist.org/| access-date=3 June 2018}}</ref> Over half of the species are threatened today with extinction, and in particular in the Pacific, a number of species have died out as a result of hunting, deforestation, and predation by invasive species.<ref name="Vincenot 2017c"/> Six flying fox species are believed to have gone extinct from 1864 to 2014: the Guam, large Palau, small Mauritian, dusky, [[Large Samoan flying fox|large Samoan]], and the [[small Samoan flying fox|small Samoan]] flying foxes.<ref name="Almeida 2014"/>

===Legal status===
All species of ''Pteropus'' are placed on [[CITES#Appendix II|Appendix II]] of [[CITES]] and 10 on [[CITES#Appendix I|Appendix I]], 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=14 April 2018}}</ref> Individual species have different legal protections from hunting and domestic trade that reflect the environmental laws of the countries where they are found.<ref name="Voigt 2016"/>

In some countries such as Bangladesh, Sri Lanka, and Thailand, flying foxes are absolutely protected from harm under the Wildlife Preservation and Security Act of 2012, Fauna and Flora Protection Ordinance of 1937, and Wildlife Protection and Reservation Act of 1992, respectively.<ref name="Voigt 2016"/> However, in Thailand, flying fox poaching and the illegal bushmeat trade still occurs outside of [[protected area]]s. The large flying fox and the small flying fox are particularly prone to poaching and roost disturbance.<ref name="Voigt 2016"/>

In other countries, such as Australia, Japan, and the United States, some species of conservation concern are protected under national environmental legislation, while others are not. In Australia, two flying foxes are listed under the [[Environment Protection and Biodiversity Conservation Act 1999|Environment Protection and Biodiversity Conservation Act of 1999]]: the grey-headed and spectacled flying foxes are listed as "vulnerable."<ref name="environment.au.gov">{{cite web| url=http://www.environment.gov.au/biodiversity/threatened/species/flying-fox-law| title=Flying-foxes| website=environment.gov.au| publisher=Commonwealth of Australia| access-date=7 June 2018}}</ref> Farmers can apply for permits to kill flying foxes when they are causing crop damage.<ref name="Williams 2012">{{cite web| last=Williams| first=Kylie| date=21 October 2012| title=Culling flying-foxes is ineffective, so why suggest slaughter?| website=The Conversation| publisher=The Conversation US | access-date=7 June 2018| url=https://theconversation.com/culling-flying-foxes-is-ineffective-so-why-suggest-slaughter-9817}}</ref>

Several flying fox species occur in Japan. The Bonin flying fox has been a [[Monuments of Japan#Natural Monuments and Special Natural Monuments|Natural Monument]] of Japan since 1969, which means that it is illegal to capture or disturb them without appropriate permits.<ref name="Vincenot 2017">{{cite iucn|last1= Vincenot| first1= C.| year= 2017| title= Pteropus pselaphon| volume=2017| page= e.T18752A22085351 |doi= 10.2305/IUCN.UK.2017-2.RLTS.T18752A22085351.en}}</ref> Two subspecies of the Ryukyu flying fox (''P.&nbsp;d.&nbsp;dasymallus'' and ''P.&nbsp;d.&nbsp;daitoensis'') are also listed as Natural Monuments.<ref name="Vincenot 2017b">{{cite iucn| last1=Vincenot| first1= C.| year= 2017| title= Pteropus dasymallus| volume=2017| page=e.T18722A22080614 |doi=10.2305/IUCN.UK.2017-2.RLTS.T18722A22080614.en}}</ref>
Flying foxes are not designated game species in Japan, and therefore cannot be legally hunted per the Wildlife Protection and Hunting Law.<ref name="env.go.jp">{{cite web| url=https://www.env.go.jp/en/nature/biodiv/law.html|title=Wildlife Protection System and Hunting Law| website=Ministry of the Environment| publisher=Ministry of the Environment Government of Japan| access-date=21 June 2018}}</ref> The Bonin flying fox and ''P.&nbsp;d.&nbsp;daitoensis'' are also listed as National Endangered Species, meaning that they cannot be killed or harmed; furthermore, the sale or transfer of live or dead individuals in whole or part is also prohibited without permits.<ref name="env.go.jp"/><ref name="Saitoh 2015">{{cite journal |doi=10.12933/therya-15-239 |title=Conservation and management of terrestrial mammals in Japan: Its system and practices |journal=Therya |volume=6 |pages=139–153 |year=2015 |last1=Saitoh |first1=Takashi |last2=Kaji |first2=Koichi |last3=Izawa |first3=Masako |last4=Yamada |first4=Fumio|doi-access=free }}</ref>

Despite not occurring in the continental United States, several species and subspecies are listed under its [[Endangered Species Act of 1973]]. ''Pteropus mariannus mariannus''&mdash;a subspecies of the Mariana fruit bat&mdash;is listed as [[Endangered Species Act of 1973#Listing status|threatened]] while the Rodrigues flying fox and Guam flying fox are listed as [[Endangered Species Act of 1973#Listing status|endangered]].<ref name="USFWS 2018">{{cite web|url=https://ecos.fws.gov/ecp0/reports/ad-hoc-species-report?kingdom=V&kingdom=I&status=E&status=T&status=EmE&status=EmT&status=EXPE&status=EXPN&status=SAE&status=SAT&fcrithab=on&fstatus=on&fspecrule=on&finvpop=on&fgroup=on&header=Listed+Animals| title=Listed Animals| website=U. S. Fish & Wildlife Service| publisher=U. S. Federal Government| access-date=7 June 2018}}</ref> Additionally, the U. S. government has been petitioned to list the Aru flying fox and Bonin flying fox as threatened or endangered.<ref name="ESA">{{cite journal| journal=Federal Register| volume=79| issue=14| date=22 January 2014| title=Review of Petition To List 15 Bat Species as Endangered or Threatened Under the Act| url=https://www.gpo.gov/fdsys/pkg/FR-2014-01-22/pdf/2014-01184.pdf}}</ref>

In such countries as India and Pakistan, flying foxes explicitly have no legal protection. In India, they are listed as "[[vermin]]" under the [[Wildlife Protection Act, 1972|Wildlife Protection Act of 1972]].<ref name="Mistry 2003">{{cite web| url=http://www.batcon.org/resources/media-education/bats-magazine/bat_article/44| last=Mistry| first=Shahroukh| title=Protecting the Bats of India: Signs of progress amid daunting challenge| website=Batcon.org| publisher=Bat Conservation International| access-date=7 June 2018}}</ref> Pakistan's only flying fox, the Indian flying fox, is listed under Schedule 4 of the Punjab Wildlife (Protection, Preservation, Conservation and Management) Act of 1974, meaning that it has no legal protections and can be hunted.<ref name="Voigt 2016"/>
In Mauritius, flying foxes were formerly protected but are now legally culled at a large scale. In 2015, the Mauritian government passed the Native Terrestrial Biodiversity And National Parks Act, which legalized culling of the Mauritian flying fox. In Mauritius, over 40,000 Mauritian flying foxes were culled in a two-year period, reducing its population by an estimated 45%.<ref name="Vincenot 2017c"/> This decision was viewed with controversy, with researchers stating "Because they spread seeds and pollinate flowers, flying foxes are vital for regenerating lost forests."<ref name="Racey 2015">{{cite web| url=https://news.nationalgeographic.com/2015/11/151111-bats-mauritius-flying-foxes-animals-science-conservation/| archive-url=https://web.archive.org/web/20151114023651/http://news.nationalgeographic.com/2015/11/151111-bats-mauritius-flying-foxes-animals-science-conservation/| url-status=dead| archive-date=November 14, 2015| title=Opinion: Killing Thousands of Flying Foxes Only Hurts the Environment| date=11 November 2015| last1=Racey| first1=P. A.| last2=Medellín| first2=R. A.| website=National Geographic| publisher=National Geographic Partners, LLC|access-date=7 June 2018}}</ref>

Legal protection can vary within a country as well, such as in [[Malaysia]]. Under the 1990 Protection of Wild Life Amendment Order, flying foxes can be hunted with a permit; each permit is good for killing up to 50 flying foxes. Permits cost U.S.$8 each. However, under the Protection of WildLife Act of 1972, flying foxes can be killed without permits if they are causing damage or if there is "reason to believe that it is about to cause serious damage" to crops.<ref name="Voigt 2016"/> In 2012, the Malaysian state of [[Terengganu]] issued a moratorium on hunting flying foxes.<ref name="Voigt 2016"/>
In [[Sarawak]], all bat species are listed as "Protected" and hunting them is not legal.<ref name="Voigt 2016">{{cite book| last1=Voigt| first1= C. C.| last2= Kingston| first2= T.| year=2016| title=Bats in the Anthropocene: conservation of bats in a changing world| publisher=Springer| doi=10.1007/978-3-319-25220-9| isbn= 978-3-319-25218-6| s2cid= 45637291| url= http://www.oapen.org/search?identifier=1001864}}</ref>

===Factors causing decline===
====Anthropogenic sources====
[[File:Greyheadedflyingfoxbabies2008canungra.jpg|thumb|upright=1.2|Abandoned [[grey-headed flying fox]] pups after a cyclone]]
Flying foxes species are declining or going extinct as a result of several [[Human impact on the environment|human impacts]] to their environments, in addition to natural phenomena.<ref name="Pierson 1992"/> Their populations are especially vulnerable to threats because the litter size is usually only individual and females generally only have one litter per year. Even when nearly every female (90%) successfully produces and raises young, if a population's [[mortality rate]] exceeds 22% annually, then it will steadily decline.<ref name="McIlwee 2002">{{cite journal|doi=10.7882/AZ.2002.008|title=On the intrinsic capacity for increase of Australian flying-foxes (Pteropus spp., Megachiroptera)|journal=Australian Zoologist|volume=32|pages=76–100|year=2002|last1=McIlwee|first1=A. P|last2=Martin|first2=L|doi-access=free}}</ref>

Invasive species, such as the [[brown tree snake]], can seriously affect populations; the brown tree snake consumes so many pups that it reduced the [[recruitment (biology)|recruitment]] of the [[Guam]] population of the Mariana fruit bat to essentially zero.<ref name="Esselstyn 2006">{{cite journal|doi=10.1353/psc.2006.0027|title=Impact of Posttyphoon Hunting on Mariana Fruit Bats (Pteropus mariannus)|journal=Pacific Science|volume=60|issue=4|pages=531|year=2006|last1=Esselstyn|first1=Jacob A|last2=Amar|first2=Arjun|last3=Janeke|first3=Dustin|s2cid=55543225}}</ref>

Many flying fox species are threatened by overhunting. While they have long been a dietary component of indigenous people, [[Population growth|expanding human population]] and more efficient weapons have resulted in population declines, [[local extinction]]s, and extinctions. Overhunting is believed to be the primary cause of extinction for the small Mauritian flying fox and the Guam flying fox.<ref name="Pierson 1992"/>

Flying foxes are also threatened with excessive culling due to conflict with farmers. They are shot, beaten to death, or poisoned to reduce their populations. Mortality also occurs via accidental entanglement into [[Bird netting|netting]] used to prevent the bats from eating fruit.<ref name="Vincenot 2015">{{cite journal|doi=10.1016/j.mambio.2015.03.003|title=Near threatened? First report of unsuspected human-driven decline factors in the Ryukyu flying fox (Pteropus dasymallus) in Japan|journal=Mammalian Biology - Zeitschrift für Säugetierkunde|volume=80|issue=4|pages=273|year=2015|last1=Vincenot|first1=Christian Ernest|last2=Koyama|first2=Lina|last3=Russo|first3=Danilo|bibcode=2015MamBi..80..273V }}</ref> Culling can dramatically reduce flying fox populations.
In Mauritius, over 40,000 Mauritian flying foxes were culled in a two-year period, reducing its population by an estimated 45%.<ref name="Vincenot 2017c">{{cite journal|doi=10.1126/science.aam7582 |pmid=28360279 |title=Can we protect island flying foxes? |journal=Science |volume=355 |issue=6332 |pages=1368–1370 |year=2017 |last1=Vincenot |first1=Christian E |last2=Florens |first2=F. B. Vincent |last3=Kingston |first3=Tigga |bibcode=2017Sci...355.1368V |s2cid=206657272 }}</ref> Flying foxes are also killed by electrocution. In one Australian orchard, it is estimated that over 21,000 bats were electrocuted to death in an 8-week period.<ref name="McIlwee 2002"/> Farmers construct electrified grids over their fruit trees to kill flying foxes before they can consume their crop. The grids are questionably effective at preventing crop loss, with one farmer who operated such a grid estimating that they still lost {{convert|100-120|t|ST}} of fruit to flying foxes in a year.<ref name="Martin 2011">{{cite book|doi=10.7882/FS.2011.039|chapter=Is the fruit you eat flying-fox friendly? The effects of orchard electrocution grids on Australian flying-foxes (Pteropus spp., Megachiroptera)|title=The Biology and Conservation of Australasian Bats|pages=380–390|year=2011|last1=Martin|first1=L|isbn=978-0-9803272-4-3}}</ref> Some electrocution deaths are also accidental, such as when bats fly into [[overhead power line]]s.<ref name="Chlopicki 2016">{{cite web| url=https://www.dailytelegraph.com.au/newslocal/macarthur/electric-wires-threaten-flying-foxes-and-their-new-babies/news-story/2fc9955551b46a552b04a1eac1043df0| title=Electric wires threaten flying foxes and their new babies| date=28 October 2016| last=Chlopicki| first=K. |website=The Daily Telegraph| access-date=26 June 2018}}</ref>

Climate change causes flying fox mortality and a source of concern for species persistence. Extreme heat waves in Australia have been responsible for the [[Australian flying fox die-offs|deaths of more than 30,000 Australian flying foxes]] from 1994 to 2008. Females and young bats are most susceptible to extreme heat, which affects a population's ability to recover.<ref name="Welbergen 2008">{{cite journal|doi=10.1098/rspb.2007.1385|pmid=18048286|pmc=2596826|title=Climate change and the effects of temperature extremes on Australian flying-foxes|journal=Proceedings of the Royal Society B: Biological Sciences|volume=275|issue=1633|pages=419–25|year=2008|last1=Welbergen|first1=J. A|last2=Klose|first2=S. M|last3=Markus|first3=N|last4=Eby|first4=P|url=https://espace.library.uq.edu.au/data/UQ_135282/A629_Welbergen.pdf?Expires=1529895373&Signature=KNW78mh7Rb7dGX1xAxbboOySf0p-H3zYnHRH5UxhGlw4H4UafYhMLbe58eNC7Pt4bSeJktE0N81n61aZDSr4Q8qYmqnGv3qJvGhid05C00FDGzNxJe-ypuk5iggMTS2LfIDaChzkyVjUYP~Rtlp8iiQjwnNiEp0Ckls-DLBYaMx0C6HeU9pr8gQCV1gPLmVglULhCX9tk1FoBDh~nsn9foWVaRb21xp145ef7ZDBFpAARYCRR3gCJLyvYVQjYlP6WtdqyUL~w8X729P6ROL-D0yQ8g-Ofkkx-2Yb~BW2A6RvygctqKvNOUsjQ4o94HmCBirZZmzq0~hsed4X7wYWMg__&Key-Pair-Id=APKAJKNBJ4MJBJNC6NLQ}}</ref> Flying foxes are threatened by sea level rise associated with climate change, as several taxa are endemic to low-lying [[atoll]]s.<ref name="Buden 2013">{{cite journal|doi=10.3897/zookeys.345.5840|pmid=24194666|pmc=3817444|title=Taxonomy, distribution, and natural history of flying foxes (Chiroptera, Pteropodidae) in the Mortlock Islands and Chuuk State, Caroline Islands|journal=ZooKeys|issue=345|pages=97–135|year=2013|last1=Buden|first1=Don|last2=Helgen|first2=Kristofer M|last3=Wiles|first3=Gary|doi-access=free|bibcode=2013ZooK..345...97B }}</ref>

====Natural sources====
Because many species are endemic to a single island, they are vulnerable to random events such as [[typhoon]]s. A 1979 typhoon halved the remaining population of the Rodrigues flying fox. Typhoons result in indirect mortality as well: Because they defoliate the trees, flying foxes are more visible and easily hunted by humans. Food resources for the bats become scarce after major storms, and flying foxes resort to riskier foraging strategies such as consuming fallen fruit off the ground. There, they are more vulnerable to depredation by domestic cats, dogs, and pigs.<ref name="Pierson 1992"/>

Flying foxes are also threatened by disease such as [[tick paralysis]]. Tick paralysis affects the spectacled flying fox, and is responsible for an estimated 1% of its annual mortality.<ref name="Buettner 2013">{{cite journal|doi=10.1371/journal.pone.0073078|pmid=24066028|pmc=3774714|title=Tick Paralysis in Spectacled Flying-Foxes (Pteropus conspicillatus) in North Queensland, Australia: Impact of a Ground-Dwelling Ectoparasite Finding an Arboreal Host|journal=PLOS ONE|volume=8|issue=9|pages=e73078|year=2013|last1=Buettner|first1=Petra G|last2=Westcott|first2=David A|last3=MacLean|first3=Jennefer|last4=Brown|first4=Lawrence|last5=McKeown|first5=Adam|last6=Johnson|first6=Ashleigh|last7=Wilson|first7=Karen|last8=Blair|first8=David|last9=Luly|first9=Jonathan|last10=Skerratt|first10=Lee|last11=Muller|first11=Reinhold|last12=Speare|first12=Richard|bibcode=2013PLoSO...873078B|doi-access=free}}</ref>

===Captive breeding===
[[File:MasoalaHalle Turm.jpg|thumb|upright=0.8|A Rodrigues flying fox flying at the [[Zürich Zoologischer Garten]]]]
Several species of endangered flying fox are bred in captivity to augment their population sizes. Critically endangered Livingstone's fruit bats were taken from the wild starting in 1995 to create a [[captive breeding]] program. All captive individuals remain the property of the [[Comoros|Comorian]] government.<ref name="Smith 2006">{{cite journal|doi=10.1644/792.1|title=Pteropus livingstonii|journal=Mammalian Species|volume=792|pages=1–5|year=2006|last1=Smith|first1=Stephanie J|last2=Leslie|first2=David M|doi-access=free}}</ref> 17 individuals were collected from the wild; with breeding, there are 71 in captivity as of 2017. Individuals are held at the [[Jersey Zoo]] and the [[Bristol Zoo]].<ref name="Richdon 2017">{{cite web| url=https://experiment.com/projects/a-bat-on-the-brink-building-family-trees-in-the-critically-endangered-livingstone-s-fruit-bat-pteropus-livingstonii| title=Building family trees in the Critically Endangered Livingstone's fruit bat| last1=Richdon| first1=Sarah| last2=Rodriguez| first2=Angelica M.| date=2017| website=experiment| access-date=20 June 2018}}</ref> Though the program has been successful in increasing the population, caretakers of the captive population have had to deal with [[Animal husbandry|husbandry]] issues such as [[obesity]] and [[cardiomyopathy]].<ref name="Thorncroft 2014">{{cite journal| last1=Thorncroft| first1= K.| last2= Wormell| first2= D.| last3= Price| first3= E.| year=2014| title= Effects of food distribution method on territoriality, activity and access to food in captive Livingstone's fruit bats (Pteropus livingstonii)| journal= Solitaire| issue=25|pages=22–26|url=https://www.researchgate.net/publication/324414066}}</ref> Relative to their wild counterparts, captive bats have a higher percentage of body fat and a lower percentage of muscle mass. The problem is pronounced in [[Dominance hierarchy|dominant]] males, which are the most [[Sedentary lifestyle|sedentary]]. Addressing these concerns involves increasing flight space so that the animals can exercise adequately.<ref name="Wormell 2018">{{cite journal| last1=Wormell| first1= Dominic| last2=Ramsay| first2=Scott| last3=Masefield| first3=Will| last4=Houston| first4=David| last5=Price| first5=Eluned |year=2018| title= Assessing muscle condition in captive Livingstone's fruit bats|journal=Solitaire| issue=27| url=https://www.researchgate.net/publication/324414151}}</ref> Keepers are also exploring ways of distributing food within enclosures to encourage exercise.<ref name="Thorncroft 2014"/>

The endangered Rodrigues flying fox has been bred in captivity with great success. By 1979, only 70&ndash;100 individuals were left in the world. In 1976, 25 individuals were removed from the wild by [[Durrell Wildlife Conservation Trust]] to begin a breeding program.<ref name="BCI 2016"/> In 1988, the breeding program was called "undoubtedly the most important chiropteran breeding project now in operation."<ref name="Carroll 2007">{{cite journal|doi=10.1111/j.1748-1090.1988.tb03198.x|title=Population management of the Rodrigues fruit bat|journal=International Zoo Yearbook|volume=27|pages=70–78|year=2007|last1=Carroll|first1=J. Bryan|last2=Mace|first2=Georgina M}}</ref> By 2016, there were 180 individuals in 16 zoos across the United States alone.<ref name="BCI 2016">{{cite web| url=http://www.batcon.org/resources/media-education/bats-magazine/bat_article/1568| title=Rodrigues Fruit Bat: Local and International Partners Work Together to Save the 'Golden Bat'| year=2016| website=batcon.org| publisher=Bat Conservation International| access-date=20 June 2018}}</ref> Worldwide, 46 zoos participate in the Rodrigues flying fox breeding program as of 2017.<ref name="Tatayah 2017">{{cite iucn| last1=Tatayah| first1= V.| last2= Jhangeer-Khan| first2= R.| last3= Bégué| first3= J.A.| last4= Jones| first4= C.A.| year= 2017| title= Pteropus rodricensis| volume= 2017| page= e.T18755A22087057| doi= 10.2305/IUCN.UK.2017-2.RLTS.T18755A22087057.en}}</ref>

==Relationship to people==
===Food===
{{See also|Bat as food}}
[[File:Paniki Rica Cabai Hijau.JPG|thumb|An Indonesian dish made with ''Paniki'' (flying fox meat)]]
Many flying foxes species are killed for [[bushmeat]]. The bushmeat harvest is often unsustainable, often resulting in severe population decline or [[local extinction]]. Flying foxes are killed and sold for bushmeat in several countries in Southeast Asia, [[South Asia]], and [[Oceania]], including [[Indonesia]], Malaysia, [[Papua New Guinea]], the [[Philippines]], Bangladesh, China,<ref name="Smith 2010">{{cite book | title=A Guide to the Mammals of China | publisher=Princeton University Press | editor1-first=Andrew T.|editor1-last= Smith| editor2-first= Yan| editor2-last= Xie| editor3-first= Robert S.| editor3-last= Hoffmann| editor4-first= Darrin| editor4-last= Lunde| editor5-first= John| editor5-last= MacKinnon| editor6-first= Don E.| editor6-last= Wilson| editor7-first= W. Chris| editor7-last= Wozencraft| year=2010 | page=222 | isbn=9781400834112|quote=[''Pteropus vampyrus'' is] heavily hunted in some areas, where its large size makes it a useful food item}}</ref> [[Fiji]], and Guam.<ref name="Mickleburgh 2009">{{cite journal|doi=10.1017/S0030605308000938|title=Bats as bushmeat: A global review|journal=Oryx |volume=43 |issue=2 |pages=217|year=2009 |last1=Mickleburgh|first1=Simon |last2=Waylen|first2=Kerry |last3=Racey |first3=Paul|doi-broken-date=18 June 2025 |doi-access=free }}</ref> Flying fox consumption is particularly common in countries with low [[food security]] and lack of environmental regulation.<ref name="Sheherazade 2015"/> In some cultures in the region, however, eating flying fox meat is [[taboo]]. In [[Namoluk]], locals are repulsed by the idea of eating flying foxes because the flying foxes urinate on themselves.<ref name="Buden 2013"/> In predominately [[Muslim]] regions such as much of Indonesia, flying foxes are rarely consumed because of [[halal]] dietary restrictions.<ref name="Sheherazade 2015"/>

[[North Sulawesi]] has the greatest demand for flying fox bushmeat. Despite being in Muslim-majority Indonesia, North Sulawesi is predominately [[Christians|Christian]]; therefore, many locals do not follow halal guidelines prohibiting flying fox consumption. In [[Manado]], most local people consume flying fox meat at least once a month. The frequency of flying fox consumption increases tenfold around holidays.
Locals believe that "unique meat" from undomesticated animals should be served on special occasions to "enliven the atmosphere." Suggestions to make the flying fox bushmeat trade more sustainable include enforcing a quota system for harvesting, encouraging hunters to release female and juvenile individuals, and providing economic alternatives to those who make a living selling flying fox bushmeat.<ref name="Sheherazade 2015">{{cite journal| doi=10.1016/j.gecco.2015.01.003| title=Quantifying the bat bushmeat trade in North Sulawesi, Indonesia, with suggestions for conservation action| journal=Global Ecology and Conservation| volume=3| pages=324–330| year=2015| last1=Sheherazade| last2=Tsang| first2=Susan M| pmid=32363220| pmc=7185848| doi-access=free| bibcode=2015GEcoC...3..324S}}</ref>

In Guam and the [[Commonwealth of the Northern Mariana Islands]], consumption of the Mariana fruit bat exposes locals to the [[neurotoxin]] [[beta-Methylamino-L-alanine]] (BMAA) which may later lead to [[Neurodegeneration|neurodegenerative diseases]]. BMAA may become [[Biomagnification|biomagnified]] in humans who consume flying foxes; flying foxes are exposed to BMAA by eating [[cycad]] fruits.<ref name="Banack 2006">{{cite journal|doi=10.1016/j.jep.2005.12.032|pmid=16457975|title=Neurotoxic flying foxes as dietary items for the Chamorro people, Marianas Islands|journal=Journal of Ethnopharmacology|volume=106|issue=1|pages=97–104|year=2006|last1=Banack|first1=Sandra Anne|last2=Murch|first2=Susan J|last3=Cox|first3=Paul Alan}}</ref><ref name="Cox 2016" >{{cite journal | author = Cox, P., Davis, D., Mash, D., Metcalf J.S., Banack, S. A. | year = 2016 | title = Dietary exposure to an environmental toxin triggers neurofibrillary tangles and amyloid deposits in the brain |journal= Proceedings of the Royal Society B |volume =  283| issue = 3 | doi = 10.1098/rspb.2015.2397|  pages=1–10 | pmid=26791617 | pmc=4795023}}</ref><ref name=Holtcamp>{{cite journal | author = Holtcamp, W. | year = 2012 | title = The emerging science of BMAA: do cyanobacteria contribute to neurodegenerative disease? | journal = Environmental Health Perspectives | volume = 120| issue = 1823 | doi = 10.1289/ehp.120-a110 | pmid=22382274 | pmc=3295368 | pages=a110–a116}}</ref>

===Medicine===
Flying foxes are killed for use in traditional medicine. The Indian flying fox, for example, has many perceived medical uses.
Some believe that its fat is a treatment for [[rheumatism]].<ref name="Walker 1999"/> Tribes in the [[Attappadi]] region of India eat the cooked flesh of the Indian flying fox to treat asthma and chest pain.<ref name="Padmanabhan 208">{{cite journal| last1=Padmanabhan| first1= P.| last2= Sujana| first2= K. A.| date= 2008| title= Animal products in traditional medicine from Attappady hills of Western Ghats| journal=Indian Journal of Traditional Knowledge| volume=7| issue=2| pages=326–329| hdl= 123456789/1595}}</ref> Healers of the Kanda tribe of Bangladesh use hair from Indian flying foxes to create treatments for "fever with shivering."<ref name="Rahmatullah">{{cite journal| last1=Rahmatullah| first1= M.| last2= Ayman| first2= U.| last3= Akter| first3= F.| last4= Sarker| first4= M.| last5= Sifa| first5= R.| last6= Sarker| first6= B.| last7=Chowdhury| first7= S. A.| date=2013| title= Medicinal formulations of a Kanda tribal healer–a tribe on the verge of disappearance in Bangladesh| journal= African Journal of Traditional, Complementary and Alternative Medicines| volume= 10| issue=2| pages= 213–222| doi= 10.4314/ajtcam.v10i2.5| pmid= 24146444| pmc= 3746568}}</ref>

===Transmitting disease===
{{See also|Bat-borne virus}}
[[File:Date Palm Sap Collection - Taki - North 24 Parganas 2015-01-13 4731.JPG|thumb|right|Date palm sap collection, a primary exposure route for Nipah virus]]
[[File:Hendra-distribution-map.jpg|thumb|right|Known henipavirus outbreaks as of 2014]]
Flying foxes are the [[natural reservoir]]s of several viruses, some of which can be [[Zoonosis|transmitted to humans]]. Notably, flying foxes can transmit [[lyssavirus]]es, which cause [[rabies]]. In Australia the [[rabies virus]] is not naturally present; [[Australian bat lyssavirus]] is the only lyssavirus present. Australian bat lyssavirus was first identified in 1996; it is very rarely transmitted to humans. Transmission occurs from the bite or scratch of an infected animal, but can also occur from getting the infected animal's saliva in a [[mucous membrane]] or an open [[wound]]. Exposure to flying fox blood, urine, or feces is not a risk of exposure to Australian bat lyssavirus.
Since 1994, there have been three records of people getting infected with it&mdash;all three were in [[Queensland]] and each case was fatal.<ref name="NSW">{{cite web| url=http://www.health.nsw.gov.au/Infectious/factsheets/Pages/rabies-australian-bat-lyssavirus-infection.aspx| title=Rabies and Australian bat lyssavirus infection fact sheet| date=30 November 2015| website=health.nsw.gov.au| publisher=State of New South Wales NSW Ministry of Health 2015| access-date=14 June 2018}}</ref>

Flying foxes are also reservoirs of [[henipavirus]]es such as [[Henipavirus#Hendra virus|Hendra virus]] and [[Henipavirus#Nipah virus|Nipah virus]]. Hendra virus was first identified in 1994; it also rarely occurs in humans. From 1994 to 2013, there have been seven reported cases of Hendra virus affecting people, four of which were fatal. The hypothesized primary route of human infection is via contact with [[horse]]s that have come into contact with flying fox [[Urination|urine]].<ref name="CDC">{{cite web| url=https://www.cdc.gov/vhf/hendra/pdf/factsheet.pdf| title=Hendra Virus Disease(HeV)| website=cdc.gov| publisher=U.S. Department of Health & Human Services| access-date=14 June 2018}}</ref>
There are no documented instances of direct transmission between flying foxes and humans.<ref name="Sanchez 2016">{{cite journal|doi=10.1371/journal.pntd.0004411|pmid=26829399|pmc=4734781|title=Disease Risk Perception and Safety Practices: A Survey of Australian Flying Fox Rehabilitators|journal=PLOS Neglected Tropical Diseases|volume=10|issue=2|pages=e0004411|year=2016|last1=Sánchez|first1=Cecilia A|last2=Baker|first2=Michelle L |doi-access=free }}</ref> As of 2012, there is a [[vaccine]] available for horses to decrease the likelihood of infection and transmission.<ref name="AVA">{{cite web| url=http://www.ava.com.au/hendra-virus-vaccine| title=The Hendra vaccine| website=ava.com| publisher=The Australian Veterinary Association Ltd (AVA)| date=2018| access-date=14 June 2018| archive-date=14 June 2018| archive-url=https://web.archive.org/web/20180614194613/http://www.ava.com.au/hendra-virus-vaccine| url-status=dead}}</ref>

Nipah virus was first identified in 1998 in Malaysia. Since 1998, there have been several Nipah outbreaks in Malaysia, [[Singapore]], [[India]], and Bangladesh, resulting in over 100 casualties. A 2018 outbreak in [[Kerala|Kerala, India]] resulted in 19 humans infected, of which 17 died.<ref name="Gulland 2018">{{cite web| url=https://www.telegraph.co.uk/news/2018/06/12/nipah-virus-control-india-britain-world-must-alert-signs-infected/ |archive-url=https://ghostarchive.org/archive/20220112/https://www.telegraph.co.uk/news/2018/06/12/nipah-virus-control-india-britain-world-must-alert-signs-infected/ |archive-date=2022-01-12 |url-access=subscription |url-status=live| title=Nipah virus 'under control' in India – but Britain and the world must be alert for signs of infected travellers| last=Gulland| first=Anne| date=12 June 2018| website=The Telegraph| publisher=Telegraph Media Group Limited 2018| access-date=14 June 2018}}{{cbignore}}</ref> The overall fatality rate is 40–75%. Humans can contract Nipah virus from direct contact with flying foxes or their fluids, through exposure to an intermediate [[host (biology)|host]] such as [[domestic pig]]s, or from contact with an infected person.<ref name="WHO">{{cite web| url=https://www.who.int/news-room/fact-sheets/detail/nipah-virus| title=Nipah virus| website=World Health Organization| publisher=WHO| date=30 May 2018| access-date=14 June 2018}}</ref> A 2014 study of the Indian flying fox and Nipah virus found that while Nipah virus outbreaks are more likely in areas preferred by flying foxes, "the presence of bats in and of itself is not considered a risk factor for Nipah virus infection." Rather, the consumption of [[date palm#Sap|date palm sap]] is a significant route of transmission. The practice of date palm sap collection involves placing collecting pots at date palm trees. Indian flying foxes have been observed licking the sap as it flows into the pots, as well as defecating and urinating in proximity to the pots. In this way, humans who drink the palm sap can be exposed to the bats' viruses.<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> The use of bamboo skirts on collecting pots lowers the risk of contamination from bat fluids.<ref name="Hahn 2014">{{cite journal| last1=Hahn| first1= M. B.| last2=Epstein |first2= J. H.| last3= Gurley| first3= E. S.| last4= Islam| first4= M. S.| last5= Luby| first5= S. P.| last6= Daszak| first6= P.| last7=Patz| first7= J. A.| date=2014| title= Roosting behaviour and habitat selection of Pteropus giganteus reveal potential links to Nipah virus epidemiology| journal=Journal of Applied Ecology| volume= 51| issue=2| pages=376–387| doi=10.1111/1365-2664.12212| pmid= 24778457| pmc=4000083| bibcode= 2014JApEc..51..376H}}</ref>

Flying foxes can transmit several non-lethal diseases as well, such as [[Menangle virus]]<ref name="Bowden 2001">{{cite journal|doi=10.1006/viro.2001.0893|pmid=11336561|title=Molecular Characterization of Menangle Virus, a Novel Paramyxovirus which Infects Pigs, Fruit Bats, and Humans|journal=Virology|volume=283|issue=2|pages=358–73|year=2001|last1=Bowden|first1=Timothy R|last2=Westenberg|first2=Marcel|last3=Wang|first3=Lin-Fa|last4=Eaton|first4=Bryan T|last5=Boyle|first5=David B|doi-access=free}}</ref> and [[Nelson Bay virus]].<ref name="Yamanaka 2014">{{cite journal|doi=10.1371/journal.pone.0092777|pmid=24667794|title=Imported Case of Acute Respiratory Tract Infection Associated with a Member of Species Nelson Bay Orthoreovirus|journal=PLOS ONE|volume=9|issue=3|pages=e92777|year=2014|last1=Yamanaka|first1=Atsushi|last2=Iwakiri|first2=Akira|last3=Yoshikawa|first3=Tomoki|last4=Sakai|first4=Kouji|last5=Singh|first5=Harpal|last6=Himeji|first6=Daisuke|last7=Kikuchi|first7=Ikuo|last8=Ueda|first8=Akira|last9=Yamamoto|first9=Seigo|last10=Miura|first10=Miho|last11=Shioyama|first11=Yoko|last12=Kawano|first12=Kimiko|last13=Nagaishi|first13=Tokiko|last14=Saito|first14=Minako|last15=Minomo|first15=Masumi|last16=Iwamoto|first16=Naoyasu|last17=Hidaka|first17=Yoshio|last18=Sohma|first18=Hirotoshi|last19=Kobayashi|first19=Takeshi|last20=Kanai|first20=Yuta|last21=Kawagishi|first21=Takehiro|last22=Nagata|first22=Noriyo|last23=Fukushi|first23=Shuetsu|last24=Mizutani|first24=Tetsuya|last25=Tani|first25=Hideki|last26=Taniguchi|first26=Satoshi|last27=Fukuma|first27=Aiko|last28=Shimojima|first28=Masayuki|last29=Kurane|first29=Ichiro|last30=Kageyama|first30=Tsutomu|display-authors=10|bibcode=2014PLoSO...992777Y|pmc=3965453|doi-access=free}}</ref> These viruses rarely affect humans and few cases have been reported.<ref name="Bowden 2001"/><ref name="Yamanaka 2014"/> While other bat species have been suspected or implicated as the reservoir of diseases such as [[Severe acute respiratory syndrome|SARS]] and [[Ebola virus disease|Ebola]], flying foxes are not suspected as hosts for either causative virus.<ref name="Smith 2013">{{cite journal|doi=10.1016/j.coviro.2012.11.006|pmid=23265969|title=Bats and their virome: An important source of emerging viruses capable of infecting humans|journal=Current Opinion in Virology|volume=3|issue=1|pages=84–91|year=2013|last1=Smith|first1=Ina|last2=Wang|first2=Lin-Fa|pmc=7102720}}</ref>

===Pests===
[[File:Protecting Grape Vines from Birds. (16371452799).jpg|thumb|Netting used to protect crops from wildlife such as flying foxes]]
Flying foxes are often considered [[pest (organism)|pests]] due to the damage they cause to [[orchard]] crops. Flying foxes have been cited as particularly destructive to [[almond]]s, [[guava]]s, and [[mango]]es in the Maldives; [[lychee]] in Mauritius; [[areca]] in India; and [[stone fruit]]s in Australia. Orchard damages from other animals are often misattributed to flying foxes, though, and economic damage can be difficult to quantify or exaggerated.<ref name="Aziz 2016">{{cite book| last1=Aziz| first1= S.A.| last2= Olival| first2= K.J.| last3= Bumrungsri| first3= S.| last4= Richards| first4= G.C.| last5= Racey| first5= P.A.| year=2016| chapter= The Conflict Between Pteropodid Bats and Fruit Growers: Species, Legislation and Mitigation| editor1-last= Voigt| editor1-first= C.| editor2-last= Kingston| editor2-first= T.|title= Bats in the Anthropocene: Conservation of Bats in a Changing World| publisher= Springer| isbn=978-3-319-25220-9}}</ref> To prevent fruit damage, farmers may legally or illegally cull flying foxes. In the 1800s, the Australian government paid farmers [[Bounty (reward)|bounties]] to kill flying foxes, though the practice has since been discontinued.<ref name="Hall 2000">{{cite book| last1=Hall| first1= L. S.| last2= Richards| first2= G.| year=2000| title= Flying foxes: fruit and blossom bats of Australia| publisher= UNSW Press| pages=52–53| isbn=978-0868405612}}</ref>
Alternatives to culling include placing barriers between the bats and fruit trees, such as netting, or harvesting fruit in a timely manner to avoid attracting as many flying foxes.<ref name="Dolbeer 1988">{{cite journal| last1=Dolbeer| first1= R. R.| last2= Fiedler| first2= L. R.| last3= Rasheed| first3= H.| date=1988| title= Management of fruit bat and rat populations in the Maldive Islands, Indian Ocean| journal=Proceedings of the Thirteenth Vertebrate Pest Conference | volume=1988| url=https://digitalcommons.unl.edu/vpcthirteen/24/}}</ref> Netting is the most effective way to prevent crop loss, though some farmers find it cost prohibitive. It costs US$4,400&ndash;44,000 to net {{cvt|1|ha|acre}} of crops.<ref name="Voigt 2016"/> Other methods of preventing fruit loss may also involve the use of scare guns, chemical deterrents, or night-time lights. Alternatively, planting [[Muntingia|Singapore cherry trees]] and other [[Trap crop|decoy crops]] next to an orchard can be effective, as flying foxes are much more attracted to their fruits than many other orchard crops.<ref name="Singaravelan 2009">{{cite journal| last1=Singaravelan| first1= N.| last2= Marimuthu| first2= G.| last3= Racey| first3= P. A.| date=2009| title= Do fruit bats deserve to be listed as vermin in the Indian Wildlife (Protection) & Amended Acts? A critical review| journal= Oryx| volume= 43| issue=4| pages= 608–613| doi=10.1017/S0030605309990391| doi-access= free}}</ref>

The location of flying fox camps can be a disturbance to humans. In [[Batemans Bay|Batemans Bay, Australia]], locals report being so disturbed by flying fox vocalizations in the morning that they lose sleep. Flying foxes can fly into power lines and cause electricity outages. Their guano and body odor are also unpleasant to smell.<ref name="AP 2016">{{cite web| url=https://www.theguardian.com/world/2016/jun/11/it-smells-so-bad-you-can-taste-it-bats-plague-australian-tourist-town| title='It smells so bad you can taste it': bats plague Australian tourist town | agency=Associated Press| date=10 June 2018| website=The Guardian| publisher=Guardian News and Media Limited| access-date=20 June 2018}}</ref> The presence of flying fox colonies can cause nearby property values to decline.<ref name="McKenzie 2016">{{cite web| url=https://www.sunshinecoastdaily.com.au/news/residents-rapt-after-nuisance-bats-evicted/3012832/|title=Residents rapt after nuisance bats evicted | last=McKenzie| first=Jessica| date=1 May 2016| website=Sunshine Coast Daily| publisher=Sunshine Coast Newspaper Company Pty Ltd| access-date=22 June 2018}}</ref>

===In culture===
[[File:Red infill flying fox - Google Art Project.jpg|thumb|left|upright=0.9|A flying fox depicted in [[Aboriginal Australians|Aboriginal]] art]]
[[File:Handbook to the ethnographical collections (1910) (14596817827).jpg|thumb|upright=0.7|Ceremonial axe decorated with flying fox fur braid in New Caledonia]]
[[File:Flying fox cord.jpg|thumb|right|upright=0.7|Flying fox braided fur cord used as currency in the Loyalty Islands]]

Flying foxes are featured in many indigenous cultures and traditions.

A [[folklore]] [[Dreamtime]] story from the [[New South Wales North Coast]] in Australia features an impatient flying fox wanting the Great Spirit to teach him how to be a bird, only to be hung upside down on a branch.<ref name="Campbell 2013">{{cite book| last1= Campbell-McLeod| first1=P.|chapter=Nallawilli-Sit Down (and Listen): The Dreamtime Stories--An Oral Tradition| editor-last=MacDonald| editor-first=M. R.| year= 2013| title=Traditional Storytelling Today: An International Sourcebook| pages=158–159| publisher=Routledge| isbn=978-1135917142}}</ref> They were also featured in Aboriginal cave art, as evinced by several surviving examples.<ref name="Parish 2012">{{cite book| last1=Parish| first1= S.| last2= Richards| first2= G.| last3= Hall| first3= L.| year=2012| title= A natural history of Australian Bats: working the night shift| publisher= Csiro Publishing| isbn=978-0643103764}}</ref>

In [[Tonga]], flying foxes are considered sacred. All flying foxes are the property of the king, meaning non-royal persons cannot harm them in any way. Tongan legend states that a colony of flying foxes at [[Kolovai]] are the descendants of a pair of flying foxes gifted to the King of Tonga by the Princess of Samoa.<ref name="Grant 1996">{{cite web| url=http://www.batcon.org/component/batmag/bat_article/746| last=Grant| first=G. S.| year=1996| title=Kingdom of Tonga: Safe Haven for Flying Foxes| website=batcon.org| publisher=Bat Conservation International| access-date=21 June 2018}}</ref>

In the Indian village of Puliangulam, a colony of Indian flying foxes roosts in a [[Banyan]] tree. Villagers believe that the flying foxes are under the protection of [[Muniandi|Muni]], and do not harm the bats. A shrine to Muni is beneath the tree.
If locals believe that they have offended Muni by failing to protect the bats, they will pray and perform [[Puja (Hinduism)|puja]] after offering sweet rice, coconut, and bananas to those attending the ceremony.<ref name="Marimuthu 1988">{{cite web| url=http://www.batcon.org/resources/media-education/bats-magazine/bat_article/324| last=Marimuthu| first=G. |year=1988|title=The Sacred Flying Fox of India| website=batcon.org|publisher=Bat Conservation International| access-date=21 June 2018}}</ref>

Flying foxes are also featured in folk stories from Papua New Guinea. Stories with flying foxes include a legend about a [[cockatoo]] stealing feathers from the flying fox, resulting in it becoming nocturnal. Another story features a flying fox that could transform into a young man; the flying fox stole a woman away from her husband to take as his wife. Another legend states that a flying fox-man was responsible for introducing [[Yam (vegetable)|yam]]s to their people.<ref name="Slone 2001">{{cite book| editor-last=Slone| editor-first= T. H.| year= 2001| title= One Thousand One Papua New Guinean Nights: Tales from 1972–1985| volume=1| publisher= Masalai Press| isbn=978-0971412712}}</ref>

Indigenous societies in Oceania used parts of flying foxes for functional and ceremonial weapons. In the Solomon Islands, people created barbs out of their bones for use in spears.<ref name="BM 1910"/> In New Caledonia, [[Ceremonial weapon|ceremonial axes]] made of [[jade]] were decorated with braids of flying fox fur.<ref name="Machray 1899">{{cite journal| last=Machray| first=Robert| year=1899| title=Strange Kinds of Money| journal=The Harmsworth Monthly Pictorial Magazine| volume=1|pages=639–641| url=https://books.google.com/books?id=HvIRAAAAYAAJ&q=flying%20fox%20fur%20cord&pg=PA640}}</ref> Flying fox wings were depicted on the war shields of the [[Asmat people]] of Indonesia; they believed that the wings offered protection to their warriors.<ref name="Werness 2003">{{cite book| last1=Werness| last2= H. B.| year=2003| title= Continuum Encyclopedia of Native Art: Worldview, Symbolism, and Culture in Africa, Oceania, and North America| publisher= A&C Black| page=31|isbn=978-0826414656}}</ref>

There are modern and historical references to flying fox byproducts used as currency. In New Caledonia, braided flying fox fur was once used as currency.<ref name="BM 1910">{{cite book| last1= British Museum. Dept. of British and Mediaeval Antiquities and Ethnography| last2= Joyce| first2= T. A.| last3=Dalton| first3= O. M.| title=Handbook to the Ethnographical Collections| year= 1910| publisher=Printed by order of the Trustees| page=125| url=https://archive.org/stream/handbooktoethnog00brit/handbooktoethnog00brit#page/125/mode/1up}}</ref> On the island of [[Makira]], which is part of the Solomon Islands, indigenous peoples still hunt flying foxes for their teeth as well as for bushmeat. The [[Canine tooth|canine teeth]] are strung together on necklaces that are used as [[currency]].<ref name="Choi 2017">{{cite web| last=Choi| first=Charles| date=16 October 2017| title=In Makira, Flying Fox Teeth Are Currency…And That Could Save the Species| website=Discover| publisher=Kalmbach Media| url=http://blogs.discovermagazine.com/d-brief/2017/10/16/flying-fox-currency/#.WxLrZyAh1PY| access-date=2 June 2018| archive-date=12 June 2018| archive-url=https://web.archive.org/web/20180612223317/http://blogs.discovermagazine.com/d-brief/2017/10/16/flying-fox-currency/#.WxLrZyAh1PY| url-status=dead}}</ref> Teeth of the [[insular flying fox]] are particularly prized, as they are usually large enough to drill holes in. The [[Makira flying fox]] is also hunted, though, despite its smaller teeth. Deterring local peoples from using flying fox teeth as currency may be detrimental to the species, with Lavery and Fasi noting, "Species that provide an important cultural resource can be highly treasured." Emphasizing [[sustainable hunting]] of flying foxes to preserve cultural currency may be more effective than encouraging the abandonment of cultural currency. Even if flying foxes were no longer hunted for their teeth, they would still be killed for bushmeat; therefore, retaining their cultural value may encourage sustainable hunting practices.<ref name="Lavery 2017">{{cite journal|doi=10.1017/S0030605317001004|title=Buying through your teeth: Traditional currency and conservation of flying foxes Pteropus spp. In Solomon Islands|journal=Oryx|volume=53|issue=3|pages=1–8|year=2017|last1=Lavery|first1=Tyrone H|last2=Fasi|first2=John|doi-access=free}}</ref> Lavery stated, "It’s a positive, not a negative, that their teeth are so culturally valuable. The practice of hunting bats shouldn’t necessarily be stopped, it needs to be managed sustainably."<ref name="Choi 2017"/>

===Other uses===
Flying foxes and other bat species in Southeast Asia are often killed and sold as "mummies". The mummified bodies or skeletons of these bats are often shipped to the United States where they are sold in souvenir or curiosity shops or online through vendors such as [[Etsy]] or [[eBay]]. From 2000 to 2013, over 100,000 dead bats were imported to the United States. Bat conservationist [[Merlin Tuttle]] wrote, "I've seen huge losses, mostly due to various kinds of over-harvesting, especially at cave entrances, either for food or for sale as mummies." Despite sometimes being advertised as "sustainable," the practice could lead to overharvesting and depletion of flying fox species, with Tuttle saying, "It is a virtual certainty that the bats you've seen advertised are not sustainably harvested."<ref name="Hugo 2017">{{cite web| url=http://www.newsweek.com/thousands-bats-slaughtered-annually-asia-end-ebay-and-etsy-artsy-americans-681147| last=Hugo|first=Kristin| date=10 October 2017| title=Thousands of Bats Slaughtered Annually in Asia End Up on eBay and Etsy for Artsy Americans| website=Newsweek| publisher=Newsweek LLC| access-date=2 June 2018}}</ref>

==References==
{{Reflist|2}}

{{Pteropodidae|P.}}
{{Taxonbar|from=Q1241332}}

{{good article}}

[[Category:Pteropus| ]]
[[Category:Bats of Asia]]
[[Category:Bats of Africa]]
[[Category:Fauna of Southeast Asia]]
[[Category:Mammals of Thailand]]
[[Category:Articles containing video clips]]
[[Category:Bat genera]]
[[Category:Taxa named by Mathurin Jacques Brisson]]
[[Category:Taxa described in 1762]]