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Pteropus vampyruslarge flying fox

By Kelsie Norton

Geographic Range

Pteropus vampyrus is found from Madagascar to Australia and in mainland Asia and Indonesia. It occurs in most of continental and insular Southeast Asia; from southern Myanmar and southern Vietnam through Malaysia to Singapore. It is present throughout most of Indonesia. This species is also found in southern Burma and southern Thailand and extends to the east Philippines, Sumatra, and Timor. Pteropus vampyrus is native to China, Indonesia, Malaysia, Myanmar, the Philippines, Singapore, Thailand, Timor-Leste, and Vietnam. (Bates, et al., 2011; Kunz and Jones, 2000; Tampa's Lowry Park Zoo, 2010; "ARKive Images of Life on Earth", 2011)

Habitat

Large flying foxes inhabit tropical forests and swamps. They occur primarily in secondary forests and use agricultural areas during forging bouts. Populations also occur on oceanic islands. During the day, groups often roost in large trees. Roost sites are often used for many years and trees become stripped of bark and foliage by the bats' sharp claws. Roosting trees are often found in mangrove forests, coconut groves, and mixed fruit orchards. Studies in Subic Bay, Philippines have shown that foraging locations range between 0.4 and 12 km from the roost. They prefer undisturbed forests in lowlands, beaches, and mangroves, for roosting and select against disturbed and agricultural areas. Large flying foxes are commonly found in riparian areas. (Tampa's Lowry Park Zoo, 2010; Bates, et al., 2011; Mildenstein, et al., 2005; Tampa's Lowry Park Zoo, 2010)

Physical Description

Pteropus vampyrus is one of the largest bats in the world. Forearm length ranges from 180 to 220 mm, mean wingspan is 1.5 m, and body mas ranges from 0.6 to 1.1 kg. It has long pointed ears and a dog-like or fox-like face and head. Pelage varies in color and texture with age and sex . Upper dorsal fur is short and stiff, with longer, woolier fur on the venter. Head and upper body are covered with a dark mantle ranging in color from mahogany-red to black, and the venter is often darker than the rest of the body. Its wings have short rounded tips. Except for parts close to the body and the edge of the wing membrane, wing membranes lack fur. Young are born with dark skin and fur but become paler as they develop. Males have thicker and stiffer pelage than females and glandular neck tufts with dark bases. Pteropus vampyrus is different from most Pteropus in that it has darker underparts and a dark mantle. For example, P. giganteus and P. lylei have pale underparts that contrast with the darker dorsal pelage. Pelage color occasionally varies as a few P. vampyrus specimens have lighter mantles, and some have a gray or silver venter. (Hollister, 1913; Kunz and Jones, 2000; "ARKive Images of Life on Earth", 2011)

Large flying foxes have robust skulls, with a nearly complete orbit and a thick, wide zygomatic arch. The postorbital processes reaches more than halfway to the zygomatic arch. The dental formula is 2/2, 1/1, 3/3, 2/3 for a total of 34 teeth. Upper canines have a prominent anterior groove and a smaller groove on the inner surface. (Hollister, 1913; Kunz and Jones, 2000; "ARKive Images of Life on Earth", 2011)

  • Sexual Dimorphism
  • male larger
  • sexes colored or patterned differently
  • Range mass
    0.6 to 1.1 kg
    1.32 to 2.42 lb
  • Average wingspan
    1.5 m
    4.92 ft

Reproduction

Pteropus vampyrus is most often polygynous, with males protecting a small harem and mating with up to ten females. Dominant males occupy the best roosting sites, and there is a social hierarchy among males. (Organization for Bat Conservation, 2011; Tampa's Lowry Park Zoo, 2010)

Unlike other pteropods, which have fused horns on the baculum, Pteropus vampyrus has a saddle-shaped baculum. It ranges from 4.5 to 8.2 mm and is wider than it is long. Females typically give birth to only one offspring per year. Synchronous birthing occurs within each population, and timing depends on local geography and seasonality. In peninsular Malaysia, mating peaks from November to January. In Thailand, birthing peaks during March and April and in the Philippines it peaks during April and May. In captive populations, birthing peaks during May and June. Mean body mass newborns is 133g (20-30% of maternal body mass) and their forearm length is around 79.5 mm. Mothers carry their young during the first few days after parturition, then leave them at the roost during foraging bouts. Young are weaned by 2 to 3 months after birth. (Kunz and Jones, 2000)

  • Breeding interval
    once yearly
  • Breeding season
    Mating season varies according to local geography and climate
  • Range number of offspring
    1 to 2
  • Average number of offspring
    1
  • Range gestation period
    140 to 192 days
  • Range weaning age
    2 to 3 months
  • Average age at sexual or reproductive maturity (female)
    2 years
  • Average age at sexual or reproductive maturity (male)
    2 years

Pteropus vampyrus offspring are born with eyes open and fully furred, but in many ways are altricial. They are carried with the mother initially, then left in the roost after a couple of days so that the mother can feed. Mothers nurse young for the first 2 to 3 months after parturition, or until weaning is complete, which can be anywhere from 3 to 5 months. All parental care is provided by the mother, though males often help protect and defend their harem. (Kunz and Jones, 2000; Organization for Bat Conservation, 2011)

  • Parental Investment
  • altricial
  • female parental care
  • pre-fertilization
    • provisioning
    • protecting
      • male
  • pre-hatching/birth
    • provisioning
      • female
    • protecting
      • male
      • female
  • pre-weaning/fledging
    • provisioning
      • female
    • protecting
      • male
      • female
  • pre-independence
    • provisioning
      • female
    • protecting
      • female

Lifespan/Longevity

Pteropus vampyrus can live 15 to 30 years in captivity, and an average of 15 years in the wild. (Kunz and Jones, 2000; Oakland Zoo, 2011; Organization for Bat Conservation, 2011; Tampa's Lowry Park Zoo, 2010)

  • Average lifespan
    Status: wild
    15 years
  • Range lifespan
    Status: captivity
    15 to 30 years

Behavior

Large flying foxes are highly social and vocal animals that live in groups sometimes numbering in the thousands. They prefer to roost in tall trees that rise above the forest canopy. Roost sites are often loud and may include several species. Large flying foxes are nocturnal, leaving the roost at around sunset and returning at dawn. Some individuals fly up to 50 km each night to reach their feeding grounds. They often fly the same route to a feeding ground, returning until all resources are exhausted. They form groups ranging from 2 to 50 at feeding grounds. They usually land on the tips of the branches and fall into a position with their head down to feed. (Kunz and Jones, 2000; Oakland Zoo, 2011; Tampa's Lowry Park Zoo, 2010)

Large flying foxes are highly territorial and communicate ownership by spreading their wings, growling, or making other vocalizations. The presence of flowers on trees appears to encourage territorial behavior. They are often met with hostile vocalizations and aggressive behavior that promotes spacing between roosting individuals. Large flying foxes rest by hanging upside down with wings wrapped around their bodies. During the warmest periods of the day, they sometimes cool themselves by fanning their wings, licking their bodies, or by panting. Roost activity increases as the day progresses and may include short flights around the roost. (Kunz and Jones, 2000; Oakland Zoo, 2011; Tampa's Lowry Park Zoo, 2010)

Home Range

There is little information available concerning the home range of Pteropus vampyrus. Satellite telemetry shows that large adult males are extremely mobile and travel hundreds of kilometers between roosting sites within a single year. (Epstein, et al., 2009)

Communication and Perception

During flight, Pteropus vampyrus tends to remain silent. During feeding, however, it is often very noisy. Returning to the roost at dawn is also associated with loud vocalizations, which are used in territorial behavior and to help maintain inter-individual spacing at roost sites. Evidence suggests that P. vampyrus vocalizes to communicate emotion as well. It uses vision rather then echolocation for in-flight navigation, and they find food using their acute senses of sight and smell. (Mohd-Azlan, et al., 2011; Oakland Zoo, 2011)

Food Habits

Large flying foxes feed on flowers, nectar, and fruit such as bananas and mangos. Common foods also include the pollen and flowers of coconut, durian, and fig trees. When foraging, they slice the rind with their teeth and extract the fruit with their long tongue, which is also used for lapping up nectar. They can carry up to 200 g of food at a time. On a daily basis, large flying foxes can eat half of their body weight in fruit. (Kunz and Jones, 2000; Oakland Zoo, 2011)

  • Plant Foods
  • leaves
  • fruit
  • nectar
  • pollen
  • flowers

Predation

Pteropus vampyrus is commonly hunted for food and sport throughout its geographic range. In peninsular Malaysia, an estimated 22,000 flying foxes are legally hunted each year. No other information exits on potential predators of this species. Its nocturnal and roosting lifestyle likely decreases risk of predation. (Ardea, 2009)

Ecosystem Roles

Flying foxes are important frugivores in tropical forests. They serve as seed dispersers and pollinators of forest trees, including durian, which produce a high-priced fruit that is considered a delicacy in southeast Asia. Members of the genus Pteropus are often the only seed dispersers or pollinators large enough to carry the large fruit they feed upon. They also pollinate canopy trees when searching for nectar. Seed dispersal by large flying foxes is thought to play an important role in the regeneration of cleared forests. They are host to the parasitic nematode Litmosa maki, which inhabits the abdominal cavity. In malaysia, ectoparasites from the families Laelapidae, Nycteribiidae, and Spinturnicidae have also been reported. (Ardea, 2009; Brown, 1997; Mohd-Azlan, et al., 2011; Muscarella and Fleming, 2007)

  • Ecosystem Impact
  • disperses seeds
  • pollinates
Commensal/Parasitic Species
  • parasitic nematode (Litmosa maki)
  • mites (Laelapidae)
  • mites (Spinturnicidae)
  • bat flies (Nycteribiidae)

Economic Importance for Humans: Positive

As a pollinator and seed disperser, Pteropus vampyrus plays an important role in maintaining forest structure and composition throughout its geographic range. Humans hunt P. vampyrus for sport, and there is a significant international market, both legal and illegal, for its meat and the various by-products that are used in traditional medicines. For example, its fat is valued in Pakistan, where some believe it helps cure rheumatism, and others claim that its meat can help cure asthma. Shipments of P. vampyrus carcasses have been confiscated on numerous occasions in Guam, and trade continues to be monitored. (Feldhamer and Drickamer, 2007; Kunz and Jones, 2000; Struebig, et al., 2007; Tampa's Lowry Park Zoo, 2010)

Economic Importance for Humans: Negative

In some areas, large flying foxes are viewed as agricultural pests, as forage sites often include fruit farms. They are also exceptionally noisy while feeding, and many farmers use flapping or whirling devices and bright lights to deter them. Large flying foxes carry a number of zoonotic diseases such as the Hendra virus and the Nipah virus. Nipah virus first appeared in humans in Malaysia in 1998, followed by cases in Bangladesh and India. Evidence suggests that large flying foxes are reservoirs for a number of different henipaviruses, including Nipah. In particular, it is suspected that this species was the reservoir hosts of the 1998 Nipah outbreak in pigs and humans. The long distance movements of large flying foxes increases its potential to transfer these diseases to other countries in the Australian and Asian regions. (Breed, et al., 2010; Hassan, et al., 2010; Kunz and Jones, 2000)

Conservation Status

The IUCN Red List of Threatened Species classifies Pteropus vampyrus. Its populations are in significant decline, and major threats include chronic over-harvesting and the destruction of its primary habitats. If harvesting continues at its current rate, P. vampyrus could disappear from western Malaysia in as little as 6 years. In the Philippines, major threats include hunting and targeting due to noise. Hunting of P. vampyrus has more than doubled since 1996. Likewise, in 2003, an estimated 4,500 large flying foxes were killed in a single location and sold into trade. Habitat destruction due to deforestation is also a major threat to their persistence in Malaysia. (Ardea, 2009; Bates, et al., 2011; Epstein, et al., 2009; Kunz and Jones, 2000; Struebig, et al., 2007)

The Convention on International Trade in Endangered Species of Fauna and Flora (CITES) lists Pteropus vampyrus under Appendix II. Efforts to conserve P. vampyrus are underway, and hunting laws have been established to reduce harvest rates. Unfortunately, anti-poaching laws and harvest limits are difficult to enforce. Currently, colony management on small islands appears to be the most effective protection of P. vampyrus. Due to its migratory behavior and its tendency to travel long distances during nocturnal foraging bouts, a more regional management approach may be necessary to successfully conserve this species. (Ardea, 2009; Bates, et al., 2011; Epstein, et al., 2009; Kunz and Jones, 2000)

Other Comments

Synonyms include for Pteropus vampyrus include Pteropus caninus, Pteropus celaeno, Pteropus edulis, Pteropus funereus, Pteropus javanicus, Pteropus kalou, Pteropus kelaarti, Pteropus kopangi, Pteropus lanensis, Pteropus malaccensis, Pteropus natunae, Pteropus nudus, Pteropus phaiops, Pteropus pluton, Pteropus pteronotus, and Pteropus sumatrensis. ("ARKive Images of Life on Earth", 2011)

Contributors

Kelsie Norton (author), University of Michigan-Ann Arbor, Phil Myers (editor), University of Michigan-Ann Arbor, John Berini (editor), Animal Diversity Web Staff.

Glossary

acoustic

uses sound to communicate

agricultural

living in landscapes dominated by human agriculture.

altricial

young are born in a relatively underdeveloped state; they are unable to feed or care for themselves or locomote independently for a period of time after birth/hatching. In birds, naked and helpless after hatching.

arboreal

Referring to an animal that lives in trees; tree-climbing.

bilateral symmetry

having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.

causes or carries domestic animal disease

either directly causes, or indirectly transmits, a disease to a domestic animal

chemical

uses smells or other chemicals to communicate

choruses

to jointly display, usually with sounds, at the same time as two or more other individuals of the same or different species

dominance hierarchies

ranking system or pecking order among members of a long-term social group, where dominance status affects access to resources or mates

drug

a substance used for the diagnosis, cure, mitigation, treatment, or prevention of disease

endothermic

animals that use metabolically generated heat to regulate body temperature independently of ambient temperature. Endothermy is a synapomorphy of the Mammalia, although it may have arisen in a (now extinct) synapsid ancestor; the fossil record does not distinguish these possibilities. Convergent in birds.

female parental care

parental care is carried out by females

food

A substance that provides both nutrients and energy to a living thing.

forest

forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.

frugivore

an animal that mainly eats fruit

herbivore

An animal that eats mainly plants or parts of plants.

iteroparous

offspring are produced in more than one group (litters, clutches, etc.) and across multiple seasons (or other periods hospitable to reproduction). Iteroparous animals must, by definition, survive over multiple seasons (or periodic condition changes).

migratory

makes seasonal movements between breeding and wintering grounds

motile

having the capacity to move from one place to another.

native range

the area in which the animal is naturally found, the region in which it is endemic.

nocturnal

active during the night

oriental

found in the oriental region of the world. In other words, India and southeast Asia.

World Map

pheromones

chemicals released into air or water that are detected by and responded to by other animals of the same species

polygynous

having more than one female as a mate at one time

rainforest

rainforests, both temperate and tropical, are dominated by trees often forming a closed canopy with little light reaching the ground. Epiphytes and climbing plants are also abundant. Precipitation is typically not limiting, but may be somewhat seasonal.

riparian

Referring to something living or located adjacent to a waterbody (usually, but not always, a river or stream).

scent marks

communicates by producing scents from special gland(s) and placing them on a surface whether others can smell or taste them

seasonal breeding

breeding is confined to a particular season

sexual

reproduction that includes combining the genetic contribution of two individuals, a male and a female

social

associates with others of its species; forms social groups.

swamp

a wetland area that may be permanently or intermittently covered in water, often dominated by woody vegetation.

tactile

uses touch to communicate

terrestrial

Living on the ground.

territorial

defends an area within the home range, occupied by a single animals or group of animals of the same species and held through overt defense, display, or advertisement

tropical

the region of the earth that surrounds the equator, from 23.5 degrees north to 23.5 degrees south.

visual

uses sight to communicate

viviparous

reproduction in which fertilization and development take place within the female body and the developing embryo derives nourishment from the female.

References

Wildscreen 2003-2011. 2011. "ARKive Images of Life on Earth" (On-line). Large flying fox (Pteropus vampyrus). Accessed March 30, 2011 at http://www.arkive.org/large-flying-fox/pteropus-vampyrus/#text=Facts.

Ardea, A. 2009. "The Telegraph" (On-line). Large flying fox: world's biggest bat being hunted to extinction. Accessed March 30, 2011 at http://www.telegraph.co.uk/earth/wildlife/6087109/Large-flying-fox-worlds-biggest-bat-being-hunted-to-extinction.html.

Bates, P., C. Francis, M. Gumal. 2011. "The IUNC Red List of Threatened Species" (On-line). Pteropus vampyrus. Accessed March 30, 2011 at http://www.iucnredlist.org/apps/redlist/details/18766/0.

Breed, A., H. Field, C. Smith, J. Meers. 2010. Bats Without Borders: Long-Distance Movements and Implications for Disease Risk Management. Ecohealth, Vol 7, Issue 2: 204-212.

Brown, M. 1997. Durio - A Bibliographic View. New Delhi, India: International Plant Genetics Resources Institute.

Epstein, J., K. Olival, J. Pulliam, C. Smith. 2009. Pteropus vampyrus, a hunted migratory species with a multinational home-range and a need for regional management. Journal of Applied Ecology, 46: 991-1002.

Feldhamer, G., L. Drickamer. 2007. Mammology: Adaptation, Diversity, Ecology. Johns Hopkins University: Johns Hopkins University Press.

Hassan, N., S. Saad, Z. Shohaimi. 2010. Characterization of Nipah virus from naturally infected pteropus vampyrus bats, Malaysia. Emerging Infecitous Diseases, 16/12: 1990.

Hollister, N. 1913. A Review of the Philippine Land mammals in the United States National Museum. Washington: Washington Government Printing Office.

Kunz, T., D. Jones. 2000. Pteropus vampryrus. Mammalian Species, No. 642: 1-6. Accessed March 30, 2011 at http://www.science.smith.edu/departments/Biology/VHAYSSEN/msi/.

Mildenstein, T., S. Stier, C. Nuevo-Diego, S. Mills. 2005. Habitat selection of endangered and endemic large flying-foxes in Subic Bay, Phillipines. Biological Conservation, 126: 93-102.

Mohd-Azlan, J., A. Zubaid, T. Kunz. 2011. Distribution, relative abundance, and conservation status of the large flying fox, Pteropus vampyrus, in peninsular Malaysia:a preliminary assessment. Acta Chiropterologica, 3(2): 149-162.

Muscarella, R., T. Fleming. 2007. The role of frugivorous bats in tropical forest succession. Biological Reviews, Vol 82, Issue 4: 573-590. Accessed April 16, 2011 at http://apps.isiknowledge.com/full_record.do?product=BIOSIS&search_mode=GeneralSearch&qid=1&SID=2CcoAkk5ePb45E12@GO&page=4&doc=31.

Oakland Zoo, 2011. "Conservation and Education; Oakland Zoo" (On-line). Malayan Flying Fox. Accessed April 03, 2011 at http://www.oaklandzoo.org/animals/mammals/bat-malayan-flying-fox.

Organization for Bat Conservation, 2011. "Organization for Bat Conservation" (On-line). Malayian Flying Fox. Accessed April 16, 2011 at http://www.batconservation.org/drupal/malayan-flying-fox.

Revkin, A., L. Kaufman. 2009. "The New York Times" (On-line). Saving the Flying Fox. Accessed March 30, 2011 at http://dotearth.blogs.nytimes.com/2009/08/25/saving-the-flying-fox/.

Stier, S., T. Meldenstein. 2005. Dietary Habits of the World's Largest Bats: The Philippine Flying Foxes, Acerodon Jubatus and Pteropus Vampyrus Lanensis. Journal of Mammology, 86(4): 719-728.

Struebig, M., M. Harrison, S. Cheyne, S. Limin. 2007. Intensive hunting of large flying foxes Pteropus vampyrus natunae in Central Kalimantan, Indonesian Borneo. Oryx, Vol 41, No 3: 390-393.

Tampa's Lowry Park Zoo, 2010. "Wallaroo Station Animal Fact Sheet" (On-line). Flying Fox Bat. Accessed April 15, 2011 at http://www.lowryparkzoo.com/bio_wallaroo_flying_fox.php.

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