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.
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.
- Habitat Regions
- tropical
- terrestrial
- Terrestrial Biomes
- forest
- rainforest
- Wetlands
- swamp
- Other Habitat Features
- agricultural
- riparian
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.
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.
- Other Physical Features
- endothermic
- homoiothermic
- bilateral symmetry
- Sexual Dimorphism
- male larger
- sexes colored or patterned differently
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.
- Mating System
- polygynous
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.
- Key Reproductive Features
- iteroparous
- seasonal breeding
- gonochoric/gonochoristic/dioecious (sexes separate)
- sexual
- viviparous
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.
- Parental Investment
- altricial
- female parental care
-
pre-fertilization
- provisioning
-
protecting
- male
-
pre-hatching/birth
-
provisioning
- female
-
protecting
- male
- female
-
provisioning
-
pre-weaning/fledging
-
provisioning
- female
-
protecting
- male
- female
-
provisioning
-
pre-independence
-
provisioning
- female
-
protecting
- female
-
provisioning
Lifespan/Longevity
Pteropus vampyrus
can live 15 to 30 years in captivity, and an average of 15 years in the wild.
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.
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.
- Key Behaviors
- arboreal
- flies
- nocturnal
- motile
- migratory
- territorial
- social
- dominance hierarchies
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.
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.
- Other Communication Modes
- choruses
- pheromones
- scent marks
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.
- 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.
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.
- Ecosystem Impact
- disperses seeds
- pollinates
- 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.
- Positive Impacts
- food
- body parts are source of valuable material
- source of medicine or drug
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.
- Negative Impacts
-
injures humans
- carries human disease
- crop pest
- causes or carries domestic animal disease
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.
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.
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
.
Additional Links
Contributors
Kelsie Norton (author), University of Michigan-Ann Arbor, Phil Myers (editor), University of Michigan-Ann Arbor, John Berini (editor), Animal Diversity Web Staff.
- oriental
-
found in the oriental region of the world. In other words, India and southeast Asia.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- tropical
-
the region of the earth that surrounds the equator, from 23.5 degrees north to 23.5 degrees south.
- terrestrial
-
Living on the ground.
- forest
-
forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.
- 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.
- swamp
-
a wetland area that may be permanently or intermittently covered in water, often dominated by woody vegetation.
- agricultural
-
living in landscapes dominated by human agriculture.
- riparian
-
Referring to something living or located adjacent to a waterbody (usually, but not always, a river or stream).
- 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.
- 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.
- polygynous
-
having more than one female as a mate at one time
- 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).
- 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
- viviparous
-
reproduction in which fertilization and development take place within the female body and the developing embryo derives nourishment from the female.
- 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.
- female parental care
-
parental care is carried out by females
- arboreal
-
Referring to an animal that lives in trees; tree-climbing.
- nocturnal
-
active during the night
- motile
-
having the capacity to move from one place to another.
- migratory
-
makes seasonal movements between breeding and wintering grounds
- 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
- social
-
associates with others of its species; forms social groups.
- dominance hierarchies
-
ranking system or pecking order among members of a long-term social group, where dominance status affects access to resources or mates
- visual
-
uses sight to communicate
- tactile
-
uses touch to communicate
- acoustic
-
uses sound 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
- pheromones
-
chemicals released into air or water that are detected by and responded to by other animals of the same species
- scent marks
-
communicates by producing scents from special gland(s) and placing them on a surface whether others can smell or taste them
- visual
-
uses sight to communicate
- tactile
-
uses touch to communicate
- acoustic
-
uses sound to communicate
- chemical
-
uses smells or other chemicals to communicate
- food
-
A substance that provides both nutrients and energy to a living thing.
- drug
-
a substance used for the diagnosis, cure, mitigation, treatment, or prevention of disease
- causes or carries domestic animal disease
-
either directly causes, or indirectly transmits, a disease to a domestic animal
- herbivore
-
An animal that eats mainly plants or parts of plants.
- frugivore
-
an animal that mainly eats fruit
References
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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.
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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 .
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