Old World fruit bats I

(Pteropus)

Class Mammalia

Order Chiroptera

Suborder Megachiroptera

Family Pteropodidae

Thumbnail description
The largest of all bats and best known of the pteropodids, with a dog-like facial appearance and very large, forward-facing eyes, hence the common name "flying foxes"; coloration ranges from light to dark brown and some have very distinctly colored mantles

Size
Head and body length 6.7–16 in (17–41 cm); forearm 3.3–9 in (8.5–23 cm); weight 0.4–3.5 lb (0.2–1.6 kg); wingspan 2–6 ft (0.6–1.8 m)

Number of genra, species
1 genus, 60 species

Habitat
Subtropical and tropical forests, caves, and swamps

Conservation status
Extinct: 5 species; Critically Endangered: 7 species; Endangered: 3 species; Vulnerable: 16 species; Lower Risk/Near Threatened: 3 species; Data Deficient: 2 species

Distribution
Islands of the Pacific and Indian Oceans, and from Pakistan across Southeast Asia to Australasia

Evolution and systematics

The family Pteropodidae is divided into two subfamilies, Macroglossinae and Pteropodinae, the latter of which includes the genus Pteropus, also known as the flying foxes. Pteropus is further divided into 17 species groups and about 60 total species, depending on authority. Fossil records of this and other chiropteran groups are scarce due to the delicate nature of the skeletal structure of bats, but pteropid fossils have been found in Europe from the middle Oligocene and Miocene periods, in Africa from the Miocene period, and in Madagascar and the East Indies from the Pleistocene period. Pteropus is assumed to have arisen in the Australo-Pacific during the early Miocene. Their closest relatives are those of the genus Acerodon, a similar group in both size and morphology.

Physical characteristics

Pteropus species are the largest of all bats, weighing up to 3.5 lb (1.6 kg) and with a wingspan of up to 6 ft (1.8 m) Fur is dense and coloration is grayish brown or black. These mammals are characterized by a yellow or grayish yellow contrasting mantle (covering portions of the head, neck, and upper shoulders). Variations among species do occur, such as in the spectacled flying fox (P. conspicillatus), which has a light ring around its eyes. The external ears are small and do not have a tragus, and the tail is absent. The second finger is independent of the third and a claw is present on the thumb. Eyes are very large, forward-facing, and highly adapted to both nocturnal and day vision—this allows flying foxes to easily recognize light colors, which assists in locating food sources.

Distribution

These mammals primarily inhabit islands of the Indian and Pacific Oceans, from Madagascar north to the Maldives and Sri Lanka, across Indonesia, and into the middle Pacific on the island groups of Caroline, Tonga, and Samoa, as far east as the Cook Islands. On the mainland, Pteropus species are found from Pakistan in the west across India and Southeast Asia to Australia.

Habitat

Flying foxes inhabit tropical coastal areas such as mangrove forests, primary and secondary growth rainforest, lowland dry forest, swamps, and occasionally caves. Most species roost high above the canopy in a ridge of emergent trees.

Behavior

Flying foxes are most active in the evening and at night. They roost in trees by day, and many of the larger species do

so in extremely large groups called "camps," which range in size from a few dozen individuals up to 250,000. When resting in the daylight hours, they hang from branches by one or both feet with wings wrapped around their bodies, though there is still sizable activity among the camp as the bats move from one spot to another. At dusk, when the time to forage arrives, Pteropus species will flap their wings until their bodies are parallel with the ground—only then do they release the branch and initiate flight.

Migration among flying foxes depends primarily on the seasonal availability of food sources. They do not migrate over particularly long distances, but instead travel between winter and summer roosts when fruits or blossoms are ready for the season. Mainland species will travel about 30 mi (50 km) to reach a new feeding site, and island groups may relocate to neighboring islands or to an accessible mainland area. Colonies will often use the same roosting sites year after year.

Flying fox vocalizations are in the range of 4–6 kHz. Vocalizations play an important role in feeding, mating, territorial disputes, and interaction with infants. In the case of the gray-headed flying fox (P. poliocephalus), at least 30 different kinds of calls have been documented.

Feeding ecology and diet

Pteropus species primarily consume fruit, nectar, and pollen. They are able to locate food using highly developed senses of vision and smell (like most fruit bats, members of this genus do not orient themselves using echolocation). Flying foxes employ optimal foraging (seeking the greatest ratio of benefit versus the amount of time and energy spent) as well as searching and handling techniques when going out to feed.

Once food is acquired, the bat will take it to a nearby roost and eat while hovering, or hang from a branch while using one foot to hold the fruit. Juice is the chief source of food for these mammals; it is consumed by compressing bits of pulp against the rigid palate of the mouth, swallowing the juice, and spitting out the pulp and seeds. If the pulp is soft, however, it may be occasionally eaten as well. They drink while traveling to or from a feeding location, skimming the surface of the water during flight. Some species drink seawater in an effort to acquire minerals that are unavailable in other food sources.

Owing to the pursuit of nectar, flying foxes have developed a working relationship with several plants within their habitat. Flower pollination and dispersal of seeds is of course beneficial to these organisms, and certain fruits and trees are specialized to attract fruit bats—some are lightly colored (sea almond trees, Terminalia catappa) while others have a strong odor (mangos, Mangfiera indiaca). Trees such as the durian (Durio zibethines) bloom only at night, easily able to attract the eye of a passing pteropid at peak feeding time.

Reproductive biology

During the summer, when fruit and blossoms are mature and in good supply, flying foxes organize camps. Mating

takes place at this time and small groups, or harems, form. Males soon become very territorial over the females and the roosts, marking their areas using a scent gland located on the throat.

Females are seasonal breeders and usually produce one young per year; they begin to breed at two years. During mating season, flying foxes will mate more than once per day and over the course of several days. Ovulation takes place from February to April, and births occur from September to November. Lactating lasts about six weeks, and most of the female's time during the remainder of the year is spent caring for the young.

The gestation period of the flying fox is six months. During that time, the sexes begin to segregate and pregnant females form a colony; each female then helps to care for the others by mutual grooming. Birth occurs during the day—when it is imminent, the female hangs by her thumbs and feet and licks her genital area until the pup's head begins to emerge—this can last up to several hours. After birth, the pup moves itself into a suckling position and attaches itself to a nipple. The mother will fly with her young for about two to three weeks.

The pup has light fur, the eyes are closed, and the ear flaps are down. The mother keeps her wings wrapped around the pup for warmth. After approximately three weeks, it becomes too heavy to carry with her and is left with the other young. Upon the mother's return, she is able to recognize her offspring by its unique vocalizations. In about a month, the young become better coordinated and begin to explore, and by January and February, they begin to form small groups near their mothers. Once they become able to care for themselves, the mother will again begin to accept the advances of a male.

Conservation Status

The IUCN Red List ranks seven species as Critically Endangered, three species as Endangered, 16 species as Vulnerable, three species as Lower Risk/Near Threatened, two species as Data Deficient, and five species as Extinct.

The most serious threat to flying fox populations is probably deforestation. The removal of primary forest not only limits habitat in the most basic sense, but it also encourages additional loss—the logging processes used in these areas tend to inhibit growth of new canopy, and the elimination of large sections of forest leaves the remaining habitat even more vulnerable to the tropical storms that frequently strike island environments. The mass conversion of mangrove swamps into shrimp farms has also had a devastating effect on certain species, most notably the Pohnpei flying fox (P. molossinus).

Some Pteropus species are also losing ground due to illegal wildlife trade, human use for food, and extermination—flying foxes are often considered by orchard growers to be destructive agricultural pests. Populations can sometimes be disturbed by predation as well. An example is the brown tree snake (Boiga irregularis), which was introduced on Guam in the 1940s and has had a significant impact on bat populations since. Other enemies include predatory birds, such as owls and falcons.

Significance to humans

All over their range, flying foxes have been considered a delicacy for centuries, and this is especially so for the Chamorro people of Guam and the nearby Northern Marianas Islands. Traditionally, the animal (fur, wings, and innards included) is boiled in coconut milk and eaten in its entirety, usually during ceremonial or otherwise special occasions. Samoan islanders use branches bound to the end of long poles to snag the animal and pull it to the ground, while aboriginal Australians have also been known to use specialized methods to hunt these mammals for food.

Flying foxes have occasionally been considered beneficial for medical use. In the 1970s, Indian flying foxes (P. giganteus) in Pakistan were harvested for their fat, which was thought to be a cure for rheumatism. Still, by far the most notable contribution is that of pollination and seed dispersal. As a whole, Pteropus plays an integral role in the survival of 300 species of plants across its range, about half of which are regularly used by humans for nourishment, materials, and medicine.

Species accounts

List of Species

Island flying fox

Pteropus hypomelanus

subfamily

Pteropodinae

taxonomy

Pteropus hypomelanus Temminck, 1853, Moluccas Islands, Indonesia.

other common names

English: Variable flying fox, small flying fox, Condoro Island flying fox.

physical characteristics

Head and body length 8–9 in (20–22 cm), forearm 5.5–6 in (14–15 cm), wingspan 3.5–4 ft (1–1.2 m). Color patterns vary. The fur pattern on the face can be black with the crown light to dark brown. Mantle can be light brown, russet brown, or red-tinged over a brown base. It is lighter in the dorsum, mixed with gray and black hairs. The stomach is a buff color, medium brown on the lower shoulders, and dark brown to black on the flanks and ventrum.

distribution

Small islands of the coast of Southeast Asia, Indonesia, New Guinea, and the Solomons.

habitat

Small to medium offshore islands, lowland and disturbed forests. It roosts in tall broadleaf trees, coconut trees (Cocos nucifera), and orchards.

behavior

This species will forage over great distances during the night. It flies approximately 20–30 mi (32–48 km) over land. During this time it will fly low or skim through the troughs of waves to lower wind resistance. It roosts singly, in small family groups, or in large colonies of up to several thousand. If it is particularly hot, these bats may use a wing to fan themselves while panting. In overcast weather, the wings are wrapped around their body and face.

feeding ecology and diet

Diet includes fruits, flowers and foliage. This includes figs (Ficus carica), sea almond, kapok (Ceiba pentandra), chico (Poaloria sapota), eucalyptus flowers (Eucalyptus globulus), bananas (Musa spp.), and paw paws (Asimina triloba).

reproductive biology

Males and females are sexually mature at 1.5 years. Gestation is 180–210 days and births take place from April to May. Polygynous.

conservation status

Not threatened.

significance to humans

Hunted for food. Performs valuable pollination and seed dispersal of plants.

Madagascar flying fox

Pteropus rufus

subfamily

Pteropodinae

taxonomy

Pteropus rufus Tiedemann, 1808, Madagascar.

other common names

English: Malagasy flying fox.

physical characteristics

Head and body length 9–10 in (23–25 cm), weight 1.2–1.5 lb (533–700 g). Fur coloration on top of the head is yellow or orange. Nose, dorsum, and ventrum are brown. Ears and nose are long and pointed.

distribution

Madagascar.

habitat

Coastal forests, low-lying central plains, and small off-shore islands.

behavior

Nocturnal, colonial, and very gregarious, with screeching vocalizations. Males tend to their territories quite vigorously and are very aggressive.

feeding ecology and diet

Various fruits, blossoms, and leaves. Tamarind (Tamarindus indica) pods are a favorite.

reproductive biology

Monogamous. Birth weight is approximately 1.5 oz (43 g). Pups are weaned by the eleventh week and volant by three to four months.

conservation status

Not threatened.

significance to humans

Hunted for food. Performs valuable pollination and seed dispersal of plants.

Blyth's flying fox

Pteropus melanotus

subfamily

Pteropodinae

taxonomy

Pteropus melanotus Blyth, 1863, Nicobar Islands, India.

other common names

English: Black-eared flying fox.

physical characteristics

Forearm 6–7 in (15–18 cm). Fur coloration on the head is brown to black. The mantle can be golden tawny, reddish, buff, or light rufous. The ventral surface colors range from

pale and light or dark brown to blackish brown. Ears are large with broad, round tips.

distribution

Andaman Islands, Nicobar Islands, Enggano, Nias Islands off western Sumatra, and Christmas Island.

habitat

Roosts in large colonies in mangrove forests, usually near a body of water.

behavior

Nothing is known.

feeding ecology and diet

Mangoes, papayas (Carica papaya), guava (Psidium guajava).

reproductive biology

Probably polygynous. Females mature rapidly, and are able to breed at at six months. Males reach maturity at 18 months. There is a single breeding season with most births taking place in February.

conservation status

Not threatened.

significance to humans

Hunted for food. Performs valuable pollination and seed dispersal of plants.

Livingstone's fruit bat

Pteropus livingstonii

subfamily

Pteropodinae

taxonomy

Pteropus livingstonii Gray, 1866, Comoro Islands.

other common names

English: Livingstone's flying fox, Comoro black flying fox.

physical characteristics

Head and body length 12 in (30 cm), weight 1.1–1.8 lb (500–800 g), wingspan 5 ft (1.5 m). Patches of golden fur on chest.

distribution

Johanna and Moheli Islands in the Comoros.

habitat

Dense mountain forest with steep-sided valleys.

behavior

There is little information on the behavior of these bats in the wild. The known social structure has small groups roosting together and forming harems. These bats fly with a slow wing beat and glide on thermals, using them to help extend their soaring time.

feeding ecology and diet

Blossoms and fruits such as figs are the major food sources.

reproductive biology

Polygynous. Females have an annual birth season from July through October.

conservation status

Critically Endangered.

significance to humans

Hunted for food. Performs valuable pollination and seed dispersal of plants.

Marianas fruit bat

Pteropus mariannus

subfamily

Pteropodinae

taxonomy

Pteropus mariannus Desmaret, 1822, Mariana Islands, Guam.

other common names

English: Micronesian flying fox, Marianas flying fox, Marianna flying fox.

physical characteristics

Dorsum and wings are brown to blackish with silver hairs. The mantle and sides are yellow to bright gold.

distribution

Okinawa, Ryukyu Islands, Guam, Mariana Islands.

habitat

Subtropical and tropical areas, primarily riparian. Marianas bats roost in lowland swamp forest and trees such as the Banyon (Ficus virens) and she-oak (Casuarina spp.) that rise above the canopy line.

behavior

This species is primarily sedentary and nonmigratory. The roosting size of the colonies are frequently 60–800 individuals, smaller groups of 10–12, smaller bachelor groups of 10–15, and solitary individuals. Marianas bats are not as vocal as other flying foxes.

feeding ecology and diet

These bats are primarily frugivorous but do eat flowers and leaves. The primary foraging area is the agroforest and pandamas savanna. They feed on 53 species of fruit, 23 species of flowers, and one species of leafy plant.

reproductive biology

Polygynous. Breeding colonies are made up of harem groups. One male to several females.

conservation status

Endangered. Habitat loss due to timber removal, invasive nonnative species, natural disasters (cyclones and tornadoes), hunting for food, and the wildlife trade market.

significance to humans

Hunted for food. Performs valuable pollination and seed dispersal of plants.

Tongan flying fox

Pteropus tonganus

subfamily

Pteropodinae

taxonomy

Pteropus tonganus Quay-Gaimard, 1830, Tonga Islands.

other common names

English: White-necked fruit bat, insular fruit bat, Tongan fruit bat, Pacific flying fox.

physical characteristics

Forearm 5–6 in (13–15 cm), wingspan up to 3 ft (0.9 m), weight 10.5–21 oz (300–600 g). The fur is black or brown with numerous white hairs on its head, dorsum, and ventrum. Mantle is red and yellow-brown with a strip on the dorsum between the wings.

distribution

Guam.

habitat

The preferred habitat is riparian. The Tongan flying fox uses coconut palm trees, broadleaf trees, and forest remnants. There are both day and night roosts.

behavior

Roosts singly in small groups or large communal groups, hanging from branches in the shade. The size and structure of the roost sizes appear to be organized by reproductive status. Bachelor males, clusters of females defended by a male, groups of females, and young. Compared to some of the other Pteropus species, it is a relatively quiet bat. Foraging begins about and hour before dark with short flights between various trees.

feeding ecology and diet

Frugivorous. Primarily feeds on fruit, flowers, nectar, and sap.

reproductive biology

Polygynous. Births can occur year-round but is most common from June to August. Gestation lasts for five months. Single or twin births can be expected. Young are weaned at three months but will stay with the mother until they reach adult size.

conservation status

Not threatened.

significance to humans

Hunted for food. Performs valuable pollination and seed dispersal of plants.

Indian flying fox

Pteropus giganteus

subfamily

Pteropodinae

taxonomy

Pteropus giganteus (Brünnich, 1782), Bengal, India.

other common names

English: Giant Indian fruit bat.

physical characteristics

Head and body length 8–12 in (20–30 cm), forearm 6–7 in (15–18 cm), wingspan 4 ft (1.2 m), weight 3.3 lb (1.5 kg).

distribution

Pakistan, Nepal, India, and the Maldives through to Myanmar. A single specimen was recorded in Tsinghai, China.

habitat

The preferred habitat is well established trees and swamp areas near large bodies of water. Some of the tree species favored for roosting sites are semal (Bombax malabaricus), peepal (Ficus bengalensis), and mango.

behavior

A colonial species that lives in large diurnal roosts. These bats form a hierarchy within the male population, defined by strength and size. Within the hierarchy, each male has a rank and his own roosting spot. Roosting takes place out in open. Colony size varies seasonally, shrinking in number during the summer and increasing during the rainy season. Colonies of these species tend to favor roosting areas near towns or villages.

feeding ecology and diet

These bats will leave the roosting site soon after sunset and return about 45 minutes after sunrise. They may break into small feeding groups after leaving and when entering a feeding area. Before feeding they will visit a lake or river to drink and they skim it while they are flying.

Diet includes fruits, flowers, and leaves during the nonfruiting season. Examples are flower buds from the silk cotton tree (Gossampinus malabarocus), tulip tree (Spathodea campunulata), guava, and the soft leaves and twigs of the tamarind trees.

reproductive biology

Polygynous. Mating takes place from July to October. Within that span, copulation will take place three times. Gestation is 140–150 days. When they are ready to give birth, females will congregate in the upper branches of the roosts. The pup is able to fly by the eleventh week and is weaned within two to six months.

conservation status

Not threatened.

significance to humans

Hunted for food. Performs valuable pollination and seed dispersal of plants.

Black flying fox

Pteropus alecto

subfamily

Pteropodinae

taxonomy

Pteropus alecto Temminck, 1837, Sulawesi, Indonesia.

other common names

English: Pygmy fruit bat, gray fruit bat.

physical characteristics

Head and body length (19–28 cm), forearm 6–7.5 in (15–19 cm), wingspan up to 3.3 ft (1 m) weight 1.1–2.2 lb (500–1,000g). The fur on the head is black, the mantle ranges from chocolate brown to reddish brown, and white hairs frequently appear over the body, including the underside.

distribution

Sulawesi, Salayer Island, Baweam and Kangean islands to the Java Sea, Lombok, Sumba and Savu islands, southern New Guinea, northern and eastern Australia.

habitat

Found in tropical and subtropical forest and woodlands.

behavior

Males will establish territory and will groom themselves on a daily basis. Groups will return to the same site to roost year after year. On occasion, they will share their roosts with gray-headed flying foxes (P. poliocephalus).

feeding ecology and diet

These groups will travel up 30 mi (50 km) to forage. Nectar, fruit, and tree blossoms are the principal diet. They do not eat citrus fruits.

reproductive biology

Polygynous. Camps congregate from early to late summer. Birthing season is October to November in southern Queensland and January to February in the Northern Territory. This difference relates to the availability of food resources. The young are carried by the mothers until approximately fours weeks and then are left at the roost site. Juvenile or immature bats do not leave but remain together and form winter camps.

conservation status

Not threatened.

significance to humans

Hunted for food. Performs valuable pollination and seed dispersal of plants.

Rodricensis flying fox

Pteropus rodricensis

subfamily

Pteropodinae

taxonomy

Pteropus rodricensis Dobson, 1878, Rodrigues, Mascarene Islands.

other common names

English: Rodrigues flying fox.

physical characteristics

Wingspan 3 ft (0.9 m), weight 0.7 lb (300 g). The fur is dark brown and it covers most of the body. Head, neck, and shoulders are a golden color.

distribution

Island of Rodriguez in the Mauritius.

habitat

Dense forests and dry woodlands.

behavior

Colonial in nature. Camps are very gregarious and vocal. They are not skilled fliers.

feeding ecology and diet

Various fruits, such as tamarind pods and mangos.

reproductive biology

Males will pick out territories and form harems during breeding season. During this time, the males will be physically aggressive with each other by biting. Young can fly by three to four months and remain with their mothers for up to a year.

conservation status

Critically Endangered.

significance to humans

Hunted for food. Performs valuable pollination and seed dispersal of plants.

Spectacled flying fox

Pteropus conspicillatus

subfamily

Pteropodinae

taxonomy

Pteropus conspicillatus Gould, 1850, Fitzroy Island, Australia.

other common names

English: Spectacled fruit bat.

physical characteristics

Head and body length 8.7–9.8 in (22–25 cm), forearm 6.3–7 in (16–18 cm), weight 0.9–1.3 lb (400–600 g). A large black flying fox with pale yellow or straw-colored fur around its eyes. The mantle is pale yellow and goes across the back, neck, and shoulders. Some specimens have been found to have pale yellow fur on the face and top of the head.

distribution

New Guinea, Irian Jaya, Indonsia, Louisiade Archipelego, D'Entrecasteaus and Trobriand Islands, and northeastern Queensland.

habitat

Spectacled flying foxes are forest dwellers. They prefer to roost in the upper canopies of rainforest. Most roost trees are sheoaks, but these bats have also been known to roost in the upper branches of broadleaf trees for protection from hunters. They have also been observed stripping the upper leaves from trees. This behavior allows better visual observation of the surrounding area.

behavior

Spectacled flying foxes roost together in mixed colonies with different species, such as the variable flying fox (Pteropus hympomelanus). The size of the camp will range from the hundreds to thousands. They are very vocal over roosting sites, food, mating, and group or individual territories. Camps can be found quite easily due to the level of noise they produce.

feeding ecology and diet

Spectacled flying foxes feed on several different species of fruits, blossoms, and foliage, as well as the occasional insect. The foraging distance for these bats is about 30 mi (50 km). As with many of the flying foxes, this species drinks seawater on the way to feeding sites.

reproductive biology

Polygynous. Breeding takes place between March and May. Females give birth to a single offspring between October and December.

conservation status

Not threatened.

significance to humans

Hunted for food. Performs valuable pollination and seed dispersal of plants.

Big-eared flying fox

Pteropus macrotis

subfamily

Pteropodinae

taxonomy

Pteropus macrotis Peters, 1867, Indonesia.

other common names

English: Black-bearded flying fox.

physical characteristics

Forearm 5–6 in (13–15 cm), weight 0.7–0.9 lb (300–400 g). The fur on the head is medium to dark brown, throat is brown, dorsum and rump are dark brown, ventrum is black or reddish brown, and the mantle is reddish to yellow.

distribution

New Guinea, Boigu Islands of Australia.

habitat

Lowland forest; inland forests.

behavior

Known to forage in drier areas than other Pteropus species.

feeding ecology and diet

Large numbers of bats will fly to the mainland of New Guinea to feed. They eat coconut palm flowers and a variety of fruit and blossoms in dry monsoon scrub. It will invade plantations to feed on coconut and sago palm (Cycas revoluta) blossoms.

reproductive biology

Nothing is known, but probably polygynous.

conservation status

Not threatened.

significance to humans

Hunted for food. Performs valuable pollination and seed dispersal of plants.

Little red flying fox

Pteropus scapulatus

subfamily

Pteropodinae

taxonomy

Pteropus scapulatus Peters, 1862, Cape York, Australia.

other common names

None known.

physical characteristics

Head and body length 5–8 in (13–20 cm), forearm 4.3–5.5 in (11–14 cm).

distribution

Extreme southern New Guinea, northern and eastern Australia, rarely on Tasmania, one record from New Zealand.

habitat

Broad range of habitat, from tropical to semiarid and monsoon forests, temperate eucalypt forests, and paperback swamps.

behavior

Roost size can range into the thousands in early summer. They are a very nomadic species and normally do not stay in a camp for long periods of time. Roosts are located near a body of water.

feeding ecology and diet

Their primary source of food is the blossoms of plants, including trees and shrubs. Other foods that may be eaten are insects, sap, and fruit.

reproductive biology

Mating occurs from November to January. Harems are then formed, two to five females with a male. Gestation is five months and birth takes places between April and May. Young are carried by the mother for the first month and then in roost while she forages and returns through the night. The young are volant at two months but females continue to care for them while they develop adult skills.

conservation status

Not threatened.

significance to humans

Hunted for food. Performs valuable pollination and seed dispersal of plants.

Resources

Books

Altringham, John D. Bats, Biology, and Behavior. New York: Oxford University Press, 1996.

Bonnacorso, Frank J. Bats of Papua New Guinea. Chicago: University of Chicago Press, 1998.

Buchmann, Stephen L., and Gary Paul Nabhan. The Forgotten Pollinators. Washington, DC: Island Press, 1997.

Crichton., Elizabeth G., and Phillip H. Krutzsch, eds. Reproductive Biology of Bats. New York: Academic Press, 2000.

Hall, Leslie, and Greg Richards. Flying Foxes, Fruit and Blossom Bats of Australia. Malabar, FL: Krieger Publishing Company, 2000.

Kunz, Thomas, and Paul Racey, eds. Bat Biology and Conservation. Washington, DC: Smithsonian Institution Press, 1998.

Mickleburgh, Simon, Anthony M. Hutson, and Paul Racey. Old World Fruit Bats: An Action Plan for Their Conservation. Gland, Switzerland: IUCN, 1992.

Neuweiler, Gerhard. The Biology of Bats. New York: Oxford University Press, 2000.

Nowak, Ronald, ed. Walker's Mammals of the World, 6th ed. Baltimore and London: The John Hopkins Unversity Press, 1999.

Taylor, Peter John. Bats of Southern Africa. Pietermaritzburg: University of Natal Press, 2000.

Periodicals

Banack, Sandra Anne. "Diet Selection and Resource Use by Flying Foxes (Genus Pteropus)." Ecology 79 (1998): 1949–1967.

Fujita, Marty. "Flying Foxes and Economics." Bats 6, no. 1 (1998): 49.

Rainey, William E. "The Flying Foxes: Becoming a Rare Commodity. Bats 8, no. 1 (1990): 69.

Other

"Family Pteropodidae." University of Michigan Animal Diversity Web. [23 June 2003]. <http://animaldiversity.ummz.umich.edu>.

Martin, Len. "The Effects of Culling the Flying Foxes, Pteropus conspicillatus in Northern Queensland, and Pteropus poliocephalus in Victoria, NSW and Southeast Queensland." [23 June 2003]. <http://www.austrop.org.au/ghff/home.htm>.

Thatcher, Oliver. "Destruction of Fruit Bat Habitat." [23 June 2003]. <http://www.biology.leeds.ac.uk/staff/dawa/bats/Fruitbats/deforest.html>.

Kate Kretschmann

Robin L. Hayes