Members of Pteropodidae are known colloquially as the flying foxes, or Old World fruit bats. The family is composed of 41 genera and about 170 species. The most species-rich genus in the family is Pteropus with 59 species, many of which are island endemics. Body and wing size ranges from small (37 mm forearm length) to large (220 mm forearm length). The family boasts the largest bats in the world. Pteropus vampyrus individuals have a wingspan of up to 1.7 m. Pteropus giganteus individuals have a comparable wingspan but a greater mass, with males weighing between 1.3 and 1.6 kg. Pteropodids are strictly vegetarian, foraging for fruits, nectar, and pollen using their sight and a sensitive olfactory system. Bats of the genus Rousettus use tongue clicks as a crude form of echolocation while navigating in the dark. Some species are migratory, covering vast distances, while others have more moderate home ranges. Eidolon helvum individuals aggregate in numbers reaching the hundreds of thousands, yet many species roost with only a few conspecifics. Members of Pteropodidae service the ecosystems they inhabit by playing important roles as pollinators and seed dispersers.
- Nowak, R. 1999. Walker's Mammals of the World. Baltimore and London: The Johns Hopkins University Press.
- Fenton, M. 2001. Bats, Revised Edition. New York, NY: Checkmark Books.
- Neuweiler, G. 2000. The Biology of Bats. New York, NY: Oxford University Press.
- Koopman, K. 1994. Handbook of Zoology, Band/Volume VIII Mammalia. Berlin, Germany: Walter de Gruyter & Co..
Pteropodidae has a tropical and subtropical distribution in the Old World (eastern hemisphere). Species are found as far north as the eastern Mediterranean, continuing along the southern coast of the Arabian Peninsula and across South Asia. Species are found as far south as South Africa, the islands of the Indian Ocean, and to the northern and western coasts of Australia. The longitudinal range reaches from the Atlantic coast of Africa to the islands of the western Pacific. Pteropodids are absent from northwest Africa, southwest Australia, a majority of the Palearctic region, and all of the Western Hemisphere.
Biogeographic Regions: palearctic (Native ); oriental (Native ); ethiopian (Native ); australian (Native ); oceanic islands (Native )
- Nowak, R. 1994. Walker's Bats of the World. Baltimore, MD: The Johns Hopkins University Press.
- Mickleburgh, S., A. Hutson, P. Racey. 1992. Old World Fruit Bats: An Action Plan for their Conservation. Gland, Switzerland: International Union for the Conservation of Nature.
The head and body length of pteropodids varies from 50 mm to 406 mm. Despite size, many characteristics are shared among genera. A relatively long rostrum (pronounced in nectarivores), large eyes, and simple external ears give members of this family a dog or fox-like appearance (hence “flying fox”). The genera Nyctimene and Paranyctimene are exceptions in that they contain tubular nostrils that project from the upper surface of the snout. On the skull, postorbital processes are present over the orbital region. The palatine extends posterior to nearly cover the presphenoid. No more than two upper and two lower incisors are present in adults, otherwise cheek and canine dentition varies between species. The tongue is highly protrusible in nectar feeding bats and often complex with terminal papillae.
The chest is robust, comprised of the down-thrusting pectoralis and serratus muscles. The articulating regions of the humerus never come into contact with the scapula, which differs from a locking mechanism that occurs in the shoulder joint of other bat groups. The second digit is relatively independent from the third digit and contains a vestigial claw that adorns the leading edge of the wing.
Modifications for hanging include a relocation of the hip socket. The acetabulum is shifted upward and dorsally, and articulates with a large headed femur for a wider range of motion. In contrast to most other mammal orders, the legs cannot be positioned in a straight line under the body. In conjunction with large claws on their feet, pteropodids use a tendon-ratchet system that allows them to hang without prolonged muscular contraction. The legs manipulate a primitive uropatagium during flight. Aside from Notopteris, most species are tailless or with just a spicule of a tail.
Several species of Pteropodidae demonstrate sexual dimorphism. Males of Hypsignathus monstrosus have rather outlandish facial features, while females have the conservative fox-like look. Males of the genus Epomops have distinctive white patches in association with a glandular membrane on their shoulders, whereas females do not. Considering the whole family, males are generally larger than females. The penis of all pteropodids is a pendant and freely movable organ, resembling that of Primates. Juveniles are typically naked or have a velvety coat that is darker than adult pelage.
Other Physical Features: endothermic ; homoiothermic; bilateral symmetry
Sexual Dimorphism: sexes alike; male larger; sexes colored or patterned differently; ornamentation
- Altringham, J. 1996. Bats: Biology and Behaviour. New York, NY: Oxford University Press.
- Myers, P. 2001. "Animal Diversity Web" (On-line). Accessed 2-11-09 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Pteropodidae.html.
Pteropodids typically occur in primary or maturing secondary forests. A few species inhabit savannah habitats where they roost in bushes and low trees. Over half of the 41 genera are made up of species that roost in trees. Gregarious species roost on the open branches of large, canopy-emergent trees. Pteropodids that roost singly or in small groups can be found in dead palm leaves, aerial roots, and even arboreal termite nests. These bats also tend to have cryptic coloration and wrap themselves with their wings in order to resemble dead leaves. In one species, Cynopterus sphinx, individuals construct tents by chewing folds in palm leaves. Caves, cliff walls, mines, and the eaves of buildings also serve as roosting locations for species in 17 genera. Most cave-dwelling species are limited to the lit areas near the opening, while members of the genus Rousettus are able to navigate the darker regions using crude echolocation.
Flowering plants are essential to the diet of pteropodid species; therefore, flying foxes mostly use woodlands or orchards for food. Canopy emergent fruiting trees, such as fig and baobab trees, are frequently used as a food source. The flowers of baobab trees have a strong fragrance and are located on the crown of the tree, which makes them easily accessible to bats (a flower syndrome known as chiropterophily). Many pteropodid species are found in coastal areas and drink salt water in order to supplement nutrients lacking in their diet.
A few species are migratory. Eidolon helvum individuals gather in large numbers and migrate hundreds of kilometers northward with seasonal rains, only to return to southern Africa at the end of the rainy season. Pteropus scapulatus populations make major, and somewhat erratic movements within Australia, following the flowering periods of Eucalyptus trees. Many species of Pteropus roost on islands and make daily migrations to the mainland for foraging. Some species range from sea level to 2500 m, yet little is known about any significant elevational migrations.
Habitat Regions: tropical ; terrestrial
Terrestrial Biomes: savanna or grassland ; forest ; rainforest
Other Habitat Features: suburban ; agricultural ; riparian ; estuarine
Pteropodids are frugivorous and nectarivorous. Some species also eat flowers of the plants they visit. Foraging habits are not well documented, though many species of the genus Pteropus rely heavily on figs. Many species rely on broad array of resources, though there may be a functional dichotomy between large species that rely heavily on canopy resources and smaller species that can use understory resources. Some larger species can use the claws on their thumbs and second digits to climb into the understory and seek out fruit that is hidden or inaccessible by flight.
Primary Diet: herbivore (Frugivore , Nectarivore )
Pteropodids provide important pollination and seed dispersal services to a wide range of plants. On islands in the south Pacific, pteropodids are the principle pollinators and dispersers of plants. Many plants have adaptations specifically for seed dispersal and pollination by bats, such as fruiting or flowering at the ends of branches and at bat accessible locations in the canopy. Eidolon dupreanum has been shown to likely be the sole pollinator of the baobab tree Adansonia suarezensis in Madagascar.
Ecosystem Impact: disperses seeds; pollinates
- Cox, P., T. Elmquist, E. Pierson, W. Rainey. 2005. Flying Foxes as Strong Interactors in South Pacific Island Ecosystems: A Conservation Hypothesis. Conservation Biology, 5/4: 448-454.
- Andriafidison, D., R. Andrianaivoarivelo, O. Ramillijaona, M. Razanahoera, J. MacKinnon, R. Jenkins, P. Racey. 2006. Nectarivory by endemic malagasy fruit bats during the dry season. Biotropica, 38/1: 85-90.
Birds of prey and carnivorous mammals, as well as snakes and large lizards may prey on pteropodids. Pteropodids tend to have fewer predators on islands. However, there have been several cases of introductions of non-native, arboreal snakes which have decimated pteropodid populations.
Known prey organisms
Based on studies in:
This list may not be complete but is based on published studies.
- J. L. Harrison, The distribution of feeding habits among animals in a tropical rain forest, J. Anim. Ecol. 31:53-63, from p. 61 (1962).
Life History and Behavior
Pteropodids rely heavily on vision and olfaction when navigating and foraging. Intraspecific communication is often vocal. In some species, such as Pteropus poliocephalus, vocal signaling may be associated with specific motor activities which enhance the meaning of the vocal signal. In species such as Eidolon helvum, sexually dimorphic sebaceous glands which are larger in males may provide olfactory behavioral cues.
Communication Channels: visual ; tactile ; acoustic ; chemical
Perception Channels: visual ; tactile ; acoustic ; ultrasound ; chemical
- Mainoya, J., K. Howell. 1979. Histology of the neck glandular skin patch in Eidolon helvum, Rousettus aegyptiacus and Rousettus angolensis chiroptera pteropodidae. African Journal of Ecology, 17: 159-164.
- Van Parijs, S., P. Corkeron. 2002. Ontogeny of vocalisations in infant black flying foxes, Pteropus alecto. Behaviour, 139/9: 1111-1124.
Pteropodids have been known to live at least 30 years, both in captivity and in the wild.
Mating behavior in pteropodids is highly variable, and much is unknown. The males of one genus (Hypsignathus) set up lekking territories twice a year and draw in females with unique vocalizations and wing-flapping displays. Male epauletted fruit bats (genus Epomophorus) often display their concealed epaulets (hair tufts near the shoulder) and emit courting calls to attract females. Many species form harems consisting of 1 dominant male and up to 37 females, while bachelor males roost separately.
Mating System: polygynous ; polygynandrous (promiscuous)
While most bats have one reproduction event per year, many pteropodids are polyestrous, with two seasonal cycles corresponding to the annual wet and dry seasons. Usually one young is born per pregnancy, but twins are not uncommon. Upon fertilization, ova implantation in the uteri can be immediate or delayed, probably in response to the environment. Development of the embryo (once implanted) may also be delayed, probably to ensure birth at a time when fruit is available during the rainy seasons. One species, Macroglossus minimus, exhibits asynchronous breeding and sperm storage, suggesting the importance of birth during an optimal (rainy) season.
Pregnant females usually leave social roosts to form nursery groups with other pregnant females. Females in nursery roosts form their own social network and take care of each other through mutual grooming. Gestation periods usually lasts 4 to 6 months, but can be longer if implantation is delayed. Birth patterns of pteropodids have been widely studied and usually occur during the wet periods both in the northern latitudes (February to April) and the southern latitudes (August to November). Species that are polyestrous will give birth during both of these rainy seasons. It is predicted that birth during these seasons yields high survival rates because lactation occurs when fruit availability is at a maximum. Birth is followed by postpartum estrous and subsequent mating. After weaning, young may stay with their mothers up to 4 months. Sexual maturity in juveniles is reached by 2 years old or earlier. Female sexual maturity is reached earlier than in males.
Key Reproductive Features: iteroparous ; seasonal breeding ; gonochoric/gonochoristic/dioecious (sexes separate); sexual ; viviparous ; delayed implantation ; post-partum estrous
Female pteropodids are primarily responsible for rearing the young. Lactation lasts anywhere from 7 weeks to 4 months, and the mother may care for her young slightly longer. In one genus (Dyacopterus), males with functional mammary glands have been reported lactating, which suggests the sharing of juvenile care among both parents.
Parental Investment: altricial ; pre-fertilization (Provisioning, Protecting: Female); pre-hatching/birth (Provisioning: Female, Protecting: Female); pre-weaning/fledging (Provisioning: Female, Protecting: Female); pre-independence (Provisioning: Female, Protecting: Female)
- Nowak, R. 1999. Walker's Mammals of the World. Baltimore and London: The Johns Hopkins University Press.
- Hood, C., J. Smith. 1989. Sperm storage in a tropical nectar-feeding bat, Macroglossus minimus (Pteropodidae). Journal of Mammalogy, 70: 404-406.
- Kofron, C. 2007. Reproduction of the dusky fruit bat Penthetor lucasi (Pteropodidae) in Brunei, Borneo. Mammalia: 166-171.
- Kretschmann, K., R. Hayes. 2004. Old World Fruit Bats I (Pteropus). Pp. 319-332 in M Hutchins, A Evans, J Jackson, eds. Grzimek's Animal Life Encyclopedia, Vol. 13: Mammals II, 2 Edition. Detroit: Gale.
- Storz, J., H. Bhat, T. Kunz. 2000. Social structure of a polygynous tent-making bat, Cynopterus sphinx (Megachiroptera). Journal of Zoology, London, 251: 151-165.
Molecular Biology and Genetics
Statistics of barcoding coverage
Specimen Records: 2200
Specimens with Sequences: 1282
Specimens with Barcodes: 1240
Species With Barcodes: 66
Public Records: 777
Public Species: 42
Public BINs: 45
Many factors threaten pteropodids throughout their range. Human activities have decimated populations of certain species directly through hunting or indirectly through habitat destruction. In Asia and Australia, deforestation is the most important contributor to pteropodid population decline. Some species are vulnerable to typhoons and hurricanes which may destroy roosting habitat on islands. The IUCN Red List of Threatened Species lists 5 species as recently extinct, 10 species as critically endangered, 19 species as endangered, 15 species as near threatened, and 39 species as vulnerable, suggesting that nearly half of all pteropodid species face significant threats to population viability.
- IUCN 2008, 2008. "2008 IUCN Red List of Threatened Species" (On-line). Accessed February 16, 2009 at http://www.iucnredlist.org/.
Relevance to Humans and Ecosystems
Many crop species are attractive food sources for pteropodids. Because cultivars are often developed from wild species, these commercial crops have the same characteristics that wild plants evolved to attract bats to their fruit. Fruit growers have experimented with visual, audio, and olfactory deterrents as well as electric wire to keep pteropodids from eating their crops. Pteropodids may also be dispersers of invasive plant species, as they consume crops introduced for cultivation and may disperse the seeds into natural habitat. Pteropodids have been indicated as reservoirs for a variety of viruses such as Ebola and other viruses in the family Paramyxoviridae. Hendra virus, Menangle virus, and Nipah virus have all been linked to pteropodids.
Negative Impacts: injures humans (carries human disease); crop pest
Larger species of pteropodids are hunted for their meat. Both subsistence and commercial hunting of Pteropus species have been reported. Consumer demand for Pteropus species on the island of Guam has been so great that it has resulted in the extinction of at least one species in the Pacific region. Flying foxes are also important in the dispersal and pollination of economically important plants. They attract tourists in some areas and produce accumulations of guano that can be used as fertilizer.
Positive Impacts: food ; ecotourism ; produces fertilizer; pollinates crops
Megabats constitute the suborder Megachiroptera, family Pteropodidae of the order Chiroptera (bats). They are also called fruit bats, old world fruit bats, or, especially the genera Acerodon and Pteropus, flying foxes. Fruit bats are restricted to the Old World in a tropical and subtropical distribution, ranging no further than the eastern Mediterranean and South Asia, and are absent from northwest Africa and southwest Australia.
Megabats, however contrary to their name, are not always large; the smallest species is 6 cm (2.4 in) long and thus smaller than some microbats. The largest attain a wingspan of 1.7 m (5.6 ft), weighing in at up to 1.6 kg (3.5 lb). Most fruit bats have large eyes, allowing them to orient themselves visually in twilight and inside caves and forests.
Their sense of smell is excellent. In contrast to the microbats, the fruit bats do not use echolocation (with one exception, the Egyptian fruit bat Rousettus egyptiacus, which uses high-pitched tongue clicks to navigate in caves).
Loss of echolocation
Megabats make up the only family (Pteropodidae) in order Chiroptera that is not capable of laryngeal echolocation, unlike the microbat. Echolocation and flight evolved early in the lineage of Chiropterans and echolocation was later lost in family Pteropodidae. Both echolocation and flight are energetically expensive processes for bats. The nature of the flight and echolocation mechanism of bats allows for creation of echolocation pulses with minimal energy use. Energetic coupling of these two processes is thought to have allowed for both energetically expensive processes to evolve in bats. It is hypothesized that the loss of echolocation is due to the uncoupling of flight and echolocation in megabats. The larger average body size of megabats compared to echolocating bats suggests that a larger body size disrupts the flight-echolocation coupling and made echolocation too energetically expensive to be conserved in megabats.
Behavior and ecology
Megabats are one of the two types of bats, the others being the insect-eating bats. The megabats are fruit bats, but most species cannot echolocate. They mostly roost in trees and shrubs, but only those that possess echolocation venture into the dark recesses of caves. Because they eat fruit, some megabat species are unpopular with orchard owners. Megabats are frugivorous or nectarivorous, i.e., they eat fruits or lick nectar from flowers. Often the fruits are crushed and only the juices are consumed. The teeth are adapted to bite through hard fruit skins. Large fruit bats must land to eat fruit, while the smaller species are able to hover with flapping wings in front of a flower or fruit.
Frugivorous bats aid the distribution of plants (and therefore, forests) by carrying the fruits with them and spitting the seeds or eliminating them elsewhere. Nectarivores actually pollinate visited plants. They bear long tongues that are inserted deep into the flower; pollen passed to the bat is then transported to the next blossom visited, thereby pollinating it. This relationship between plants and bats is a form of mutualism known as chiropterophily. Examples of plants that benefit from this arrangement include the baobabs of the genus Adansonia and the sausage tree (Kigelia).
As disease reservoirs
Researchers tested fruit bats for the presence of the Ebola virus between 2001 and 2003. Three species of bats tested positive for Ebola, but had no symptoms of the virus. This indicates the bats may be acting as a reservoir for the virus. Of the infected animals identified during these field collections, immunoglobulin G (IgG) specific for Ebola virus was detected in Hypsignathus monstrosus (Hammer-Headed Bat), Epomops franqueti (Franquet's Epauletted Fruit Bat), and Myonycteris torquata (Little Collared Fruit Bat).
Other viral diseases which can be carried by fruit bats include Australian bat lyssavirus and Henipavirus (notably Hendra virus and Nipah virus), both of which can prove fatal to humans. These bats have been shown to infect other species (specifically horses) with Hendra virus in Australian regions. Later, humans became infected with Hendra virus after being exposed to horse body fluids and excretions.
Fruit bats are considered a delicacy by South Pacific Islanders as well as in Micronesia. Consumption has been suggested as a cause of Lytico-Bodig disease on the Micronesian island of Guam, through bioaccumulation of a plant toxin that the bats are immune to.
Bats are usually thought to belong to one of two monophyletic groups, a view that is reflected in their classification into two suborders (Megachiroptera and Microchiroptera). According to this hypothesis, all living megabats and microbats are descendants of a common ancestor species that was already capable of flight.
However, there have been other views, and a vigorous debate persists to this date. For example, in the 1980s and 1990s, some researchers proposed (based primarily on the similarity of the visual pathways) that the Megachiroptera were in fact more closely affiliated with the primates than the Microchiroptera, with the two groups of bats having therefore evolved flight via convergence (see Flying primates theory). However, a recent flurry of genetic studies confirms the more longstanding notion that all bats are indeed members of the same clade, the Chiroptera. Other studies have recently suggested that certain families of microbats (possibly the horseshoe bats, mouse-tailed bats and the false vampires) are evolutionarily closer to the fruit bats than to other microbats.
List of species
- Subfamily Nyctimeninae
- Genus Nyctimene – tube-nosed fruit bats
- Broad-striped Tube-nosed Fruit Bat, Nyctimene aello
- Common Tube-nosed Fruit Bat, Nyctimene albiventer
- Pallas's Tube-nosed Fruit Bat, Nyctimene cephalotes
- Dark Tube-nosed Fruit Bat, Nyctimene celaeno
- Mountain Tube-nosed Fruit Bat, Nyctimene certans
- Round-eared Tube-nosed Fruit Bat, Nyctimene cyclotis
- Dragon Tube-nosed Fruit Bat, Nyctimene draconilla
- Keast's Tube-nosed Fruit Bat, Nyctimene keasti
- Island Tube-nosed Fruit Bat, Nyctimene major
- Malaita Tube-nosed Fruit Bat, Nyctimene malaitensis
- Demonic Tube-nosed Fruit Bat, Nyctimene masalai
- Lesser Tube-nosed Bat, Nyctimene minutus
- Philippine Tube-nosed Fruit Bat, Nyctimene rabori
- Eastern Tube-nosed Bat, Nyctimene robinsoni
- Nendo Tube-nosed Fruit Bat, Nyctimene sanctacrucis (early 20th century †)
- Umboi Tube-nosed Fruit Bat, Nyctimene vizcaccia
- Genus Paranyctimene
- Genus Nyctimene – tube-nosed fruit bats
- Subfamily Cynopterinae
- Genus Aethalops – pygmy fruit bats
- Genus Alionycteris
- Genus Balionycteris
- Genus Chironax
- Genus Cynopterus – dog-faced fruit bats or short-nosed fruit bats
- Lesser Short-nosed Fruit Bat, Cynopterus brachyotis
- Horsfield’s Fruit Bat, Cynopterus horsfieldii
- Peters’s Fruit Bat, Cynopterus luzoniensis
- Minute Fruit Bat, Cynopterus minutus
- Nusatenggara Short-nosed Fruit Bat, Cynopterus nusatenggara
- Greater Short-nosed Fruit Bat, Cynopterus sphinx
- Indonesian Short-nosed Fruit Bat, Cynopterus titthaecheilus
- Genus Dyacopterus – Dayak fruit bats
- Genus Haplonycteris
- Genus Latidens
- Genus Megaerops
- Genus Otopteropus
- Genus Penthetor
- Genus Ptenochirus – musky fruit bats
- Genus Sphaerias
- Genus Thoopterus
- Subfamily Harpiyonycterinae
- Genus Aproteles
- Genus Dobsonia – bare-backed fruit bats
- Andersen's Bare-backed Fruit Bat, Dobsonia anderseni
- Beaufort's Naked-backed Fruit Bat, Dobsonia beauforti
- Philippine Bare-backed Fruit Bat, Dobsonia chapmani
- Halmahera Naked-backed Fruit Bat, Dobsonia crenulata
- Biak Naked-backed Fruit Bat, Dobsonia emersa
- Sulawesi Naked-backed Fruit Bat, Dobsonia exoleta
- Solomon's Naked-backed Fruit Bat, Dobsonia inermis
- Lesser Naked-backed Fruit Bat, Dobsonia minor
- Moluccan Naked-backed Fruit Bat, Dobsonia moluccensis
- Panniet Naked-backed Fruit Bat, Dobsonia pannietensis
- Western Naked-backed Fruit Bat, Dobsonia peroni
- New Britain Naked-backed Fruit Bat, Dobsonia praedatrix
- Greenish Naked-backed Fruit Bat, Dobsonia viridis
- Genus Harpyionycteris
- Subfamily Macroglossinae
- Genus Macroglossus – long-tongued fruit bats
- Genus Melonycteris
- Genus Notopteris – long-tailed fruit bats
- Genus Syconycteris – blossom bats
- Subfamily Pteropodinae
- Genus Acerodon
- Genus Desmalopex
- Genus Eidolon – straw-coloured fruit bats
- Genus Mirimiri
- Fijian Monkey-faced Bat, Mirimiri acrodonta
- Genus Neopteryx
- Genus Pteralopex
- Genus Pteropus – flying foxes
- P. alecto species group
- P. caniceps species group
- P. chrysoproctus species group
- P. conspicillatus species group
- P. livingstonii species group
- P. mariannus species group
- P. melanotus species group
- P. molossinus species group
- P. neohibernicus species group
- P. niger species group
- P. personatus species group
- P. poliocephalus species group
- P. pselaphon species group
- Chuuk Flying Fox, Pteropus insularis
- Temotu Flying Fox, Pteropus nitendiensis
- Large Palau Flying Fox, Pteropus pilosus (19th century †)
- Bonin Flying Fox, Pteropus pselaphon
- Guam Flying Fox, Pteropus tokudae (1970s †)
- Insular Flying Fox, Pteropus tonganus
- Vanikoro Flying Fox, Pteropus tuberculatus
- New Caledonia Flying Fox, Pteropus vetulus
- P. samoensis species group
- P. scapulatus species group
- P. subniger species group
- Admiralty Flying Fox, Pteropus admiralitatum
- Dusky Flying Fox, Pteropus brunneus (19th century †)
- Ryukyu Flying Fox, Pteropus dasymallus
- Nicobar Flying Fox, Pteropus faunulus
- Gray Flying Fox, Pteropus griseus
- Ontong Java Flying Fox, Pteropus howensis
- Small Flying Fox, Pteropus hypomelanus
- Ornate Flying Fox, Pteropus ornatus
- Little Golden-mantled Flying Fox, Pteropus pumilus
- Philippine Gray Flying Fox, Pteropus speciosus
- Small Mauritian Flying Fox, Pteropus subniger (19th century †)
- P. vampyrus species group
- incertae sedis
- Genus Styloctenium
- Subfamily Rousettinae
- Genus Eonycteris – dawn fruit bats
- Genus Rousettus – rousette fruit bats
- Subgenus Boneia
- Subgenus Rousettus
- Geoffroy's Rousette, Rousettus amplexicaudatus
- Sulawesi Rousette, Rousettus celebensis
- Egyptian Rousette (Egyptian Fruit Bat), Rousettus aegyptiacus
- Leschenault's Rousette, Rousettus leschenaulti
- Linduan Rousette, Rousettus linduensis
- Comoro Rousette, Rousettus obliviosus
- Bare-backed Rousette, Rousettus spinalatus
- Subgenus Stenonycteris
- Subfamily Epomophorinae
- Tribe Epomophorini
- Genus Epomophorus – epauletted fruit bats
- Angolan Epauletted Fruit Bat, Epomophorus angolensis
- Ansell's Epauletted Fruit Bat, Epomophorus anselli
- Peters's Epauletted Fruit Bat, Epomophorus crypturus
- Gambian Epauletted Fruit Bat, Epomophorus gambianus
- Lesser Angolan Epauletted Fruit Bat, Epomophorus grandis
- Ethiopian Epauletted Fruit Bat, Epomophorus labiatus
- East African Epauletted Fruit Bat, Epomophorus minimus
- Minor Epauletted Fruit Bat, Epomophorus minor
- Wahlberg's Epauletted Fruit Bat, Epomophorus wahlbergi
- Genus Epomops – epauletted bats
- Genus Hypsignathus
- Genus Micropteropus – dwarf epauletted bats
- Genus Nanonycteris
- Genus Epomophorus – epauletted fruit bats
- Tribe Myonycterini
- Genus Lissonycteris
- Genus Megaloglossus
- Genus Myonycteris – little collared fruit bats
- Tribe Plerotini
- Tribe Scotonycterini
- Tribe Epomophorini
- Mickleburgh, Hutson and Racey. "Old World Fruit Bats:Introduction". International Union for Conservation of Nature. Retrieved July 19, 2013.
- Kenneth Cody Luzynski; Emily Margaret Sluzas; Megan Marie Wallen. "PteropodidaeOld World fruit bats". Animal Diversity Web/University of Michigan.
- Charles H. Smith. "PTEROPODIDAE (Fruit Bats/Flying Foxes)". MAMMFAUN/University of Kentucy. (includes range map)
- E.g., the Mauritian Tomb Bat. See Garbutt, Nick. "Mauritian Tomb Bat." Mammals of Madagascar: A Complete Guide. Yale University Press. 2007. p. 67. 
- Nowak, R. M., editor (1999). Walker's Mammals of the World. Vol. 1. 6th edition. pp. 264–271. ISBN 0-8018-5789-9
- Matti Airas. "Echolocation in bats". HUT, Laboratory of Acoustics and Audio Signal Processing. p. 4. Retrieved July 19, 2013.
- Springer, M.S., E.C. Teeling, O. Madsen, M.J. Stanhope, and W.W. de Jong (2001). "Integrated fossil and molecular data reconstruct bat echolocation". Proceedings of the National Academy of Sciences 98 (11): 6241–6246. Bibcode:2001PNAS...98.6241S. doi:10.1073/pnas.111551998.
- Speakman, J.R., and P.A. Racey (1991). "No cost of echolocation for bats in flight". Nature 350 (6317): 421–423. Bibcode:1991Natur.350..421S. doi:10.1038/350421a0. PMID 2011191.
- Lancaster, W.C., O.W. Henson, and A.W. Keating (1995). "Respiratory muscle activity in relation to vocalization in flying bats". Journal of Experimental Biology 198 (Pt 1): 175–191. PMID 7891034.
- Altringham, J.D. (2011). Echolocation and other senses. New York: Oxford University Press.
- Hutcheon, J.M., and T.J. Garland (1995). "Are megabats big?". Journal of Mammalian Evolution 11 (3/4): 257–277. doi:10.1023/B:JOMM.0000047340.25620.89.
- National Geographic, October 2007. "Deadly Contact," David Quammen, pp. 78–105.
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- Bat World Sanctuary
- Rodrigues Fruit Bats[dead link]
- Bat Conservation International
- Criticism of the molecular evidence for bat monophyly
- Brief history of Megachiroptera / Megabats
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