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In , Danish mammalogist Knud Andersen divided Pteropodidae into three subfamilies: Macroglossinae, Pteropinae corrected to Pteropodinae , and Harpyionycterinae.
Other subfamilies and tribes within Pteropodidae have also undergone changes since Andersen's publication. Three of the subfamilies proposed in based on morphology received support: Cynopterinae, Harpyionycterinae, and Nyctimeninae.
All species formerly included in Epomophorinae were moved to Rousettinae, which was subdivided into additional tribes. The genus Eidolon , formerly in the tribe Rousettini of Rousettinae, was moved to its own subfamily, Eidolinae.
In , an additional pteropodid subfamily, Propottininae, was proposed, representing one extinct species described from a fossil discovered in Africa, Propotto leakeyi.
The fossil record for pteropodid bats is the most incomplete of any bat family. Several factors could explain why so few pteropodid fossils have been discovered: tropical regions where their fossils might be found are undersampled relative to Europe and North America; conditions for fossilization are poor in the tropics, which could lead to fewer fossils overall; and fossils may have been created, but they may have been destroyed by subsequent geological activity.
Pteropodidae split from the superfamily Rhinolophoidea which contains all the other families of the suborder Yinpterochiroptera approximately 58 Mya million years ago.
The family Pteropodidae likely originated in Australasia based on biogeographic reconstructions. Megabats reached Africa in at least four distinct events.
The four proposed events are represented by 1 Scotonycteris , 2 Rousettus , 3 Scotonycterini, and 4 the "endemic Africa clade", which includes Stenonycterini, Plerotini, Myonycterini, and Epomophorini, according to a study.
It is unknown when megabats reached Africa, but several tribes Scotonycterini, Stenonycterini, Plerotini, Myonycterini, and Epomophorini were present by the Late Miocene.
How megabats reached Africa is also unknown. It has been proposed that they could have arrived via the Middle East before it became more arid at the end of the Miocene.
Conversely, they could have reached the continent via the Gomphotherium land bridge , which connected Africa and the Arabian Peninsula to Eurasia.
The genus Pteropus flying foxes , which is not found on mainland Africa, is proposed to have dispersed from Melanesia via island hopping across the Indian Ocean ;  this is less likely for other megabat genera, which have smaller body sizes and thus have more limited flight capabilities.
Megabats are the only family of bats incapable of laryngeal echolocation. It is unclear whether the common ancestor of all bats was capable of echolocation, and thus echolocation was lost in the megabat lineage, or multiple bat lineages independently evolved the ability to echolocate the superfamily Rhinolophoidea and the suborder Yangochiroptera.
This unknown element of bat evolution has been called a "grand challenge in biology". This evidence supports that laryngeal echolocation evolved once among bats, and was lost in pteropodids, rather than evolving twice independently.
Both echolocation and flight are energetically expensive processes. Instead of pressurizing a bolus of air for the production of sound, laryngeally echolocating bats likely use the force of the downbeat of their wings to pressurize the air, cutting energetic costs by synchronizing wingbeats and echolocation.
The family Pteropodidae is divided into six subfamilies represented by 46 genera :  . Megabats are so called for their larger weight and size; the largest, the great flying fox Pteropus neohibernicus weighs up to 1.
In reality, these genera are outliers, creating a misconception of the true size of most megabat species. Megabats can be distinguished from microbats in appearance by their dog-like faces, by the presence of claws on the second digit see Megabat Postcrania , and by their simple ears.
Megabats of the genus Nyctimene appear less dog-like, with shorter faces and tubular nostrils. These include countershading in four percent of species, a neck band or mantle in five percent of species, stripes in ten percent of species, and spots in nineteen percent of species.
Unlike microbats, megabats have a greatly reduced uropatagium , which is an expanse of flight membrane that runs between the hind limbs.
In Dobsonia species, the wings attach nearer the spine, giving them the common name of "bare-backed" or "naked-backed" fruit bats.
Megabats have large orbits , which are bordered by well-developed postorbital processes posteriorly. The postorbital processes sometimes join to form the postorbital bar.
The snout is simple in appearance and not highly modified, as is seen in other bat families. The premaxilla is well-developed and usually free,  meaning that it is not fused with the maxilla ; instead, it articulates with the maxilla via ligaments , making it freely movable.
In genera with shorter faces Penthetor , Nyctimene , Dobsonia , and Myonycteris , the skull has little to no bending.
The number of teeth varies among megabat species; totals for various species range from 24 to All species have two upper and lower canine teeth.
The number of premolars is variable, with four or six each of upper and lower premolars. The first upper and lower molars are always present, meaning that all megabats have at least four molars.
The remaining molars may be present, present but reduced, or absent. Like most mammals, megabats are diphyodont , meaning that the young have a set of deciduous teeth milk teeth that falls out and is replaced by permanent teeth.
For most species, there are 20 deciduous teeth. As is typical for mammals,  the deciduous set does not include molars. The scapulae shoulder blades of megabats have been described as the most primitive of any chiropteran family.
The primitive insertion of the omohyoid muscle from the clavicle collarbone to the scapula is laterally displaced more towards the side of the body —a feature also seen in the Phyllostomidae.
The shoulder also has a well-developed system of muscular slips narrow bands of muscle that augment larger muscles that anchor the tendon of the occipitopollicalis muscle muscle in bats that runs from base of neck to the base of the thumb  to the skin.
While microbats only have claws on the thumbs of their forelimbs, most megabats have a clawed second digit as well;  only Eonycteris , Dobsonia , Notopteris , and Neopteryx lack the second claw.
The second digit is incapable of flexion. Megabats' hindlimbs have the same skeletal components as humans. Most megabat species have an additional structure called the calcar , a cartilage spur arising from the calcaneus.
The structure exists to stabilize the uropatagium, allowing bats to adjust the camber of the membrane during flight. Megabats lacking the calcar or spur include Notopteris , Syconycteris , and Harpyionycteris.
All five digits of the foot flex in the direction of the sagittal plane , with no digit capable of flexing in the opposite direction, as in the feet of perching birds.
Flight is very energetically expensive, requiring several adaptations to the cardiovascular system. During flight, bats can raise their oxygen consumption by twenty times or more for sustained periods; human athletes can achieve an increase of a factor of twenty for a few minutes at most.
Among these two species, the gray-headed flying fox Pteropus poliocephalus and the Egyptian fruit bat Rousettus aegyptiacus , maximum heart rates in flight varied between beats per minute gray-headed flying fox and beats per minute Egyptian fruit bat.
The maximum number of breaths per minute ranged from gray-headed flying fox to straw-colored fruit bat. While terrestrial mammals such as shrews have a lung volume of 0.
Megabats have rapid digestive systems, with a gut transit time of half an hour or less. There is no distinct difference between the small and large intestine, nor a distinct beginning of the rectum.
They have very high densities of intestinal microvilli , which creates a large surface area for the absorption of nutrients. Like all bats, megabats have much smaller genomes than other mammals.
A study of 43 megabat species found that their genomes ranged from 1. All values were much lower than the mammalian average of 3.
Megabats have even smaller genomes than microbats, with a mean weight of 2. It was speculated that this difference could be related to the fact that the megabat lineage has experienced an extinction of the LINE1 —a type of long interspersed nuclear element.
With very few exceptions, megabats do not echolocate , and therefore rely on sight and smell to navigate.
A study that examined the eyes of 18 megabat species determined that the common blossom bat Syconycteris australis had the smallest eyes at a diameter of 5.
At high brightness levels, megabat visual acuity is poorer than that of humans; at low brightness it is superior.
Pteropus bats are dichromatic , possessing two kinds of cone cells. The other three genera, with their lack of S-cones, are monochromatic , unable to see color.
All genera had very high densities of rod cells, resulting in high sensitivity to light, which corresponds with their nocturnal activity patterns.
In Pteropus and Rousettus , measured rod cell densities were ,—, per square millimeter, equal to or exceeding other nocturnal or crepuscular animals such as the house mouse , domestic cat , and domestic rabbit.
Megabats use smell to find food sources like fruit and nectar. The secretions of these glands vary by species—of the 65 chemical compounds isolated from the glands of four species, no compound was found in all species.
Megabats possess the TAS1R2 gene, meaning they have the ability to detect sweetness in foods. This gene is present among all bats except vampire bats.
Like all other bats, megabats cannot taste umami , due to the absence of the TAS1R1 gene. Among other mammals, only giant pandas have been shown to lack this gene.
Megabats, like all bats, are long-lived relative to their size for mammals. Some captive megabats have had lifespans exceeding thirty years.
Different species of megabats have reproductive adaptations that lengthen the period between copulation and giving birth. Some species such as the straw-coloured fruit bat have the reproductive adaptation of delayed implantation , meaning that copulation occurs in June or July, but the zygote does not implant into the uterine wall until months later in November.
The litter size of all megabats is usually one. At birth, megabat offspring are, on average, This is the smallest offspring-to-mother ratio for any bat family; across all bats, newborns are Megabat offspring are not easily categorized into the traditional categories of altricial helpless at birth or precocial capable at birth.
Species such as the greater short-nosed fruit bat are born with their eyes open a sign of precocial offspring , whereas the Egyptian fruit bat offspring's eyes do not open until nine days after birth a sign of altricial offspring.
As with nearly all bat species, males do not assist females in parental care. Many megabat species are highly gregarious or social.
Megabats will vocalize to communicate with each other, creating noises described as "trill-like bursts of sound",  honking,  or loud, bleat-like calls  in various genera.
At least one species, the Egyptian fruit bat, is capable of a kind of vocal learning called vocal production learning, defined as "the ability to modify vocalizations in response to interactions with conspecifics".
It has been postulated that these dialect differences may result in individuals of different colonies communicating at different frequencies, for instance.
Megabat social behavior includes using sexual behaviors for more than just reproduction. Evidence suggests that female Egyptian fruit bats take food from males in exchange for sex.
Paternity tests confirmed that the males from which each female scrounged food had a greater likelihood of fathering the scrounging female's offspring.
Megabats are mostly nocturnal and crepuscular , though some have been observed flying during the day. Diurnal taxa include a subspecies of the black-eared flying fox Pteropus melanotus natalis , the Mauritian flying fox Pteropus niger , the Caroline flying fox Pteropus molossinus , a subspecies of Pteropus pelagicus P.
A summary of forty-one megabat genera noted that twenty-nine are tree-roosting genera. A further eleven genera roost in caves, and the remaining six genera roost in other kinds of sites human structures, mines, and crevices, for example.
Tree-roosting species can be solitary or highly colonial , forming aggregations of up to one million individuals. Cave-roosting species form aggregations ranging from ten individuals up to several thousand.
Highly colonial species often exhibit roost fidelity, meaning that their trees or caves may be used as roosts for many years.
Solitary species or those that aggregate in smaller numbers have less fidelity to their roosts. Most megabats are primarily frugivorous.
Megabats fly to roosting and foraging resources. They typically fly straight and relatively fast for bats; some species are slower with greater maneuverability.
Most megabats have below-average aspect ratios ,  which is measurement relating wingspan and wing area. Megabats play an important role in seed dispersal.
As a result of their long evolutionary history, some plants have evolved characteristics compatible with bat senses, including fruits that are strongly scented, brightly colored, and prominently exposed away from foliage.
The bright colors and positioning of the fruit may reflect megabats' reliance on visual cues and inability to navigate through clutter.
In a study that examined the fruits of more than forty fig species, only one fig species was consumed by both birds and megabats; most species are consumed by one or the other.
Bird-consumed figs are frequently red or orange, while megabat-consumed figs are often yellow or green. This heightens megabats' capacity to disperse seeds far from parent trees.
Megabats, especially those living on islands, have few native predators: species like the small flying fox Pteropus hypomelanus have no known natural predators.
The mangrove monitor , which is a native predator for some megabat species but an introduced predator for others, opportunistically preys on megabats, as it is a capable tree climber.
The island is now considered a sink for the Mariana fruit bat, as its population there relies on bats immigrating from the nearby island of Rota to bolster it rather than successful reproduction.
Megabats are the hosts of several parasite taxa. Known parasites include Nycteribiidae and Streblidae species "bat flies" ,   as well as mites of the genus Demodex.
Megabats are widely distributed in the tropics of the Old World , occurring throughout Africa, Asia, Australia, and throughout the islands of the Indian Ocean and Oceania.
Of those twenty-eight species, twenty-four are only found in tropical or subtropical climates. The remaining four species are mostly found in the tropics, but their ranges also encompass temperate climates.
In respect to habitat types, eight are exclusively or mostly found in forested habitat; nine are found in both forests and savannas ; nine are found exclusively or mostly in savannas; and two are found on islands.
Only one African species, the long-haired rousette Rousettus lanosus , is found mostly in montane ecosystems , but an additional thirteen species' ranges extend into montane habitat.
Outside of Southeast Asia, megabats have relatively low species richness in Asia. The Egyptian fruit bat is the only megabat whose range is mostly in the Palearctic realm ;  it and the straw-colored fruit bat are the only species found in the Middle East.
In China, only six species of megabat are considered resident, while another seven are present marginally at the edge of their ranges , questionably due to possible misidentification , or as accidental migrants.
Other habitat types include human-modified land 66 species , caves 23 species , savanna 7 species , shrubland 4 species , rocky areas 3 species , grassland 2 species , and desert 1 species.
In Australia, five genera and eight species of megabat are present. In Oceania, the countries of Palau and Tonga have the fewest megabat species, with one each.
Papua New Guinea has the greatest number of species with thirty-six. Other habitat types include human-modified land 42 species , caves 9 species , savanna 5 species , shrubland 3 species , and rocky areas 3 species.
Megabats are killed and eaten as bushmeat throughout their range. Bats are consumed extensively throughout Asia, as well as in islands of the West Indian Ocean and the Pacific, where Pteropus species are heavily hunted.
In continental Africa where no Pteropus species live, the straw-coloured fruit bat, the region's largest megabat, is a preferred hunting target.
In Guam, consumption of the Mariana fruit bat exposes locals to the neurotoxin beta-Methylamino-L-alanine BMAA which may later lead to neurodegenerative diseases.
Megabats are the reservoirs of several viruses that can affect humans and cause disease. Species that have tested positive for the presence of EBOV include Franquet's epauletted fruit bat Epomops franqueti , the hammer-headed fruit bat, and the little collared fruit bat.
Additionally, antibodies against EBOV have been found in the straw-coloured fruit bat, Gambian epauletted fruit bat Epomophorus gambianus , Peters's dwarf epauletted fruit bat Micropteropus pusillus , Veldkamp's dwarf epauletted fruit bat Nanonycteris veldkampii , Leschenault's rousette, and the Egyptian fruit bat.
Scientists hypothesize that humans initially become infected through contact with an infected animal such as a megabat or non-human primate. Other megabats implicated as disease reservoirs are primarily Pteropus species.
Notably, flying foxes can transmit Australian bat lyssavirus , which, along with the rabies virus , causes rabies. Australian bat lyssavirus was first identified in ; it is very rarely transmitted to humans.
Transmission occurs from the bite or scratch of an infected animal but can also occur from getting the infected animal's saliva in a mucous membrane or an open wound.
Exposure to flying fox blood, urine, or feces cannot cause infections of Australian bat lyssavirus. Since , there have been three records of people becoming infected with it in Queensland —each case was fatal.
Flying foxes are also reservoirs of henipaviruses such as Hendra virus and Nipah virus. Hendra virus was first identified in ; it rarely occurs in humans.
From to , there have been seven reported cases of Hendra virus affecting people, four of which were fatal.
The hypothesized primary route of human infection is via contact with horses that have come into contact with flying fox urine.
Nipah virus was first identified in in Malaysia. Since , there have been several Nipah outbreaks in Malaysia, Singapore , India, and Bangladesh, resulting in over casualties.
A outbreak in Kerala, India resulted in 19 humans becoming infected—17 died. Humans can contract Nipah virus from direct contact with flying foxes or their fluids, through exposure to an intermediate host such as domestic pigs , or from contact with an infected person.
The practice of date palm sap collection involves placing collecting pots at date palm trees. Indian flying foxes have been observed licking the sap as it flows into the pots, as well as defecating and urinating in proximity to the pots.
In this way, humans who drink the palm sap can be exposed to the bats' viruses. The use of bamboo skirts on collecting pots lowers the risk of contamination from bat fluids.
Flying foxes can transmit several non-lethal diseases as well, such as Menangle virus  and Nelson Bay virus. Megabats, particularly flying foxes, are featured in indigenous cultures and traditions.
Folk stories from Australia and Papua New Guinea feature them. Indigenous societies in Oceania used parts of flying foxes for functional and ceremonial weapons.
In the Solomon Islands, people created barbs out of their bones for use in spears. There are modern and historical references to flying fox byproducts used as currency.
In New Caledonia, braided flying fox fur was once used as currency. The canine teeth are strung together on necklaces that are used as currency.
The Makira flying fox Pteropus cognatus is also hunted, despite its smaller teeth. Deterring people from using flying fox teeth as currency may be detrimental to the species, with Lavery and Fasi noting, "Species that provide an important cultural resource can be highly treasured.
Even if flying foxes were no longer hunted for their teeth, they would still be killed for bushmeat; therefore, retaining their cultural value may encourage sustainable hunting practices.
The practice of hunting bats shouldn't necessarily be stopped, it needs to be managed sustainably. As of , the International Union for Conservation of Nature IUCN evaluated a quarter of all megabat species as threatened , which includes species listed as critically endangered , endangered , and vulnerable.
Megabats are substantially threatened by humans, as they are hunted for food and medicinal uses. Additionally, they are culled for actual or perceived damage to agriculture, especially to fruit production.
The status breakdown is as follows: . Megabats are threatened by habitat destruction by humans. Deforestation of their habitats has resulted in the loss of critical roosting habitat.
Deforestation also results in the loss of food resource, as native fruit-bearing trees are felled. Habitat loss and resulting urbanization leads to construction of new roadways, making megabat colonies easier to access for overharvesting.
Additionally, habitat loss via deforestation compounds natural threats, as fragmented forests are more susceptible to damage from typhoon -force winds.
Guano mining is a livelihood in some countries within their range, bringing people to caves. Caves are also disturbed by mineral mining and cave tourism.
Megabats are also killed by humans, intentionally and unintentionally. Half of all megabat species are hunted for food, in comparison to only eight percent of insectivorous species,  while human persecution stemming from perceived damage to crops is also a large source of mortality.
Some megabats have been documented to have a preference for native fruit trees over fruit crops, but deforestation can reduce their food supply, causing them to rely on fruit crops.
Mortality also occurs via accidental entanglement in netting used to prevent the bats from eating fruit.
In one Australian orchard, it is estimated that over 21, bats were electrocuted to death in an eight-week period.
Climate change causes flying fox mortality and is a source of concern for species persistence. Extreme heat waves in Australia have been responsible for the deaths of more than 30, flying foxes from to Females and young bats are most susceptible to extreme heat, which affects a population's ability to recover.
Because many species are endemic to a single island, they are vulnerable to random events such as typhoons.
A typhoon halved the remaining population of the Rodrigues flying fox Pteropus rodricensis. Typhoons result in indirect mortality as well: because typhoons defoliate the trees, they make megabats more visible and thus more easily hunted by humans.
Food resources for the bats become scarce after major storms, and megabats resort to riskier foraging strategies such as consuming fallen fruit off the ground.
There, they are more vulnerable to depredation by domestic cats, dogs, and pigs. Flying foxes, including the endangered Mariana fruit bat,   have been nearly exterminated from the island of Anatahan following a series of eruptions beginning in From Wikipedia, the free encyclopedia.
For other uses, see Fruit bat disambiguation. For other species known as fruit-eating bats, see Leaf-nosed bat. Family of relatively large flying mammals fruit bats.
Megabats of various subfamilies. Clockwise from upper left: greater short-nosed fruit bat Cynopterinae , Indian flying fox Pteropodinae , Egyptian fruit bat Rousettinae , eastern tube-nosed bat Nyctimeninae.
Main article: List of fruit bats. Play media. Classification of mammals: above the species level. Columbia University Press.
Acta Chiropterologica. London Medical Repository 25 : United States National Museum Bulletin. Retrieved 22 May Taxonomy of Australian Mammals.
Csiro Publishing. Proceedings of the National Academy of Sciences. Bibcode : PNAS Scientific Reports. Bibcode : NatSR Current Biology.
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