Temporal range: Holocene
|A large flying fox (Pteropus vampyrus)|
|Worldwide distribution of flying foxes|
Pteropus (suborder Yinpterochiroptera) is a genus of bats which are among the largest in the world. They are commonly known as fruit bats or flying foxes, among other colloquial names. They live in the tropics and subtropics of Asia (including the Indian subcontinent), Australia, East Africa, and some oceanic islands in the Indian and Pacific Oceans. There are at least 60 extant species in the genus.
Flying foxes eat fruit and other plant matter, and occasionally consume insects as well. They locate resources with their keen sense of smell. Most, but not all, are nocturnal. They navigate with keen eyesight, as they cannot echolocate. They have long life spans and low reproductive outputs, with females of most species producing only one offspring per year. Their slow life history makes their populations vulnerable to threats such as overhunting, culling, and natural disasters. Six flying fox species have been made extinct in modern times by overhunting. Flying foxes are often persecuted for their real or perceived role in damaging crops. They are ecologically beneficial by assisting in the regeneration of forests via seed dispersal. They benefit ecosystems and human interests by pollinating plants.
Flying foxes are relevant to humans as a source of disease, as they are the reservoirs of rare but fatal disease agents including Australian bat lyssavirus, which causes rabies, and Hendra virus; seven known human deaths have resulted from these two diseases. Nipah virus is also transmitted by flying foxes—it affects more people, with over 100 attributed fatalities. They have cultural significance to indigenous people, with appearances in traditional art, folklore, and weaponry. Their fur and teeth were used as currency in the past. Some cultures still use their teeth as currency today.
- 1 Taxonomy and etymology
- 2 Description
- 3 Evolution
- 4 Biology and ecology
- 5 Conservation
- 6 Relationship to people
- 7 Species
- 8 References
Taxonomy and etymology
The genus name Pteropus was coined by French zoologist Mathurin Jacques Brisson in 1762. Prior to 1998, genus authority was sometimes given to German naturalist Johann Christian Polycarp Erxleben. Although the Brisson publication (1762) predated the Erxleben publication (1777), thus giving him preference under the Principle of Priority, some authors gave preference to Erxleben as genus authority because Brisson's publication did not consistently use binomial nomenclature. In 1998, the International Commission on Zoological Nomenclature (ICZN) decided that Brisson's 1762 publication was a "rejected work" for nomenclatural purposes. Despite rejecting the majority of the publication, the ICZN decided to conserve a dozen generic names from the work and retain Brisson as authority, including Pteropus.
The type species of the genus is the Mauritian flying fox, Pteropus niger (described as Vespertilio vampyrus niger by Robert Kerr in 1792). The decision to designate P. niger as the type species was made by the ICZN through their plenary powers over biological nomenclature. "Pteropus" comes from Ancient Greek "pterón" meaning "wing" and "poús" meaning "foot." The phrase "flying fox" has been used to refer to Pteropus bats since at least 1759.
Flying fox species vary in body weight, ranging from 120–1,600 g (0.26–3.53 lb). Across all species, males are usually larger than females. The large flying fox has the longest forearm length and reported wingspan of any species, but some species exceed it in weight. Its wingspan is up to 1.5 m (4 ft 11 in), and it can weigh up to 1.1 kg (2.4 lb). The Indian and great flying foxes are heavier, at 1.6 kg (3.5 lb) and 1.45 kg (3.2 lb), respectively. Outside this genus, the giant golden-crowned flying fox (genus Acerodon) is the only bat with similar dimensions.
Most flying fox species are considerably smaller and generally weigh less than 600 g (21 oz). Smaller species such as the masked, Temminck's, Guam, and dwarf flying foxes all weigh less than 170 g (6.0 oz).
The pelage is long and silky with a dense underfur. In many species, individuals have a "mantle" of contrasting fur color on the back of their head, the shoulders, and the upper back. They lack tails. As the common name "flying fox" suggests, their heads resemble that of a small fox because of their small ears and large eyes. Females have one pair of mammae located in the chest region. Their ears are long and pointed at the tip and lack tragi. The outer margin of each ear forming an unbroken ring. The toes have sharp, curved claws. While microbats only have a claw on each thumb of their forelimbs, flying foxes additionally have a claw on each index finger.
Skull and dentition
A flying fox skull is composed of 24 bones. The snout is made of 7, the cranium of 16, and the mandible is a single bone. It has a large and bulbous braincase. Like all mammals, flying foxes have three middle ear ossicles which assist in transmitting sound to the brain. Flying fox skulls continue to develop after they are born. Compared to adults, young flying foxes have very short snouts; as they reach maturity, the maxilla elongates, gaining bone between the zygomatic processes and the canine teeth.
Based on the grey-headed flying fox's development, pups are born with some milk teeth already erupted: canines and incisors. By 9 days old, all the milk teeth have emerged, with a dental formula of 22.214.171.124 and a total of 20 teeth. By 140 days old (4.6 months), all the milk teeth have fallen and been replaced by permanent teeth. The canines are usually replaced first, followed by the premolars, incisors, and then molars. The adult dental formula is 126.96.36.199 for a total of 34 teeth. The occlusal surface of the molars is generally smooth but with longitudinal furrows.
Flying foxes have simple digestive tracts; the time between ingestion and excretion is as short as 12 minutes. They lack both a cecum and an appendix. The stomach has marked cardiac and fundic regions.
The megabats, including flying foxes, have the greatest encephalization quotient (brain size relative to body size) of any bat family at 1.20. This value is equivalent to that of domestic dogs. Flying foxes display behaviors that indicate a reliance on long-term information storage. Though they have wide-ranging movements and cover thousands of square kilometers annually, they are consistently able to locate the same resource patches and roosts. They will visit these resource patches consistently in a strategy known as trap-lining. They can also be conditioned to perform behaviors, such as one study where spectacled flying foxes were trained to pull a lever using juice as a reinforcement. In a follow-up to the initial study, individuals who had learned to pull the lever to receive juice still did so 3.5 years later.
Flying foxes rely heavily on their sense of smell. They have large olfactory bulbs to process scents. They use scent to locate food, for mothers to locate their pups, and for mates to locate each other. Males have enlarged androgen-sensitive sebaceous glands on their shoulders that they use for scent-marking their territories, particularly during the mating season. 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. Males also engage in "urine washing", meaning that they coat themselves in their own urine.
Flying foxes do not echolocate, and therefore rely on sight to navigate. Their eyes are relatively large and positioned on the front of their heads, giving them binocular vision. Like most mammals, though not primates, they are dichromatic. They have both rods and cones; they have "blue" cones that detect short-wavelength light and "green" cones that detect medium-to-long-wavelengths. The rods greatly outnumber the cones, however, as cones comprise only 0.5% of photoreceptors. Flying foxes are adapted to seeing in low-light conditions.
|Phylogeny of Pteropodidae|
Flying foxes are poorly represented in the fossil record. Relative to the current number of extant species, Pteropodidae has one of the most incomplete fossil records of any bat group. As of 2014, there are no flying fox fossils from before the Holocene. Many flying foxes live in the Tropics, however, where conditions for fossilization are poor. Based on molecular evolution, flying foxes diverged from a common ancestor with Rousettus 28–18 million years ago and from their sister taxa Neopteryx and Acerodon 6.6–10.6 million years ago. Neopteryx, Acerodon, Desmalopex, Melonycteris, Mirimiri, Pteralopex, and Styloctenium are all relatively closely related to the flying foxes, as they are the other members of its subfamily Pteropodinae.
Phylogenetic analysis indicates that flying foxes diversified rapidly in an explosive evolutionary radiation, creating many taxa in a relatively short time frame. Most flying fox lineages emerged after the Zanclean, with two major clades created: one consisting of the Indian Ocean species and the other of the Melanesian, Micronesian, Australian, and insular Southeast Asian species. Flying foxes likely originated on mainland Asia; molecular data suggests that there were at least three colonization events into the Indian Ocean. One event resulted in the Livingstone's fruit bat and the Pemba flying fox, which are the westernmost flying foxes. A second colonization event resulted in the Rodrigues flying fox to Rodrigues Island; while a third event resulted in several species diverging to Mauritius, Seychelles, Madagascar, and Aldabra.
With one possible exception of the masked flying fox (P. personatus), flying foxes are likely monophyletic. There are over 60 extant species of flying fox. Flying foxes are now present from the western Indian Ocean midway through the Pacific Ocean as far east as the Cook Islands. They are found in tropical and subtropical climates.
Biology and ecology
Reproduction and life cycle
Many species of flying fox are polygynandrous, meaning that each individual will mate with several other individuals. The Samoa flying fox is a notable exception because it is monogamous. Flying fox sexual behaviors include oral sex in addition to intercourse, with fellatio and cunnilingus observed between opposite sexes, as well as homosexual fellatio in at least one species, the Bonin flying fox. Opposite-sex oral sex is associated with increased duration of intercourse, while same-sex fellatio is hypothesized to encourage colony formation of otherwise-antagonistic males in colder climates.
Flying fox gestation length varies among species; gestation length is 140–190 days (4.6–6.3 months). Females have a litter size of one young at a time, called a pup. Twins have been occasionally documented in some species, however. Twins can be fraternal, identical, or the result of superfetation. Pups are altricial and sparsely furred at birth. Pups are relatively small at birth, weighing approximately 12% of the mother's weight. Bats in other genera can have pups that weigh as much as 30% of the mother's weight at birth. They cling to their mothers' abdomens, gripping her fur with their thumb claws and teeth; females carry the pups for the first several weeks of life. After this, the females may leave the pups behind at the roost at night while they forage. As with nearly all bat species, males do not assist females in parental care. While male flying foxes of at least one species, the Bismarck masked flying fox, can lactate, it is unclear if the lactation is functional and males actually nurse pups or if it is a result of stress or malnutrition. Pups fledge beginning at 3 months old, but may not be weaned until 4–6 months old. Pups may stay with their mothers until age one. Flying foxes do not reach sexual maturity until 1.5–2 years old. Females can have up to two litters annually, though one is the norm due to the long weaning period. Most flying foxes are seasonal breeders and give birth in the spring, though the Mariana fruit bat seems to have aseasonal breeding with new pups documented throughout the year. Females remain fertile with no decrease in reproductive capability for at least the first 12 or 13 years of life.
Flying foxes, like all bats, are long-lived relative to their size. In the wild, average lifespans are likely 15 years. However, individuals part of populations that face excessive disturbance may have lifespans as short as 7.1 years. In captivity, individuals can live approximately 20–28 years. The longest-lived flying fox was an Indian flying fox who lived in captivity. It lived to age 31 years and 5 months.
Most flying fox species are gregarious and form large aggregations of individuals called colonies or "camps." The large flying fox forms colonies of up to 15,000 individuals, while the little red flying fox forms colonies of up to 100,000 individuals. A few species and subspecies, such as Orii's flying fox (P. dasymallus inopinatus) and the Ceram fruit bat, are solitary.
Colony size varies throughout the year in response to biological needs. The grey-headed flying fox forms harems during the breeding season consisting of one male and up to six females. These colonies break up after the breeding season is over. In the Bonin flying fox, colony formation is based on both the sex and age of individuals, as well as the season. In the winter breeding season, adult females will form colonies that include a few adult males (likely harems). Adult males who do not roost with females will form colonies with other adult and subadult males. Subadults will form mixed-sex "subadult groups" with each other. In the summer, however, individuals are solitary, with the exception of nursing females, who roost with their pups.
Diet and foraging
Flying foxes consume 25-35% of their body weight daily. They are generalists that will consume a variety of items to meet their nutritional needs. Food items include fruit, flowers, nectar, and leaves. They will sometimes deliberately consume insects such as cicadas as well. In Australia, eucalypt blossoms and pollen are preferred food sources, followed by Melaleuca and Banksia flowers  They feed on a wide variety of crops as well, causing conflicts with farmers. Crops eaten by flying foxes include sisal, cashew, pineapple, areca, breadfruit, jackfruit, neem, papaya, citrus, fig, mango, banana, avocado, guava, sugar cane, tamarind, grapes, and more.
In captivity, the recommended diet for flying foxes consists of two-thirds hard fruits like pears and apples and one-third soft fruits. Bananas and other high-fiber fruits should only be offered occasionally, as flying foxes are not adapted to high-fiber diets. Protein supplements are recommended for captive flying foxes; other supplements such as vitamin C, calcium, chondroitin sulfate, and glucosamine can be recommended periodically.
The majority of flying fox species are nocturnal and forage at night. A few island species and subspecies are diurnal, however, hypothesized as a response to a lack of predators. Diurnal taxa include P. melanotus natalis, the Mauritian flying fox, the Caroline flying fox, P. p. insularis, and the Seychelles fruit bat. Foraging resources are often far from roosts, with individuals traveling up to 40–60 km (25–37 mi) to reach them. Flying foxes can travel at 6 m/s (13 mph) for three hours or more, and can reach top speeds of 8.6 m/s (19 mph). Some colonial species will forage in groups, especially when resources are abundant. Less social species will forage alone. When they land on a tree with food, they will hang onto the branch with their clawed hind feet and use their clawed thumbs to pull branches bearing flowers or fruits towards them. As they forage on fruit, flying foxes will compress the fruit against the palate with the tongue to squeeze out and consume the juices. The rest of the fruit is then discarded in "ejecta pellets."
Role in ecosystems
Flying foxes have important roles as seed dispersers and pollinators. They help spread the seeds in the fruit they eat by discarding them in ejecta pellets or through their guano. In Madagascar, fig seeds have better germination success if they have passed through the gut of a flying fox, which is important because fig trees are a vital pioneer species in regenerating lost forest. Even though flying foxes can have a gut transit time as fast as 12 minutes, seeds can be retained in the gut for as long as 20 hours. As the flying foxes travel large distances, seeds can be deposited up to 20 km (12 mi) from the parent tree. They are particularly important in fragmented forests, as many other frugivores are terrestrial and often confined to forest fragments. Flying foxes have the capability to spread seeds beyond the forest fragments through flight.
Flying foxes pollinate a variety of plants, including the economically valuable durian. They forage on its nectar in such a way that the flowers (and eventual fruit production) are not usually harmed. Flying fox pollination has a positive effect on durian reproductive success, suggesting that both flying foxes and durian trees benefit from this relationship.
Of the 62 flying fox species evaluated by the IUCN as of 2018, 3 are considered critically endangered— the Aru flying fox, Livingstone's fruit bat, and the Vanikoro flying fox. Another 7 species are listed as endangered; 20 are listed as vulnerable, 6 as near threatened, 14 as least concern, and 8 as data deficient. A further 4 are listed as extinct—the dusky flying fox, the large Palau flying fox, the small Mauritian flying fox, and the Guam flying fox. Over half of the species are threatened today with extinction, and in particular in the Pacific, a number of species have died out as a result of hunting, deforestation, and predation by invasive species. Six flying fox species are believed to have gone extinct from 1864 to 2014—the Guam, large Palau, small Mauritian, dusky, large Samoan, and the small Samoan flying foxes.
All species of Pteropus are placed on Appendix II of CITES and ten on Appendix I, which restricts international trade. Individual species have different legal protections from hunting and domestic trade that reflect the environmental laws of the countries where they are found.
In some countries such as Bangladesh, Sri Lanka, and Thailand, flying foxes are absolutely protected from harm under the Wildlife Preservation and Security Act of 2012, Fauna and Flora Protection Ordinance of 1937, and Wildlife Protection and Reservation Act of 1992, respectively. However, in Thailand, flying fox poaching and illegal bushmeat trade still occurs outside of protected areas. The large flying fox and the small flying fox are particularly prone to poaching and roost disturbance.
In other countries, such as Australia, Japan, and the United States, some species of conservation concern are protected under national environmental legislation while others are not. In Australia, two flying foxes are listed under the Environment Protection and Biodiversity Conservation Act of 1999: the grey-headed and spectacled flying foxes are listed as "vulnerable." Farmers can apply for permits to kill flying foxes when they are causing crop damage.
Several flying fox species occur in Japan. The Bonin flying fox has been a Natural Monument of Japan since 1969, which means that it is illegal to capture or disturb them without appropriate permits. Two subspecies of the Ryukyu flying fox (P. d. dasymallus and P. d. daitoensis) are also listed as Natural Monuments. Flying foxes are not designated game species in Japan, and therefore cannot be legally hunted per the Wildlife Protection and Hunting Law. The Bonin flying fox and P. d. daitoensis are also listed as National Endangered Species, meaning that they cannot be killed or harmed; furthermore, the sale or transfer of live or dead individuals in whole or part is also prohibited without permits.
Despite not occurring in the United States, several species and subspecies are listed under its Endangered Species Act of 1973. Pteropus mariannus mariannus—a subspecies of the Mariana fruit bat—is listed as threatened while the Rodrigues flying fox and Guam flying fox are listed as endangered. Additionally, the U. S. Government has been petitioned to list the Aru flying fox and Bonin flying fox as threatened or endangered.
In countries such as India and Pakistan, flying foxes explicitly have no legal protection. In India, they are listed as "vermin" under the Wildlife Protection Act of 1972. Pakistan's only flying fox, the Indian flying fox, is listed under Schedule 4 of the Punjab Wildlife (Protection, Preservation, Conservation and Management) Act of 1974, meaning that it has no legal protections and can be hunted. In Mauritius, flying foxes were formerly protected but are now legally culled at a large scale. In 2015, the Mauritian government passed the Native Terrestrial Biodiversity And National Parks Act, which legalized culling of the Mauritian flying fox. In Mauritius, over 40,000 Mauritian flying foxes were culled in a two-year period, reducing its population by an estimated 45%. This decision was viewed with controversy, with researchers stating "Because they spread seeds and pollinate flowers, flying foxes are vital for regenerating lost forests."
Legal protection can vary within a country as well, such as in Malaysia. Under the 1990 Protection of Wild Life Amendment Order, flying foxes can be hunted with a permit; each permit is good for killing up to 50 flying foxes. Permits cost US$8 each. However, under the Protection of WildLife Act of 1972, flying foxes can be killed without permits if they are causing damage or if there is "reason to believe that it is about to cause serious damage" to crops. In 2012, the Malaysian state of Terengganu issued a moratorium on hunting flying foxes. In Sarawak, all bat species are listed as "Protected" and hunting them is not legal.
Factors causing decline
Flying foxes species are declining or going extinct as a result of several human impacts to their environments, in addition to natural phenomena. Their populations are especially vulnerable to threats because the litter size is usually only individual and females generally only have one litter per year. Even when nearly every female (90%) successfully produces and raises young, if a population's mortality rate exceeds 22% annually, then it will steadily decline.
Invasive species such as the brown tree snake can seriously impact populations; the brown tree snake consumes so many pups that it reduced the recruitment of the Guam population of the Mariana fruit bat to essentially zero.
Many flying fox species are threatened by overhunting. While they have long been a dietary component of indigenous people, expanding human population and more efficient weapons have resulted in population declines, local extinctions, and extinctions. Overhunting is believed to be the primary cause of extinction for the Small Mauritian flying fox and the Guam flying fox.
Flying foxes are also threatened with excessive culling due to conflict with farmers. They are shot, beaten to death, or poisoned to reduce their populations. Mortality also occurs via accidental entanglement into netting used to prevent the bats from eating fruit. Culling can dramatically reduce flying fox populations. In Mauritius, over 40,000 Mauritian flying foxes were culled in a two-year period, reducing its population by an estimated 45%. Flying foxes are also killed by electrocution. In one Australian orchard, it is estimated that over 21,000 bats were electrocuted to death in an 8-week period. Farmers construct electrified grids over their fruit trees to kill flying foxes before they can consume their crop. The grids are questionably effective at preventing crop loss, with one farmer who operated such a grid estimating that they still lost 100–120 tonnes (220,000–260,000 lb) of fruit to flying foxes in a year. Some electrocution deaths are also accidental, such as when bats fly into overhead power lines.
Climate change causes flying fox mortality and a source of concern for species persistence. Extreme heat waves in Australia have been responsible for the deaths of more than 30,000 flying foxes from 1994 to 2008. Females and young bats are most susceptible to extreme heat, which affects a population's ability to recover. Flying foxes are threatened by sea level rise associated with climate change, as several taxa are endemic to low-lying atolls.
Because many species are endemic to a single island, they are vulnerable to random events such as typhoons. A 1979 typhoon halved the remaining population of the Rodrigues flying fox. Typhoons result in indirect mortality as well: Because they defoliate the trees, flying foxes are more visible and easily hunted by humans. Food resources for the bats become scarce after major storms, and flying foxes 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.
Several species of endangered flying fox are bred in captivity to augment their population sizes. Critically endangered Livingstone's fruit bats were taken from the wild starting in 1995 to create a captive breeding program. All captive individuals remain the property of the Comorian government. 17 individuals were collected from the wild; with breeding, there are 71 in captivity as of 2017. Individuals are held at Jersey Zoo and Bristol Zoo. Though the program has been successful in increasing the population, caretakers of the captive population have had to deal with husbandry issues such as obesity and cardiomyopathy. Relative to their wild counterparts, captive bats have a higher percentage of body fat and a lower percentage of muscle mass. The problem is pronounced in dominant males, which are the most sedentary. Addressing these concerns involves increasing flight space so that the animals can exercise adequately. Keepers are also exploring ways of distributing food within enclosures to encourage exercise.
The endangered Rodrigues flying fox has been bred in captivity with great success. By 1979, only 70–100 individuals were left in the world. In 1976, 25 individuals were removed from the wild by Durrell Wildlife Conservation Trust to begin a breeding program. In 1988, the breeding program was called "undoubtedly the most important chiropteran breeding project now in operation." By 2016, there were 180 individuals in 16 zoos across the United States alone. Worldwide, 46 zoos participate in the Rodrigues flying fox breeding program as of 2017.
Relationship to people
Many flying foxes species are killed for bushmeat. The bushmeat harvest is often unsustainable, often resulting in severe population decline or local extinction. Flying foxes are killed and sold for bushmeat in several countries in Southeast Asia, South Asia, and Oceania, including Indonesia, Malaysia, Papua New Guinea, the Philippines, Bangladesh, India, Fiji, and Guam. Flying fox consumption is particularly common in countries with low food security and lack of environmental regulation. In some cultures in the region, however, eating flying fox meat is taboo. In Namoluk, locals are repulsed by the idea of eating flying foxes because the flying foxes urinate on themselves. In predominately Muslim regions such as much of Indonesia, flying foxes are rarely consumed because of halal dietary restrictions.
North Sulawesi has the greatest demand for flying fox bushmeat. Despite being in Muslim-majority Indonesia, North Sulawesi is predominately Christian; therefore, many locals do not follow halal guidelines prohibiting flying fox consumption. In Manado, most local people consume flying fox meat at least once a month. The frequency of flying fox consumption increases tenfold around holidays. Locals believe that "unique meat" from undomesticated animals should be served on special occasions to "enliven the atmosphere." Suggestions to make the flying fox bushmeat trade more sustainable include enforcing a quota system for harvesting, encouraging hunters to release female and juvenile individuals, and providing economic alternatives to those who make a living selling flying fox bushmeat.
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. BMAA may become biomagnified in humans who consume flying foxes; flying foxes are exposed to BMAA by eating cycad fruits.
Flying foxes are killed for use in traditional medicine. The Indian flying fox, for example, has many perceived medical uses. Some believe that its fat is a treatment for rheumatism. Tribes in the Attappadi region of India eat the cooked flesh of the Indian flying fox to treat asthma and chest pain. Healers of the Kanda Tribe of Bangladesh use hair from Indian flying foxes to create treatments for "fever with shivering."
Flying foxes are the natural reservoirs of several viruses, some of which can be transmitted to humans. Notably, flying foxes can transmit lyssaviruses, which cause rabies. In Australia the rabies virus is not naturally present; Australian bat lyssavirus is the only lyssavirus present. Australian bat lyssavirus was first identified in 1996; 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 is not a risk of exposure to Australian bat lyssavirus. Since 1994, there have been three records of people getting infected with it—all three were in Queensland and each case was fatal.
Flying foxes are also reservoirs of henipaviruses such as Hendra virus and Nipah virus. Hendra virus was first identified in 1994; it also rarely occurs humans. From 1994 to 2013, 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. There are no documented instances of direct transmission between flying foxes and humans. As of 2012, there is a vaccine available for horses to decrease the likelihood of infection and transmission.
Nipah virus was first identified in 1998 in Malaysia. Since 1998, there have been several Nipah outbreaks in Malaysia, Singapore, India, and Bangladesh, resulting in over 100 casualties. A 2018 outbreak in Kerala, India resulted in 19 humans infected, of which 17 died. The overall fatality rate is 40-75%. 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. A 2014 study of the Indian flying fox and Nipah virus found that while Nipah virus outbreaks are more likely in areas preferred by flying foxes, "the presence of bats in and of itself is not considered a risk factor for Nipah virus infection." Rather, the consumption of date palm sap is a significant route of transmission. 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. These viruses rarely affect humans and few cases have been reported. While other bat species have been suspected or implicated as the reservoir of diseases such as SARS and Ebola, flying foxes are not suspected as hosts for either causative virus.
Flying foxes are often considered pests due to the damage they cause to orchard crops. Flying foxes have been cited as particularly destructive to almonds, guavas, and mangoes in the Maldives; lychee in Mauritius; areca in India; and stone fruits in Australia. Orchard damages from other animals are often misattributed to flying foxes, though, and economic damage can be difficult to quantify or exaggerated. To prevent fruit damage, farmers may legally or illegally cull flying foxes. In the 1800s, the Australian government paid farmers bounties to kill flying foxes, though the practice has since been discontinued. Alternatives to culling include placing barriers between the bats and fruit trees, such as netting, or harvesting fruit in a timely manner to avoid attracting as many flying foxes. Netting is the most effective way to prevent crop loss, though some farmers find it cost prohibitive. It costs US$4,400–44,000 to net 1 ha (2.5 acres) of crops. Other methods of preventing fruit loss may also involve the use of scare guns, chemical deterrents, or night-time lights. Alternatively, planting Singapore cherrry trees and other decoy crops next to an orchard can be effective, as flying foxes are much more attracted to their fruits than many other orchard crops.
The location of flying fox camps can be a disturbance to humans. In Batemans Bay, Australia, locals report being so disturbed by flying fox vocalizations in the morning that they lose sleep. Flying foxes can fly into power lines and cause electricity outages. Their guano and body odor are also unpleasant to smell. The presence of flying fox colonies can cause nearby property values to decline.
Flying foxes are featured in indigenous cultures and traditions. A folklore Dreamtime story from the New South Wales North Coast in Australia features an impatient flying fox wanting the Great Spirit to teach him how to be a bird, only to be hung upside down on a branch. They were also featured in aboriginal cave art, as evinced by several surviving examples.
In Tonga, flying foxes are considered sacred. All flying foxes are the property of the king, meaning non-royal persons cannot harm them in any way. Tongan legend states that a colony of flying foxes at Kolovai are the descendants of a pair of flying foxes gifted to the King of Tonga by the Princess of Samoa.
In the Indian village of Puliangulam, a colony of Indian flying foxes roosts in a Banyan tree. Villagers believe that the flying foxes are under the protection of Muni, and do not harm the bats. A shrine to Muni is beneath the tree. If locals believe that they have offended Muni by failing to protect the bats, they will pray and perform puja after offering sweet rice, coconut, and bananas to those attending the ceremony.
Flying foxes are also featured in folk stories from Papua New Guinea. Stories with flying foxes include a legend about a cockatoo stealing feathers from the flying fox, resulting in it becoming nocturnal. Another story features a flying fox that can transform into a young man; the flying fox stole a woman away from her husband to take as his wife. Another legend states that a flying fox-man was responsible for introducing yams to their people.
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. In New Caledonia, ceremonial axes made of jade were decorated with braids of flying fox fur. Flying fox wings were depicted on the war shields of the Asmat people of Indonesia; they believed that the wings offered protection to their warriors.
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. On the island of Makira, which is part of the Solomon Islands, indigenous peoples still hunt flying foxes for their teeth as well as for bushmeat. The canine teeth are strung together on necklaces that are used as currency. Teeth of the insular flying fox are particularly prized, as they are usually large enough to drill holes in. The Makira flying fox is also hunted, though, despite its smaller teeth. Deterring local peoples 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." Emphasizing sustainable hunting of flying foxes to preserve cultural currency may be more effective than encouraging the abandonment of cultural currency. 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. Lavery stated, "It’s a positive, not a negative, that their teeth are so culturally valuable. The practice of hunting bats shouldn’t necessarily be stopped, it needs to be managed sustainably."
Flying foxes and other bat species in Southeast Asia are often killed and sold as "mummies". The mummified bodies or skeletons of these bats are often shipped to the United States where they are sold in souvenir or curiosity shops or online through vendors such as Etsy or eBay. From 2000 to 2013, over 100,000 dead bats were imported to the United States. Bat conservationist Merlin Tuttle wrote, "I've seen huge losses, mostly due to various kinds of over-harvesting, especially at cave entrances, either for food or for sale as mummies." Despite sometimes being advertised as "sustainable," the practice could lead to overharvesting and depletion of flying fox species, with Tuttle saying, "It is a virtual certainty that the bats you've seen advertised are not sustainably harvested."
|Phylogeny of Pteropus based on mitochondrial cytochrome b and rRNA12S genes.|
- "Pteropus Brisson 1762 (flying fox)". Fossilworks. Retrieved 2 June 2018.
- ICZN (1998). "Opinion 1894. Regnum Animale ..., Ed. 2 (M.J. Brisson, 1762): rejected for nomenclatural purposes, with the conservation of the mammalian generic names Philander (Marsupialia), Pteropus (Chiroptera), Glis, Cuniculus, and Hydrochoerus (Rodentia), Meles, Lutra and Hyaena (Carnivora), Tapirus (Perissodactyla), Tragulus and Giraffa (Artiodactyla)". Bulletin of Zoological Nomenclature. 55 (1): 64–71.
- Nowak, R. M., ed. (1999). Walker's Mammals of the World. 1 (6 ed.). pp. 264–271. ISBN 978-0-8018-5789-8.
- Simmons, N.B. (2005). "Genus Pteropus". In Wilson, D.E.; Reeder, D.M. Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. pp. 334–346. ISBN 978-0-8018-8221-0. OCLC 62265494.
- Brisson, A. D. (1762). Regnum Animale in Classes IX distributum sive synopsis methodica. Lugduni Batavorum :Apud Theodorum Haak,1762. pp. 153–155.
- Erxleben, J. C. P. (1777). Io. Christ. Polyc. Erxleben Phil. D. ... Systema regni animalis per classes, ordines, genera, species, varietates : cvm synonymia et historia animalivm : Classis I. Mammalia. Lipsiae :Impensis Weygandianis,1777. pp. 130–141.
- Jackson, S.; Jackson, S. M.; Groves, C. (2015). Taxonomy of Australian mammals. Csiro Publishing. p. 238. ISBN 978-1486300136.
- Kerr, R.; Gmelin, J. F. (1792). The Animal Kingdom Or Zoological System of the Celebrated Sir Charles Linnaeus: Class I. Mammalia (class II. the Birds): Containing a Complete Systematic Description, arrangement, and nomenclature, of all the known species and varieties of the mammalia, or animals which give suck to their young. Edinburgh :Printed for A. Strahan, and T. Cadell, London, and W. Creech, Edinburgh,1792. pp. 90–91.
- "Flying Fox". Merriam-Webster.com. Merriam-Webster, n.d. Retrieved 5 June 2018.
- McNab, Brian K; Armstrong, Monica I (2001). "Sexual Dimorphism and Scaling of Energetics in Flying Foxes of the Genus Pteropus". Journal of Mammalogy. 82 (3): 709. Bibcode:2007JMamm..88..275L. doi:10.1644/1545-1542(2001)082<0709:SDASOE>2.0.CO;2.
- Kunz, Thomas H; Jones, Deborah P (2000). "Pteropus vampyrus". Mammalian Species. 642: 1. doi:10.1644/1545-1410(2000)642<0001:PV>2.0.CO;2.
- Flannery, T. (1995). Mammals of New Guinea. Cornell University Press. pp. 376–377. ISBN 978-0801431494.
- Flannery, T. (1995). Mammals of the South-West Pacific & Moluccan Islands. Cornell University Press. pp. 245–303. ISBN 978-0801431500.
- Nelson, John E. Fauna of Australia (PDF) (Report). 1B. Australian Government Department of the Environment and Energy.
- Giannini, Norberto P; Wible, John R; Simmons, Nancy B (2006). "On the Cranial Osteology of Chiroptera. I. Pteropus (Megachiroptera: Pteropodidae)". Bulletin of the American Museum of Natural History. 295: 1–134. doi:10.1206/0003-0090(2006)295[0001:OTCOOC]2.0.CO;2.
- Divljan, A.; Parry-Jones, K.; Wardle, G. M. (2011). "One hundred and forty days in the life of a flying-fox tooth-fairy: estimating the age of pups using tooth eruption and replacement". In Law, B.; Eby, P.; Lunney, D.; Lumsden, L. The Biology and Conservation of Australasian bats. Royal Zoological Society of New South Wales. pp. 97–105. ISBN 978-0-9803272-4-3.
- Parish, S.; Richards, G.; Hall, L. (2012). A natural history of Australian Bats: working the night shift. Csiro Publishing. ISBN 978-0643103764.
- Tedman, RA; Hall, LS (1985). "The Morphology of the Gastrointestinal Tract and Food Transit Time in the Fruit Bats Pteropus Alecto and P. Poliocephalus (Megachiroptera)". Australian Journal of Zoology. 33 (5): 625. doi:10.1071/ZO9850625.
- Eisenberg, John F; Wilson, Don E (1978). "Relative Brain Size and Feeding Strategies in the Chiroptera". Evolution. 32 (4): 740. doi:10.1111/j.1558-5646.1978.tb04627.x. PMID 28567922.
- Deamer, D. (5 November 2009). "Calculating Animal Intelligence". Science2.0. ION Publications LLC. Retrieved 24 June 2018.
- Flick, B.; Spencer, H.; Van Der Zwan, R. (2011). "Are hand-raised flying-foxes (Pteropus conspicillatus) better learners than wild-raised ones in an operant conditioning situation?". In Law, B.; Eby, P.; Lunney, D.; Lumsden, L. The Biology and Conservation of Australasian bats. Royal Zoological Society of New South Wales. pp. 86–91. doi:10.7882/FS.2011.011. ISBN 978-0-9803272-4-3.
- Wood, W. F.; Walsh, A.; Seyjagat, J.; Weldon, P. J. (2005). "Volatile Compounds in Shoulder Gland Secretions of Male Flying Foxes, Genus Pteropus (Pteropodidae, Chiroptera)" (PDF). Z Naturforsch C. 60 (9–10).
- Wagner, J. (2008). "Glandular secretions of male Pteropus (Flying foxes): preliminary chemical comparisons among species". SIT Digital Collections. Independent Study Project (ISP) Collection.
- Shen, Yong-Yi; Lim, Burton K; Liu, He-Qun; Liu, Jie; Irwin, David M; Zhang, Ya-Ping (2012). "Multiple Episodes of Convergence in Genes of the Dim Light Vision Pathway in Bats". PLOS One. 7 (4): e34564. Bibcode:2012PLoSO...734564S. doi:10.1371/journal.pone.0034564. PMC 3324491. PMID 22509324.
- Melin, Amanda D; Danosi, Christina F; McCracken, Gary F; Dominy, Nathaniel J (2014). "Dichromatic vision in a fruit bat with diurnal proclivities: The Samoan flying fox (Pteropus samoensis)". Journal of Comparative Physiology A. 200 (12): 1015–22. doi:10.1007/s00359-014-0951-x. PMID 25319538.
- Max-Planck-Gesellschaft (16 June 2007). "Fruit Bats Are Not 'Blind As A Bat'". ScienceDaily. ScienceDaily. Retrieved 25 June 2018.
- Giannini, Norberto P; Simmons, Nancy B (2003). "A phylogeny of megachiropteran bats (Mammalia: Chiroptera: Pteropodidae) based on direct optimization analysis of one nuclear and four mitochondrial genes". Cladistics. 19 (6): 496. Bibcode:2002clad.book.....S. doi:10.1111/j.1096-0031.2003.tb00385.x.
- Eiting, Thomas P; Gunnell, Gregg F (2009). "Global Completeness of the Bat Fossil Record". Journal of Mammalian Evolution. 16 (3): 151. doi:10.1007/s10914-009-9118-x.
- Almeida, Francisca C; Giannini, Norberto P; Simmons, Nancy B; Helgen, Kristofer M (2014). "Each flying fox on its own branch: A phylogenetic tree for Pteropus and related genera (Chiroptera: Pteropodidae)". Molecular Phylogenetics and Evolution. 77: 83–95. doi:10.1016/j.ympev.2014.03.009. PMID 24662680.
- Teeling, E. C; Springer, M. S; Madsen, O; Bates, P; O'Brien, S. J; Murphy, W. J (2005). "A Molecular Phylogeny for Bats Illuminates Biogeography and the Fossil Record". Science. 307 (5709): 580–4. Bibcode:2005Sci...307..580T. doi:10.1126/science.1105113. PMID 15681385.
- Almeida, Francisca C; Giannini, Norberto P; Desalle, Rob; Simmons, Nancy B (2011). "Evolutionary relationships of the old world fruit bats (Chiroptera, Pteropodidae): Another star phylogeny?". BMC Evolutionary Biology. 11: 281. doi:10.1186/1471-2148-11-281. PMC 3199269. PMID 21961908.
- O'Brien, John; Mariani, Carol; Olson, Link; Russell, Amy L; Say, Ludovic; Yoder, Anne D; Hayden, Tom J (2009). "Multiple colonisations of the western Indian Ocean by Pteropus fruit bats (Megachiroptera: Pteropodidae): The furthest islands were colonised first". Molecular Phylogenetics and Evolution. 51 (2): 294. doi:10.1016/j.ympev.2009.02.010. PMID 19249376.
- Maruthupandian, Jayabalan; Marimuthu, Ganapathy (2013). "Cunnilingus Apparently Increases Duration of Copulation in the Indian Flying Fox, Pteropus giganteus". PLOS One. 8 (3): e59743. Bibcode:2013PLoSO...859743M. doi:10.1371/journal.pone.0059743. PMID 23544092.
- Sugita, Norimasa; Inaba, Makoto; Ueda, Keisuke (2009). "Roosting Pattern and Reproductive Cycle of Bonin Flying Foxes (Pteropus pselaphon)". Journal of Mammalogy. 90: 195–202. Bibcode:2007JMamm..88..275L. doi:10.1644/07-MAMM-A-368.1.
- Sugita, Norimasa (2016). "Homosexual Fellatio: Erect Penis Licking between Male Bonin Flying Foxes Pteropus pselaphon". PLOS One. 11 (11): e0166024. Bibcode:2016PLoSO..1166024S. doi:10.1371/journal.pone.0166024. PMC 5100941. PMID 27824953.
- Tan, Min; Jones, Gareth; Zhu, Guangjian; Ye, Jianping; Hong, Tiyu; Zhou, Shanyi; Zhang, Shuyi; Zhang, Libiao (2009). "Fellatio by Fruit Bats Prolongs Copulation Time". PLOS One. 4 (10): e7595. Bibcode:2009PLoSO...4.7595T. doi:10.1371/journal.pone.0007595. PMID 19862320.
- Pierson, E. D.; Rainey, W. E. (1992). "The biology of flying foxes of the genus Pteropus: a review" (PDF). Biological Report. 90 (23).
- Fox, Samantha; Spencer, Hugh; O'Brien, Gemma M (2008). "Analysis of twinning in flying-foxes (Megachiroptera) reveals superfoetation and multiple-paternity". Acta Chiropterologica. 10 (2): 271. doi:10.3161/150811008X414845.
- Orr, R. T. (2012). "Development: Prenatal and Postnatal". In Wimsatt, W. Biology of Bats. Elsevier. pp. 221–222. ISBN 978-0323151191.
- Kunz, T. H.; Kurta, A. (1987). "Size of bats at birth and maternal investment during pregnancy" (PDF). Symposia of the Zoological Society of London. 57.
- Safi, Kamran (2008). "Social Bats: The Males' Perspective". Journal of Mammalogy. 89 (6): 1342–1350. Bibcode:2007JMamm..88..275L. doi:10.1644/08-MAMM-S-058.1.
- Kunz, Thomas H; Hosken, David J (2009). "Male lactation: Why, why not and is it care?". Trends in Ecology & Evolution. 24 (2): 80–5. doi:10.1016/j.tree.2008.09.009. PMID 19100649.
- McIlwee, A. P; Martin, L (2002). "On the intrinsic capacity for increase of Australian flying-foxes (Pteropus spp., Megachiroptera)". Australian Zoologist. 32: 76–100. doi:10.7882/AZ.2002.008.
- Ma, Siming; Yim, Sun Hee; Lee, Sang-Goo; Kim, Eun Bae; Lee, Sang-Rae; Chang, Kyu-Tae; Buffenstein, Rochelle; Lewis, Kaitlyn N; Park, Thomas J; Miller, Richard A; Clish, Clary B; Gladyshev, Vadim N (2015). "Organization of the Mammalian Metabolome according to Organ Function, Lineage Specialization, and Longevity". Cell Metabolism. 22 (2): 332. doi:10.1016/j.cmet.2015.07.005. PMID 26244935.
- Vincenot, Christian E; Florens, F. B. Vincent; Kingston, Tigga (2017). "Can we protect island flying foxes?". Science. 355 (6332): 1368. Bibcode:2017Sci...355.1368V. doi:10.1126/science.aam7582. PMID 28360279.
- Tidemann, Christopher R; Nelson, John E (2011). "Life Expectancy, Causes of Death and Movements of the Grey-Headed Flying-Fox (Pteropus poliocephalus) Inferred from Banding". Acta Chiropterologica. 13 (2): 419. doi:10.3161/150811011X624901.
- Reeder, Deeann M; Kosteczko, Nicole S; Kunz, Thomas H; Widmaier, Eric P (2006). "The hormonal and behavioral response to group formation, seasonal changes, and restraint stress in the highly social Malayan Flying Fox (Pteropus vampyrus) and the less social Little Golden-mantled Flying Fox (Pteropus pumilus) (Chiroptera: Pteropodidae)". Hormones and Behavior. 49 (4): 484. doi:10.1016/j.yhbeh.2005.11.001. PMID 16380123.
- Epstein, Jonathan H; Olival, Kevin J; Pulliam, Juliet R.C; Smith, Craig; Westrum, Justin; Hughes, Tom; Dobson, Andrew P; Zubaid, Akbar; Rahman, Sohayati Abdul; Basir, Misliah Mohamad; Field, Hume E; Daszak, Peter (2009). "Pteropus vampyrus, a hunted migratory species with a multinational home-range and a need for regional management". Journal of Applied Ecology. 46 (5): 991. doi:10.1111/j.1365-2664.2009.01699.x.
- Nakamoto, Atsushi; Kinjo, Kazumitsu; Izawa, Masako (2012). "Ranging Patterns and Habitat Use of a Solitary Flying Fox (Pteropus dasymallus) on Okinawa-jima Island, Japan". Acta Chiropterologica. 14 (2): 387. doi:10.3161/150811012X661701.
- Tsang, Susan M; Wiantoro, Sigit; Simmons, Nancy B (2015). "New Records of Flying Foxes (Chiroptera:Pteropussp.) from Seram, Indonesia, with Notes on Ecology and Conservation Status". American Museum Novitates. 3842 (3842): 1. doi:10.1206/3842.1. hdl:2246/6622.
- Klose, S. M; Welbergen, J. A; Kalko, E. K. V (2009). "Testosterone is associated with harem maintenance ability in free-ranging grey-headed flying-foxes, Pteropus poliocephalus". Biology Letters. 5 (6): 758. doi:10.1098/rsbl.2009.0563. PMID 19689980.
- Department of Environment and Natural Resources. Guidelines for the Captive Management of Flying Foxes in South Australia (Report). South Australian Government. pp. 18–19.
- Marshall, Adrian G (1985). "Old World phytophagous bats (Megachiroptera) and their food plants: A survey". Zoological Journal of the Linnean Society. 83 (4): 351–369. doi:10.1111/j.1096-3642.1985.tb01181.x.
- Aziz, Sheema Abdul; Clements, Gopalasamy Reuben; Peng, Lee Yin; Campos-Arceiz, Ahimsa; McConkey, Kim R; Forget, Pierre-Michel; Gan, Han Ming (2017). "Elucidating the diet of the island flying fox (Pteropus hypomelanus) in Peninsular Malaysia through Illumina Next-Generation Sequencing". PeerJ. 5: e3176. doi:10.7717/peerj.3176. PMC 5391789. PMID 28413729.
- Clulow, Simon; Blundell, Adam T (2011). "Deliberate Insectivory by the Fruit Bat Pteropus poliocephalusby Aerial Hunting". Acta Chiropterologica. 13: 201–205. doi:10.3161/150811011X578750.
- NWC - NSW Wildlife Council (June 2010). Flying-foxes (PDF) (Report).
- Aziz, S.A.; Olival, K.J.; Bumrungsri, S.; Richards, G.C.; Racey, P.A. (2016). "The Conflict Between Pteropodid Bats and Fruit Growers: Species, Legislation and Mitigation". In Voigt, C.; Kingston, T. Bats in the Anthropocene: Conservation of Bats in a Changing World. Springer. ISBN 978-3-319-25220-9.
- Carpenter, R. E. (1985). "Flight physiology of flying foxes, Pteropus poliocephalus" (PDF). Journal of Experimental Biology. 114 (1): 619–647.
- Banack, Sandra Anne (1998). "Diet Selection and Resource Use by Flying Foxes (Genus: Pteropus)". Ecology. 79 (6): 1949–1967. doi:10.1890/0012-9658(1998)079[1949:DSARUB]2.0.CO;2.
- Oleksy, Ryszard; Giuggioli, Luca; McKetterick, Thomas J; Racey, Paul A; Jones, Gareth (2017). "Flying foxes create extensive seed shadows and enhance germination success of pioneer plant species in deforested Madagascan landscapes". PLOS One. 12 (9): e0184023. Bibcode:2017PLoSO..1284023O. doi:10.1371/journal.pone.0184023. PMID 28877248.
- Aziz, Sheema A; Clements, Gopalasamy R; McConkey, Kim R; Sritongchuay, Tuanjit; Pathil, Saifful; Abu Yazid, Muhammad Nur Hafizi; Campos-Arceiz, Ahimsa; Forget, Pierre-Michel; Bumrungsri, Sara (2017). "Pollination by the locally endangered island flying fox (Pteropus hypomelanus) enhances fruit production of the economically important durian (Durio zibethinus)". Ecology and Evolution. 7 (21): 8670. doi:10.1002/ece3.3213. PMID 29152168.
- "The IUCN Red List of Threatened Species Version 2017-3". Retrieved 3 June 2018.
- "Appendices I, II and III". CITES. 4 October 2017. Retrieved 14 April 2018.
- Voigt, C. C.; Kingston, T. (2016). Bats in the Anthropocene: conservation of bats in a changing world. Springer. doi:10.1007/978-3-319-25220-9. ISBN 978-3-319-25218-6.
- "Flying-foxes". environment.gov.au. Commonwealth of Australi. Retrieved 7 June 2018.
- Williams, Kylie (21 October 2012). "Culling flying-foxes is ineffective, so why suggest slaughter?". The Conversation. The Conversation US. Retrieved 7 June 2018.
- Vincenot, C. (2017). "Pteropus pselaphon". The IUCN Red List of Threatened Species. 2017: e.T18752A22085351. doi:10.2305/IUCN.UK.2017-2.RLTS.T18752A22085351.en.
- Vincenot, C. (2017). "Pteropus dasymallus". The IUCN Red List of Threatened Species. 2017: e.T18722A22080614. doi:10.2305/IUCN.UK.2017-2.RLTS.T18722A22080614.en.
- "Wildlife Protection System and Hunting Law". Ministry of the Environment. Ministry of the Environment Government of Japan. Retrieved 21 June 2018.
- Saitoh, Takashi; Kaji, Koichi; Izawa, Masako; Yamada, Fumio (2015). "Conservation and management of terrestrial mammals in Japan: Its system and practices". Therya. 6: 139–153. doi:10.12933/therya-15-239.
- "Listed Animals". U. S. Fish & Wildlife Service. U. S. Federal Government. Retrieved 7 June 2018.
- "Review of Petition To List 15 Bat Species as Endangered or Threatened Under the Act" (PDF). Federal Register. 79 (14). 22 January 2014.
- Mistry, Shahroukh. "Protecting the Bats of India: Signs of progress amid daunting challenge". Batcon.org. Bat Conservation International. Retrieved 7 June 2018.
- Racey, P. A.; Medellín, R. A. (11 November 2015). "Opinion: Killing Thousands of Flying Foxes Only Hurts the Environment". National Geographic. National Geographic Partners, LLC. Retrieved 7 June 2018.
- Esselstyn, Jacob A; Amar, Arjun; Janeke, Dustin (2006). "Impact of Posttyphoon Hunting on Mariana Fruit Bats (Pteropus mariannus)". Pacific Science. 60 (4): 531. doi:10.1353/psc.2006.0027.
- Vincenot, Christian Ernest; Koyama, Lina; Russo, Danilo (2015). "Near threatened? First report of unsuspected human-driven decline factors in the Ryukyu flying fox (Pteropus dasymallus) in Japan". Mammalian Biology - Zeitschrift für Säugetierkunde. 80 (4): 273. doi:10.1016/j.mambio.2015.03.003.
- Martin, L (2011). "Is the fruit you eat flying-fox friendly? The effects of orchard electrocution grids on Australian flying-foxes (Pteropus spp., Megachiroptera)". The Biology and Conservation of Australasian Bats. pp. 380–390. doi:10.7882/FS.2011.039. ISBN 978-0-9803272-4-3.
- Chlopicki, K. (28 October 2016). "Electric wires threaten flying foxes and their new babies". The Daily Telegraph. News Pty Ltd. Retrieved 26 June 2018.
- Welbergen, J. A; Klose, S. M; Markus, N; Eby, P (2008). "Climate change and the effects of temperature extremes on Australian flying-foxes" (PDF). Proceedings of the Royal Society B: Biological Sciences. 275 (1633): 419–25. doi:10.1098/rspb.2007.1385. PMC 2596826. PMID 18048286.
- Buden, Don; Helgen, Kristofer M; Wiles, Gary (2013). "Taxonomy, distribution, and natural history of flying foxes (Chiroptera, Pteropodidae) in the Mortlock Islands and Chuuk State, Caroline Islands". ZooKeys. 345 (345): 97–135. doi:10.3897/zookeys.345.5840. PMC 3817444. PMID 24194666.
- Buettner, Petra G; Westcott, David A; MacLean, Jennefer; Brown, Lawrence; McKeown, Adam; Johnson, Ashleigh; Wilson, Karen; Blair, David; Luly, Jonathan; Skerratt, Lee; Muller, Reinhold; Speare, Richard (2013). "Tick Paralysis in Spectacled Flying-Foxes (Pteropus conspicillatus) in North Queensland, Australia: Impact of a Ground-Dwelling Ectoparasite Finding an Arboreal Host". PLOS One. 8 (9): e73078. Bibcode:2013PLoSO...873078B. doi:10.1371/journal.pone.0073078. PMID 24066028.
- Smith, Stephanie J; Leslie, David M (2006). "Pteropus livingstonii". Mammalian Species. 792: 1–5. doi:10.1644/792.1.
- Richdon, Sarah; Rodriguez, Angelica M. (2017). "Building family trees in the Critically Endangered Livingstone's fruit bat". experiment. Experiment. Retrieved 20 June 2018.
- Thorncroft, K.; Wormell, D.; Price, E. (2014). "Effects of food distribution method on territoriality, activity and access to food in captive Livingstone's fruit bats (Pteropus livingstonii)" (PDF). Solitaire (25): 22–26.
- Wormell, Dominic; Ramsay, Scott; Masefield, Will; Houston, David; Price, Eluned (2018). "Assessing muscle condition in captive Livingstone's fruit bats". Solitaire (27).
- "Rodrigues Fruit Bat: Local and International Partners Work Together to Save the 'Golden Bat'". batcon.org. Bat Conservation International. 2016. Retrieved 20 June 2018.
- Carroll, J. Bryan; Mace, Georgina M (2007). "Population management of the Rodrigues fruit bat". International Zoo Yearbook. 27: 70–78. doi:10.1111/j.1748-1090.1988.tb03198.x.
- Tatayah, V.; Jhangeer-Khan, R.; Bégué, J.A.; Jones, C.A. (2017). "Pteropus rodricensis". The IUCN Red List of Threatened Species. 2017: e.T18755A22087057. doi:10.2305/IUCN.UK.2017-2.RLTS.T18755A22087057.en.
- Mickleburgh, Simon; Waylen, Kerry; Racey, Paul (2009). "Bats as bushmeat: A global review". Oryx. 43 (2): 217. doi:10.1017/S0030605308000938.
- Sheherazade; Tsang, Susan M (2015). "Quantifying the bat bushmeat trade in North Sulawesi, Indonesia, with suggestions for conservation action". Global Ecology and Conservation. 3: 324–330. doi:10.1016/j.gecco.2015.01.003.
- Banack, Sandra Anne; Murch, Susan J; Cox, Paul Alan (2006). "Neurotoxic flying foxes as dietary items for the Chamorro people, Marianas Islands". Journal of Ethnopharmacology. 106 (1): 97–104. doi:10.1016/j.jep.2005.12.032. PMID 16457975.
- Cox, P., Davis, D., Mash, D., Metcalf J.S., Banack, S. A. (2016). "Dietary exposure to an environmental toxin triggers neurofibrillary tangles and amyloid deposits in the brain". Proceedings of the Royal Society B. 283 (3): 1–10. doi:10.1098/rspb.2015.2397. PMC 4795023. PMID 26791617.CS1 maint: Multiple names: authors list (link)
- Holtcamp, W. (2012). "The emerging science of BMAA: do cyanobacteria contribute to neurodegenerative disease?". Environmental Health Perspectives. 120 (1823): a110–a116. doi:10.1289/ehp.120-a110. PMC 3295368. PMID 22382274.
- Padmanabhan, P.; Sujana, K. A. (2008). "Animal products in traditional medicine from Attappady hills of Western Ghats". Indian Journal of Traditional Knowledge. 7 (2): 326–329. hdl:123456789/1595.
- Rahmatullah, M.; Ayman, U.; Akter, F.; Sarker, M.; Sifa, R.; Sarker, B.; Chowdhury, S. A. (2013). "Medicinal formulations of a Kanda tribal healer–a tribe on the verge of disappearance in Bangladesh". African Journal of Traditional, Complementary and Alternative Medicines. 10 (2): 213–222.
- "Rabies and Australian bat lyssavirus infection fact sheet". health.nsw.gov.au. State of New South Wales NSW Ministry of Health 2015. 30 November 2015. Retrieved 14 June 2018.
- "Hendra Virus Disease(HeV)" (PDF). cdc.gov. U.S. Department of Health & Human Services. Retrieved 14 June 2018.
- Sánchez, Cecilia A; Baker, Michelle L (2016). "Disease Risk Perception and Safety Practices: A Survey of Australian Flying Fox Rehabilitators". PLOS Neglected Tropical Diseases. 10 (2): e0004411. doi:10.1371/journal.pntd.0004411. PMID 26829399.
- "The Hendra vaccine". ava.com. The Australian Veterinary Association Ltd (AVA). 2018. Retrieved 14 June 2018.
- Gulland, Anne (12 June 2018). "Nipah virus 'under control' in India – but Britain and the world must be alert for signs of infected travellers". The Telegraph. Telegraph Media Group Limited 2018. Retrieved 14 June 2018.
- "Nipah virus". World Health Organization. WHO. 30 May 2018. Retrieved 14 June 2018.
- Hahn, M. B.; Epstein, J. H.; Gurley, E. S.; Islam, M. S.; Luby, S. P.; Daszak, P.; Patz, J. A. (2014). "Roosting behaviour and habitat selection of Pteropus giganteus reveal potential links to Nipah virus epidemiology". Journal of Applied Ecology. 51 (2): 376–387. doi:10.1111/1365-2664.12212. PMC 4000083. PMID 24778457.
- Bowden, Timothy R; Westenberg, Marcel; Wang, Lin-Fa; Eaton, Bryan T; Boyle, David B (2001). "Molecular Characterization of Menangle Virus, a Novel Paramyxovirus which Infects Pigs, Fruit Bats, and Humans". Virology. 283 (2): 358. doi:10.1006/viro.2001.0893. PMID 11336561.
- Yamanaka, Atsushi; Iwakiri, Akira; Yoshikawa, Tomoki; Sakai, Kouji; Singh, Harpal; Himeji, Daisuke; Kikuchi, Ikuo; Ueda, Akira; Yamamoto, Seigo; Miura, Miho; Shioyama, Yoko; Kawano, Kimiko; Nagaishi, Tokiko; Saito, Minako; Minomo, Masumi; Iwamoto, Naoyasu; Hidaka, Yoshio; Sohma, Hirotoshi; Kobayashi, Takeshi; Kanai, Yuta; Kawagishi, Takehiro; Nagata, Noriyo; Fukushi, Shuetsu; Mizutani, Tetsuya; Tani, Hideki; Taniguchi, Satoshi; Fukuma, Aiko; Shimojima, Masayuki; Kurane, Ichiro; et al. (2014). "Imported Case of Acute Respiratory Tract Infection Associated with a Member of Species Nelson Bay Orthoreovirus". PLOS One. 9 (3): e92777. Bibcode:2014PLoSO...992777Y. doi:10.1371/journal.pone.0092777. PMC 3965453. PMID 24667794.
- Smith, Ina; Wang, Lin-Fa (2013). "Bats and their virome: An important source of emerging viruses capable of infecting humans". Current Opinion in Virology. 3 (1): 84–91. doi:10.1016/j.coviro.2012.11.006. PMID 23265969.
- Hall, L. S.; Richards, G. (2000). Flying foxes: fruit and blossom bats of Australia. UNSW Press. pp. 52–53. ISBN 978-0868405612.
- Dolbeer, R. R.; Fiedler, L. R.; Rasheed, H. (1988). "Management of fruit bat and rat populations in the Maldive Islands, Indian Ocean". Proceedings of the Thirteenth Vertebrate Pest Conference. 1988.
- Singaravelan, N.; Marimuthu, G.; Racey, P. A. (2009). "Do fruit bats deserve to be listed as vermin in the Indian Wildlife (Protection) & Amended Acts? A critical review". Oryx. 43 (4): 608–613. doi:10.1017/S0030605309990391.
- Associated Press (10 June 2018). "'It smells so bad you can taste it': bats plague Australian tourist town". The Guardian. Guardian News and Media Limited. Retrieved 20 June 2018.
- McKenzie, Jessica (1 May 2016). "Residents rapt after nuisance bats evicted". Sunshine Coast Daily. Sunshine Coast Newspaper Company Pty Ltd. Retrieved 22 June 2018.
- Campbell-McLeod, P. (2013). "Nallawilli-Sit Down (and Listen): The Dreamtime Stories--An Oral Tradition". In MacDonald, M. R. Traditional Storytelling Today: An International Sourcebook. pp. 158–159. ISBN 978-1135917142.
- Grant, G. S. (1996). "Kingdom of Tonga: Safe Haven for Flying Foxes". batcon.org. Bat Conservation International. Retrieved 21 June 2018.
- Marimuthu, G. (1988). "The Sacred Flying Fox of India". batcon.org. Bat Conservation International. Retrieved 21 June 2018.
- Slone, T. H., ed. (2001). One Thousand One Papua New Guinean Nights: Tales from 1972–1985. 1. Masalai Press. ISBN 978-0971412712.
- British Museum. Dept. of British and Mediaeval Antiquities and Ethnography; Joyce, T. A.; Dalton, O. M. Handbook to the Ethnographical Collections. Printed by order of the Trustees. p. 125.
- Machray, Robert (1899). "Strange Kinds of Money". The Harmsworth Monthly Pictorial Magazine. 1: 639–641.
- Werness; H. B. (2003). Continuum Encyclopedia of Native Art: Worldview, Symbolism, and Culture in Africa, Oceania, and North America. A&C Black. p. 31. ISBN 978-0826414656.
- Choi, Charles (16 October 2017). "In Makira, Flying Fox Teeth Are Currency…And That Could Save the Species". Discover. Kalmbach Media. Retrieved 2 June 2018.
- Lavery, Tyrone H; Fasi, John (2017). "Buying through your teeth: Traditional currency and conservation of flying foxes Pteropus spp. In Solomon Islands". Oryx: 1–8. doi:10.1017/S0030605317001004.
- Hugo, Kristin (10 October 2017). "Thousands of Bats Slaughtered Annually in Asia End Up on eBay and Etsy for Artsy Americans". Newsweek. Newsweek LLC. Retrieved 2 June 2018.