Flying Foxes
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''Pteropus'' (suborder
Yinpterochiroptera The Yinpterochiroptera (or Pteropodiformes) is a suborder of the Chiroptera, which includes taxa formerly known as megabats and five of the microbat families: Rhinopomatidae, Rhinolophidae, Hipposideridae, Craseonycteridae, and Megadermatidae ...
) is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of
megabat Megabats constitute the 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. They are the only member of the su ...
s which are among the largest bats in the world. They are commonly known as fruit bats or flying foxes, among other colloquial names. They live in
South Asia South Asia is the southern subregion of Asia, which is defined in both geographical and ethno-cultural terms. The region consists of the countries of Afghanistan, Bangladesh, Bhutan, India, Maldives, Nepal, Pakistan, and Sri Lanka.;;;;;;;; ...
,
Southeast Asia Southeast Asia, also spelled South East Asia and South-East Asia, and also known as Southeastern Asia, South-eastern Asia or SEA, is the geographical United Nations geoscheme for Asia#South-eastern Asia, south-eastern region of Asia, consistin ...
,
Australia Australia, officially the Commonwealth of Australia, is a Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous List of islands of Australia, sma ...
,
East Africa East Africa, Eastern Africa, or East of Africa, is the eastern subregion of the African continent. In the United Nations Statistics Division scheme of geographic regions, 10-11-(16*) territories make up Eastern Africa: Due to the historical ...
, and some oceanic islands in the Indian and Pacific Oceans. There are at least 60
extant Extant is the opposite of the word extinct. It may refer to: * Extant hereditary titles * Extant literature, surviving literature, such as ''Beowulf'', the oldest extant manuscript written in English * Extant taxon, a taxon which is not extinct, ...
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 Nocturnality is an animal behavior characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnal meaning the opposite. Nocturnal creatures generally have highly developed sens ...
. 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 Overexploitation, also called overharvesting, refers to harvesting a renewable resource to the point of diminishing returns. Continued overexploitation can lead to the destruction of the resource, as it will be unable to replenish. The term app ...
,
culling In biology, culling is the process of segregating organisms from a group according to desired or undesired characteristics. In animal breeding, it is the process of removing or segregating animals from a breeding stock based on a specific tr ...
, and natural disasters. Six flying fox species have been made
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
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 In Spermatophyte plants, seed dispersal is the movement, spread or transport of seeds away from the parent plant. Plants have limited mobility and rely upon a variety of dispersal vectors to transport their seeds, including both abiotic vectors, ...
. They benefit ecosystems and human interests by
pollinating Pollination is the transfer of pollen from an anther of a plant to the stigma of a plant, later enabling fertilisation and the production of seeds, most often by an animal or by wind. Pollinating agents can be animals such as insects, birds, a ...
plants. Like other bats, flying foxes are relevant to humans as a source of disease, as they are the
reservoirs A reservoir (; from French ''réservoir'' ) is an enlarged lake behind a dam. Such a dam may be either artificial, built to store fresh water or it may be a natural formation. Reservoirs can be created in a number of ways, including control ...
of rare but fatal disease agents including
Australian bat lyssavirus ''Australian bat lyssavirus'' (''ABLV''), originally named ''Pteropid lyssavirus'' (''PLV''), is a zoonotic virus closely related to the rabies virus. It was first identified in a 5-month-old juvenile black flying fox (''Pteropus alecto'') col ...
, which causes
rabies Rabies is a viral disease that causes encephalitis in humans and other mammals. Early symptoms can include fever and tingling at the site of exposure. These symptoms are followed by one or more of the following symptoms: nausea, vomiting, vi ...
, and
Hendra virus Hendra virus (HeV), scientific name ''Hendra henipavirus'', is a bat-borne virus that is associated with a highly fatal infection in horses and humans. Numerous disease outbreaks in Australia among horses have been caused by Hendra virus. The Hend ...
; seven known human deaths have resulted from these two diseases.
Nipah virus Nipah virus, scientific name ''Nipah henipavirus'', is a bat-borne virus that causes Nipah virus infection in humans and other animals, a disease with a high mortality rate. Numerous disease outbreaks caused by Nipah virus have occurred in Sout ...
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.


Taxonomy and etymology

The genus name ''Pteropus'' was coined by French zoologist
Mathurin Jacques Brisson Mathurin Jacques Brisson (; 30 April 1723 – 23 June 1806) was a French zoologist and natural philosopher. Brisson was born at Fontenay-le-Comte. The earlier part of his life was spent in the pursuit of natural history; his published works ...
in 1762. Prior to 1998, genus authority was sometimes given to German naturalist
Johann Christian Polycarp Erxleben Johann Christian Polycarp Erxleben was a German natural history, naturalist from Quedlinburg. Erxleben was professor of physics and veterinary medicine at the University of Göttingen. He wrote ''Anfangsgründe der Naturlehre'' (1772) and ''Sy ...
. Although the Brisson publication (1762) predated the Erxleben publication (1777), thus giving him preference under the
Principle of Priority 270px, '' valid name. Priority is a fundamental principle of modern botanical nomenclature and zoological nomenclature. Essentially, it is the principle of recognising the first valid application of a name to a plant or animal. There are two asp ...
, some authors gave preference to Erxleben as genus authority because Brisson's publication did not consistently use
binomial nomenclature In taxonomy, binomial nomenclature ("two-term naming system"), also called nomenclature ("two-name naming system") or binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, bot ...
. In 1998, the
International Commission on Zoological Nomenclature The International Commission on Zoological Nomenclature (ICZN) is an organization dedicated to "achieving stability and sense in the scientific naming of animals". Founded in 1895, it currently comprises 26 commissioners from 20 countries. Orga ...
(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 In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen ...
of the genus is the
Mauritian flying fox The Mauritian flying fox (''Pteropus niger''), also known as Greater Mascarene flying fox or Mauritius fruit bat is a large megabat species endemic to Mauritius and La Réunion. Description The Mauritian flying fox can reach a wingspan of 80&nb ...
, ''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 power A plenary power or plenary authority is a complete and absolute power to take action on a particular issue, with no limitations. It is derived from the Latin term ''plenus'' ("full"). United States In United States constitutional law, plenary p ...
s over biological nomenclature. "''
Pteropus ''Pteropus'' (suborder Yinpterochiroptera) is a genus of megabats which are among the largest bats in the world. They are commonly known as fruit bats or flying foxes, among other colloquial names. They live in South Asia, Southeast Asia, Austra ...
''" comes from
Ancient Greek Ancient Greek includes the forms of the Greek language used in ancient Greece and the ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Dark Ages (), the Archaic peri ...
"''pterón''" meaning "wing" and "''poús''" meaning "foot." The phrase "flying fox" has been used to refer to ''Pteropus'' bats since at least 1759.


Species


Description


External characteristics

Flying fox species vary in body weight, ranging from . Across all species, males are usually larger than females. The
large flying fox The large flying fox (''Pteropus vampyrus'', formerly ''Pteropus giganteus''), also known as the greater flying fox, Malayan flying fox, Malaysian flying fox, large fruit bat, kalang, or kalong, is a southeast Asian species of megabat in the fam ...
has the longest forearm length and reported wingspan of any bat species, but some bat species exceed it in weight. Its wingspan is up to , and it can weigh up to . The
Indian Indian or Indians may refer to: Peoples South Asia * Indian people, people of Indian nationality, or people who have an Indian ancestor ** Non-resident Indian, a citizen of India who has temporarily emigrated to another country * South Asia ...
and
great Great may refer to: Descriptions or measurements * Great, a relative measurement in physical space, see Size * Greatness, being divine, majestic, superior, majestic, or transcendent People * List of people known as "the Great" *Artel Great (born ...
flying foxes are heavier, at , respectively. Outside this genus, the
giant golden-crowned flying fox The giant golden-crowned flying fox (''Acerodon jubatus''), also known as the golden-capped fruit bat, is a species of megabat endemic to the Philippines. Since its description in 1831, three subspecies of the giant golden-crowned flying fox hav ...
(genus ''
Acerodon ''Acerodon'' (meaning: Sharp-tooth) is a genus of bats in the family Pteropodidae containing five species, all native to forests in Southeast Asia, and all considered threatened. They are closely related to ''Pteropus''. Species Genus ''Acerod ...
'') is the only bat with similar dimensions. Most flying fox species are considerably smaller and generally weigh less than . Smaller species such as the masked, Temminck's,
Guam Guam (; ch, Guåhan ) is an organized, unincorporated territory of the United States in the Micronesia subregion of the western Pacific Ocean. It is the westernmost point and territory of the United States (reckoned from the geographic cent ...
, and
dwarf Dwarf or dwarves may refer to: Common uses *Dwarf (folklore), a being from Germanic mythology and folklore * Dwarf, a person or animal with dwarfism Arts, entertainment, and media Fictional entities * Dwarf (''Dungeons & Dragons''), a humanoid ...
flying foxes all weigh less than . 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 A mammary gland is an exocrine gland in humans and other mammals that produces milk to feed young offspring. Mammals get their name from the Latin word ''mamma'', "breast". The mammary glands are arranged in organs such as the breasts in primat ...
located in the chest region. Their ears are long and pointed at the tip and lack
tragi The tragus is a small pointed eminence of the external ear, situated in front of the concha, and projecting backward over the meatus. It also is the name of hair growing at the entrance of the ear. Its name comes the Ancient Greek (), meaning 'g ...
. The outer margin of each ear forming an unbroken ring. The toes have sharp, curved claws. While
microbat Microbats constitute the suborder Microchiroptera within the order Chiroptera (bats). Bats have long been differentiated into Megachiroptera (megabats) and Microchiroptera, based on their size, the use of echolocation by the Microchiroptera a ...
s only have a claw on each thumb of their forelimbs, flying foxes additionally have a claw on each index finger.


Skull and dentition

The skulls of ''Pteropus'' species are composed of 24 bones, the
snout A snout is the protruding portion of an animal's face, consisting of its nose, mouth, and jaw. In many animals, the structure is called a muzzle, rostrum, or proboscis. The wet furless surface around the nostrils of the nose of many mammals is c ...
is made of 7, the
cranium The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, the ...
of 16 and the
mandible In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...
is a single bone. It has a large and bulbous
braincase In human anatomy, the neurocranium, also known as the braincase, brainpan, or brain-pan is the upper and back part of the skull, which forms a protective case around the brain. In the human skull, the neurocranium includes the calvaria or skul ...
. Like all mammals, flying foxes have three
middle ear ossicles The ossicles (also called auditory ossicles) are three bones in either middle ear that are among the smallest bones in the human body. They serve to transmit sounds from the air to the fluid-filled labyrinth (inner ear), labyrinth (cochlea). The ...
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 The maxilla (plural: ''maxillae'' ) in vertebrates is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The t ...
elongates, gaining bone between the
zygomatic process The zygomatic processes are three processes (protrusions) from other bones of the skull which each articulate with the zygomatic bone. The three processes are: * Zygomatic process of frontal bone from the frontal bone * Zygomatic process of ma ...
es and the
canine teeth In mammalian oral anatomy, the canine teeth, also called cuspids, dog teeth, or (in the context of the upper jaw) fangs, eye teeth, vampire teeth, or vampire fangs, are the relatively long, pointed teeth. They can appear more flattened howeve ...
. Based on the
grey-headed flying fox The grey-headed flying fox (''Pteropus poliocephalus'') is a megabat native to Australia. The species shares mainland Australia with three other members of the genus ''Pteropus'': the little red ''Pteropus scapulatus, P. scapulatus'', spectacle ...
's development, pups are born with some
milk teeth Deciduous teeth or primary teeth, also informally known as baby teeth, milk teeth, or temporary teeth,Illustrated Dental Embryology, Histology, and Anatomy, Bath-Balogh and Fehrenbach, Elsevier, 2011, page 255 are the first set of teeth in the ...
already erupted: canines and
incisor Incisors (from Latin ''incidere'', "to cut") are the front teeth present in most mammals. They are located in the premaxilla above and on the mandible below. Humans have a total of eight (two on each side, top and bottom). Opossums have 18, whe ...
s. By 9 days old, all the milk teeth have emerged, with a
dental formula Dentition pertains to the development of teeth and their arrangement in the mouth. In particular, it is the characteristic arrangement, kind, and number of teeth in a given species at a given age. That is, the number, type, and morpho-physiolo ...
of 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 Permanent teeth or adult teeth are the second set of teeth formed in diphyodont mammals. In humans and old world simians, there are thirty-two permanent teeth, consisting of six maxillary and six mandibular molars, four maxillary and four mandibul ...
. The canines are usually replaced first, followed by the
premolar The premolars, also called premolar teeth, or bicuspids, are transitional teeth located between the canine and molar teeth. In humans, there are two premolars per quadrant in the permanent set of teeth, making eight premolars total in the mouth ...
s, incisors, and then
molars The molars or molar teeth are large, flat teeth at the back of the mouth. They are more developed in mammals. They are used primarily to grind food during chewing. The name ''molar'' derives from Latin, ''molaris dens'', meaning "millstone to ...
. The adult dental formula is for a total of 34 teeth. The occlusal surface of the molars is generally smooth but with longitudinal furrows.


Internal systems

Flying foxes have large hearts and a relatively fast
heart rate Heart rate (or pulse rate) is the frequency of the heartbeat measured by the number of contractions (beats) of the heart per minute (bpm). The heart rate can vary according to the body's physical needs, including the need to absorb oxygen and excr ...
: resting individuals have a heart rate of 100-400 beats per minute. Flying foxes have simple
digestive tracts The gastrointestinal tract (GI tract, digestive tract, alimentary canal) is the tract or passageway of the digestive system that leads from the mouth to the anus. The GI tract contains all the major organs of the digestive system, in humans and ...
; the time between ingestion and excretion is as short as 12 minutes. They lack both a
cecum The cecum or caecum is a pouch within the peritoneum that is considered to be the beginning of the large intestine. It is typically located on the right side of the body (the same side of the body as the appendix (anatomy), appendix, to which i ...
and an appendix. The stomach has marked cardiac and fundic regions.


Intelligence

The megabats, including flying foxes, have the greatest
encephalization quotient Encephalization quotient (EQ), encephalization level (EL), or just encephalization is a relative brain size measure that is defined as the ratio between observed to predicted brain mass for an animal of a given size, based on nonlinear regressi ...
(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 In ethology and behavioral ecology, trap-lining or traplining is a feeding strategy in which an individual visits food sources on a regular, repeatable sequence, much as trappers check their lines of traps. Traplining is usually seen in species ...
. They can also be conditioned to perform behaviors, such as one study where
spectacled flying fox The spectacled flying fox (''Pteropus conspicillatus''), also known as the spectacled fruit bat, is a megabat that lives in Australia's north-eastern regions of Queensland. It is also found in New Guinea and on the offshore islands including Wood ...
es were trained to pull a lever using juice as a
reinforcement In behavioral psychology, reinforcement is a consequence applied that will strengthen an organism's future behavior whenever that behavior is preceded by a specific antecedent stimulus. This strengthening effect may be measured as a higher freq ...
. 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.


Senses


Smell

Flying foxes rely heavily on their sense of smell. They have large
olfactory bulb The olfactory bulb (Latin: ''bulbus olfactorius'') is a grey matter, neural structure of the vertebrate forebrain involved in olfaction, the sense of odor, smell. It sends olfactory information to be further processed in the amygdala, the orbitof ...
s 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 An androgen (from Greek ''andr-'', the stem of the word meaning "man") is any natural or synthetic steroid hormone that regulates the development and maintenance of male characteristics in vertebrates by binding to androgen receptors. This inc ...
-sensitive
sebaceous gland A sebaceous gland is a microscopic exocrine gland in the skin that opens into a hair follicle to secrete an oily or waxy matter, called sebum, which lubricates the hair and skin of mammals. In humans, sebaceous glands occur in the greatest number ...
s on their shoulders that they use for
scent-marking In ethology, territory is the sociographical area that an animal consistently defends against conspecific competition (or, occasionally, against animals of other species) using agonistic behaviors or (less commonly) real physical aggression. A ...
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.


Sight

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 In biology, binocular vision is a type of vision in which an animal has two eyes capable of facing the same direction to perceive a single three-dimensional image of its surroundings. Binocular vision does not typically refer to vision where an ...
. Like most mammals, though not
primate Primates are a diverse order of mammals. They are divided into the strepsirrhines, which include the lemurs, galagos, and lorisids, and the haplorhines, which include the tarsiers and the simians (monkeys and apes, the latter including huma ...
s, they are dichromatic. They have both rods and
cones A cone is a three-dimensional geometric shape that tapers smoothly from a flat base (frequently, though not necessarily, circular) to a point called the apex or vertex. A cone is formed by a set of line segments, half-lines, or lines conn ...
; 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.


Evolutionary history

Flying foxes are poorly represented in the fossil record. Relative to the current number of extant species, the Pteropodidae has one of the most incomplete fossil records of any bat group. As of 2014, no flying fox fossils are known from before the
Holocene The Holocene ( ) is the current geological epoch. It began approximately 11,650 cal years Before Present (), after the Last Glacial Period, which concluded with the Holocene glacial retreat. The Holocene and the preceding Pleistocene togethe ...
. Many flying foxes live in the
tropics The tropics are the regions of Earth surrounding the Equator. They are defined in latitude by the Tropic of Cancer in the Northern Hemisphere at N and the Tropic of Capricorn in the Southern Hemisphere at S. The tropics are also referred to ...
, where conditions for fossilization are poor. Based on
molecular evolution Molecular evolution is the process of change in the sequence composition of cellular molecules such as DNA, RNA, and proteins across generations. The field of molecular evolution uses principles of evolutionary biology and population genetics ...
, flying foxes diverged from a common ancestor with ''
Rousettus ''Rousettus'' is a genus of Old World fruit bats or megabats, referred to as rousette bats. The genus is a member of the family Pteropodidae. The genus consists of seven species that range over most of Africa to southeast Asia, and the islands o ...
'' 28–18 million years ago and from their
sister taxa In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and t ...
''
Neopteryx The small-toothed fruit bat or small-toothed Flying fox (''Neopteryx frosti'') is a species of megabat in the family Pteropodidae. It is monotypic, the only species within the genus ''Neopteryx''. It is endemic to central Indonesia. It is known o ...
'' and ''Acerodon'' 6.6–10.6 million years ago. ''Neopteryx'', ''Acerodon'', ''
Desmalopex ''Desmalopex'' (Known as the White-winged Flying Fox or mottled-winged Flying Fox) is a genus of megabats in the family Pteropodidae. It has historically been included in the genus ''Pteropus'' and occurs only in the Philippines. It comprises t ...
'', ''
Melonycteris ''Melonycteris'' (dark blossom bat) is a genus of megabat in the family Pteropodidae. Members are found in the Solomon Islands or in the case of the black-bellied fruit bat, in Papua New Guinea. It contains the following species: * Fardoulis's ...
'', ''
Mirimiri The Fijian monkey-faced bat (''Mirimiri acrodonta'') Also known as Fijian flying fox or Fijian flying monkey, is a megabat endemic to Fiji. It was discovered in old-growth cloud forest on Des Vœux Peak, the second highest mountain peak (1,195 m ...
'', ''
Pteralopex ''Pteralopex'' is a genus of large megabats in the family Pteropodidae. Species in this genus are commonly known as "monkey-faced bats". They are restricted to Solomon Islands rain forests in Melanesia, and all species are seriously threatened, b ...
'', and ''
Styloctenium ''Styloctenium'' (known as stripe-faced fruit bat or stripe-faced flying fox) is a genus of stripe-faced fruit bat Bats are mammals of the order Chiroptera.''cheir'', "hand" and πτερόν''pteron'', "wing". With their forelimbs adapte ...
'' are all relatively closely related to the flying foxes, as they are the other members of its
subfamily In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end subfamily botanical names with "-oideae", and zoologi ...
Pteropodinae The Pteropodinae are a subfamily of megabats. Taxa within this subfamily are: * Genus ''Acerodon'' **Sulawesi flying fox, ''A. celebensis'' **Talaud flying fox, ''A. humilis'' **Giant golden-crowned flying fox, ''A. jubatus'' **Palawan fruit bat ...
. Phylogenetic analysis indicates that flying foxes diversified rapidly in an explosive
evolutionary radiation An evolutionary radiation is an increase in taxonomic diversity that is caused by elevated rates of speciation, that may or may not be associated with an increase in morphological disparity. Radiations may affect one clade or many, and be rapid ...
, creating many taxa in a relatively short time frame. Most flying fox lineages emerged after the
Zanclean The Zanclean is the lowest stage or earliest age on the geologic time scale of the Pliocene. It spans the time between 5.332 ± 0.005 Ma (million years ago) and 3.6 ± 0.005 Ma. It is preceded by the Messinian Age of the Miocene Epoch, and fol ...
, with two major
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
s created: one consisting of the
Indian Ocean The Indian Ocean is the third-largest of the world's five oceanic divisions, covering or ~19.8% of the water on Earth's surface. It is bounded by Asia to the north, Africa to the west and Australia to the east. To the south it is bounded by th ...
species and the other of the
Melanesia Melanesia (, ) is a subregion of Oceania in the southwestern Pacific Ocean. It extends from Indonesia's New Guinea in the west to Fiji in the east, and includes the Arafura Sea. The region includes the four independent countries of Fiji, Va ...
n,
Micronesia Micronesia (, ) is a subregion of Oceania, consisting of about 2,000 small islands in the western Pacific Ocean. It has a close shared cultural history with three other island regions: the Philippines to the west, Polynesia to the east, and ...
n, Australian, and insular
Southeast Asia Southeast Asia, also spelled South East Asia and South-East Asia, and also known as Southeastern Asia, South-eastern Asia or SEA, is the geographical United Nations geoscheme for Asia#South-eastern Asia, south-eastern region of Asia, consistin ...
n 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
Livingstone's fruit bat Livingstone's fruit bat (''Pteropus livingstonii''), also called the Comoro flying fox, is a megabat in the genus ''Pteropus''. It is an Old World fruit bat found only in the Anjouan and Mohéli islands in the Union of the Comoros in the western I ...
and the Pemba flying fox, which are the westernmost flying foxes. A second colonization event resulted in the
Rodrigues flying fox The Rodrigues flying fox or Rodrigues fruit bat (''Pteropus rodricensis'') is a species of bat in the family Pteropodidae, the flying foxes or fruit bats. It is endemic to Rodrigues, an island in the Indian Ocean belonging to Mauritius. Its nat ...
to
Rodrigues Rodrigues (french: Île Rodrigues, link=yes ; Creole: ) is a autonomous outer island of the Republic of Mauritius in the Indian Ocean, about east of Mauritius. It is part of the Mascarene Islands, which include Mauritius and Réunion. Rodr ...
Island; while a third event resulted in several species diverging to
Mauritius Mauritius ( ; french: Maurice, link=no ; mfe, label=Mauritian Creole, Moris ), officially the Republic of Mauritius, is an island nation in the Indian Ocean about off the southeast coast of the African continent, east of Madagascar. It incl ...
, the
Seychelles Seychelles (, ; ), officially the Republic of Seychelles (french: link=no, République des Seychelles; Creole: ''La Repiblik Sesel''), is an archipelagic state consisting of 115 islands in the Indian Ocean. Its capital and largest city, V ...
,
Madagascar Madagascar (; mg, Madagasikara, ), officially the Republic of Madagascar ( mg, Repoblikan'i Madagasikara, links=no, ; french: République de Madagascar), is an island country in the Indian Ocean, approximately off the coast of East Africa ...
, and
Aldabra Aldabra is the world's second-largest coral atoll, lying south-east of the continent of Africa. It is part of the Aldabra Group of islands in the Indian Ocean that are part of the Outer Islands of the Seychelles, with a distance of 1,120 k ...
. With one possible exception - the masked flying fox (''P. personatus''), flying foxes are likely
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
. There are over 60 extant species of flying fox. Flying foxes are now present from the western Indian Ocean midway through the
Pacific Ocean The Pacific Ocean is the largest and deepest of Earth's five oceanic divisions. It extends from the Arctic Ocean in the north to the Southern Ocean (or, depending on definition, to Antarctica) in the south, and is bounded by the continen ...
as far east as the
Cook Islands ) , image_map = Cook Islands on the globe (small islands magnified) (Polynesia centered).svg , capital = Avarua , coordinates = , largest_city = Avarua , official_languages = , lan ...
. They are found in tropical and subtropical climates.


Biology and ecology


Reproduction and life cycle

Many species of flying fox are
polygynandrous Polygynandry is a mating system in which both males and females have multiple mating partners during a breeding season. In sexually reproducing diploid animals, different mating strategies are employed by males and females, because the cost of ga ...
, meaning that each individual will mate with several other individuals. The
Samoa flying fox The Samoa flying fox or Samoan flying fox (''Pteropus samoensis'') is a species of flying fox in the family Pteropodidae. It is found in American Samoa, Fiji, and Samoa (where it is known as ''pe'a'' and ''pe'a vao''). Its natural habitat is sub ...
is a notable exception because it is
monogamous Monogamy ( ) is a form of Dyad (sociology), dyadic Intimate relationship, relationship in which an individual has only one Significant other, partner during their lifetime. Alternately, only one partner at any one time (Monogamy#Serial monogamy, ...
. Flying fox sexual behaviors include oral sex in addition to intercourse, with
fellatio Fellatio (also known as fellation, and in slang as blowjob, BJ, giving head, or sucking off) is an oral sex act involving a person stimulating the penis of another person by using the mouth, throat, or both. Oral stimulation of the scrotum may ...
and
cunnilingus Cunnilingus is an oral sex act performed by a person on the vulva or vagina of another person. The clitoris is the most sexually sensitive part of the human female genitalia, and its stimulation may result in a woman becoming sexually aroused ...
observed between opposite sexes, as well as homosexual fellatio in at least one species, the
Bonin flying fox The Bonin flying fox, Bonin fruit bat (''Pteropus pselaphon''), or in Japanese is a species of flying fox in the family Pteropodidae. It is endemic to four islands (Chichijima, Hahajima, North Iwo Jima, and South Iwo Jima) in Ogasawara Islands, ...
. 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 Two things are identical if they are the same, see Identity (philosophy). Identical may also refer to: * ''Identical'' (Hopkins novel), a 2008 young adult novel by Ellen Hopkins * ''Identical'' (Turow novel), a 2013 legal drama novel by Scott T ...
, or the result of
superfetation Superfetation (also spelled superfoetation – see fetus) is the simultaneous occurrence of more than one stage of developing offspring in the same animal. In mammals, it manifests as the formation of an embryo from a different menstrual cycle wh ...
. Pups are
altricial In biology, altricial species are those in which the young are underdeveloped at the time of birth, but with the aid of their parents mature after birth. Precocial species are those in which the young are relatively mature and mobile from the mome ...
and sparsely furred at birth, thereby dependent on their mothers for care. 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 The Bismarck masked flying fox (''Pteropus capistratus'') is a species of flying fox in the family Pteropodidae found in Papua New Guinea and named after the Bismarck Archipelago. It was once considered a subspecies of '' Pteropus temminckii'' b ...
, can lactate, it is unclear if the lactation is functional and males actually nurse pups or if it is a result of
stress Stress may refer to: Science and medicine * Stress (biology), an organism's response to a stressor such as an environmental condition * Stress (linguistics), relative emphasis or prominence given to a syllable in a word, or to a word in a phrase ...
or
malnutrition Malnutrition occurs when an organism gets too few or too many nutrients, resulting in health problems. Specifically, it is "a deficiency, excess, or imbalance of energy, protein and other nutrients" which adversely affects the body's tissues a ...
. Pups
fledge Fledging is the stage in a flying animal's life between hatching or birth and becoming capable of flight. This term is most frequently applied to birds, but is also used for bats. For altricial birds, those that spend more time in vulnerable c ...
beginning at 3 months old, but may not be
weaned Weaning is the process of gradually introducing an infant human or another mammal to what will be its adult diet while withdrawing the supply of its mother's milk. The process takes place only in mammals, as only mammals produce milk. The infan ...
until 4–6 months old. Pups may stay with their mothers until age one. Flying foxes do not reach
sexual maturity Sexual maturity is the capability of an organism to reproduce. In humans it might be considered synonymous with adulthood, but here puberty is the name for the process of biological sexual maturation, while adulthood is based on cultural definitio ...
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 breeder Seasonal breeders are animal species that successfully mate only during certain times of the year. These times of year allow for the optimization of survival of young due to factors such as ambient temperature, food and water availability, and cha ...
s and give birth in the
spring Spring(s) may refer to: Common uses * Spring (season), a season of the year * Spring (device), a mechanical device that stores energy * Spring (hydrology), a natural source of water * Spring (mathematics), a geometric surface in the shape of a ...
, though the
Mariana fruit bat The Mariana fruit bat (''Pteropus mariannus''), also known as the Mariana flying fox, and the ''fanihi'' in Chamorro, is a megabat found only in the Mariana Islands and Ulithi (an atoll in the Caroline Islands). Habitat loss has driven it to enda ...
seems to have aseasonal breeding with new pups documented throughout the year. Females remain
fertile Fertility is the capability to produce offspring through reproduction following the onset of sexual maturity. The fertility rate is the average number of children born by a female during her lifetime and is quantified demographically. Fertilit ...
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 Disturbance and its variants may refer to: Math and science * Disturbance (ecology), a temporary change in average environmental conditions that causes a pronounced change in an ecosystem * Disturbance (geology), linear zone of faults and folds ...
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 named Statler, who was a resident at Bat World Sanctuary for his last few years. He was born at a zoo in 1987, and was 34 years old at the time of his death.


Social systems

Most flying fox species are
gregarious Sociality is the degree to which individuals in an animal population tend to associate in social groups (gregariousness) and form cooperative societies. Sociality is a survival response to evolutionary pressures. For example, when a mother wasp ...
and form large aggregations of individuals called
colonies In modern parlance, a colony is a territory subject to a form of foreign rule. Though dominated by the foreign colonizers, colonies remain separate from the administration of the original country of the colonizers, the '' metropolitan state'' ...
or "camps." The large flying fox forms colonies of up to 15,000 individuals, while the
little red flying fox The little red flying-fox (''Pteropus scapulatus'') is a megachiropteran bat native to northern and eastern Australia. The species weighs about half a kilogram, one US pound, and is the smallest species of ''Pteropus'' in mainland Australia. '' ...
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 The Ceram fruit bat or Seram flying fox (''Pteropus ocularis'') is a species of megabat in the family Pteropodidae. It is endemic to the mountainous forests of two Maluku islands, Buru and Seram, including the Manusela National Park on Seram. T ...
, are solitary. Colony size varies throughout the year in response to biological needs. The grey-headed flying fox forms
harems Harem (Persian: حرمسرا ''haramsarā'', ar, حَرِيمٌ ''ḥarīm'', "a sacred inviolable place; harem; female members of the family") refers to domestic spaces that are reserved for the women of the house in a Muslim family. A hare ...
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 A juvenile is an individual organism that has not yet reached its adult form, sexual maturity or size. Juveniles can look very different from the adult form, particularly in colour, and may not fill the same niche as the adult form. In many org ...
males. Subadults will form mixed-sex "subadult groups" with each other. In the summer, however, individuals are solitary, with the exception of
nursing Nursing is a profession within the health care sector focused on the care of individuals, families, and communities so they may attain, maintain, or recover optimal health and quality of life. Nurses may be differentiated from other health ...
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
cicada The cicadas () are a superfamily, the Cicadoidea, of insects in the order Hemiptera (true bugs). They are in the suborder Auchenorrhyncha, along with smaller jumping bugs such as leafhoppers and froghoppers. The superfamily is divided into two ...
s as well. In Australia,
eucalypt Eucalypt is a descriptive name for woody plants with capsule fruiting bodies belonging to seven closely related genera (of the tribe Eucalypteae) found across Australasia: ''Eucalyptus'', '' Corymbia'', '' Angophora'', ''Stockwellia'', ''Allosyn ...
blossoms and pollen are preferred food sources, followed by ''
Melaleuca ''Melaleuca'' () is a genus of nearly 300 species of plants in the myrtle family, Myrtaceae, commonly known as paperbarks, honey-myrtles or tea-trees (although the last name is also applied to species of '' Leptospermum''). They range in size ...
'' and ''
Banksia ''Banksia'' is a genus of around 170 species in the plant family Proteaceae. These Australian wildflowers and popular garden plants are easily recognised by their characteristic flower spikes, and fruiting "cones" and heads. ''Banksias'' range i ...
'' flowers They feed on a wide variety of
crop A crop is a plant that can be grown and harvested extensively for profit or subsistence. When the plants of the same kind are cultivated at one place on a large scale, it is called a crop. Most crops are cultivated in agriculture or hydroponic ...
s as well, causing conflicts with farmers. Crops eaten by flying foxes include
sisal Sisal (, ) (''Agave sisalana'') is a species of flowering plant native to southern Mexico, but widely cultivated and naturalized in many other countries. It yields a stiff fibre used in making rope and various other products. The term sisal ma ...
,
cashew The cashew tree (''Anacardium occidentale'') is a tropical evergreen tree native to South America in the genus ''Anacardium'' that produces the cashew seed and the cashew apple accessory fruit. The tree can grow as tall as , but the dwarf cult ...
,
pineapple The pineapple (''Ananas comosus'') is a tropical plant with an edible fruit; it is the most economically significant plant in the family Bromeliaceae. The pineapple is indigenous to South America, where it has been cultivated for many centuri ...
,
areca ''Areca'' is a genus of 51 species of palms in the family Arecaceae, found in humid tropical forests from the islands of the Philippines, Malaysia and India, across Southeast Asia to Melanesia. The generic name ''Areca'' is derived from a name u ...
,
breadfruit Breadfruit (''Artocarpus altilis'') is a species of flowering tree in the mulberry and jackfruit family (Moraceae) believed to be a domesticated descendant of ''Artocarpus camansi'' originating in New Guinea, the Maluku Islands, and the Philippi ...
,
jackfruit The jackfruit (''Artocarpus heterophyllus''), also known as jack tree, is a species of tree in the fig, mulberry, and breadfruit family (Moraceae). Its origin is in the region between the Western Ghats of southern India, all of Bangladesh, ...
,
neem ''Azadirachta indica'', commonly known as neem, nimtree or Indian lilac, is a tree in the mahogany family Meliaceae. It is one of two species in the genus ''Azadirachta'', and is native to the Indian subcontinent and most of the countries in Afr ...
,
papaya The papaya (, ), papaw, () or pawpaw () is the plant species ''Carica papaya'', one of the 21 accepted species in the genus ''Carica'' of the family Caricaceae. It was first domesticated in Mesoamerica, within modern-day southern Mexico and ...
,
citrus ''Citrus'' is a genus of flowering plant, flowering trees and shrubs in the rue family, Rutaceae. Plants in the genus produce citrus fruits, including important crops such as Orange (fruit), oranges, Lemon, lemons, grapefruits, pomelos, and lim ...
,
fig The fig is the edible fruit of ''Ficus carica'', a species of small tree in the flowering plant family Moraceae. Native to the Mediterranean and western Asia, it has been cultivated since ancient times and is now widely grown throughout the world ...
,
mango A mango is an edible stone fruit produced by the tropical tree ''Mangifera indica''. It is believed to have originated in the region between northwestern Myanmar, Bangladesh, and northeastern India. ''M. indica'' has been cultivated in South a ...
,
banana A banana is an elongated, edible fruit – botanically a berry – produced by several kinds of large herbaceous flowering plants in the genus ''Musa''. In some countries, bananas used for cooking may be called "plantains", distinguis ...
,
avocado The avocado (''Persea americana'') is a medium-sized, evergreen tree in the laurel family (Lauraceae). It is native to the Americas and was first domesticated by Mesoamerican tribes more than 5,000 years ago. Then as now it was prized for i ...
,
guava Guava () is a common tropical fruit cultivated in many tropical and subtropical regions. The common guava ''Psidium guajava'' (lemon guava, apple guava) is a small tree in the myrtle family ( Myrtaceae), native to Mexico, Central America, the ...
,
sugar cane Sugarcane or sugar cane is a species of (often hybrid) tall, perennial grass (in the genus ''Saccharum'', tribe Andropogoneae) that is used for sugar production. The plants are 2–6 m (6–20 ft) tall with stout, jointed, fibrous stalks t ...
,
tamarind Tamarind (''Tamarindus indica'') is a Legume, leguminous tree bearing edible fruit that is probably indigenous to tropical Africa. The genus ''Tamarindus'' is monotypic taxon, monotypic, meaning that it contains only this species. It belongs ...
,
grape A grape is a fruit, botanically a berry, of the deciduous woody vines of the flowering plant genus ''Vitis''. Grapes are a non- climacteric type of fruit, generally occurring in clusters. The cultivation of grapes began perhaps 8,000 years ago, ...
s, 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 Vitamin C (also known as ascorbic acid and ascorbate) is a water-soluble vitamin found in citrus and other fruits and vegetables, also sold as a dietary supplement and as a topical 'serum' ingredient to treat melasma (dark pigment spots) an ...
,
calcium Calcium is a chemical element with the symbol Ca and atomic number 20. As an alkaline earth metal, calcium is a reactive metal that forms a dark oxide-nitride layer when exposed to air. Its physical and chemical properties are most similar to ...
,
chondroitin sulfate Chondroitin sulfate is a sulfated glycosaminoglycan (GAG) composed of a chain of alternating sugars ( N-acetylgalactosamine and glucuronic acid). It is usually found attached to proteins as part of a proteoglycan. A chondroitin chain can have ove ...
, and
glucosamine Glucosamine (C6H13NO5) is an amino sugar and a prominent precursor in the biochemical synthesis of glycosylated proteins and lipids. Glucosamine is part of the structure of two polysaccharides, chitosan and chitin. Glucosamine is one of the most ...
can be recommended periodically. The majority of flying fox species are
nocturnal Nocturnality is an animal behavior characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnal meaning the opposite. Nocturnal creatures generally have highly developed sens ...
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 The Caroline flying fox or Pohnpei flying fox (''Pteropus molossinus'') is a species of megabat in the genus ''Pteropus'', endemic to Micronesia. Its natural habitat is subtropical or tropical dry forests. Prior to a ban on commercial exploitatio ...
, '' P. p. insularis'', and the
Seychelles fruit bat The Seychelles fruit bat or Seychelles flying fox (''Pteropus seychellensis'') is a megabat found on the granitic islands of Seychelles, and on the Comoros and Mafia Island. It is a significant component of the ecosystems for the islands, disper ...
. Foraging resources are often far from roosts, with individuals traveling up to to reach them. Flying foxes can travel at for three hours or more, and can reach top speeds of . 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 The palate () is the roof of the mouth in humans and other mammals. It separates the oral cavity from the nasal cavity. A similar structure is found in crocodilians, but in most other tetrapods, the oral and nasal cavities are not truly sepa ...
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
pollinator A pollinator is an animal that moves pollen from the male anther of a flower to the female stigma of a flower. This helps to bring about fertilization of the ovules in the flower by the male gametes from the pollen grains. Insects are the maj ...
s. They help spread the seeds in the fruit they eat by discarding them in ejecta pellets or through their
guano Guano (Spanish from qu, wanu) is the accumulated excrement of seabirds or bats. As a manure, guano is a highly effective fertilizer due to the high content of nitrogen, phosphate, and potassium, all key nutrients essential for plant growth. G ...
. 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 Pioneer species are hardy species that are the first to colonize barren environments or previously biodiverse steady-state ecosystems that have been disrupted, such as by wildfire. Pioneer flora Some lichens grow on rocks without soil, so m ...
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 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 The durian (, ) is the edible fruit of several tree species belonging to the genus ''Durio''. There are 30 recognised ''Durio'' species, at least nine of which produce edible fruit. ''Durio zibethinus'', native to Borneo and Sumatra, is the onl ...
. 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.


Conservation


Conservation status

Of the 62 flying fox species evaluated by the
IUCN The International Union for Conservation of Nature (IUCN; officially International Union for Conservation of Nature and Natural Resources) is an international organization working in the field of nature conservation and sustainable use of natu ...
as of 2018, 3 are considered critically endangered: the
Aru flying fox The Aru flying fox (''Pteropus aruensis'') is a Critically Endangered species of megabat found in the Aru Islands in Indonesia. It was described by Wilhelm Peters in 1867. It was formerly considered a subspecies of the black-bearded flying fox.Si ...
, Livingstone's fruit bat, and the
Vanikoro flying fox The Vanikoro flying fox (''Pteropus tuberculatus''), also known locally as the basapine, is a species of bat in the family Pteropodidae. It has only been found in the Vanikoro island group located in the southern Solomon Islands. The species a ...
. Another 7 species are listed as
endangered An endangered species is a species that is very likely to become extinct in the near future, either worldwide or in a particular political jurisdiction. Endangered species may be at risk due to factors such as habitat loss, poaching and inva ...
; 20 are listed as vulnerable, 6 as
near threatened A near-threatened species is a species which has been categorized as "Near Threatened" (NT) by the International Union for Conservation of Nature as that may be vulnerable to endangerment in the near future, but it does not currently qualify fo ...
, 14 as
least concern A least-concern species is a species that has been categorized by the International Union for Conservation of Nature (IUCN) as evaluated as not being a focus of species conservation because the specific species is still plentiful in the wild. T ...
, and 8 as
data deficient A data deficient (DD) species is one which has been categorized by the International Union for Conservation of Nature (IUCN) as offering insufficient information for a proper assessment of conservation status to be made. This does not necessaril ...
. A further 4 are listed as
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
: the dusky flying fox, the
large Palau flying fox The large Palau flying fox (''Pteropus pilosus'') is an extinct species of medium-sized megabats from the Palau Islands in Micronesia. It had brownish fur with long, silvery hairs on its belly, and a wingspan of about 60 cm. It probably be ...
, the
small Mauritian flying fox The small Mauritian flying fox or dark flying fox (''Pteropus subniger''), known as a ''rougette'' to early French travelers, is an extinct species of megabat. It lived on the islands of Réunion and Mauritius in the Mascarene Islands of the Indi ...
, 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.


Legal status

All species of ''Pteropus'' are placed on
Appendix II CITES (shorter name for the Convention on International Trade in Endangered Species of Wild Fauna and Flora, also known as the Washington Convention) is a multilateral treaty to protect endangered plants and animals from the threats of interna ...
of
CITES CITES (shorter name for the Convention on International Trade in Endangered Species of Wild Fauna and Flora, also known as the Washington Convention) is a multilateral treaty to protect endangered plants and animals from the threats of interna ...
and 10 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 the illegal bushmeat trade still occurs outside of
protected area Protected areas or conservation areas are locations which receive protection because of their recognized natural, ecological or cultural values. There are several kinds of protected areas, which vary by level of protection depending on the ena ...
s. 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 A natural monument is a natural or natural/cultural feature of outstanding or unique value because of its inherent rarity, representative of aesthetic qualities or cultural significance. Under World Commission on Protected Areas guidelines, nat ...
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 continental United States, several species and subspecies are listed under its
Endangered Species Act of 1973 The Endangered Species Act of 1973 (ESA or "The Act"; 16 U.S.C. § 1531 et seq.) is the primary law in the United States for protecting imperiled species. Designed to protect critically imperiled species from extinction as a "consequence of ec ...
. ''Pteropus mariannus mariannus''—a subspecies of the Mariana fruit bat—is listed as
threatened Threatened species are any species (including animals, plants and fungi) which are vulnerable to endangerment in the near future. Species that are threatened are sometimes characterised by the population dynamics measure of ''critical depensat ...
while the Rodrigues flying fox and Guam flying fox are listed as
endangered An endangered species is a species that is very likely to become extinct in the near future, either worldwide or in a particular political jurisdiction. Endangered species may be at risk due to factors such as habitat loss, poaching and inva ...
. 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 Vermin (colloquially varmint(s) or varmit(s)) are pests or nuisance animals that spread diseases or destroy crops or livestock. Since the term is defined in relation to human activities, which species are included vary by region and enterpr ...
" under the
Wildlife Protection Act of 1972 The Wild Life (Protection) Act, 1972 is an Act of the Parliament of India enacted for protection of plants and animal species. Before 1972, India had only five designated national parks. Among other reforms, the Act established scheduled pr ...
. 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 Malaysia ( ; ) is a country in Southeast Asia. The federation, federal constitutional monarchy consists of States and federal territories of Malaysia, thirteen states and three federal territories, separated by the South China Sea into two r ...
. 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 U.S.$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 Terengganu (; Terengganu Malay: ''Tranung'', Jawi: ), formerly spelled Trengganu or Tringganu, is a sultanate and constitutive state of federal Malaysia. The state is also known by its Arabic honorific, ''Dāru l- Īmān'' ("Abode of Faith"). ...
issued a moratorium on hunting flying foxes. In
Sarawak Sarawak (; ) is a States and federal territories of Malaysia, state of Malaysia. The largest among the 13 states, with an area almost equal to that of Peninsular Malaysia, Sarawak is located in northwest Borneo Island, and is bordered by the M ...
, all bat species are listed as "Protected" and hunting them is not legal.


Factors causing decline


Anthropogenic sources

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 Mortality rate, or death rate, is a measure of the number of deaths (in general, or due to a specific cause) in a particular population, scaled to the size of that population, per unit of time. Mortality rate is typically expressed in units of de ...
exceeds 22% annually, then it will steadily decline. Invasive species such as the
brown tree snake The brown tree snake (''Boiga irregularis''), also known as the brown catsnake, is an arboreal rear-fanged colubrid snake native to eastern and northern coastal Australia, eastern Indonesia (Sulawesi to Papua), Papua New Guinea, and many islands ...
can seriously impact populations; the brown tree snake consumes so many pups that it reduced the
recruitment Recruitment is the overall process of identifying, sourcing, screening, shortlisting, and interviewing candidates for jobs (either permanent or temporary) within an organization. Recruitment also is the processes involved in choosing individual ...
of the
Guam Guam (; ch, Guåhan ) is an organized, unincorporated territory of the United States in the Micronesia subregion of the western Pacific Ocean. It is the westernmost point and territory of the United States (reckoned from the geographic cent ...
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 extinction Local extinction, also known as extirpation, refers to a species (or other taxon) of plant or animal that ceases to exist in a chosen geographic area of study, though it still exists elsewhere. Local extinctions are contrasted with global extinct ...
s, 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 In law, set-off or netting are legal techniques applied between persons or businesses with mutual rights and liabilities, replacing gross positions with net positions. It permits the rights to be used to discharge the liabilities where cross cla ...
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 of fruit to flying foxes in a year. Some electrocution deaths are also accidental, such as when bats fly into
overhead power line An overhead power line is a structure used in electric power transmission and distribution to transmit electrical energy across large distances. It consists of one or more uninsulated electrical cables (commonly multiples of three for three-p ...
s. 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 Australian 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
atoll An atoll () is a ring-shaped island, including a coral rim that encircles a lagoon partially or completely. There may be coral islands or cays on the rim. Atolls are located in warm tropical or subtropical oceans and seas where corals can gr ...
s.


Natural sources

Because many species are endemic to a single island, they are vulnerable to random events such as
typhoon A typhoon is a mature tropical cyclone that develops between 180° and 100°E in the Northern Hemisphere. This region is referred to as the Northwestern Pacific Basin, and is the most active tropical cyclone basin on Earth, accounting for a ...
s. 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. Flying foxes are also threatened by disease such as
tick paralysis Tick paralysis is the only tick-borne disease that is not caused by an infectious organism. The illness is caused by a neurotoxin produced in the tick's salivary gland. After prolonged attachment, the engorged tick transmits the toxin to its host. ...
. Tick paralysis affects the spectacled flying fox, and is responsible for an estimated 1% of its annual mortality.


Captive breeding

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 Captive breeding, also known as captive propagation, is the process of plants or animals in controlled environments, such as wildlife reserves, zoos, botanic gardens, and other conservation facilities. It is sometimes employed to help species that ...
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 the
Jersey Zoo Jersey Zoo (formerly Durrell Wildlife Park) is a zoological park established in 1959 on the island of Jersey in the English Channel by naturalist and writer Gerald Durrell (1925–1995). It is operated by the Durrell Wildlife Conservation Trust. ...
and the
Bristol Zoo Bristol Zoo was a zoo in the city of Bristol in South West England. The zoo's stated mission was to "maintain and defend” biodiversity through breeding endangered species, conserving threatened species and habitats and promoting a wider unders ...
. Though the program has been successful in increasing the population, caretakers of the captive population have had to deal with
husbandry Animal husbandry is the branch of agriculture concerned with animals that are raised for meat, fibre, milk, or other products. It includes day-to-day care, selective breeding, and the raising of livestock. Husbandry has a long history, starti ...
issues such as
obesity Obesity is a medical condition, sometimes considered a disease, in which excess body fat has accumulated to such an extent that it may negatively affect health. People are classified as obese when their body mass index (BMI)—a person's we ...
and
cardiomyopathy Cardiomyopathy is a group of diseases that affect the heart muscle. Early on there may be few or no symptoms. As the disease worsens, shortness of breath, feeling tired, and swelling of the legs may occur, due to the onset of heart failure. A ...
. 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 Sedentary lifestyle is a lifestyle type, in which one is physically inactive and does little or no physical movement and or exercise. A person living a sedentary lifestyle is often sitting or lying down while engaged in an activity like soci ...
. 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 Durrell Wildlife Conservation Trust is a conservation organization with a mission to save species from extinction. Gerald Durrell founded the Jersey Wildlife Preservation Trust as a charitable institution in 1963 with the dodo as its symbol. The ...
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


Food

Many flying foxes species are killed for
bushmeat Bushmeat is meat from wildlife species that are hunted for human consumption, most often referring to the meat of game in Africa. Bushmeat represents a primary source of animal protein and a cash-earning commodity for inhabitants of humid tropi ...
. The bushmeat harvest is often unsustainable, often resulting in severe population decline or
local extinction Local extinction, also known as extirpation, refers to a species (or other taxon) of plant or animal that ceases to exist in a chosen geographic area of study, though it still exists elsewhere. Local extinctions are contrasted with global extinct ...
. Flying foxes are killed and sold for bushmeat in several countries in Southeast Asia,
South Asia South Asia is the southern subregion of Asia, which is defined in both geographical and ethno-cultural terms. The region consists of the countries of Afghanistan, Bangladesh, Bhutan, India, Maldives, Nepal, Pakistan, and Sri Lanka.;;;;;;;; ...
, and
Oceania Oceania (, , ) is a region, geographical region that includes Australasia, Melanesia, Micronesia, and Polynesia. Spanning the Eastern Hemisphere, Eastern and Western Hemisphere, Western hemispheres, Oceania is estimated to have a land area of ...
, including
Indonesia Indonesia, officially the Republic of Indonesia, is a country in Southeast Asia and Oceania between the Indian and Pacific oceans. It consists of over 17,000 islands, including Sumatra, Java, Sulawesi, and parts of Borneo and New Guine ...
, Malaysia,
Papua New Guinea Papua New Guinea (abbreviated PNG; , ; tpi, Papua Niugini; ho, Papua Niu Gini), officially the Independent State of Papua New Guinea ( tpi, Independen Stet bilong Papua Niugini; ho, Independen Stet bilong Papua Niu Gini), is a country i ...
, the
Philippines The Philippines (; fil, Pilipinas, links=no), officially the Republic of the Philippines ( fil, Republika ng Pilipinas, links=no), * bik, Republika kan Filipinas * ceb, Republika sa Pilipinas * cbk, República de Filipinas * hil, Republ ...
, Bangladesh, China,
Fiji Fiji ( , ,; fj, Viti, ; Fiji Hindi: फ़िजी, ''Fijī''), officially the Republic of Fiji, is an island country in Melanesia, part of Oceania in the South Pacific Ocean. It lies about north-northeast of New Zealand. Fiji consists ...
, and Guam. Flying fox consumption is particularly common in countries with low
food security Food security speaks to the availability of food in a country (or geography) and the ability of individuals within that country (geography) to access, afford, and source adequate foodstuffs. According to the United Nations' Committee on World F ...
and lack of environmental regulation. In some cultures in the region, however, eating flying fox meat is
taboo A taboo or tabu is a social group's ban, prohibition, or avoidance of something (usually an utterance or behavior) based on the group's sense that it is excessively repulsive, sacred, or allowed only for certain persons.''Encyclopædia Britannica ...
. In
Namoluk Namoluk (meaning "lagoon in the middle") is an atoll and municipality in the state of Chuuk, Federated States of Micronesia. It is the smallest atoll in Micronesia, with a total surface of only 13 km2 and a land mass of less than 1 km2. Nam ...
, locals are repulsed by the idea of eating flying foxes because the flying foxes urinate on themselves. In predominately
Muslim Muslims ( ar, المسلمون, , ) are people who adhere to Islam, a monotheistic religion belonging to the Abrahamic tradition. They consider the Quran, the foundational religious text of Islam, to be the verbatim word of the God of Abrah ...
regions such as much of Indonesia, flying foxes are rarely consumed because of
halal ''Halal'' (; ar, حلال, ) is an Arabic word that translates to "permissible" in English. In the Quran, the word ''halal'' is contrasted with ''haram'' (forbidden). This binary opposition was elaborated into a more complex classification kno ...
dietary restrictions.
North Sulawesi North Sulawesi ( id, Sulawesi Utara) is a province of Indonesia. It is located on the Minahasa Peninsula of Sulawesi, south of the Philippines and southeast of Sabah, Malaysia. It borders the Philippine province of Davao Occidental and Soccsks ...
has the greatest demand for flying fox bushmeat. Despite being in Muslim-majority Indonesia, North Sulawesi is predominately
Christian Christians () are people who follow or adhere to Christianity, a monotheistic Abrahamic religion based on the life and teachings of Jesus Christ. The words ''Christ'' and ''Christian'' derive from the Koine Greek title ''Christós'' (Χρι ...
; therefore, many locals do not follow halal guidelines prohibiting flying fox consumption. In
Manado Manado () is the capital City status in Indonesia, city of the Indonesian Provinces of Indonesia, province of North Sulawesi. It is the second largest city in Sulawesi after Makassar, with the 2020 Census giving a population of 451,916 distribu ...
, 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 and the
Commonwealth of the Northern Mariana Islands The Northern Mariana Islands, officially the Commonwealth of the Northern Mariana Islands (CNMI; ch, Sankattan Siha Na Islas Mariånas; cal, Commonwealth Téél Falúw kka Efáng llól Marianas), is an unincorporated territory and commonwea ...
, consumption of the Mariana fruit bat exposes locals to the
neurotoxin Neurotoxins are toxins that are destructive to nerve tissue (causing neurotoxicity). Neurotoxins are an extensive class of exogenous chemical neurological insultsSpencer 2000 that can adversely affect function in both developing and mature ner ...
beta-Methylamino-L-alanine β-Methylamino--alanine, or BMAA, is a non-proteinogenic amino acid produced by cyanobacteria. BMAA is a neurotoxin and its potential role in various neurodegenerative disorders is the subject of scientific research. Structure and properties ...
(BMAA) which may later lead to
neurodegenerative diseases A neurodegenerative disease is caused by the progressive loss of structure or function of neurons, in the process known as neurodegeneration. Such neuronal damage may ultimately involve cell death. Neurodegenerative diseases include amyotrophic ...
. BMAA may become biomagnified in humans who consume flying foxes; flying foxes are exposed to BMAA by eating
cycad Cycads are seed plants that typically have a stout and woody (ligneous) trunk (botany), trunk with a crown (botany), crown of large, hard, stiff, evergreen and (usually) pinnate leaves. The species are dioecious, that is, individual plants o ...
fruits.


Medicine

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 Rheumatism or rheumatic disorders are conditions causing chronic, often intermittent pain affecting the joints or connective tissue. Rheumatism does not designate any specific disorder, but covers at least 200 different conditions, including art ...
. Tribes in the
Attappadi Attappady (HQ:Agali) is a tribal taluk in Kerala state covering an area of . It is carved out from Mannarkkad taluk in Palakkad district on February 2021. Attappady Reserve Forest is a protected area comprising 249 km2 of land area in ...
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."


Transmitting disease

Flying foxes are the
natural reservoir In infectious disease ecology and epidemiology, a natural reservoir, also known as a disease reservoir or a reservoir of infection, is the population of organisms or the specific environment in which an infectious pathogen naturally lives and rep ...
s of several viruses, some of which can be transmitted to humans. Notably, flying foxes can transmit
lyssavirus ''Lyssavirus'' (from the Greek ''lyssa'' "rage, fury, rabies" and the Latin '' vīrus'') is a genus of RNA viruses in the family ''Rhabdoviridae'', order ''Mononegavirales''. Mammals, including humans, can serve as natural hosts. The genus ' ...
es, which cause
rabies Rabies is a viral disease that causes encephalitis in humans and other mammals. Early symptoms can include fever and tingling at the site of exposure. These symptoms are followed by one or more of the following symptoms: nausea, vomiting, vi ...
. In Australia the
rabies virus Rabies virus, scientific name ''Rabies lyssavirus'', is a neurotropic virus that causes rabies in humans and animals. Rabies transmission can occur through the saliva of animals and less commonly through contact with human saliva. ''Rabies lyss ...
is not naturally present;
Australian bat lyssavirus ''Australian bat lyssavirus'' (''ABLV''), originally named ''Pteropid lyssavirus'' (''PLV''), is a zoonotic virus closely related to the rabies virus. It was first identified in a 5-month-old juvenile black flying fox (''Pteropus alecto'') col ...
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 A mucous membrane or mucosa is a membrane that lines various cavities in the body of an organism and covers the surface of internal organs. It consists of one or more layers of epithelial cells overlying a layer of loose connective tissue. It is ...
or an open
wound A wound is a rapid onset of injury that involves laceration, lacerated or puncture wound, punctured skin (an ''open'' wound), or a bruise, contusion (a ''closed'' wound) from blunt force physical trauma, trauma or compression. In pathology, a '' ...
. 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 ) , nickname = Sunshine State , image_map = Queensland in Australia.svg , map_caption = Location of Queensland in Australia , subdivision_type = Country , subdivision_name = Australia , established_title = Before federation , established_ ...
and each case was fatal. Flying foxes are also reservoirs of
henipavirus ''Henipavirus'' is a genus of negative-strand RNA viruses in the family ''Paramyxoviridae'', order ''Mononegavirales'' containing six established species, and numerous others still under study. Henipaviruses are naturally harboured by several sp ...
es such as
Hendra virus Hendra virus (HeV), scientific name ''Hendra henipavirus'', is a bat-borne virus that is associated with a highly fatal infection in horses and humans. Numerous disease outbreaks in Australia among horses have been caused by Hendra virus. The Hend ...
and
Nipah virus Nipah virus, scientific name ''Nipah henipavirus'', is a bat-borne virus that causes Nipah virus infection in humans and other animals, a disease with a high mortality rate. Numerous disease outbreaks caused by Nipah virus have occurred in Sout ...
. 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
horse The horse (''Equus ferus caballus'') is a domesticated, one-toed, hoofed mammal. It belongs to the taxonomic family Equidae and is one of two extant subspecies of ''Equus ferus''. The horse has evolved over the past 45 to 55 million y ...
s that have come into contact with flying fox
urine Urine is a liquid by-product of metabolism in humans and in many other animals. Urine flows from the kidneys through the ureters to the urinary bladder. Urination results in urine being excretion, excreted from the body through the urethra. Cel ...
. There are no documented instances of direct transmission between flying foxes and humans. As of 2012, there is a
vaccine A vaccine is a biological Dosage form, preparation that provides active acquired immunity to a particular infectious disease, infectious or cancer, malignant disease. The safety and effectiveness of vaccines has been widely studied and verifie ...
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 Singapore (), officially the Republic of Singapore, is a sovereign island country and city-state in maritime Southeast Asia. It lies about one degree of latitude () north of the equator, off the southern tip of the Malay Peninsula, borde ...
,
India India, officially the Republic of India (Hindi: ), is a country in South Asia. It is the seventh-largest country by area, the second-most populous country, and the most populous democracy in the world. Bounded by the Indian Ocean on the so ...
, and Bangladesh, resulting in over 100 casualties. A 2018 outbreak in
Kerala, India Kerala ( ; ) is a States and union territories of India, state on the Malabar Coast of India. It was formed on 1 November 1956, following the passage of the States Reorganisation Act, by combining Malayalam-speaking regions of the erstwhile ...
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 A host is a person responsible for guests at an event or for providing hospitality during it. Host may also refer to: Places * Host, Pennsylvania, a village in Berks County People *Jim Host (born 1937), American businessman * Michel Host ...
such as
domestic pig The pig (''Sus domesticus''), often called swine, hog, or domestic pig when distinguishing from other members of the genus '' Sus'', is an omnivorous, domesticated, even-toed, hoofed mammal. It is variously considered a subspecies of ''Sus s ...
s, 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 ''Phoenix dactylifera'', commonly known as date or date palm, is a flowering plant species in the palm family, Arecaceae, cultivated for its edible sweet fruit called dates. The species is widely cultivated across northern Africa, the Middle Eas ...
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 ''Menangle pararubulavirus'', also called Menangle virus, is a virus that infects pigs, humans and bats. History Menangle virus was first identified in 1997 after a piggery in Menangle near Sydney, NSW, Australia experienced a high number o ...
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 Severe acute respiratory syndrome (SARS) is a viral respiratory disease of zoonotic origin caused by the severe acute respiratory syndrome coronavirus (SARS-CoV or SARS-CoV-1), the first identified strain of the SARS coronavirus species, ''sever ...
and
Ebola Ebola, also known as Ebola virus disease (EVD) and Ebola hemorrhagic fever (EHF), is a viral hemorrhagic fever in humans and other primates, caused by ebolaviruses. Symptoms typically start anywhere between two days and three weeks after becom ...
, flying foxes are not suspected as hosts for either causative virus.


Pests

Flying foxes are often considered
pests PESTS was an anonymous American activist group formed in 1986 to critique racism, tokenism, and exclusion in the art world. PESTS produced newsletters, posters, and other print material highlighting examples of discrimination in gallery represent ...
due to the damage they cause to
orchard An orchard is an intentional plantation of trees or shrubs that is maintained for food production. Orchards comprise fruit- or nut-producing trees which are generally grown for commercial production. Orchards are also sometimes a feature of larg ...
crops. Flying foxes have been cited as particularly destructive to
almond The almond (''Prunus amygdalus'', syn. ''Prunus dulcis'') is a species of tree native to Iran and surrounding countries, including the Levant. The almond is also the name of the edible and widely cultivated seed of this tree. Within the genus ...
s,
guava Guava () is a common tropical fruit cultivated in many tropical and subtropical regions. The common guava ''Psidium guajava'' (lemon guava, apple guava) is a small tree in the myrtle family ( Myrtaceae), native to Mexico, Central America, the ...
s, and
mango A mango is an edible stone fruit produced by the tropical tree ''Mangifera indica''. It is believed to have originated in the region between northwestern Myanmar, Bangladesh, and northeastern India. ''M. indica'' has been cultivated in South a ...
es in the Maldives;
lychee Lychee (US: ; UK: ; ''Litchi chinensis''; ) is a Monotypic taxon, monotypic taxon and the sole member in the genus ''Litchi'' in the Sapindus, soapberry family, ''Sapindaceae''. It is a tropical tree native to Southeast and Southwest China (t ...
in Mauritius;
areca ''Areca'' is a genus of 51 species of palms in the family Arecaceae, found in humid tropical forests from the islands of the Philippines, Malaysia and India, across Southeast Asia to Melanesia. The generic name ''Areca'' is derived from a name u ...
in India; and
stone fruit In botany, a drupe (or stone fruit) is an indehiscent fruit in which an outer fleshy part (exocarp, or skin, and mesocarp, or flesh) surrounds a single shell (the ''pit'', ''stone'', or ''pyrena'') of hardened endocarp with a seed (''kernel'') ...
s 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 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 cherry 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.


In culture

Flying foxes are featured in many indigenous cultures and traditions. A
folklore Folklore is shared by a particular group of people; it encompasses the traditions common to that culture, subculture or group. This includes oral traditions such as tales, legends, proverbs and jokes. They include material culture, ranging ...
Dreamtime The Dreaming, also referred to as Dreamtime, is a term devised by early anthropologists to refer to a religio-cultural worldview attributed to Australian Aboriginal mythology, Australian Aboriginal beliefs. It was originally used by Francis Ja ...
story from the
New South Wales North Coast New South Wales North Coast or NSW North Coast, an interim Australian bioregion, is located in New South Wales ) , nickname = , image_map = New South Wales in Australia.svg , map_caption = Location of New South Wales in AustraliaCoord ...
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 Tonga (, ; ), officially the Kingdom of Tonga ( to, Puleʻanga Fakatuʻi ʻo Tonga), is a Polynesian country and archipelago. The country has 171 islands – of which 45 are inhabited. Its total surface area is about , scattered over in ...
, 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 Kolovai is a village on the Tongan island of Tongatapu. Its 2006 population was 4,098. The village is notable for its lakalaka, the national dance of Tonga. A national monument has been proposed to preserve the site of the koka tree where members ...
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 A banyan, also spelled "banian", is a fig that develops accessory trunks from adventitious prop roots, allowing the tree to spread outwards indefinitely. This distinguishes banyans from other trees with a strangler habit that begin life as a ...
tree. Villagers believe that the flying foxes are under the protection of
Muni Muni may refer to: Municipal * A common US abbreviation for municipal, municipal services, and the like *Municipal bond *Municipal Bridge, the former name of the George Rogers Clark Memorial Bridge in Louisville, Kentucky *"Muni", slang for a mu ...
, 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 A cockatoo is any of the 21 parrot species belonging to the family Cacatuidae, the only family in the superfamily Cacatuoidea. Along with the Psittacoidea (true parrots) and the Strigopoidea (large New Zealand parrots), they make up the ord ...
stealing feathers from the flying fox, resulting in it becoming nocturnal. Another story features a flying fox that could 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 Jade is a mineral used as jewellery or for ornaments. It is typically green, although may be yellow or white. Jade can refer to either of two different silicate minerals: nephrite (a silicate of calcium and magnesium in the amphibole group of ...
were decorated with braids of flying fox fur. Flying fox wings were depicted on the war shields of the
Asmat people The Asmat are an ethnic group of New Guinea, residing in the province of South Papua, Indonesia. The Asmat inhabit a region on the island's southwestern coast bordering the Arafura Sea, with lands totaling approximately 18,000 km2 (7,336&n ...
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 The island of Makira (also known as San Cristobal and San Cristóbal) is the largest island of Makira-Ulawa Province in the Solomon Islands. It is third most populous island after Malaita and Guadalcanal, with a population of 55,126 as of 2020 ...
, which is part of the Solomon Islands, indigenous peoples still hunt flying foxes for their teeth as well as for bushmeat. The
canine teeth In mammalian oral anatomy, the canine teeth, also called cuspids, dog teeth, or (in the context of the upper jaw) fangs, eye teeth, vampire teeth, or vampire fangs, are the relatively long, pointed teeth. They can appear more flattened howeve ...
are strung together on necklaces that are used as
currency A currency, "in circulation", from la, currens, -entis, literally meaning "running" or "traversing" is a standardization of money in any form, in use or circulation as a medium of exchange, for example banknotes and coins. A more general def ...
. Teeth of the
insular flying fox The insular flying fox or Pacific flying fox (''Pteropus tonganus'') is a species of flying fox in the family Pteropodidae. It is geographically widespread, the most widespread flying fox in the Pacific: it is found in American Samoa, the Cook I ...
are particularly prized, as they are usually large enough to drill holes in. The
Makira flying fox The Makira flying fox (''Pteropus cognatus'') is a species of megabat in the genus ''Pteropus'', found in the Solomon Islands.D.E. Wilson & D.M. Reeder, 2005: ''Mammal Species of the World: A Taxonomic and Geographic Reference''. Third Edition. Th ...
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."


Other uses

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 Etsy, Inc. is an American e-commerce company focused on handmade or vintage items and craft supplies. These items fall under a wide range of categories, including jewelry, bags, clothing, home décor and furniture, toys, art, as well as craft su ...
or
eBay eBay Inc. ( ) is an American multinational e-commerce company based in San Jose, California, that facilitates consumer-to-consumer and business-to-consumer sales through its website. eBay was founded by Pierre Omidyar in 1995 and became a ...
. From 2000 to 2013, over 100,000 dead bats were imported to the United States. Bat conservationist
Merlin Tuttle Merlin Devere Tuttle (born August 26, 1941) is an American ecologist, conservationist, writer and wildlife photographer who has specialized in bat ecology, behavior, and conservation. He is credited with protecting the Austin Congress Avenue Br ...
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."


References

{{good article Bats of Asia Bats of Africa Fauna of Southeast Asia Mammals of Thailand Articles containing video clips Bat genera Taxa named by Mathurin Jacques Brisson