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The [[saltwater crocodile]], a native species, is a known predator of megabats, based on analysis of crocodile stomach contents in northern Australia.<ref>{{cite journal|doi=10.1371/journal.pone.0197159|pmid=29874276|pmc=5991389|title=Estuarine crocodiles in a tropical coastal floodplain obtain nutrition from terrestrial prey|journal=PLOS ONE|volume=13|issue=6|pages=e0197159|year=2018|last1=Adame|first1=Maria Fernanda|last2=Jardine|first2=Timothy D.|last3=Fry|first3=Brian|last4=Valdez|first4=Dominic|last5=Lindner|first5=Garry|last6=Nadji|first6=Jonathan|last7=Bunn|first7=Stuart E.|bibcode=2018PLoSO..1397159A}}</ref>
The [[saltwater crocodile]], a native species, is a known predator of megabats, based on analysis of crocodile stomach contents in northern Australia.<ref>{{cite journal|doi=10.1371/journal.pone.0197159|pmid=29874276|pmc=5991389|title=Estuarine crocodiles in a tropical coastal floodplain obtain nutrition from terrestrial prey|journal=PLOS ONE|volume=13|issue=6|pages=e0197159|year=2018|last1=Adame|first1=Maria Fernanda|last2=Jardine|first2=Timothy D.|last3=Fry|first3=Brian|last4=Valdez|first4=Dominic|last5=Lindner|first5=Garry|last6=Nadji|first6=Jonathan|last7=Bunn|first7=Stuart E.|bibcode=2018PLoSO..1397159A}}</ref>
During extreme heat events, megabats like the [[little red flying fox]] (''Pteropus scapulatus'') must cool off and rehydrate by drinking from waterways, making them susceptible to opportunistic depredation by [[freshwater crocodiles]].<ref>{{cite AV media| url=https://www.youtube.com/watch?v=wi30w-Mk2yQ| date=10 April 2015|medium=video| publisher=BBC Earth|title=Flying Foxes Vs Freshwater Crocodile|access-date=22 May 2019}}</ref>
During extreme heat events, megabats like the [[little red flying fox]] (''Pteropus scapulatus'') must cool off and rehydrate by drinking from waterways, making them susceptible to opportunistic depredation by [[freshwater crocodiles]].<ref>{{cite AV media| url=https://www.youtube.com/watch?v=wi30w-Mk2yQ| date=10 April 2015|medium=video| publisher=BBC Earth|title=Flying Foxes Vs Freshwater Crocodile|access-date=22 May 2019}}</ref>

Megabats are the hosts of several parasite taxa. Known parasites include [[Nycteribiidae]] and [[Streblidae]] species ("bat flies").<ref>{{cite journal|doi=10.1186/s13071-017-2582-x|pmid=29284533|pmc=5747079|title=Hidden diversity of Nycteribiidae (Diptera) bat flies from the Malagasy region and insights on host-parasite interactions|journal=Parasites & Vectors|volume=10|issue=1|pages=630|year=2017|last1=Ramasindrazana|first1=Beza|last2=Goodman|first2=Steven M.|last3=Gomard|first3=Yann|last4=Dick|first4=Carl W.|last5=Tortosa|first5=Pablo}}</ref><ref>{{cite journal|doi=10.1186/s13071-018-2918-1|pmid=29859123|pmc=5984742|title=Rates of hematophagous ectoparasite consumption during grooming by an endemic Madagascar fruit bat|journal=Parasites & Vectors|volume=11|issue=1|pages=330|year=2018|last1=Ramanantsalama|first1=Riana V.|last2=Andrianarimisa|first2=Aristide|last3=Raselimanana|first3=Achille P.|last4=Goodman|first4=Steven M.}}</ref>
Blood parasites of the family [[Haemoproteidae]] also affect megabat species.<ref>{{cite journal|doi=10.1051/parasite/2012192137|pmid=22550624|pmc=3671437|title=The haemosporidian parasites of bats with description of ''Sprattiella'' alectogen. Nov., sp. Nov|journal=Parasite|volume=19|issue=2|pages=137–146|year=2012|last1=Landau|first1=I.|last2=Chavatte|first2=J.M.|last3=Karadjian|first3=G.|last4=Chabaud|first4=A.|last5=Beveridge|first5=I.}}</ref>


==Conservation==
==Conservation==

Revision as of 00:51, 23 May 2019

"Fruit bat" redirects here. For other uses, see Fruit bat (disambiguation).
"Fruit-eating bat" redirects here. For other species known as fruit-eating bats, see Leaf-nosed bat.

Megabat
Little red flying foxes, Pteropus scapulatus
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Chiroptera
Superfamily: Pteropodoidea
Family: Pteropodidae
Gray, 1821
Subfamilies
Distribution of megabats
Synonyms

Pteropidae Gray, 1821[1]
Pteropodina C. L. Bonaparte, 1837[1]

Megabats constitute the family Pteropodidae of the order Chiroptera (bats). They are also called fruit bats, Old World fruit bats,[2] or, especially the genera Acerodon and Pteropus, flying foxes. The evolution of megabats has been determined primarily by genetic data, as the fossil record for this family is the worst of all bats. Megabats likely evolved in Australasia, with the common ancestor of all living pteropodids existing approximately 31 million years ago. Many megabat lineages likely originated in Melanesia, then dispersed to mainland Asia, the Mediterranean, and Africa over time. Megabats are found in tropical and subtropical areas of Eurasia, Africa, and Oceania.[3][4][5]

Compared to insectivorous bats, fruit bats are relatively large, and with some exceptions, do not navigate by echolocation. Most species are primarily frugivores and rely on their keen senses of sight and smell to locate food.[6] They reach sexual maturity slowly, and have a low reproductive output. Most species have one offspring at a time after a pregnancy of 4–6 months. This low reproductive output means that, after a population loss, their populations are slow to rebound, making them more susceptible to threats. A quarter of all megabat species are listed as threatened, with key responsible factors as habitat destruction and overhunting. Megabats are a popular food source in some areas, leading to population declines and extinction. While megabats can be a useful food resource, they are also of interest to public health as the natural reservoirs of several viruses than can affect humans.

Taxonomy

The family Pteropodidae was first described in 1821 by British zoologist John Edward Gray. Gray named the family "Pteropidae" (after the genus Pteropus) and placed it within the now-defunct order Fructivorae.[7] However, Gray's spelling was possibly based on a misunderstanding of the suffix of "Pteropus", and was subsequently changed to "Pteropodidae".[8] The Greek word pous of Pteropus is from the stem word pod-; therefore, Latinizing Pteropus correctly results in the prefix "Pteropod-". French biologist Charles Lucien Bonaparte was the first to use the corrected spelling Pteropodidae in 1838.[9] As of 2011, there were 186 species of megabat,[10] around a third of which are flying foxes of the genus Pteropus.[11]

In 1875, Irish zoologist George Edward Dobson was the first to split the order Chiroptera (bats) into two suborders: Megachiroptera (sometimes listed as Macrochiroptera) and Microchiroptera, which are commonly abbreviated to megabats and microbats.[12] Dobson selected these names to allude to the body size differences of these two groups, with many fruit-eating bats being larger than insect-eating bats. Pteropodidae was the only family he included within Megachiroptera.[8][12]

A 2001 study by Springer et al. found that the dichotomy of megabats and microbats did not accurately reflect their evolutionary lineages, however. Instead of Megachiroptera and Microchiroptera, they proposed the new suborders Yinpterochiroptera and Yangochiroptera. Yinpterochiroptera contained species formerly included in Megachiroptera (all of Pteropodidae), as well as several families formerly included in Microchiroptera: Megadermatidae, Rhinolophidae, Nycteridae, Craseonycteridae, and Rhinopomatidae.[13] Two superfamilies comprise Yinpterochiroptera: Rhinolophoidea—containing the above families formerly in Microchiroptera—and Pteropodoidea, which only contains Pteropodidae.[14]

In 1917, Danish mammalogist Knud Andersen divided Pteropodidae into three subfamilies: Macroglossinae, Pteropinae (corrected to Pteropodinae), and Harpyionycterinae.[15] However, a 1995 study found that Macroglossinae as previously defined (Eonycteris, Notopteris, Macroglossus, Syconycteris, Melonycteris, and Megaloglossus) was paraphyletic, meaning that the subfamily did not group all of the descendants of a common ancestor.[16] Subsequent publications consider Macroglossini as a tribe within Pteropodinae that contains only Macroglossus and Syconycteris.[17][10] Eonycteris and Melonycteris are within other tribes in Pteropodinae,[18][10] Megaloglossus was placed in Rousettinae:Myonycterini, and Notopteris is of uncertain placement.[10]

Pteropodidae
Internal relationships of African Pteropodidae based on combined evidence of mitochondrial and nuclear DNA.[19]

Other subfamilies and tribes within Pteropodidae have also undergone changes recently.[10] In 1997, Bergmans et al. classified the pteropodids into 6 subfamilies and 9 tribes based on their morphology, or physical characteristics.[10] A 2011 DNA study concluded that not all of these subfamilies were clades, or consisting of all the descendants of a common ancestor, and therefore they did not accurately depict the relationships between megabat species. Three of Bergmans's subfamilies received support: Cynopterinae, Harpyionycterinae, and Nyctimeninae. The three other clades recovered in this study consisted of Macroglossini, Epomophorinae + Rousettini, and Pteropodini + Melonycteris.[10] A 2016 DNA study focused only on African pteropodids (Harpyionycterinae, Rousettinae, and Epomophorinae) also challenged the 1997 Bergmans classification. All species formerly included in Epomophorinae were moved to Rousettinae, which was subdivided into additional tribes. The genus Eidolon, formerly in the tribe Rousettini of Rousettinae, was moved to its own subfamily, Eidolinae.[18] With these changes, the internal relationships of Pteropodidae are as follows:[10][18]

Megabats of various subfamilies. Clockwise from upper left: Greater short-nosed fruit bat (Cynopterinae), Indian flying fox (Pteropodinae), Egyptian fruit bat (Rousettinae), Eastern tube-nosed bat (Nyctimeninae).
  • Subfamily Pteropodinae
    • Tribe Pteropodini
    • Tribe Macroglossini
    • Tribe Notopterini
  • Subfamily Nyctimeninae
  • Subfamily Harpyionyterinae (expanded to include Boneia)
  • Subfamily Rousettinae (expanded)
    • Tribe Rousettini (revised—now only includes Rousettus; formerly, Rousettini included Eidolon and Eonycteris)
    • Tribe Eonycterini (new tribe)
    • Tribe Scotonycterini
    • Tribe Epomophorini
    • Tribe Stenonycterini (new tribe)
    • Tribe Myonycterini
    • Tribe Plerotini
  • Subfamily Cynopterinae
  • Subfamily Eidolinae (new subfamily)

In 1984, Butler proposed an additional pteropodid subfamily, Propottininae, representing one extinct species described from a fossil discovered in Africa, Propotto leakeyi Simpson, 1967.[20] However, in 2018 the fossils were reexamined and determined to actually represent a lemur.[21]

Description

Spectacled flying fox (Pteropus conspicillatus)

Megabats are so called for their larger weight and size, weighing up to one kilogram (two pounds) with wingspans reaching over one meter (several feet) in length. The tails are short if present, but are often effectively absent. The digestive system is structured to a herbivorous diet, sometimes restricted to soft fruit or nectar, and is shorter than those of the insectivorous microchiropterans. The tendency while resting amongst this group is to hang by the rear limb with the wings cloaking the body and its head facing forward.[22] The length of the digestive system is short for a herbivore, as the fibrous content is mostly separated by the action of the palate, tongue and teeth and then discarded. The ingested plant material is largely nectar, juices and pollen, the small seeds that are passed are often important to the dispersal of the tree species.[22]

Despite the fact that large body size is included as a defining characteristic of megabats, not all species of megabat are larger than microbats. Megabats range in size from 14.2 g (0.50 oz) in the spotted-winged fruit bat (Balionycteris maculata) to 1,075 g (37.9 oz) in the giant golden-crowned flying fox (Acerodon jubatus).[23] Megabats as a whole are often represented by the flying foxes of Pteropus and Acerodon; these species are outliers in their body size, leading some to overestimate the size of megabats.[8] One review showed that 28% of megabat species weigh less than 50 g (1.8 oz).[23]

Skeleton

Skull and dentition

Skull of the black-eared flying fox

The number of teeth a megabat has is dependent on the species, with a range of 24-34 teeth possible. All megabats have two or four each of upper and lower incisors, with the exception Bulmer's fruit bat (Aproteles bulmerae), which completely lacks incisors,[24] and the São Tomé collared fruit bat (Myonycteris brachycephala), which has two upper incisors and three lower incisors.[25] This makes it the only mammal species with an asymmetrical dental formula.[25]

All species have two upper and lower canine teeth. The number of premolars is variable, with four or six each of upper and lower premolars. The first upper and lower molars are always present, meaning that all megabats have at least four molars. The remaining molars may be present, present but reduced, or absent.[24] Megabat molars and premolars are simplified, with a reduction in the cusps and ridges resulting in a more flattened crown.[26]

Like most mammals, megabats are diphyodont, meaning that the young have a set of deciduous teeth (milk teeth) that falls out and is replaced by permanent teeth. For most species, there are 20 deciduous teeth. The deciduous set does not include molars.[26]

Postcrania

Flying fox skeleton

The scapulae of megabats are described as the most primitive of any chiropteran family. Within Pteropus, the scapula is narrow and has a large supraspinatous fossa. Additionally, the humerus is not as modified as in other families, lacking the same enlargement of the greater tuberosity and the pit at the anterior edge of the head. The shaft of the humerus is curved. While other bats only have claws on the thumbs of their forelimbs, most megabats possess a clawed second digit as well;[26] only Eonycteris, Dobsonia, Notopteris, and Neopteryx lack the second claw.[27] The first digit is the shortest, while the third digit is the longest. The second digit is incapable of flexion.[26]

Megabats' hindlimbs have all the same skeletal components as do humans, with the exception of an additional structure called the calcar, which is a cartilage spur arising from the calcaneus. The entire leg is rotated at the hip compared to normal mammal orientation, meaning that the knees face posteriorly. All five of the digits of the foot flex in the direction of the sagittal plane, with no digit capable of flexing in opposition (as seen in the feet of perching birds).[28]

Internal systems

Internal anatomy of the hammer-headed bat, H. monstrosus

Flight is very energetically expensive, requiring several adaptations to the cardiovascular system. During flight, bats can raise their oxygen consumption by twenty times or more for sustained periods; top human athletes can reach a maximum of 15-20 times oxygen consumption for a few minutes at most.[29] Based on the straw-coloured fruit bat (Eidolon helvum) and hammer-headed bat (Hypsignathus monstrosus), megabats have a mean respiratory exchange ratio (carbon dioxide produced:oxygen used) of approximately 0.78. With the two above species as well as the gray-headed flying fox (Pteropus poliocephalus) and the Egyptian fruit bat (Rousettus aegyptiacus), maximum heart rates of the four species in flight varied between 476 beats per minute (P. poliocephalus) and 728 beats per minute (R. aegyptiacus). Maximum number of breaths per minute also varied, with P. poliocephalus with 163 breaths per minute and E. helvum with up to 316 breaths per minute.[30] Additionally, megabats have exceptionally large lung volumes relative to their sizes. While terrestrial mammals such as shrews have a lung volume of 0.03 cm3 per gram of body weight, species such as the Wahlberg's epauletted fruit bat (Epomophorus wahlbergi) have lung volumes 4.3 times greater at 0.13 cm3 of lung volume per gram of body weight.[29]

Megabats have rapid digestive systems, with a gut transit time of half an hour or less.[31] Many megabats have U-shaped stomachs. There is no distinct difference between the small and large intestine, nor a distinct beginning of the rectum. They have very high densities of intestinal microvilli, which creates a large surface area for the absorption of nutrients.[32]

Senses

Sight

The reddish-orange eyes of the Fijian monkey-faced bat (Mirimiri acrodonta)

With very few exceptions, megabats do not echolocate, and therefore rely on sight and smell to navigate.[33] They have large eyes that are positioned at the front of their heads.[34] Their eyes are larger than those of the common ancestor of all bats, with one study suggesting a trend of increasing eye size among pteropodids. A study that examined the eyes of 18 megabat species determined that the common blossom bat had the smallest eyes at a diameter of 5.03 mm (0.198 in), while the largest eyes were those of large flying fox at 12.34 mm (0.486 in) in diameter.[35] Megabat eyes are usually brown, though they can be red or orange as seen in some species of the following genera: Desmalopex, Mirimiri, Pteralopex, and some Pteropus.[36]

At high brightness levels, their visual acuity is poorer than humans'; at low brightness, however, their visual acuity exceeds humans'.[34] One study that examined the eyes of some Rousettus, Epomophorus, Eidolon, and Pteropus species determined that the former three genera possess a tapetum lucidum, while the Pteropus species do not. All species examined had retinae with both rod cells and cone cells, but only the Pteropus species had S-cones, which detect the shortest wavelengths of light (blue and/or ultraviolet); Pteropus bats are dichromatic, or possessing two kinds of cone cells. The other three genera, with their lack of S-cones, are monochromatic, or color blind. All genera had very high densities of rod cells, which corresponds with their nocturnal activity patterns, as increasing numbers of rod cells result in increased sensitivity to light. In Pteropus and Rousettus, measured rod cell densities were 350,000 – 800,000 per square millimeter, equivalent to or exceeding animals such as the house mouse, domestic cat, and domestic rabbit.[33]

Smell

Megabats use smell to find food sources such as fruit and nectar.[37] They have keen senses of smell that rival those of the domestic dog.[38] Tube-nosed fruit bats such as the eastern tube-nosed bat (Nyctimene robinsoni) have stereo olfaction, meaning that they are able to map and follow odor plumes three-dimensionally.[39] Along with most (or perhaps all) other bat species, megabats also use scent for mothers and offspring to recognize each other, as well as for recognition of individuals.[37] In flying foxes, males have enlarged androgen-sensitive sebaceous glands on their shoulders that they use for scent-marking their territories, particularly during the mating season. The secretions of these glands vary by species—of the 65 chemical compounds isolated from the glands of four species, no compound was found in all species.[40] Males also engage in "urine washing", meaning that they coat themselves in their own urine.[40][41]

Taste

Megabats possess the TAS1R2 gene, meaning that they have the ability to detect sweetness in foods. This gene is present among all bats except vampire bats. Like all bats, megabats cannot taste umami, based on the absence of the TAS1R1 gene. Outside of bats, only giant pandas have been shown to also lack this gene.[37] Megabats also possess multiple TAS2R genes, indicating an ability to taste bitterness.[42]

Evolution

Fossil record and divergence times

The fossil record for pteropodid bats is the most incomplete of any bat family. Several factors could explain why so few pteropodid fossils have been discovered: tropical regions where their fossils might be found are undersampled relative to Europe and North America; conditions for fossilization are poor in the tropics, which could lead to fewer fossils overall; and fossils may have been created, but they may have been destroyed by subsequent geological activity.[43] It is estimated that more than 98% of the pteropodid fossil record is missing.[44] However, even without fossils, the age and divergence times of the family can still be estimated by using computational phylogenetics. Pteropodidae split from the superfamily Rhinolophoidea (which contains all the other families of the suborder Yinpterochiroptera) approximately 58 Mya (million years ago).[44] The ancestor of the crown group of Pteropodidae, or all living species, lived approximately 31 Mya.[45]

Biogeography

Melanesia, where many megabat subfamilies likely originated

The family Pteropodidae likely originated in Australasia.[18] The Melanesian Islands, including New Guinea, are a plausible candidate for the origin of most megabat subfamilies, with the exception of Cynopterinae;[18] the cynopterines likely originated on the Sunda Shelf.[45] From these regions, pteropodids were able to colonize other areas, including continental Asia and Africa. Megabats reached Africa in at least four distinct events. The four proposed events are represented by (1) Scotonycteris, (2) Rousettus, (3) Scotonycterini, and (4) the "endemic Africa clade", which includes Stenonycterini, Plerotini, Myonycterini, and Epomophorini as proposed by Almeida et al. 2016. It is unknown when megabats reached Africa, but several tribes (Scotonycterini, Stenonycterini, Plerotini, Myonycterini, and Epomophorini) were present by the Late Miocene. How megabats reached Africa is also unknown. It has been proposed that they could have arrived via the Middle East before it became more arid at the end of the Miocene. Conversely, they could have reached the continent via the Gomphotherium land bridge, which connected Africa/the Arabian Peninsula to Eurasia. The genus Pteropus (flying foxes), which is not found on mainland Africa, is proposed to have dispersed from Melanesia via island hopping across the Indian Ocean,[46] though this is less likely for other megabat genera, which have smaller body size and thus have more limited flight capabilities.[18]

Loss of echolocation

Megabats make up the only family (Pteropodidae) in order Chiroptera that is not capable of laryngeal echolocation. Echolocation and flight evolved early in the lineage of chiropterans. Although echolocation was later lost in family Pteropodidae,[47] bats in the genus Rousettus are capable of primitive echolocation through clicking their tongues,[48] and some species have been shown to create clicks similar to those of echolocating bats using their wings.[49][50]

Both echolocation and flight are energetically expensive processes.[51] The nature of the flight and echolocation mechanism of bats allows for creation of echolocation pulses with minimal energy use.[52] Energetic coupling of these two processes is thought to have allowed for both energetically expensive processes to evolve in bats. The loss of echolocation may be due to the uncoupling of flight and echolocation in megabats.[53] The larger average body size of megabats compared to echolocating bats[54] suggests that a larger body size disrupts the flight-echolocation coupling and made echolocation too energetically expensive to be conserved in megabats.[53]

Biology and ecology

Reproduction and life cycle

Flying fox with offspring

Relative to their sizes, megabats have low reproductive outputs and delayed sexual maturity, with females of most species not giving birth until the age of one or two.[55]: 6  Some megabats appear to be able to breed throughout the year, but the majority of species are likely seasonal breeders.[27] Gestation length is variable. The Fischer's pygmy fruit bat (Haplonycteris fischeri) has the longest gestation length of any bat species, taking up to 11.5 months to carry a pregnancy to term.[56] Shorter gestation lengths are found in the greater short-nosed fruit bat (Cynopterus sphinx) with a period of three months.[57] Among all species, a gestation length of 4-6 months is common. Some species such as the straw-coloured fruit bat have the reproductive adaptation of delayed implantation, meaning that copulation occurs in June or July, but the zygote does not implant into the uterine wall until months later in November. The Fischer's pygmy fruit bat has the adaptation of post-implantation delay, meaning that development of the embryo is suspended for up to eight months after implantation in the uterine wall. This is responsible for its very long pregnancies. The litter size of all megabats is usually one offspring.[55]: 6  Megabats, like all bats, are long-lived relative to their size. Some captive megabats have had lifespans exceeding thirty years.[27]

Diet and foraging

Most megabats are primarily frugivorous, meaning that they mostly consume fruit.[58] Throughout the family, a diverse array of fruit is consumed from nearly 188 plant genera.[59] Some species are also nectarivorous, meaning that they also drink nectar from flowers.[58] Other food resources include leaves, shoots, buds, flowers, pollen, seed pods, sap, cones, bark, and twigs.[60] They are prodigious eaters, and can consume up to 2.5 times their own body weight in fruit per night.[59]

Predators and parasites

Megabats have few native predators, especially those living on islands: species like the small flying fox (Pteropus hypomelanus) have no known natural predators.[61] Non-native predators of flying foxes include domestic cats and rats. The mangrove monitor, which is a native predator for some megabat species but an introduced predator for others, opportunistically preys on megabats, as it is a capable tree climber.[62] Another invasive species, the brown tree snake can seriously impact megabat populations; the brown tree snake consumes so many offspring that it reduced the recruitment of the Guam population of the Mariana fruit bat (Pteropus mariannus) to essentially zero.[63] Native predators of megabats include crocodilians. The saltwater crocodile, a native species, is a known predator of megabats, based on analysis of crocodile stomach contents in northern Australia.[64] During extreme heat events, megabats like the little red flying fox (Pteropus scapulatus) must cool off and rehydrate by drinking from waterways, making them susceptible to opportunistic depredation by freshwater crocodiles.[65]

Megabats are the hosts of several parasite taxa. Known parasites include Nycteribiidae and Streblidae species ("bat flies").[66][67] Blood parasites of the family Haemoproteidae also affect megabat species.[68]

Conservation

Status

The small Mauritian flying fox (Pteropus subniger), which was driven to extinction by overhunting[69]

As of 2014, the IUCN evaluated a quarter of all megabat species as threatened with endangerment, which includes species listed as critically endangered, endangered, and vulnerable. Megabats are substantially threatened by humans, as they are hunted for food and medicinal uses. Additionally, they are culled for actual or perceived damage to agriculture, especially to fruit production.[70] As of 2019, the IUCN has evaluations for 187 megabat species. The status breakdown is as follows:[71]

  • Extinct: 4 species (2.1%)
  • Critically endangered: 8 species (4.3%)
  • Endangered: 16 species (8.6%)
  • Vulnerable: 37 species (19.8%)
  • Near-threatened: 13 species (7.0%)
  • Least-concern: 89 species (47.6%)
  • Data deficient: 20 species (10.7%)

Factors causing decline

Anthropogenic sources

An electrocuted megabat on an electrical wire in Australia

Megabats are threatened by habitat destruction by humans, which takes several forms. Deforestation of their habitats has resulted in the loss of critical roosting habitat. Deforestation also results in the loss of food resource, as native fruit-bearing trees are felled. Habitat loss and resulting urbanization leads to construction of new roadways, making megabat colonies easier to access for overharvesting. Additionally, habitat loss via deforestation compounds natural threats, as fragmented forests are more susceptible to damage from typhoon-force winds.[55]: 7  Cave-roosting megabats are threatened by human disturbance at their roost sites. Guano mining is a livelihood in some countries within their range, bringing people to caves. Caves are also disturbed via mineral mining and cave tourism.[55]: 8 

Megabats are also intentionally killed by humans for several reasons. A large source of mortality is by human persecution stemming from agricultural conflicts. Though some megabats have been documented to have a preference for native fruit trees over fruit crops, deforestation can reduce the megabat food supply, causing them to rely on fruit crops.[55]: 8  They are shot, beaten to death, or poisoned to reduce their populations. Mortality also occurs via accidental entanglement into netting used to prevent the bats from eating fruit.[72] Culling can dramatically reduce megabat populations. In Mauritius, over 40,000 Mauritian flying foxes were culled in a two-year period, reducing its population by an estimated 45%.[73] Megabats 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.[74] Farmers construct electrified grids over their fruit trees to kill megabats before they can consume their crop. The grids are questionably effective at preventing crop loss, with one farmer who operated such a grid estimating that they still lost 100–120 tonnes (220,000–260,000 lb) of fruit to flying foxes in a year.[75] Some electrocution deaths are also accidental, such as when bats fly into overhead power lines.[76]

Climate change causes flying fox mortality and a source of concern for species persistence. Extreme heat waves in Australia have been responsible for the deaths of more than 30,000 flying foxes from 1994 to 2008. Females and young bats are most susceptible to extreme heat, which affects a population's ability to recover.[77] Megabats are threatened by sea level rise associated with climate change, as several species are endemic to low-lying atolls.[78]

Natural sources

The Anatahan volcano eruption of 2003

Because many species are endemic to a single island, they are vulnerable to random events such as typhoons. A 1979 typhoon halved the remaining population of the Rodrigues flying fox. Typhoons result in indirect mortality as well: Because they defoliate the trees, megabats are more visible and easily hunted by humans. Food resources for the bats become scarce after major storms, and megabats resort to riskier foraging strategies such as consuming fallen fruit off the ground. There, they are more vulnerable to depredation by domestic cats, dogs, and pigs.[69] As many megabat species are located in the tectonically active Ring of Fire, they are also threatened by volcanic eruptions. Flying foxes have been nearly exterminated from the island of Anatahan following a series of eruptions beginning in 2003.[79]

Relationship to people

Food

Main articles: Bat as food and Pteropus § Food

Megabats are killed and consumed as bushmeat throughout their range. Bats are consumed extensively throughout Asia, as well as in islands of the West Indian Ocean and the Pacific, where Pteropus species are heavily hunted. In continental Africa, where no Pteropus species live, its largest megabat, the straw-coloured fruit bat, is a preferred hunting target.[80]

In Guam, consumption of the Mariana fruit bat exposes locals to the neurotoxin beta-Methylamino-L-alanine (BMAA) which may later lead to neurodegenerative diseases. BMAA may become biomagnified in humans who consume flying foxes; flying foxes are exposed to BMAA by eating cycad fruits.[81][82][83]

As disease reservoirs

The Egyptian fruit bat (Rousettus aegyptiacus), which has tested positive for Marburg virus and antibodies against the Ebola virus (though not the virus itself)
Henipavirus outbreaks overlaid on flying fox distribution map, with Nipah virus as blue icons and Hendra virus as red icons.

Megabats are the reservoirs of several viruses that can affect humans and cause disease. They can carry filoviruses, including the Ebola virus (EBOV) and Marburgvirus. Species that have tested positive for the presence of EBOV include Franquet's epauletted fruit bat, the hammer-headed fruit bat, and the little collared fruit bat. Additionally, antibodies against EBOV have been found in the straw-coloured fruit bat, Gambian epauletted fruit bat, Peters's dwarf epauletted fruit bat, Veldkamp's dwarf epauletted fruit bat, Leschenault's rousette, and the Egyptian fruit bat. Marburgvirus has been confirmed in one species, the Egyptian fruit bat.[84][85][86]

Other megabats implicated as disease reservoirs are primarily Pteropus species. Notably, flying foxes can transmit lyssaviruses, which cause rabies. In Australia the rabies virus is not naturally present; Australian bat lyssavirus is the only lyssavirus present. Australian bat lyssavirus was first identified in 1996; it is very rarely transmitted to humans. Transmission occurs from the bite or scratch of an infected animal, but can also occur from getting the infected animal's saliva in a mucous membrane or an open wound. Exposure to flying fox blood, urine, or feces is not a risk of exposure to Australian bat lyssavirus. Since 1994, there have been three records of people getting infected with it—all three were in Queensland and each case was fatal.[87]

Flying foxes are also reservoirs of henipaviruses such as Hendra virus and Nipah virus. Hendra virus was first identified in 1994; it also rarely occurs humans. From 1994 to 2013, there have been seven reported cases of Hendra virus affecting people, four of which were fatal. The hypothesized primary route of human infection is via contact with horses that have come into contact with flying fox urine.[88] There are no documented instances of direct transmission between flying foxes and humans.[89] As of 2012, there is a vaccine available for horses to decrease the likelihood of infection and transmission.[90]

Nipah virus was first identified in 1998 in Malaysia. Since 1998, there have been several Nipah outbreaks in Malaysia, Singapore, India, and Bangladesh, resulting in over 100 casualties. A 2018 outbreak in Kerala, India resulted in 19 humans infected, of which 17 died.[91] The overall fatality rate is 40-75%. Humans can contract Nipah virus from direct contact with flying foxes or their fluids, through exposure to an intermediate host such as domestic pigs, or from contact with an infected person.[92] A 2014 study of the Indian flying fox and Nipah virus found that while Nipah virus outbreaks are more likely in areas preferred by flying foxes, "the presence of bats in and of itself is not considered a risk factor for Nipah virus infection." Rather, the consumption of date palm sap is a significant route of transmission. The practice of date palm sap collection involves placing collecting pots at date palm trees. Indian flying foxes have been observed licking the sap as it flows into the pots, as well as defecating and urinating in proximity to the pots. In this way, humans who drink the palm sap can be exposed to the bats' viruses. The use of bamboo skirts on collecting pots lowers the risk of contamination from bat fluids.[93]

Flying foxes can transmit several non-lethal diseases as well, such as Menangle virus[94] and Nelson Bay virus.[95] These viruses rarely affect humans and few cases have been reported.[94][95] While other bat species have been suspected or implicated as the reservoir of diseases such as SARS and Ebola, flying foxes are not suspected as hosts for either causative virus.[96]

List of genera

Main article: List of fruit bats
The spotted-winged fruit bat (Balionycteris maculata)
The straw-coloured fruit bat (Eidolon helvum)
Long-tongued nectar bat (Macroglosus minimus)
Wahlberg's epauletted fruit bat (Epomophorus wahlbergi)

The family Pteropodidae is divided into seven subfamilies represented by 44–46 genera:

Family Pteropodidae

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