Megabat

Megabat
The image depicts a group of large bats hanging from a tree
A colony of little red flying foxes, Pteropus scapulatus
Scientific classification e
Kingdom:Animalia
Phylum:Chordata
Class:Mammalia
Order:Chiroptera
Superfamily:Pteropodoidea
Family:Pteropodidae
Gray, 1821
Subfamilies
Worldwide distribution of Pteropodidae.jpg
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, or—especially the genera Acerodon and Pteropusflying foxes. The evolution of megabats has been determined primarily by genetic data, as the fossil record for this family is the most fragmented 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 over time to mainland Asia, the Mediterranean, and Africa. Megabats are found in tropical and subtropical areas of Eurasia, Africa, and Oceania.

Compared to insectivorous bats, megabats 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. They reach sexual maturity slowly and have a low reproductive output. Most species have one offspring at a time after a pregnancy of four to six months. This low reproductive output means that after a population loss their numbers are slow to rebound, making them more susceptible to threats. A quarter of all megabat species are listed as threatened, with habitat destruction and overhunting key factors responsible for this. 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 those involved in public health; they are the natural reservoirs of several viruses that can affect humans.

Taxonomy

The family Pteropodidae was first described in 1821 by British zoologist John Edward Gray. He named the family "Pteropidae" (after the genus Pteropus) and placed it within the now-defunct order Fructivorae.[2] However, Gray's spelling was possibly based on a misunderstanding of the suffix of "Pteropus" and was subsequently changed to "Pteropodidae".[3] "Pteropus" comes from Ancient Greek "pterón" meaning "wing" and "poús" meaning "foot".[4] 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.[5] As of 2011, there were 186 described species of megabat,[6] around a third of which are flying foxes of the genus Pteropus.[7]

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.[8] Dobson selected these names to allude to the body size differences of the two groups, with many fruit-eating bats being larger than insect-eating bats. Pteropodidae was the only family he included within Megachiroptera.[3][8]

A 2001 study, however, found that the dichotomy of megabats and microbats did not accurately reflect their evolutionary relationships. Instead of Megachiroptera and Microchiroptera, they proposed the new suborders Yinpterochiroptera and Yangochiroptera.[9] This classification scheme has been verified several times subsequently and remains widely supported as of 2019.[10][11][12][13] 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.[9] 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 the descendants of a common ancestor.[16] Subsequent publications consider Macroglossini as a tribe within Pteropodinae that contains only Macroglossus and Syconycteris.[17][6] Eonycteris and Melonycteris are within other tribes in Pteropodinae,[18][6] Megaloglossus was placed in the tribe Myonycterini of the subfamily Rousettinae, and Notopteris is of uncertain placement.[6]

Pteropodidae

Pteropodinae

Nyctimeninae

Cynopterinae

Eidolinae

Rousettinae

Scotonycterini

Eonycterini

Rousettini

Stenonycterini

Plerotini

Myonycterini

Epomophorini

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.[6] In 1997, the pteropodids were classified into six subfamilies and nine tribes based on their morphology or physical characteristics.[6] A 2011 DNA study concluded that not all of these subfamilies were clades, or consisted of all the descendants of a common ancestor, and therefore they did not accurately depict the relationships between megabat species. Three of the subfamilies proposed in 1997 based on morphology received support: Cynopterinae, Harpyionycterinae, and Nyctimeninae. The other three clades recovered in this study consisted of Macroglossini, Epomophorinae + Rousettini, and Pteropodini + Melonycteris.[6] 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:[6][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, an additional pteropodid subfamily, Propottininae, was proposed, 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 represent a lemur.[21]

List of genera

A small brown bat with black wings is hanging upside down on a tree branch. Its wings have small, pinkish spots.
The spotted-winged fruit bat (Balionycteris maculata)
A bat with large eyes and a dog-like face in profile. Its fur is a tawny yellow, while the side of its neck is bright yellow.
The straw-coloured fruit bat (Eidolon helvum)
A small bat clings upside down to a rope. It has brown fur and a narrow snout.
Long-tongued nectar bat (Macroglosus minimus)
A bat with its wings wrapped around its body. Its eyes are tawny brown and prominent, and the sun shines through its ear membranes.
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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