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. However, Gray's spelling was possibly based on a misunderstanding of the suffix of "Pteropus" and was subsequently changed to "Pteropodidae". "Pteropus" comes from Ancient Greek "pterón" meaning "wing" and "poús" meaning "foot". 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. As of 2011, there were 186 described species of megabat, around a third of which are flying foxes of the genus Pteropus.
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. 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.
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. This classification scheme has been verified several times subsequently and remains widely supported as of 2019.
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. Two superfamilies comprise Yinpterochiroptera: Rhinolophoidea—containing the above families formerly in Microchiroptera—and Pteropodoidea, which only contains Pteropodidae.
In 1917, Danish mammalogist Knud Andersen divided Pteropodidae into three subfamilies: Macroglossinae, Pteropinae (corrected to Pteropodinae), and Harpyionycterinae. 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. Subsequent publications consider Macroglossini as a tribe within Pteropodinae that contains only Macroglossus and Syconycteris. Eonycteris and Melonycteris are within other tribes in Pteropodinae, Megaloglossus was placed in the tribe Myonycterini of the subfamily Rousettinae, and Notopteris is of uncertain placement.
Other subfamilies and tribes within Pteropodidae have also undergone changes recently. In 1997, the pteropodids were classified into six subfamilies and nine tribes based on their morphology or physical characteristics. 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. 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. With these changes, the internal relationships of Pteropodidae are as follows:
- 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. However, in 2018 the fossils were reexamined and determined to represent a lemur.
List of genera
The family Pteropodidae is divided into seven subfamilies represented by 44–46 genera:
- genus Eidolon – straw-coloured fruit bats
- subfamily Pteropodinae