Epomophorinae + Rousettini, and Pteropodini + Melonycteris. A 2016 genetic study focused only on African pteropodids (Harpyionycterinae, Rousettinae, and Epomophorinae) also challenged the 1997 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.

In 1984, an additional