ABSTRACT The fossil Afrillonura namibensis gen. et sp. nov. from the late Middle Miocene (12–13 Ma) site of Berg Aukas I, Namibia was a medium-sized emballonurid with a gracile dentition. It shows a mosaic combination of dental features of extant Emballonurini and distinct plesiomorphies suggesting its basal phylogenetic position close to the root of the extant clades of the tribe. It shares many odontological characteristics with Coleura and Paremballonura but also with some species of Emballonura. Afrillonura namibensis gen. et sp. nov. differs from most members of the tribe Emballonurini particularly in the presence of a vestigial paracrista on M1, resembling the state in the Palaeogene genus Dhofarella. Together with other available fossils, the new material from Namibia supports the hypothesis of the African origin of the family Emballonuridae. Further phylogenetic conclusions and biogeographic scenarios of emballonurid ancestral distribution are discussed.