Morphology and phylogenetic relationships of the earliest known hippopotamids (Cetartiodactyla, Hippopotamidae, Kenyapotaminae)

The earliest known hippopotamids, attributed to the subfamily Kenyapotaminae, are known essentially from dental remains of two species. The first was found in the middle Miocene of Kenya, the second at the beginning of the upper Miocene in eastern and northern Africa. The exact affinities of the Kenyapotaminae are critical for resolving the long debated origin of the Hippopotamidae, as part of the wider question of cetacean affinities within artiodactyls. We performed the first detailed comparative description, character by character, of kenyapotamines, using the following putatively related taxa: Hippopotaminae, Suoidea, and Anthracotheriidae. The development of an improved nomenclature for the dentition facilitates comparisons amongst a wide array of cetartiodactyls. This has permitted the first assessment of the phylogenetic position of Kenyapotaminae using a cladistic analysis. This work provides an emendation of kenyapotamine taxonomy and diversity. Kenyapotaminae are indeed closely related to Hippopotaminae and should be kept within Hippopotamidae, the late Miocene kenyapotamines not necessarily being the forerunners of the first known hippopotamines. Hippopotamidae are deeply nested within anthracotheriids, with crown bothiodontines as sister group, and any close affinities with Suoidea should be rejected. This reinforces a scenario linking cetaceans to Hippopotamoidea (Hippopotamidae + Anthracotheriidae), possibly via other early Palaeogene artiodactyls.