New dicynodonts (Therapsida, Anomodontia) and updated tetrapod stratigraphy of the Permian Ruhuhu Formation (Songea Group, Ruhuhu Basin) of southern Tanzania

Permian tetrapod fossils were discovered in the Tanzanian Ruhuhu Formation in 1963, but they have received far less attention than the tetrapods of the overlying Usili (formerly Kawinga) Formation. Here, we describe two dicynodonts collected in the Ruhuhu Formation in 2008. Abajudon kaayai, gen. et sp. nov., is represented by a partial skull and mandible and is characterized by autapomorphic upper teeth that are triangular in cross-section, have procurved tips, and bear a deep groove on the mesial surface. Although it shows similarities to taxa such as Endothiodon and Chelydontops, the exact relationships between A. kaayai and other dicynodonts are uncertain. The second specimen also consists of a partial skull and mandible. We refer it to cf. Endothiodontia based on the medial placement of the long maxillary tooth rows, the presence of depressions on the palatine pad, a long posterior dentary sulcus, and similarities of the mandibular dentition. Tetrapods occur in three fossiliferous horizons in the Ruhuhu Formation. The likely Middle Permian lower horizon includes dinocephalians, temnospondyls, and the dicynodonts described here. The middle horizon includes a new, tusked species of Endothiodon and at least one other dicynodont. The upper horizon appears to sample an assemblage similar to that of the Usili Formation and therefore may be of Late Permian age. The discovery of Middle Permian fossils in Tanzania and Zambia provides the opportunity to test whether southern Gondwana was characterized by a cosmopolitan tetrapod fauna for an extended period of time before the biogeographic restructuring caused by the end-Permian mass extinction.