Cranial morphology and phylogenetic relationships of the Middle Permian pareiasaur Embrithosaurus schwarzi from the Karoo Basin of South Africa

Abstract

Pareiasaurs were globally distributed, abundant, herbivorous parareptiles of the Middle to Late Permian, with the basal-most members found in the Middle Permian of South Africa. These basal taxa were particularly abundant and went extinct at the end of the Gaudalupian (Capitanian) at the top of the Tapinocephalus Assemblage Zone. Currently four taxa are recognized in this group: Bradysaurus seeleyi, B. baini, Nochelesaurus alexanderi and Embrithosaurus schwarzi, but they are all poorly understood. We here present the first detailed cranial description and updated diagnosis for Embrithosaurus schwarzi. No cranial autapomorphies were identified. However, Embrithosaurus schwarzi is a distinct taxon in this group, based on its unique dentition and using a combination of cranial features. It has nine marginal cusps on all maxillary and mandibular teeth, and wider maxillary teeth than in the co-occurring taxa, due to the marginal cusps being arranged more regularly around the crown, and the apex of the crown lacking the long, central, three-cusped trident. Our updated phylogenetic analysis recovers the four Middle Permian South African taxa as a monophyletic group for the first time, which we call Bradysauria, comprising a clade including Embrithosaurus, Bradysaurus baini and a polytomy including Nochelesaurus and Bradysaurus seeleyi.