Cranial morphology of the middle Permian pareiasaur Nochelesaurus alexanderi from the Karoo Basin of South Africa

ABSTRACT Pareiasaurs were globally distributed, abundant, herbivorous parareptiles with the basal-most members found only in the mid-Permian of South Africa. These basal forms form a monophyletic group and were locally abundant and became extinct at the top of the Tapinocephalus Assemblage Zone at the end of the Guadalupian. Four species of basal pareiasaurs are currently recognised: Bradysaurus baini, B. seeleyi, Embrithosaurus schwarzi and Nochelesaurus alexanderi, but they are all poorly understood and there remains historic uncertainty as to their validity. In this paper, our second contribution designed to improve understanding of the basal group, we present the first detailed cranial description and updated diagnosis for Nochelesaurus alexanderi and demonstrate that it is a distinct taxon based on one cranial autapomorphy, a large transversely wide postparietal, and a combination of cranial characters. Within the local group of mid-Permian pareiasaurs, we recognise new dental features of Nochelesaurus alexanderi: non-symmetrical marginal cusp arrangements on upper and lower teeth resulting from an extra basal mesial cusp; an incipient horizontal cingulum on lower jaw teeth, sometimes with one or two tiny medial cingular cusps; and up to ten marginal cusps. Our study demonstrates that tooth morphology and orientation, cranial ornamentation, morphology of the cheek bosses, shape of the postfrontal and postparietal, and morphology of the distal paroccipital process of the opisthotic are the most useful to identify South African mid-Permian pareiasaurs.