The rich fossil vertebrate record from the Beaufort Group, Main Karoo Basin, provides a global standard for mid‐Permian to Mid‐Triassic continental faunas. However, recent studies have demonstrated variability in the composition of contemporaneous faunas across Gondwana. This raises the question of how much the vertebrate faunas differ within the Karoo, where the taxonomic composition of vertebrate assemblage zones (AZs) is mostly considered to be uniform. Although fossil material is known from across the outcrop of the Beaufort Group, the lowest Beaufort strata have received little attention, particularly north of S31°10′. Here, we report two fossil tetrapod assemblages from the lowest Beaufort Group in the southern Free State Province, which represent the northernmost point at which the lowest Beaufort has been targeted for collecting. The lower assemblage is characterized by an abundance of the small dicynodont Eosimops and can thus be attributed to the Tapinocephalus AZ (Guadalupian), but the absence of dinocephalian or pareiasaurian material is unlike contemporaneous assemblages found further south. This suggests that the Tapinocephalus AZ was not uniform across the entire basin and highlights that the abundance, distribution and taxonomic composition of Karoo biozones may vary more than currently appreciated. The upper assemblage, characterized by the dicynodonts Oudenodon, Aulacephalodon and Dinanomodon, is attributable to the upper Cistecephalus AZ to lower Daptocephalus AZ. The juxtaposition of the lower Tapinocephalus AZ and upper Cistecephalus\lower Daptocephalus AZ in the southern Free State implies a stratigraphic gap from the Middle to Late Permian of up to 6 million years.
Pareiasaurs were relatively abundant, globally distributed, herbivorous parareptiles of the mid to late Permian. The basal-most forms, all members of the Bradysauria, are restricted to the Guadalupian (mid-Permian) of South Africa and went extinct in the late Capitanian near the top of the Tapinocephalus Assemblage Zone. Currently four species are recognised in 2 this group: Bradysaurus seeleyi, B. baini, Embrithosaurus schwarzi and Nochelesaurus alexanderi. Those taxa have historically been poorly defined and based on a limited number of specimens, leaving the taxonomic diversity of the group open to doubt and limiting their utility in biostratigraphy. Here we present our fourth and final contribution designed to improve the understanding of this group of pareiasaurs by providing a taxonomic and phylogenetic review, updated stratigraphic ranges and updated diagnoses for each taxon of the Bradysauria. Bradysaurus seeleyi is synonymised with Bradysaurus baini, resulting in three valid mid-Permian pareiasaur taxa: Bradysaurus baini, Embrithosaurus schwarzi and Nochelesaurus alexanderi. Our cladistic analysis of cranial and postcranial characters supports the monophyly of Bradysauria with five synapomorphies. Embrithosaurus schwarzi is recovered as the sister taxon to a clade containing Bradysaurus baini and Nochelesaurus alexanderi. By identifying 157 pareiasaur specimens in fossil collections we show that the Bradysauria are stratigraphically restricted to the Abrahamskraal Formation of the Beaufort Group and suggest a staggered appearance. Bradysaurus baini is first to appear, followed by Nochelesaurus alexanderi, and lastly by Embrithosaurus schwarzi. All three taxa perished during the Capitanian mass extinction, and have their highest occurrences near the top of the Abrahamskraal Formation.
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