Extending the footprint record of Pareiasauromorpha to the Cisuralian: earlier appearance and wider palaeobiogeography of the group

Pareiasauromorpha is one of the most important tetrapod groups of the Permian. Skeletal evidence suggests a late Kungurian origin in North America, whereas the majority of occurrences come from the Guadalupian and Lopingian of South Africa and Russia. However, Pareiasauromorpha footprints include the ichnogenus Pachypes, which is unknown from strata older than late Guadalupian. A revision of several Pachypes-like footprints from the Cisuralian–Guadalupian of Europe and North America confirm the occurrence of this ichnogenus and of the ichnospecies Pachypes ollieri comb. nov. beginning in the Artinskian. This is the earliest known occurrence of Pachypes and it coincides with the Artinskian reptile radiation. Based on a synapomorphy-based track–trackmaker correlation, P. ollieri can be attributed to nycteroleter pareiasauromorphs such as Macroleter. Therefore, the earliest occurrences of pareiasauromorph footprints precede by at least 10 myr the earliest occurrence of this group in the skeletal record. Moreover, the palaeobiogeography of the group is extended to the Cisuralian and Guadalupian of western Europe.