Permian-Triassic vertebrate footprints from South Africa: Ichnotaxonomy, producers and biostratigraphy through two major faunal crises

The fossil record of mid to late Permian terrestrial vertebrates in the South African Karoo Basin is regarded as the most abundant and diverse in the world. Despite the extensive research on body fossils, to-date the vertebrate footprint sites have not been subjected to an anatomy-consistent ichnotaxonomic investigation. Here we present a comprehensive ichnotaxonomic revision of Permian-Early Triassic tracksites in the main Karoo Basin of South Africa. Furthermore, a track-trackmaker correlation for all Permian synapsid groups is provided for the first time, based on the analysis of the functional morphology of potential producers. The following ichnotaxa and their proposed trackmakers are recognized: Brontopus giganteus (dinocephalians), cf. Capitosauroides isp. ( therocephalians), cf. Dicynodontipus isp. (cynodonts), Dolomitipes accordii (small dicynodonts), Dolomitipes icelsi n. comb. (large dicynodonts), Karoopes gansfonteinensis n. igen. n. isp. (gorgonopsids), Procolophonichnium nopcsai (procolophonids) and Rhynchosauroides isp. (non-archosauriform diapsids). Three different footprint assemblages (FA I-III) are proposed for footprint biostratigraphy: FA I (lower Tapinocephalus AZ), a Guadalupian assemblage dominated by dinocephalian tracks; FA II (topmost Tapinocephalus-Cistecephalus AZ), a latest Guadalupian-Wuchiapingian assemblage dominated by gorgonopsid and dicynodont tracks in association with subordinate therocephalian tracks and FA Ill (lower Lystrasaurus AZ), an Induan assemblage with dicynodont, cynodont, procolophonid and diapsid tracks. The lower FA II includes the earliest ichnofauna with Lopingian affinity all over the world (topmost Tapinocephalus-Pristemgnathus AZ, similar to 260-259 Ma) and could indicate an early recovery phase after the end-Guadalupian mass extinction, because of the high abundance of large gorgonopsid tracks and absence of dinocephalian tracks. Thi s footprint record may also predate the body fossil record, suggesting an earlier gorgonopsid radiation. FA III represents the earliest and most complete post end-Permian extinction ichnofauna, which includes an early phase of abundant small dicynodont tracks, potentially indicating a stressed post-event community. Nevertheless, this ichnofauna looks very similar to pre-extinction ichnofaunas from Europe, in agreement with the skeletal record at the Daptocephalus-Lystrosaurus AZ transition. (C) 2019 International Association for Gondwana Research. Published by Elsevier B.V. All rights reserved.