A tale of two Tweefonteins: What physical correlation, geochronology, magnetic polarity stratigraphy, and palynology reveal about the end-Permian terrestrial extinction paradigm in South Africa

The contact between the Daptocephalus to Lystrosaurus declivis (previously Lystrosaurus) Assemblage Zones (AZs) described from continental deposits of the Karoo Basin was commonly interpreted to represent an extinction crisis associated with the end-Permian mass-extinction event at ca. 251.901 +/- 0.024 Ma. This terrestrial extinction model is based on several sections in the Eastern Cape and Free State Provinces of South Africa. Here, new stratigraphic and paleontologic data are presented for the Eastern Cape Province, in geochronologic and magnetostratigraphic context, wherein lithologic and biologic changes are assessed over a physically correlated stratigraphy exceeding 4.5 km in distance. Spatial variation in lithofacies demonstrates the gradational nature of lithostratigraphic boundaries and depositional trends. This pattern is mimicked by the distribution of vertebrates assigned to the Daptocephalus and L. declivis AZs where diagnostic taxa of each co-occur as lateral equivalents in landscapes dominated by a Glossopteris flora. High-precision U-Pb zircon (chemical abrasion-isotope dilution-thermal ionization mass spectrometry) age results indicate maximum Changhsingian depositional dates that can be used as approximate tie points in our stratigraphic framework, which is supported by a magnetic polarity stratigraphy. The coeval nature of diagnostic pre- and post-extinction vertebrate taxa demonstrates that the L. declivis AZ did not replace the Daptocephalus AZ stratigraphically, that a biotic crisis and turnover likely is absent, and a reevaluation is required for the utilization of these biozones here and globally. Based on our data set, we propose a multidisciplinary approach to correlate the classic Upper Permian localities of the Eastern Cape Province with the Free State Province localities, which demonstrates their time-transgressive nature.