Michael O. Day

When and how did the terrestrial mid-Permian mass extinction occur? Evidence from the tetrapod record of the Karoo Basin, South Africa

A mid-Permian (Guadalupian epoch) extinction event at approximately 260 Ma has been mooted for two decades. This is based primarily on invertebrate biostratigraphy of Guadalupian–Lopingian marine carbonate platforms in southern China, which are temporally constrained by correlation to the associated Emeishan Large Igneous Province (LIP). Despite attempts to identify a similar biodiversity crisis in the terrestrial realm, the low resolution of mid-Permian tetrapod biostratigraphy and a lack of robust geochronological constraints have until now hampered both the correlation and quantification of terrestrial extinctions. Here we present an extensive compilation of tetrapod-stratigraphic data analysed by the constrained optimization (CONOP) algorithm that reveals a significant extinction event among tetrapods within the lower Beaufort Group of the Karoo Basin, South Africa, in the latest Capitanian. Our fossil dataset reveals a 74–80% loss of generic richness between the upper Tapinocephalus Assemblage Zone (AZ) and the mid-Pristerognathus AZ that is temporally constrained by a U–Pb zircon date (CA-TIMS method) of 260.259 ± 0.081 Ma from a tuff near the top of the Tapinocephalus AZ. This strengthens the biochronology of the Permian Beaufort Group and supports the existence of a mid-Permian mass extinction event on land near the end of the Guadalupian. Our results permit a temporal association between the extinction of dinocephalian therapsids and the LIP volcanism at Emeishan, as well as the marine end-Guadalupian extinctions.

A reevaluation of Brachyprosopus broomi and Chelydontops altidentalis, dicynodonts (Therapsida, Anomodontia) from the middle Permian Tapinocephalus Assemblage Zone of the Karoo Basin, South Africa

ABSTRACT Brachyprosopus broomi was described in 1937 based on a specimen from the Tapinocephalus Assemblage Zone (Karoo Basin, South Africa), but it was largely overlooked by subsequent workers. We have identified several new specimens that show that Brachyprosopus is a valid taxon. An autapomorphy for the taxon is a curled lateral edge of the squamosal that forms a lateral wall of the external adductor fossa. Other important characters are absence of anterior median palatal ridges; maxillary tooth rows bounded laterally by a shelf; unfused vomers; raised margins of the interpterygoid vacuity; broad intertemporal region; pineal boss; dentary tables; and a long, wide posterior dentary sulcus that extends posterior to the dentary teeth. Chelydontops altidentalis is a junior synonym of B. broomi. A phylogenetic analysis places Brachyprosopus among basal dicynodonts, not as a close relative of Endothiodon. It is noteworthy that some characters, such as well-developed medial maxillary tooth rows and the shape of the palatines, are shared by Brachyprosopus, Pristerodon, Endothiodon, and Niassodon, hinting that a final resolution of relationships among toothed dicynodonts remains to be achieved. Most Brachyprosopus specimens are from the upper Abrahamskraal Formation (Moordenaars and Karelskraal members). One specimen is from low in the Koonap Formation, just above the lowermost maroon mudrocks of the Beaufort Group, and an exact correlation between this level and the strata of the Abrahamskraal Formation is uncertain. Therefore, it is likely that the stratigraphic range of Brachyprosopus extends throughout the Tapinocephalus zone and possibly down into the Eodicynodon Assemblage Zone.

Olson's Extinction and the latitudinal biodiversity gradient of tetrapods in the Permian

The terrestrial vertebrate fauna underwent a substantial change in composition between the lower and middle Permian. The lower Permian fauna was characterized by diverse and abundant amphibians and pelycosaurian-grade synapsids. During the middle Permian, a therapsid-dominated fauna, containing a diverse array of parareptiles and a considerably reduced richness of amphibians, replaced this. However, it is debated whether the transition is a genuine event, accompanied by a mass extinction, or whether it is merely an artefact of the shift in sampling from the palaeoequatorial latitudes to the palaeotemperate latitudes. Here we use an up-to-date biostratigraphy and incorporate recent discoveries to thoroughly review the Permian tetrapod fossil record. We suggest that the faunal transition represents a genuine event; the lower Permian temperate faunas are more similar to lower Permian equatorial faunas than middle Permian temperate faunas. The transition was not consistent across latitudes; the turnover occurred more rapidly in Russia, but was delayed in North America. The argument that the mass extinction is an artefact of a latitudinal biodiversity gradient and a shift in sampling localities is rejected: sampling correction demonstrates an inverse latitudinal biodiversity gradient was prevalent during the Permian, with peak diversity in the temperate latitudes.

Advances in Nonmarine Karoo Biostratigraphy: Significance for Understanding Basin Development

The nonmarine Permo-Jurassic deposits of the Karoo Supergroup of South Africa have long been a world standard for tetrapod biostratigraphy. Recent and ongoing research is revising the palaeoflora and palaeofauna of these sedimentary strata with an unprecedented level of stratigraphic precision. This work has shown that: Permian palynomorphs are useful for correlating time-equivalent lithostratigraphic units in different sectors of the basin; that there is a marked end-Guadalupian diversity drop in tetrapods; that the Dicynodon Assemblage Zone can be subdivided, and should be renamed as the Daptocephalus Assemblage Zone; that the Cynognathus Assemblage Zone has a robust threefold subdivision; and that the name Euskelosaurus for the Euskelosaurus Range Zone is invalid and should be replaced. This work, together with new radiometric dates from the Karoo Supergroup, has dramatically enhanced our understanding of the timing of major evolutionary events in terrestrial ecosystems and provides strong evidence for tectonic controls on accommodation and sedimentation in the Karoo Basin during the Permo-Jurassic, within an overall flexural basinal setting.

A New Mid-Permian Burnetiamorph Therapsid from the Main Karoo Basin of South Africa and a Phylogenetic Review of Burnetiamorpha

Discoveries of burnetiamorph therapsids in the last decade and a half have increased their known diversity but they remain a minor constituent of middle—late Permian tetrapod faunas. In the Main Karoo Basin of South Africa, from where the clade is traditionally best known, specimens have been reported from all of the Permian biozones except the Eodicynodon and Pristerognathus assemblage zones. Although the addition of new taxa has provided more evidence for burnetiamorph synapomorphies, phylogenetic hypotheses for the clade remain incongruent with their appearances in the stratigraphic column. Here we describe a new burnetiamorph specimen (BP/1/7098) from the Pristerognathus Assemblage Zone and review the phylogeny of the Burnetiamorpha through a comprehensive comparison of known material. Phylogenetic analysis suggests that BP/1/7098 is closely related to the Russian species Niuksenitia sukhonensis. Remarkably, the supposed mid-Permian burnetiids Bullacephalus and Pachydectes are not recovered as burnetiids and in most cases are not burnetiamorphs at all, instead representing an earlier-diverging clade of biarmosuchians that are characterised by their large size, dentigerous transverse process of the pterygoid and exclusion of the jugal from the lateral temporal fenestra. The evolution of pachyostosis therefore appears to have occurred independently in these genera. The resulting biarmosuchian tree is significantly more congruent with the stratigraphic appearance of its constituent taxa than in previous phylogenetic hypotheses and, consequently, does not necessarily constrain the diversification of the Burnetiamorpha to before the Capitanian.

Evolutionary rates of mid-Permian tetrapods from South Africa and the role of temporal resolution in turnover reconstruction

Abstract. The Main Karoo Basin of South Africa contains a near-continuous sequence of continental deposition spanning ∼80 Myr from the mid-Permian to the Early Jurassic. The terrestrial vertebrates of this sequence provide a high-resolution stratigraphic record of regional origination and extinction, especially for the mid-late Permian. Until now, data have only been surveyed at coarse stratigraphic resolution using methods that are biased by nonuniform sampling rates, limiting our understanding of the dynamics of diversification through this important time period. Here, we apply robust methods (gap-filler and modified gap-filler rates) for the inference of patterns of species richness, origination rates, and extinction rates to a subset of 1321 reliably-identified fossil occurrences resolved to approximately 50m stratigraphic intervals. This data set provides an approximate time resolution of 0.3–0.6 Myr and shows that extinction rates increased considerably in the upper 100m of the mid-Permian Abrahamskraal Formation, corresponding to the latest part of the Tapinocephalus Assemblage Zone (AZ). Origination rates were only weakly elevated in the same interval and were not sufficient to compensate for these extinctions. Subsampled species richness estimates for the lower part of the overlying Teekloof Formation (corresponding to the Pristerognathus and Tropidostoma AZs) are low, showing that species richness remained low for at least 1.5–3 million years after the main extinction pulse. A high unevenness of the taxon abundance-frequency distribution, which is classically associated with trophically unstable postextinction faunas, in fact developed shortly before the acme of elevated extinction rates due to the appearance and proliferation of the dicynodont Diictodon. Our findings provide strong support for a Capitanian (“end-Guadalupian”) extinction event among terrestrial vertebrates and suggest that further high-resolution quantitative studies may help resolve the lack of consensus among paleobiologists regarding this event.

Comparative ecological dynamics of Permian-Triassic communities from the Karoo, Luangwa, and Ruhuhu basins of southern Africa

ABSTRACT The Permian-Triassic mass extinction (PTME) was one of the transformative events of the Phanerozoic, marked by extinction, post-Permian transformation of surviving ecosystems, and the formation of new communities. The South African Karoo Basin has served as the primary source of data on the terrestrial component of these events, but its global applicability remains poorly known. Here, we compare Permian-Triassic communities of the Karoo Basin with those from the Luangwa and Ruhuhu basins of Zambia and Tanzania, respectively, analyzing their functional structures and simulating dynamic responses to environmental perturbation. Results show that compositional similarities of late Permian communities among the basins underlie similarities in their dynamics and resistance to secondary extinction. The Karoo Basin ecosystem also displays evidence of a transformation to increased resistance during the late Permian. Although the Karoo Basin ecosystem was reduced significantly by the PTME, structural features of that resistance persisted into the Early Triassic, facilitated by a greater susceptibility to extinction of small-body-sized amniotes and large carnivorous amniotes. It was undone by the initial stages of postextinction restructuring. Continued evolution of the Triassic ecosystem led to a recovery of resistance, but in a community compositionally dissimilar from its Permian antecedents. Likewise, the upper part of the Lifua Member of the Manda Beds (Middle Triassic) of Tanzania was structurally distinct from the Karoo Basin communities but displayed similar dynamics. The recurrence and convergence of communities with different histories toward similar dynamics suggest that there are taxon-independent norms of community assembly and function operating on geological timescales.