Taphonomy of drought afflicted tetrapods in the Early Triassic Karoo Basin, South Africa

Abstract

The sedimentology and taphonomy of in-situ fossils from earliest Triassic strata (Induan) in the southern Karoo Basin of South Africa is presented as evidence for episodes of drought-induced mass death of the resident tetrapods. Abundant skeletons are preserved in a 2 m-thick tabular silty-sandstone capping a multi-storeyed low-sinuosity channel sandstone interpreted as a wide shallow channel that became progressively abandoned, with more ephemeral flow regime than in the underlying channels and subjected to intermittent flows of low-density sediment-laden floodwaters. Stratigraphic and planimetric distribution of 170 in-situ tetrapod fossils shows several clusters of up to eight closely-spaced articulated Lystrosaurus skeletons preserved in prone and spread-eagled body position. These are interpreted as drought-stricken carcasses that collapsed and died of starvation in and alongside dried-up water sources. Two of the specimens display an unusual micritic envelope with a distinctive pustular texture interpreted as permineralised mummified skin indicative of rapid desiccation after death. Bonebeds of disarticulated bones of multiple juvenile Lystrosaurus occur in shallow depressions within the rubified mudstones. Layering of different skeletal elements suggests some hydraulic sorting but the initial aggregation was likely a behavioural response to drought. Osteohistology of spread-eagled Lystrosaurus (L. declivis and L. murrayi species) skeletons show that they represent early juvenile stage which is in accordance with previous findings that throughout Pangaea Early Triassic Lystrosaurus died relatively young due to environmental stressors. Our results support the hyperthermal hypothesis that ∼252 Mya increased continental aridity, already a consequence of the coalescence of Pangaea, was critically intensified by volcanogenic greenhouse gasses from the Siberian traps. We propose that in the aftermath of the End-Permian mass extinction event, a succession of climatic drying episodes orchestrated a series of fully-functioning terrestrial ecosystems that were markedly different to those of the pre-extinction, and likely had a profound and lasting influence on the evolution of tetrapods.