Recovery and diversification of marine communities following the late Permian mass extinction event in the western Palaeotethys

The recovery of benthic invertebrates following the late Permian mass extinction event is often described as occurring in the Middle Triassic associated with the return of Early Triassic Lazarus taxa, increased body sizes, platform margin metazoan reefs, and increased tiering. Most quantitative palaeoecological studies, however, are limited to the Early Triassic and the timing of the final phase of recovery is rarely quantified. Here, quantitative abundance data of benthic invertebrates were collected from the Middle Triassic (Anisian) succession of the Mecsek Mountains (Hungary), and analysed with univariate and multivariate statistics to investigate the timing of recovery following the late Permian mass extinction. These communities lived in a mixed siliciclastic-carbonate ramp setting on the western margin of the Palaeotethys Ocean. The new data presented here is combined with the previously studied Lower Triassic succession of the Aggtelek Karst (Hungary), which records deposition of comparable facies and in the same region of the Palaeotethys Ocean. The Middle Triassic benthic fauna can be characterised by three distinct ecological states. The first state is recorded in the Viganvár Limestone Formation representing mollusc-dominated communities restricted to above wave base, which are comparable to the lower and mid-Spathian Szin Marl Formation faunas. The second state is recorded in the Lapis Limestone Formation and records extensive bioturbation that is not limited to wave base and is comparable to the upper Spathian Szinpetri Limestone Formation. The third ecological state occurs in the Zuhánya Limestone Formation which was deposited in the Pelsonian Binodosus Zone, and has a more 'Palaeozoic' structure with sessile brachiopods dominating assemblages for the first time in the Mesozoic. The return of community-level characteristics to pre-extinction levels and the diversification of invertebrates suggests that the final stages of recovery and the radiation of the benthos in ramp settings following the late Permian mass extinction occurred in the upper Pelsonian Zuhánya Limestone Formation, approximately 8 million years after the extinction event.