An unpredictable body size response to the Permo–Triassic climate crisis

A predictive ecological response to both the present and past climate crises is that marine ectotherm species will become smaller before going extinct or fluctuate in abundance and size with environmental conditions. The problem with studying past climate events with high rates and magnitude of warming, which may serve as analogues for projected climate change, is that very few species, or even genera, survived such events. Here, we utilized one of the few records of marine bivalves that spans the Permian–Triassic climate crisis with specimen-level data and at a high resolution. These measurements come from the Bellerophon and Werfen formations of the Dolomites in Italy, representing relatively shallow marine environments. At the species-level, there is almost a complete turnover, and the newly evolved species are typically significantly smaller, but not unusually small, whereas the three surviving species do not show a significant body size change. Our results clarify that the observed temporary size reduction at the genus-level is primarily driven by the preferential evolution of smaller species after the extinction, rather than, as often assumed, by a size decrease within existing species; this challenges the universal validity of the 'Lilliput effect' in the sense of direct intra-species dwarfing, but confirms it as a consequence of faunal turnover. Subsequently, there are two pulses of genus-level body size recovery determined by different mechanisms. The first phase (late Griesbachian) is driven by the size-increase of the existing species, whereas the second phase (early Spathian) is also due to the evolution of larger species. The effects of abiotic and biotic factors in controlling these body size dynamics are superimposed during the Early Triassic. These results suggest a mechanism to explain size reductions during climate crises, but does not find a species-level body size reduction to be a forecastable response to extreme climate warming.