Exceptional vertebrate biotas from the Triassic of China, and the expansion of marine ecosystems after the Permo-Triassic mass extinction

The Triassic was a time of turmoil, as life recovered from the most devastating of all mass extinctions, the Permo-Triassic event 252. million years ago. The Triassic marine rock succession of southwest China provides unique documentation of the recovery of marine life through a series of well dated, exceptionally preserved fossil assemblages in the Daye, Guanling, Zhuganpo, and Xiaowa formations. New work shows the richness of the faunas of fishes and reptiles, and that recovery of vertebrate faunas was delayed by harsh environmental conditions and then occurred rapidly in the Anisian. The key faunas of fishes and reptiles come from a limited area in eastern Yunnan and western Guizhou provinces, and these may be dated relative to shared stratigraphic units, and their palaeoenvironments reconstructed. The Luoping and Panxian biotas, both from the Guanling Formation, are dated as Anisian (Pelsonian) on the basis of conodonts and radiometric dates, the former being slightly older than the latter. The Xingyi biota is from the Zhuganpo Formation, and is Ladinian or early Carnian, while the Guanling biota is from the overlying Xiaowa Formation, dated as Carnian. The first three biotas include extensive benthos and burrowing in the sediments, and they were located in restricted basins close to shore. Further, even though the Luoping and Panxian biotas are of similar age, their faunas differ significantly, reflecting perhaps palaeogeographically isolated basins. Between the time of the Xingyi and Guanling biotas, there was a major transgression, and the Guanling biota is entirely different in character from the other three, being dominated by pelagic forms such as large floating crinoids attached to logs, very large ichthyosaurs and thalattosaurs, and pseudoplanktonic bivalves, with no benthos and no burrowing. Phylogenetic study of the fishes and marine reptiles shows apparently explosive diversification among 20 actinopterygian lineages very early in the Early Triassic, but a later expansion of marine reptile groups, in the late Olenekian and early Anisian. This offset in diversification patterns is matched by comparisons of feeding guild categories and body size data. New research tools will shed considerable light on the phylogenetic and ecological implications of recovery of mairne vertebrates in the Triassic.