Reconstructing the diversity of early terrestrial herbivorous tetrapods

Terrestrial herbivorous tetrapods first appear in the fossil record during the Late Carboniferous (306.5 Ma). The diversification of herbivores is a key aspect of the transition to the modern trophic structure of terrestrial vertebrate ecosystems, because it allowed tetrapods to exploit terrestrial (i.e. non-aquatic) primary productivity. However, the palaeodiversity dynamics of the earliest terrestrial vertebrate herbivores have received relatively little attention, apart from a few studies that focus on specific clades. A new dataset containing 287 species occurrences of herbivorous tetrapods including the major Palaeozoic and Early Triassic clades Anomodontia, Archosauromorpha, Bolosauridae, Captorhinidae, Caseidae, Cynodontia, Dinocephalia, Diadectomorpha, Edaphosauridae, Pareiasauria, Poposauroidea, Procolophonoidea, Rhynchosauria, Silesauridae and Therocephalia is used to analyse palaeodiversity from the Late Carboniferous to the Middle Triassic (~ 306.5–~ 236 Ma), taking into account the effects of potential sampling biases by using the number of tetrapod-bearing formations as a proxy. The results support a gradual increase in taxic diversity from the Late Carboniferous to the Wordian, followed by a dip in diversity during the Guadalupian (Middle Permian), and an increase to a peak in the Late Permian at the Wuchiapingian/Changhsingian boundary. Herbivorous tetrapods were strongly affected by the end-Permian mass extinction with both the herbivorous Pareiasauria and Captorhinidae becoming extinct and the observed number of anomodont species decreasing by up to 80%. The drop in observed diversity at the end Permian is dampened slightly because of the radiation of new herbivorous forms during the Early Triassic. A strong biological signal is apparent even after correcting for sampling.