Complex macroevolution of pterosaurs.

Pterosaurs, the earliest flying tetrapods, are the subject of some recent quantitative macroevolutionary analyses from different perspectives. Here, we use an integrative approach involving newly assembled phylogenetic and body size datasets, net diversification rates, morphological rates, and morphological disparity to gain a holistic understanding of the pterosaur macroevolution. The first two parameters are important in quantitative analyses of macroevolution, but they have been rarely used in previous pterosaur studies. Our study reveals an ∼115-Ma period-from Early Triassic to Early Cretaceous-of multi-wave increasing net diversification rates and disparity, as well as high morphological rates, followed by an ∼65-Ma period-from Early Cretaceous to the end of the Cretaceous-of mostly negative net diversification rates, decreasing disparity, and relatively low morphological rates in pterosaur evolution. Our study demonstrates the following: (1) body size plays an important role in pterosaur lineage diversification during nearly their whole evolutionary history, and the evolution of locomotion, trophic, and ornamental structures also plays a role in different periods; (2) birds, the other major flying tetrapod group at the time, might have affected pterosaur macroevolution for ∼100 Ma; and (3) different mass extinction events might have affected pterosaur evolution differently. Particularly, the revealed decline in pterosaur biodiversity during the Middle and Late Cretaceous periods provides further support for the possible presence of a biodiversity decline of large-sized terrestrial amniotes starting in the mid-Cretaceous, which may have been caused by multiple factors including a global land area decrease during these periods.