evolved in a terrestrial context.

Padian further hypothesized that pterosaurs originated from small bipedal archosaurs, an idea anticipated in a different form by Friedrich von Huene and later supported by phylogenetic studies, including work by Davide Foffa, Sterling Nesbitt, and colleagues in 2022 and 2023. His interpretations of pterosaur anatomy and posture built on earlier comparative approaches dating back to Georges Cuvier.