Pterosaur tracks and the terrestrial ability of pterosaurs

Unusual tracks of a quadrupedal animal with a three-digit (occasionally four-digit) manus print and four-digit pes print were first interpreted as those of pterosaurs in the 1950s. In the 1980s these tracks were reinterpreted as crocodilian, but new material shows that the original identification was correct. Two features: evidence for elongate penultimate phalanges in digits two to four of the pes, and manus trackways up to three times the width of pes trackways, can only be attributed to pterosaurs. Recent improvements in understanding of pterosaur anatomy and functional morphology explain remaining difficulties regarding the interpretation of ich-nites such as the orientation of the manus digits and the absence of some expected ichnological features. Pteraichnus and Pteraichnus-like tracks show that, when grounded, some, perhaps all, pterosaurs were plantigrade, quadrupedal, and had a semi-erect stance and gait. This is consistent with some functional interpretations of pterosaur anatomy and resolves a long-running debate regarding the terrestrial ability of this group.