The first non-pterodactyloid pterosaurian trackways and the terrestrial ability of non-pterodactyloid pterosaurs

New discoveries on the ichnological site known as “the Pterosaur Beach of Crayssac” (lower Tithonian, Upper Jurassic; south-western France) answer the question of terrestrial capabilities of non-pterodactyloid pterosaurs. If the terrestrial type of locomotion of pterodactyloid pterosaurs has been solved from ichnological evidence for more than twenty years, no tracks and trackways referable to non-pterodactyloid pterosaurs have ever been described. Thus, the debate on terrestrial capabilities of these non-pterodactyloids was based on morpho-functional studies, with the main conclusion that those pterosaurs were arboreal dwellers and bad walkers. Six trackways referable to three non-pterodactyloid new ichnotaxa, maybe closely related to Rhamphorhynchidae, are described in this work. Their study leads to the conclusion that grounded non-pterodatyloids, at least during the Late Jurassic, were quadrupedal with digitigrade manus and plantigrade to digitigrade pes. They were clearly good walkers, even if hindlimbs are supposed to be hampered by the uropatagium, what could have constrained the terrestrial agility of these animals. Thus, from ichnological evidence and contrary to the current hypotheses, non-pterodactyloid pterosaurs seem to have been good walkers even though their trackways are very rare or unidentified to date. This rarity could be due to behaviour rather than to functional capacities, many non-pterodactyloids being considered both littoral fishers and arboreal or cliff dwellers. However, the concept of non-pterodactyloid “good climbers and bad walkers” has to be modified to “good climbers and rare walkers”, unless many non-pterodactyloid ichnites have yet to be discovered.