Pterosaur Footprints and Ghost Lineages

A 2025 study found more variation in so called Pteraichnus footprints, identifying three variants associated with the most terrestrial pterosaur lineages: Ctenochasmatoidea, Dsungaripteroidea and Neoazhdarchia. They accurately pin point the diversity of these pterosaurs across time, including confirming an early Cretaceous extinction for ctenochasmatoids and dsungaripteroids.

However, the paper also identifies a large amount of pterosaur footprints not assigned to Neoazhdarchia in the Late Cretaceous as the diagram above showsx, many of which in terrestrial settings (so partially ruling out pteranodontids and nyctosaurids). Another study (not addressed in this paper) also found numerous footprints of Germanodactylus-like pterosaurs in a Late Cretaceous quarry; this raises the possibility that many of the unidentified tracks may belong to similar pterosaurs.

Overall we’re limited by a parse fossil reccord, and I hope future discoveries further clarify the diversity of terminal Cretaceous pterosaurs.