A manus dominated pterosaur track assemblage from Gansu, China: implications for behavior

The Yangouxia dinosaur tracksites are well known for a diverse assemblage of tetrapod tracks preserved as natural impressions (concave epireliefs) on large bedding planes, representing a locally widespread surface marking the transition from a sand- to a mud-dominated sequence in the Hekou Group. Previous ichnological studies at these large sites have focused on the morphology and ichnotaxonomy of the tracks, including a single trackway representing the first pterosaur tracks reported from China. Here, we report a distinctly different assemblage associated with minor sandstones in the mud-dominated sequence 20 m above the main tracksite level. This assemblage consists of at least 20 pterosaur manus track casts attributed to a single ichnotaxon (Pteraichnus). No pes tracks have been identified. These tracks mostly occur in random orientations, although one possible trackway segment is inferred, to represent walking progression. Manus-only pterosaur track assemblages are common and likely reflect differential registration depths of manus and pes and/or sub optimal preservation conditions. The tracks are associated with distinctive invertebrate traces including Cochlichnus, Spongeliomorpha and Paleophycus and suggest the pterosaurs were likely feeding on the invertebrate tracemakers.