The oldest fossil bird-like footprints from the upper Triassic of southern Africa

Footprint morphology reflects the anatomy of the trackmaker's foot and is direct evidence for the animal's behaviour. Consequently, fossil tracks can be used to infer ancient diversity, ethology, and evolutionary trends. This is particularly useful for deep-time intervals during which the early history of an animal group is reliant upon limited fossil skeletal material. Fossil tracks of early birds and theropods, the co-existing dinosaurian ancestors of birds, co-occur in the rock record since the Early Cretaceous. However, the evolutionary transition from dinosaur to bird and the timing of the birds' origin are still contested. Skeletal remains of the basal-most birds Aurornis, Anchiornis, Archaeopteryx and Xiaotingia are Middle to Late Jurassic, while tracks with tentative bird affinities, attributed to dinosaurs, are known from as early as the Late Triassic. Here, we present numerous, well-provenanced, Late Triassic and Early Jurassic tridactyl tracks from southern Africa, with demonstrable bird-like affinities, predating basal bird body fossils by c. 60 million years.