Theropod trackways as indirect evidence of pre-avian aerial behavior.

Body fossils set limits on feasible reconstructions of functional capacity and behavior in theropod dinosaurs, but do not document in-life behaviors. In contrast, trace fossils such as footprints preserve in-life behaviors that can potentially test and enhance existing reconstructions. Here, we demonstrate how theropod trackways can be used as indirect evidence of pre-avian aerial behavior, expanding the approaches available to study vertebrate flight origins. This involved exploring the behavioral implications of a two-toed Cretaceous-aged theropod trackway produced by a small, bird-like microraptorine moving at high speed. Applying first principle running biomechanics, we were able to conclude that the trackway is atypical, indirectly evidencing pre-avian aerial behavior. This trackway documents the evidence of wing-assisted aerodynamic force production during locomotion, supporting a broader distribution of this behavior than currently known. These findings support previously proposed aerial behavior in early bird-like theropods, showing how trackways will help to deepen our understanding of theropod flight origins.