Abstract
Locomotion has influenced the ecology, evolution, and extinction of species throughout history, yet studying locomotion in the fossil record is challenging. Computational biomechanics can provide novel insight by mechanistically relating observed anatomy to whole-animal function and behavior. Here, we leverage optimal control methods to generate the first fully predictive, three-dimensional, muscle-driven simulations of locomotion in an extinct terrestrial vertebrate, the bipedal non-avian theropod dinosaur Coelophysis. Unexpectedly, our simulations involved pronounced lateroflexion movements of the tail. Rather than just being a static counterbalance, simulations indicate that the tail played a crucial dynamic role, with lateroflexion acting as a passive, physics-based mechanism for regulating angular momentum and improving locomotor economy, analogous to the swinging arms of humans. We infer this mechanism to have existed in many other bipedal non-avian dinosaurs as well, and our methodology provides new avenues for exploring the functional diversity of dinosaur tails in the future.