Tail use improves soft substrate performance in models of early vertebrate land locomotors

Abstract

In the evolutionary transition from an aquatic to a terrestrial environment, ancient vertebrates (e.g. early tetrapods) faced the challenges of terrestrial locomotion on flowable substrates (e.g. sand and mud) of variable stiffness and incline. While morphology and ranges of motion of appendages can be revealed in fossils, biological and robophysical studies of modern taxa demonstrate that movement on such substrates can be sensitive to small changes in appendage use. Using a biological model (the mudskipper), a physical model (a robot), granular drag measurements, and theoretical tools from geometric mechanics, we demonstrate how tail use can improve robustness to variable limb use and substrate conditions. We hypothesize that properly coordinated tail movements may have provided a substantial benefit for the earliest vertebrates to move on land.