Multi-functional Foot Use While Running Across Complex Terrain

Abstract

Running on land is fraught with challenges. Changes in topography, material properties, and the general heterogeneity of terrain necessitate the ability to sense, process, and compensate for changes quickly and repeatedly. Failure to do so successfully has potentially fatal consequences on an animal’s survival. I used lizards as an experimental model to examine how animals move successfully across complex terrain, because they are found in a vast range of habitat types and have a foot shape conserved across many families. I found that sprawled running appears to confer surprising robustness against large surface drops without any decrease in running speed, through the interplay between changes in body and limb posture. Increased combinations of foot placement and positioning result in more stable solutions than are otherwise possible among animals with erect limb postures. Using high-speed x-ray to visualize subsurface foot motion while running on sand, I discovered that lizards often found on fine sand spread their toes to distances known to maximize particle-particle interactions for force production on sand. Finally, I used a hopping robot outfitted with 3-D printed bio-inspired feet to show that the toe spacing pattern used by sand specialist lizards increases jump height for both stiff and flexible foot models. In summary, this work illuminates how adaptations for complex terrain may not be largely driven by selection for foot shape, but rather through modifications of behavior and functional morphology which likely confer robustness during locomotion across a variety of terrain.