It's just sand between the toes: how particle size and shape variation affect running performance and kinematics in a...

Animals must cope with and be able to move effectively on a variety of substrates. Substrates composed of granular media, such as sand and gravel, are extremely common in nature, and vary tremendously in particle size and shape. Despite many studies of the properties of granular media and comparisons of locomotion between granular and solid substrates, the effects of systematically manipulating these media on locomotion is poorly understood. We studied granular media ranging over four orders of magnitude in particle size, and differing in the amount of particle shape variation, to determine how these factors affected substrate physical properties and sprinting in the generalist lizard Eremias arguta We found that media with intermediate particle sizes had high bulk densities, low angles of stability and low load-bearing capacities. Rock substrates with high shape variation had higher values for all three properties than glass bead substrates with low shape variation. We found that E. arguta had the highest maximum velocities and accelerations on intermediate size particles, and higher velocities on rock than glass beads. Lizards had higher stride frequencies and lower duty factors on intermediate particle size substrates, but their stride lengths did not change with substrate. Our findings suggest that sand and gravel may represent different locomotor challenges for animals. Sand substrates provide animals with an even surface for running, but particles shift underfoot. In contrast, gravel particles are heavy, so move far less underfoot, yet provide the animal with an uneven substrate.