Fossilized footprints contain information about the dynamics of gait, but their interpretation is difficult, as they are the combined result of foot anatomy, gait dynamics, and substrate properties. We explore how footprints are generated in modern humans. Sixteen healthy subjects walked on a solid surface and in a layer of fine-grained sand. In each condition, 3D kinematics of the leg and foot were analyzed for three trials at preferred speed, using an infrared camera system. Additionally, calibrated plantar pressures were recorded. After each trial in sand, the depth of the imprint was measured under specific sites. When walking in sand, subjects showed greater toe clearance during swing and a 7° higher knee yield during stance. Maximal pressure was the most influential factor for footprint depth under the heel. For other foot zones, a combination of factors correlates with imprint depth, with pressure impulse (the pressure-time integral) gaining importance distally, at the metatarsal heads and the hallux. We conclude that footprint topology cannot be related to a single variable, but that different zones of the footprint reflect different aspects of the kinesiology of walking. Therefore, an integrated approach, combining anatomical, kinesiological, and substrate-mechanical insights, is necessary for a correct interpretation. Am J Phys Anthropol, 2010. © 2009 Wiley-Liss, Inc.