Can skeletal surface area predict in vivo foot surface area?

The surface area of feet in contact with the ground is a key morphological feature that influences animal locomotion. Underfoot pressures (and consequently stresses experienced by the foot), as well as stability of an animal during locomotion, depend on the size and shape of this area. Here we tested whether the area of a skeletal foot could predict in vivo soft tissue foot surface area. Computed tomography scans of 29 extant tetrapods (covering mammals, reptiles, birds and amphibians) were used to produce models of both the soft tissues and the bones of their feet. Soft tissue models were oriented to a horizontal plane, and their outlines projected onto a surface to produce two-dimensional silhouettes. Silhouettes of skeletal models were generated either from bones in CT pose or with all autopodial bones aligned to the horizontal plane. Areas of these projections were calculated using alpha shapes (mathematical tight-fitting outline). Underfoot area of soft tissue was approximately 1.67 times that of skeletal tissue area (~ 2 times for manus, ~ 1.6 times for pes, if analysed separately). This relationship between skeletal foot area and soft tissue area, while variable in some of our study taxa, could provide information about the size of the organisms responsible for fossil trackways, suggest what size of tracks might be expected from potential trackmakers known only from skeletal remains, and aid in soft tissue reconstruction of skeletal remains for biomechanical modelling.