Abstract
Microscopic papillae on polar bear paw pads are considered adaptations for increased friction on ice/snow, yet this assertion is based on a single study of one species. The lack of comparative data from species that exploit different habitats renders the ecomorphological associations of papillae unclear. Here, we quantify the surface roughness of the paw pads of four species of bear over five orders of magnitude by calculating their surface roughness power spectral density. We find that interspecific variation in papillae base diameter can be explained by paw pad width, but that polar bear paw pads have 1.5 times taller papillae and 1.3 times more true surface area than paw pads of the American black bear and brown bear. Based on friction experiments with three-dimensional printed model surfaces and snow, we conclude that these factors increase the frictional shear stress of the polar bear paw pad on snow by a factor of 1.3-1.5 compared with the other species. Absolute frictional forces, however, are estimated to be similar among species once paw pad area is accounted for, suggesting that taller papillae may compensate for frictional losses resulting from the relatively smaller paw pads of polar bears compared with their close relatives.