SUMMARY
Small mammals must negotiate terrains that consist of numerous substrates that vary in diameter, surface structure, rigidity and orientation. Most studies on mammals have focused on the effects of substrate diameter during horizontal locomotion, especially in small- to medium-sized primates and marsupials. Locomotion across sloped arboreal substrates, however, is poorly understood. Here, in order to determine which locomotor parameters a terrestrial mammal, the rat, and a tree-dwelling mammal, the European red squirrel, modify in response to differences in substrate orientation, three-dimensional kinematics were examined using biplanar videoradiography as the animals moved on 30 and 60 deg inclined branches. Our results revealed that to maintain stability and friction as well as balance during inclined branch locomotion, these species utilize comparable locomotor adjustments despite significant differences in travel speed and gait. Rats and European red squirrels increased limb flexion and retraction in order to bring the center of mass as close as possible to the substrate surface and to achieve maximum propulsion. Additionally, forelimbs were placed more laterally and underneath the branch whereas the hindlimbs were placed approximately on the top of the branch. These locomotor adjustments, which have also been observed in primates and marsupials, are independent of speed, morphological adaptations and limb proportions and thus might be strategies used by early mammals. Our results also suggest that mammals that lack, or have reduced, grasping abilities try to maintain the locomotor mode used during horizontal branch locomotion on inclined branches for as long as possible.