Biodynamics of climbing: effects of substrate orientation on the locomotion of a highly arboreal lizard (Chamaeleo calyptratus).

Arboreal substrates differ not only in diameter, but also in continuity and orientation. To gain more insight into the dynamics of small-branch locomotion in tetrapods we studied the veiled chameleon walking on inclined and declined perches of up to 60 deg slope. We found that forelimbs and hindlimbs contribute equally to the body's progression along inclines and declines. The higher-positioned limb's vertical impulses decreased with slope. And although vertical impulses in the lower-positioned limb increased with substrate slope, peak vertical forces decreased. The decrease in peak vertical forces in the lower-positioned limb can be explained by a considerable increase of tensile forces in the higher-positioned limb as the slope increases. In addition, limbs were more crouched on slopes whereas no changes in forward and backward reach were observed. Mediolateral impulses were the smallest amongst the force components, and lateral impulses (medially directed limb forces) exceeded medial impulses (laterally directed limb forces). On inclines and declines, limb placement was more variable than on level substrates. The tail never contacted the substrate during level locomotion; however, on inclines and declines, the tail was held closer to the substrate, with short substrate contacts in one-third of the analyzed trials. Regardless of substrate orientation the tail was always held straight above the branch; therefore, rotational moments induced by the tail were minimized.