Gait dynamics of Cebus apella during quadrupedalism on different substrates.

Primates are distinguished from many mammals by emphasizing arboreal lifestyles. Primate arboreal adaptations include specializations for enhancing balance and manipulative skills. Compliant gait and diagonal sequence (DS) footfalls are hypothesized mechanisms for improving balance during arboreal quadrupedalism (AQ), while simultaneously permitting vertical peak force reductions sustained by limbs, particularly forelimbs (FLs). Capuchin monkeys (Cebus apella) are arboreally-adapted quadrupeds that use both lateral sequence (LS) and DS footfalls. As tool-users, capuchins experience selective pressures for FL manipulative capabilities, which seemingly conflict with encountering substantial locomotor stresses. We evaluate kinetic and 3-D kinematic data from 172 limb contacts of two adult males on terrestrial and arboreal substrates to address questions about C. apella gait compliancy, kinematics of LS and DS footfalls during quadrupedalism on different substrates, and whether capuchins reduce FL vertical peak forces relative to hind limb (HL) forces more than other primates that use tools or those that do not. Lower vertical peak forces during AQ are consistent with compliant gait, but mixed kinematic results obscure how the reduction occurs. Forearm adduction angle is one consistent kinematic difference between terrestrial and arboreal quadrupedalism, which may implicate frontal plane movements in gait compliancy. Major differences between DS and LS gaits were not observed in kinetic or kinematic comparisons. Capuchins exhibit low FL/HL vertical peak force ratios like several anthropoids, including tool-users (e.g., chimpanzees), and species not considered tool-users in free-ranging conditions (e.g., spider monkeys). Additional selective pressures besides simply tool use appear responsible for the relative reduction in primate forelimb forces.