Pump and sway: Wild primates use compliant supports as a tool to augment leaping in the canopy.

Abstract

Despite qualitative observations of wild primates pumping branches before leaping across gaps in the canopy, most studies have suggested that support compliance increases the energetic cost of arboreal leaping, thus limiting leaping performance. In this study, we quantified branch pumping behavior and tree swaying in wild primates to test the hypothesis that these behaviors improve leaping performance. We recorded wild colobine monkeys crossing gaps in the canopy and quantitatively tracked the kinematics of both the monkey and the compliant support during behavioral sequences. We also empirically measured the compliance of a sample of locomotor supports in the monkeys' natural habitat, allowing us to quantify the resonant properties of substrates used during leaping. Analyses of three recordings show that adult red colobus monkeys (Piliocolobus tephrosceles) use branch compliance to their advantage by actively pumping branches before leaping, augmenting their vertical velocity at take-off. Quantitative modeling of branch resonance periods, based on empirical measurements of support compliance, suggests that monkeys specifically employed branch pumping on relatively thin branches with protracted periods of oscillation. Finally, an additional four recordings show that both red colobus and black and white colobus monkeys (Colobus guereza) utilize tree swaying to cross large gaps, augmenting horizontal velocity at take-off. This deliberate branch manipulation to produce a mechanical effect for stronger propulsion is consistent with the framework of instrumental problem-solving. To our knowledge, this is the first study of wild primates which quantitatively shows how compliant branches can be used advantageously to augment locomotor performance.