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Background  Previous analyses of factors influencing footfall timings and gait selection in quadrupeds have focused on the implications for energetic cost or gait mechanics separately. Here we present a model for symmetrical walking gaits in quadrupedal mammals that combines both factors, and aims to predict the substrate contexts in which animals will select certain ranges of footfall timings that (1) minimize energetic cost, (2) minimize rolling and pitching moments, or (3) balance the two. We hypothesize that energy recovery will be a priority on all surfaces, and will be the dominant factor determining footfall timings on flat, ground-like surfaces. The ability to resist pitch and roll, however, will play a larger role in determining footfall choice on narrower and more complex branch-like substrates. As a preliminary test of the expectations of the model, we collected sample data on footfall timings in a primate with relatively high flexibility in footfall timings – the squirrel monkey ( Saimiri sciureus ) – walking on a flat surface, straight pole, and a pole with laterally-projecting branches to simulate simplified ground and branch substrates. We compare limb phase values on these supports to the expectations of the model.    Results  As predicted, walking steps on the flat surface tended towards limb phase values that promote energy exchange. Both pole substrates induced limb phase values predicted to favor reduced pitching and rolling moments.    Conclusions  These data provide novel insight into the ways in which animals may choose to adjust their behavior in response to movement on flat versus complex substrates and the competing selective factors that influence footfall timing in mammals. These data further suggest a pathway for future investigations using this perspective.

Limb phase flexibility in walking: a test case in the squirrel monkey (Saimiri sciureus)