Models of benthic bipedalism.

Walking is a common bipedal and quadrupedal gait and is often associated with terrestrial and aquatic organisms. Inspired by recent evidence of the neural underpinnings of primitive aquatic walking in the little skate , we introduce a theoretical model of aquatic walking that reveals robust and efficient gaits with modest requirements for body morphology and control. The model predicts undulatory behaviour of the system body with a regular foot placement pattern, which is also observed in the animal, and additionally predicts the existence of gait bistability between two states, one with a large energetic cost for locomotion and another associated with almost no energetic cost. We show that these can be discovered using a simple reinforcement learning scheme. To test these theoretical frameworks, we built a bipedal robot and show that its behaviours are similar to those of our minimal model: its gait is also periodic and exhibits bistability, with a low efficiency mode separated from a high efficiency mode by a 'jump' transition. Overall, our study highlights the physical constraints on the evolution of walking and provides a guide for the design of efficient biomimetic robots.