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 Leucoraja erinacea, 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.