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Synopsis Underwater walking was a crucial step in the evolutionary transition from water to land. Underwater walkers use fins and/or limbs to interact with the benthic substrate and produce propulsive forces. The dynamics of underwater walking remain poorly understood due to the lack of a sufficiently sensitive and waterproof system to measure substrate reaction forces (SRFs). Using an underwater force plate (described in our companion paper), we quantify SRFs during underwater walking in axolotls (Ambystoma mexicanum) and Spot prawn (Pandalus platyceros), synchronized with videography. The horizontal propulsive forces were greater than the braking forces in both species to overcome hydrodynamic drag. In axolotls, potential energy (PE) fluctuations were far smaller than kinetic energy (KE) fluctuations due to high buoyant support (97%), whereas the magnitudes were similar in the prawn due to lower buoyant support (93%). However, both species show minimal evidence of exchange between KE and PE, which, along with the effects of hydrodynamic drag, is incompatible with inverted pendulum dynamics. Our results show that, despite their evolutionary links, underwater walking has fundamentally different dynamics compared with terrestrial walking and emphasize the substantial consequences of differences in body plan in underwater walking. 

Synopsis The study of underwater walking presents major challenges because the small forces applied during underwater walking are difficult to measure due to the lack of a sufficiently sensitive force plate that functions underwater. Understanding the force interaction between the underwater walker and the substrate may lead to better understanding of the evolution, ecology, and biomechanics of underwater walking. The shift from aquatic to terrestrial life was a crucial transition in animal evolution where, underwater walking preceded the invasion of land and combines mechanics from terrestrial locomotion (substrate reaction forces) and aquatic swimming (buoyancy and drag). In this work, we describe our design of a low-cost underwater force plate made using 3D printed multi axis load cells equipped with commercial strain gauges amplified with a custom circuit board, and custom code to gather force data. The use of 3D printed sensors allows customization of the material and thickness of the shear beam load cell to accommodate the loads for a wide range of study species. We show that our design can detect loads as small as 1 mN (filtered) with minimal noise and present sample live animal trials of several species. The 3D multiaxial load cells, circuit design, and custom code are open-source and available online.