The stance phase of walking is when forces are applied to the environment to support, propel, and maneuver the body. Unlike solid surfaces, deformable substrates yield under load, allowing the foot to sink to varying degrees. For bipedal birds and their dinosaurian ancestors, a shared response to walking on these substrates has been identified in the looping path the digits follow underground. Because a volume of substrate preserves a 3-D record of stance phase in the form of footprints or tracks, understanding how the bipedal stride cycle relates to this looping motion is critical for building a track-based framework for the study of walking in extinct taxa. Here we used biplanar X-ray imaging to record and analyze 161 stance phases from 81 trials of three Helmeted Guineafowl (Numida meleagris) walking on radiolucent substrates of different consistency (solid, dry granular, firm to semi-liquid muds). Across all substrates, the feet sank to a range of depths up to 78% of hip height. With increasing substrate hydration, the majority of foot motion shifted from above to below ground. Walking kinematics sampled across all stride cycles revealed six sequential gait-based events originating from both feet, conserved throughout the spectrum of substrate consistencies during normal alternating walking. On all substrates that yielded, five sub-phases of gait were drawn out in space and formed a loop of varying shape. We describe the two-footed coordination and weight distribution that likely contributed to the observed looping patterns of an individual foot. Given such complex subsurface foot motion during normal alternating walking and some atypical walking behaviors, we discuss the definition of "stance phase" on deformable substrates. We also discuss implications of the gait-based origins of subsurface looping on the interpretation of locomotory information preserved in fossil dinosaur tracks.
Keywords: XROMM; biped; foot; guineafowl; kinematics; walking.
© The Author(s) 2022. Published by Oxford University Press on behalf of the Society for Integrative and Comparative Biology.