Active Adaptive Strategies of Mallard Feet in Response to Changes in Wetness and Compactness of the Sand Terrain.

Mallards (Anas platyrhynchos) exhibit exceptional locomotive abilities in diverse terrains, such as beaches, swamps, and tidal flats. This capability is primarily attributed to their unique webbed toe structure and cooperative locomotion posture of their feet. Therefore, this study aims to further delve into the active adaptive strategies of mallard feet in response to diverse external environmental conditions. Six adult male mallards were selected for this research. Their locomotion on sandy surfaces with differing wetness levels and varying degrees of compaction were captured using a high-speed camera, and analysis of instantaneous and continuous changes in the primary joint angles of the mallards' feet, including the toe-webbed opening and closing angles, the tarsometatarsal-phalangeal joint (TMTPJ), and the intertarsal joint (ITJ). It was found that on loose sandy surfaces, increasing wetness expanded the ground contact area of the mallards' feet. This led to greater flexion at the TMTPJ joint during mid-stance, accompanied by decreased flexion of the ITJ during touch-down and mid-stance. Conversely, on compacted sand, increasing wetness resulted in a reduced foot effect area and lessened ITJ flexion at both touch-down and mid-stance. Furthermore, on looser sand, the ground contact area of the mallards' feet decreased, with an increase in ITJ buckling at touch-down. During the swing phase, sand wetness and compactness effected minimally on the feet of the mallards. On dry and loose sand ground, mallards will contract their second and fourth toes with webbing upon ground contact, covering and compacting the sand beneath, while increasing ITJ flexion to mitigate sinking. This adaptation reduces the energy expended on sand and enhances body stability. In wet and compacted sand conditions, mallards expand their second and fourth toes upon ground contact and reduce ITJ flexion. Therefore, this coordinated foot and ITJ locomotion offers mallards a natural advantage when moving on various environmental media.