Wading through water: effects of water depth and speed on the drag and kinematics of walking Chilean flamingos, Phoenicopterus chilensis.

Wading behaviours, in which an animal walks while partially submerged in water, are present in a variety of taxa including amphibians, reptiles, mammals and birds. Despite the ubiquity of wading behaviours, few data are available to evaluate how animals adjust their locomotion to accommodate changes in water depth. Because drag from water might impose additional locomotor costs, wading animals might be expected to raise their feet above the water up to a certain point until such behaviours lead to awkward steps and are abandoned. To test for such mechanisms, we measured drag on models of the limbs of Chilean flamingos (Phoenicopterus chilensis) and measured their limb and body kinematics as they walked and waded through increasing depths of water in a zoo enclosure. Substantial drag was incurred by models of both open- and closed-toed feet, suggesting that flamingos could avoid some locomotor costs by stepping over water, rather than through it, during wading. Step height was highest while wading through intermediate water depths and while wading at a faster speed. Stride length increased with increasing water depth and velocity, and the limb joints generally flexed more while moving through intermediate water depths. However, movements of the head and neck were not strongly correlated with water depth or velocity. Our results show a wide range of kinematic changes that occur to allow wading birds to walk through different water depths, and have implications for better understanding the locomotor strategies employed by semi-aquatic species.