Kinematics and center of mass mechanics during terrestrial locomotion in northern lapwings (Vanellus vanellus, Charadriiformes).

Avian bipedalism is best studied in derived walking/running specialists. Here, we use kinematics and center of mass (CoM) mechanical energy patterns to investigate gait transitions of lapwings-migratory birds that forage on the ground, and therefore may need a trade-off between the functional demands of terrestrial locomotion and long distance flights. The animals ran on a treadmill while high-speed X-ray videos were recorded within the sustainable speed range. Instantaneous CoM mechanics were computed from integrating kinematics and body segment properties. Lapwings exhibit similar locomotor characteristics to specialized walking/running birds, but have less distinct gaits. At slow speeds no clear separation between vaulting (i.e., walking) and bouncing (i.e., running) energy patterns exists. Mechanical energy recovery of non-bouncing gaits correlates poorly with speed and suggests inefficient use of the inverted pendulum mechanism. Speed ranges of gaits overlap considerably, especially those of grounded running, a gait with CoM mechanics indicative of running but without an aerial phase, and aerial phase running, with no preferential gait at most speeds. Compliant limb morphology and grounded running in birds can be regarded as an evolutionary constraint, but lapwings effectively make use of advantages offered by this gait for a great fraction of their speed range. Thus, effective usage of grounded running during terrestrial locomotion is suggested generally to be a part of striding avian bipedalism-even in species not specialized in walking/running locomotion.