Fig 3. Changes to a landing bat’s body orientation, roll, ψ, pitch, θ, and yaw, ϕ, are prompted by pronounced changes in its wing kinematics.

The top two frames indicate body orientation and angular velocity (about body-fixed axes). The lower three frames show simplified wing kinematics: instantaneous wing extension, e, wing stroke angle, ϕ _w, and protraction-retraction angle, θ _w. The departure from symmetric left-right wing motion coincides with changes in the body orientation. The five moments in time indicated by dashed vertical lines correspond to the images shown in Fig 2.