Across all inclinations, upstroke contributions increase with body angle. During takeoff (A) and landing (B), the legs exert vertical (solid line) and horizontal forces (dashed line) on the perch to accelerate and decelerate. Takeoff angle increases with inclination, whereas landing angle decreases. In contrast, takeoff speed remains relatively constant across 40- and 75-cm flights (inset; table S1). (C) Net impulse is the integrated vertical leg force [(A) and (B)] minus bodyweight. Impulse transfer shifts from landing to takeoff for ascending flight, and vice versa for descending. (D) During the first wingbeats after toe-off (see Materials and Methods for wingbeat selection criteria), bodyweight is primarily supported by downstroke impulse (dashed boundaries, end of downstroke). (E) Just before touchdown, the upstroke contribution to bodyweight support increases. Bird avatars show how body angle and actuator disc area increase from takeoff to landing wingbeats. (F) The upstroke to downstroke vertical impulse ratio increases with body angle, regardless of flight inclination. (A to C) Panels show mean results for each flight variation with N = 4, n = 5. (D to F) Panels show mean results for 75-cm variations with N = 4, n = 3. SDs are shown by shaded regions in (A), (B), (D), and (E) and by error bars in (C) and (F).