Birds morph their wing shape to accomplish extraordinary manoeuvres^1–4, which are governed by avian-specific equations of motion. Solving these equations requires information about a bird’s aerodynamic and inertial characteristics⁵. Avian flight research to date has focused on resolving aerodynamic features, whereas inertial properties including centre of gravity and moment of inertia are seldom addressed. Here we use an analytical method to determine the inertial characteristics of 22 species across the full range of elbow and wrist flexion and extension. We find that wing morphing allows birds to substantially change their roll and yaw inertia but has a minimal effect on the position of the centre of gravity. With the addition of inertial characteristics, we derived a novel metric of pitch agility and estimated the static pitch stability, revealing that the agility and static margin ranges are reduced as body mass increases. These results provide quantitative evidence that evolution selects for both stable and unstable flight, in contrast to the prevailing narrative that birds are evolving away from stability⁶. This comprehensive analysis of avian inertial characteristics provides the key features required to establish a theoretical model of avian manoeuvrability.

A gliding bird's ability to stabilize its flight path is as critical as its ability to produce sufficient lift. In flight, birds often morph the shape of their wings, but the consequences of avian wing morphing on flight stability are not well understood. Here, we investigate how morphing the gull elbow joint in gliding flight affects their static pitch stability. First, we combined observations of freely gliding gulls and measurements from gull wing cadavers to identify the wing configurations used during gliding flight. These measurements revealed that, as wind speed and gusts increased, gulls flexed their elbows to adopt wing shapes characterized by increased spanwise camber. To determine the static pitch stability characteristics of these wing shapes, we prepared gull wings over the anatomical elbow range and measured the developed pitching moments in a wind tunnel. Wings prepared with extended elbow angles had low spanwise camber and high passive stability, meaning that mild perturbations could be negated without active control. Wings with flexed elbow angles had increased spanwise camber and reduced static pitch stability. Collectively, these results demonstrate that gliding gulls can transition across a broad range of static pitch stability characteristics using the motion of a single joint angle.

An emergent property of wing morphology bears signature of adaptation to flight ecology more strongly than morphology itself.

Birds dynamically adapt to disparate flight behaviours and unpredictable environments by actively manipulating their skeletal joints to change their wing shape. This in-flight adaptability has inspired many unmanned aerial vehicle (UAV) wings, which predominately morph within a single geometric plane. By contrast, avian joint-driven wing morphing produces a diverse set of non-planar wing shapes. Here, we investigated if joint-driven wing morphing is desirable for UAVs by quantifying the longitudinal aerodynamic characteristics of gull-inspired wing-body configurations. We used a numerical lifting-line algorithm (MachUpX) to determine the aerodynamic loads across the range of motion of the elbow and wrist, which was validated with wind tunnel tests using three-dimensional printed wing-body models. We found that joint-driven wing morphing effectively controls lift, pitching moment and static margin, but other mechanisms are required to trim. Within the range of wing extension capability, specific paths of joint motion (trajectories) permit distinct longitudinal flight control strategies. We identified two unique trajectories that decoupled stability from lift and pitching moment generation. Further, extension along the trajectory inherent to the musculoskeletal linkage system produced the largest changes to the investigated aerodynamic properties. Collectively, our results show that gull-inspired joint-driven wing morphing allows adaptive longitudinal flight control and could promote multifunctional UAV designs.