Birds can transition between stable and unstable states via wing morphing.

Birds morph their wing shape to accomplish extraordinary manoeuvres1-4, 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.

Individual variation and the biomechanics of maneuvering flight in hummingbirds.

An animal's maneuverability will determine the outcome of many of its most important interactions. A common approach to studying maneuverability is to force the animal to perform a specific maneuver or to try to elicit maximal performance. Recently, the availability of wider-field tracking technology has allowed for high-throughput measurements of voluntary behavior, an approach that produces large volumes of data. Here, we show how these data allow for measures of inter-individual variation that are necessary to evaluate how performance depends on other traits, both within and among species. We use simulated data to illustrate best practices when sampling a large number of voluntary maneuvers. Our results show how the sample average can be the best measure of inter-individual variation, whereas the sample maximum is neither repeatable nor a useful metric of the true variation among individuals. Our studies with flying hummingbirds reveal that their maneuvers fall into three major categories: simple translations, simple rotations and complex turns. Simple maneuvers are largely governed by distinct morphological and/or physiological traits. Complex turns involve both translations and rotations, and are more subject to inter-individual differences that are not explained by morphology. This three-part framework suggests that different wingbeat kinematics can be used to maximize specific aspects of maneuverability. Thus, a broad explanatory framework has emerged for interpreting hummingbird maneuverability. This framework is general enough to be applied to other types of locomotion, and informative enough to explain mechanisms of maneuverability that could be applied to both animals and bio-inspired robots.

Range of motion in the avian wing is strongly associated with flight behavior and body mass.

Avian wing shape is highly variable across species but only coarsely associated with flight behavior, performance, and body mass. An underexplored but potentially explanatory feature is the ability of birds to actively change wing shape to meet aerodynamic and behavioral demands. Across 61 species, we found strong associations with flight behavior and mass for range of motion traits but not wing shape and strikingly different associations for different aspects of motion capability. Further, static morphology exhibits high phylogenetic signal, whereas range of motion shows greater evolutionary lability. These results suggest a new framework for understanding the evolution of avian flight: Rather than wing morphology, it is range of motion, an emergent property of morphology, that is predominantly reshaped as flight strategy and body size evolve.

Wing morphing allows gulls to modulate static pitch stability during gliding.

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.

Allometry of hummingbird lifting performance.

Vertical lifting performance in 67 hummingbird species was studied across a 4000 m elevational gradient. We used the technique of asymptotic load-lifting to elicit maximum sustained muscle power output during loaded hovering flight. Our analysis incorporated direct measurements of maximum sustained load and simultaneous wingbeat kinematics, together with aerodynamic estimates of mass-specific mechanical power output, all within a robust phylogenetic framework for the Trochilidae. We evaluated key statistical factors relevant to estimating slopes for allometric relationships by performing analyses with and without phylogenetic information, and incorporating species-specific measurement error. We further examined allometric relationships at different elevations because this gradient represents a natural experiment for studying physical challenges to animal flight mechanics. Maximum lifting capacity (i.e. vertical force production) declined with elevation, but was either isometric or negatively allometric with respect to both body and muscle mass, depending on elevational occurrence of the corresponding taxa. Maximum relative muscle power output exhibited a negative allometry with respect to muscle mass, supporting theoretical predictions from muscle mechanics.

Hovering performance of hummingbirds in hyperoxic gas mixtures.

Hummingbirds evolved during a period of decline in atmospheric oxygen concentration and currently encounter varying levels of oxygen availability along their elevational distribution. We tested the hypothesis that inspiration of hyperoxic gas increases hummingbird hovering performance when birds are simultaneously challenged aerodynamically. We measured the maximum duration of hovering flight while simultaneously monitoring the rate of oxygen consumption of ruby-throated hummingbirds (Archilochus colubris) in low-density heliox that was either normoxic (21% O2) or hyperoxic (35% O2). As air density decreased below 0.85 kg x m(-3), hummingbirds hovered significantly longer in hyperoxia than in normoxia, but the air density at which the birds could no longer sustain hovering flight was independent of oxygen concentration. At low air densities in hyperoxia flight trials, hummingbirds appeared to increase their rate of oxygen consumption relative to flight sequences at equivalent densities in normoxia trials, but these differences were not significant. We tested the hypothesis that hummingbirds can discriminate between environments that differ in oxygen concentration. In another density-reduction experiment, hummingbirds were allowed to choose between artificial feeders infused with either normoxic or hyperoxic gases. The hypothesis was not supported because birds failed to associate oxygen concentration with a particular feeder independently of air density. Supplemental oxygen thus yields increased hovering duration at intermediate air densities, but the minimum density at which birds can fly is limited exclusively by aerodynamic considerations.

Maximal horizontal flight performance of hummingbirds: effects of body mass and molt.

Hovering and fast forward flapping represent two strenuous types of flight that differ in aerodynamic power requirement. Maximal capabilities of ruby-throated hummingbirds (Archilochus colubris) in hovering and forward flight were compared under varying body mass and wing area. The capability to hover in low-density gas mixtures was adversely affected by body mass, whereas the capability to fly in a wind tunnel did not show any adverse mass effect. Molting birds that lost primary flight feathers and reduced wing area also displayed mass loss and loss of aerodynamic power and flight speed. This suggests that maximal flight speed is insensitive to short-term perturbations of body mass but that molting is stressful and reduces the birds' speed and capacity for chase and escape. Hummingbirds' flight behavior in confined space was also investigated. Birds reduced their speeds flying through a narrow tube to approximately one-fifth of that in the wind tunnel and did not display differences under varying body mass and wing area. Hence, performance in the flight tube was submaximal and did not correlate with performance variation in the wind tunnel. For ruby-throated hummingbirds, both maximal mass-specific aerodynamic power derived from hovering performance in low-density air media and maximal flight velocity measured in the wind tunnel were invariant with body mass.