No abstract

Birds morph their wings during a single wingbeat, across flight speeds and among flight modes. Such morphing may allow them to maximize aerodynamic performance, but this assumption remains largely untested. We tested the aerodynamic performance of swept and extended wing postures of 13 raptor species in three families (Accipitridae, Falconidae and Strigidae) using a propeller model to emulate mid-downstroke of flapping during take-off and a wind tunnel to emulate gliding. Based on previous research, we hypothesized that (1) during flapping, wing posture would not affect maximum ratios of vertical and horizontal force coefficients (C V :C H ), and that (2) extended wings would have higher maximum C V :C H when gliding. Contrary to each hypothesis, during flapping, extended wings had, on average, 31% higher maximum C V :C H ratios and 23% higher C V than swept wings across all biologically relevant attack angles (α), and, during gliding, maximum C V :C H ratios were similar for the two postures. Swept wings had 11% higher C V than extended wings in gliding flight, suggesting flow conditions around these flexed raptor wings may be different from those in previous studies of swifts (Apodidae). Phylogenetic affiliation was a poor predictor of wing performance, due in part to high intrafamilial variation. Mass was only significantly correlated with extended wing performance during gliding. We conclude that wing shape has a greater effect on force per unit wing area during flapping at low advance ratio, such as takeoff, than during gliding.

Wing tip slots are a distinct morphological trait broadly expressed across the avian clade, but are generally perceived to be unique to soaring raptors. These slots are the result of emarginations on the distal leading and trailing edges of primary feathers, and allow the feathers to behave as individual airfoils. Research suggests these emarginate feathers are an adaptation to increase glide efficiency by mitigating induced drag in a manner similar to aircraft winglets. If so, we might expect birds known for gliding and soaring to exhibit emarginate feather morphology; however, that is not always the case. Here, we explore emargination across the avian clade, and examine associations between emargination and ecological and morphological variables. Pelagic birds exhibit pointed, high-aspect ratio wings without slots, whereas soaring terrestrial birds exhibit prominent wing-tip slots. Thus, we formed four hypotheses: (1) Emargination is segregated according to habitat (terrestrial, coastal/freshwater, pelagic). (2) Emargination is positively correlated with mass. (3) Emargination varies inversely with aspect ratio and directly with wing loading and disc loading. (4) Emargination varies according to flight style, foraging style, and diet. We found that emargination falls along a continuum that varies with habitat: Pelagic species tend to have zero emargination, coastal/freshwater birds have some emargination, and terrestrial species have a high degree of emargination. Among terrestrial and coastal/freshwater species, the degree of emargination is positively correlated with mass. We infer this may be the result of selection to mitigate induced power requirements during slow flight that otherwise scale adversely with increasing body size. Since induced power output is greatest during slow flight, we hypothesize that emargination may be an adaptation to assist vertical take-off and landing rather than glide efficiency as previously hypothesized.

The environmental conditions animals experience during development can have sustained effects on morphology, physiology, and behavior. Exposure to elevated levels of stress hormones (glucocorticoids, GCs) during development is one such condition that can have long‐term effects on animal phenotype. Many of the phenotypic effects of GC exposure during development (developmental stress) appear negative. However, there is increasing evidence that developmental stress can induce adaptive phenotypic changes. This hypothesis can be tested by examining the effect of developmental stress on fitness‐related traits. In birds, flight performance is an ideal metric to assess the fitness consequences of developmental stress. As fledglings, mastering takeoff is crucial to avoid bodily damage and escape predation. As adults, takeoff can contribute to mating and foraging success as well as escape and, thus, can affect both reproductive success and survival. We examined the effects of developmental stress on flight performance across life‐history stages in zebra finches (Taeniopygia guttata). Specifically, we examined the effects of oral administration of corticosterone (CORT, the dominant avian glucocorticoid) during development on ground‐reaction forces and velocity during takeoff. Additionally, we tested for associations between flight performance and reproductive success in adult male zebra finches. Developmental stress had no effect on flight performance at all ages. In contrast, brood size (an unmanipulated variable) had sustained, negative effects on takeoff performance across life‐history stages with birds from small broods performing better than birds from large broods. Flight performance at 100 days posthatching predicted future reproductive success in males; the best fliers had significantly higher reproductive success. Our results demonstrate that some environmental factors experienced during development (e.g. clutch size) have stronger, more sustained effects than others (e.g. GC exposure). Additionally, our data provide the first link between flight performance and a direct measure of reproductive success.

Bird feathers are complex structures that passively deflect as they interact with air to produce aerodynamic force. Newtonian theory suggests that feathers should be stiff to effectively utilize this force. Observations of flying birds indicate that feathers respond to aerodynamic loading via spanwise bending, twisting, and sweeping. These deflections are hypothesized to optimize flight performance, but this has not yet been tested. We measured deflection of isolated feathers in a wind tunnel to explore how flexibility altered aerodynamic forces in emulated gliding flight. Using primary feathers from seven raptors and a rigid airfoil, we quantified bending, sweep, and twisting, as well as α (attack angle) and slip angle. We predicted that (1) feathers would deflect under aerodynamic load, (2) bending would result in lateral redirection of force, (3) twisting would alter spanwise α 'washout' and delay the onset of stall, and (4) flexural stiffness of feathers would exhibit positive allometry. The first three predictions were supported by our results, but not the fourth. We found that bending resulted in the redirection of lateral forces more toward the base of the feather on the order of ∼10% of total lift. In comparison to the airfoil which stalled at α = 13.5 • , all feathers continued to increase lift production with increasing angle of attack to the limit of our range of measurements (α = 27.5 • ). We observed that feather stiffness exhibited positive allometry (∝ mass 1.1±0.3 ), however this finding is not statistically different from other hypothesized scaling relationships such as geometric similarity (∝ mass 1.67 ). These results demonstrate that feather flexibility may provide passive roll stability and delay stall by twisting to reduce local α at the feather tip. Our findings are the first to measure forces due to feather deflection under aerodynamic loading and can inform future models of avian flight as well as biomimetic morphing-wing technology.