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Animals must tune their physical performance to changing environmental conditions, and the breadth of environmental tolerance may contribute to delineating the geographic range of a species. A common environmental challenge that flying animals face is the reduction of air density at high elevation and the reduction in the effectiveness of lift production that accompanies it. As a species, turkey vultures (Cathartes aura) inhabit a >3000 m elevation range, and fly considerably higher, necessitating that they accommodate for a 27% change in air density (0.890 to 1.227 kg m-3) through behavior, physiology or biomechanics. We predicted that birds flying at high elevation would maintain aerodynamic lift performance behaviorally via higher flight speeds, rather than increases in power output or local phenotypic adaptation. We used three-dimensional videography to track turkey vultures flying at three elevations, and data supported the hypothesized negative relationship between median airspeed and air density. Additionally, neither the ratio of horizontal speed to sinking speed nor flapping behavior varied with air density., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2024. Published by The Company of Biologists Ltd.)

Wind-hovering birds exhibit remarkable steadiness in flight, achieved through the morphing of their wings and tail. We analysed the kinematics of two nankeen kestrels (Falco cenchroides) engaged in steady wind-hovering flights in a smooth flow wind tunnel. Motion-tracking cameras were used to capture the movements of the birds as they maintained their position. The motion of the birds' head and body, and the morphing motions of their wings and tail were tracked and analysed using correlation methods. The results revealed that wing sweep, representing the flexion/extension movement of the wing, played a significant role in wing motion. Additionally, correlations between different independent degrees of freedom (DoF), including wing and tail coupling, were observed. These kinematic couplings indicate balancing of forces and moments necessary for steady wind hovering. Variation in flight behaviour between the two birds highlighted the redundancy of DoF and the versatility of wing morphing in achieving control. This study provides insights into fixed-wing craft flight control from the avian world and may inspire novel flight control strategies for future fixed-wing aircraft., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2024. Published by The Company of Biologists Ltd.)

Birds, and especially raptors, are highly visual animals. Some of them have the highest spatial resolving power known in the animal kingdom, allowing prey detection at distance. While many raptors visually track fast-moving and manoeuvrable prey, requiring high temporal resolution, this aspect of their visual system has never been studied before. In this study, we estimated how fast raptors can see, by measuring the flicker fusion frequency of three species with different lifestyles. We found that flicker fusion frequency differed among species, being at least 129 Hz in the peregrine falcon, Falco peregrinus , 102 Hz in the saker falcon, Falco cherrug , and 81 Hz in the Harris's hawk, Parabuteo unicinctus We suggest a potential link between fast vision and hunting strategy, with high temporal resolution in the fast-flying falcons that chase fast-moving, manoeuvrable prey and a lower resolution in the Harris's hawk, which flies more slowly and targets slower prey., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2020. Published by The Company of Biologists Ltd.)

Birds primarily create and control the forces necessary for flight through changing the shape and orientation of their wings and tail. Their wing geometry is characterised by complex variation in parameters such as camber, twist, sweep and dihedral. To characterise this complexity, a multi-view stereo-photogrammetry setup was developed for accurately measuring surface geometry in high resolution during free flight. The natural patterning of the birds was used as the basis for phase correlation-based image matching, allowing indoor or outdoor use while being non-intrusive for the birds. The accuracy of the method was quantified and shown to be sufficient for characterising the geometric parameters of interest, but with a reduction in accuracy close to the wing edge and in some localised regions. To demonstrate the method's utility, surface reconstructions are presented for a barn owl ( Tyto alba ) and peregrine falcon ( Falco peregrinus ) during three instants of gliding flight per bird. The barn owl flew with a consistent geometry, with positive wing camber and longitudinal anhedral. Based on flight dynamics theory, this suggests it was longitudinally statically unstable during these flights. The peregrine falcon flew with a consistent glide angle, but at a range of air speeds with varying geometry. Unlike the barn owl, its glide configuration did not provide a clear indication of longitudinal static stability/instability. Aspects of the geometries adopted by both birds appeared to be related to control corrections and this method would be well suited for future investigations in this area, as well as for other quantitative studies into avian flight dynamics., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2019. Published by The Company of Biologists Ltd.)

Many large birds rely on thermal soaring flight to travel cross-country. As such, they are under selective pressure to minimise the time spent gaining altitude in thermal updrafts. Birds should be able to maximise their climb rates by maintaining a position close to the thermal core through careful selection of bank angle and airspeed; however, there have been few direct measurements of either parameter. Here, we apply a novel methodology to quantify the bank angles selected by soaring birds using on-board magnetometers. We couple these data with airspeed measurements to parameterise the soaring envelope of two species of Gyps vulture, from which it is possible to predict 'optimal' bank angles. Our results show that these large birds respond to the challenges of gaining altitude in the initial phase of the climb, where thermal updrafts are weak and narrow, by adopting relatively high, and conserved, bank angles (25-35 deg). The bank angle decreased with increasing altitude, in a manner that was broadly consistent with a strategy of maximising the rate of climb. However, the lift coefficients estimated in our study were lower than those predicted by theoretical models and wind-tunnel studies. Overall, our results highlight how the relevant currency for soaring performance changes within individual climbs: when thermal radius is limiting, birds vary bank angle and maintain a constant airspeed, but speed increases later in the climb in order to respond to decreasing air density., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2018. Published by The Company of Biologists Ltd.)

Birds, particularly raptors, are believed to forage primarily using visual cues. However, raptor foraging tactics are highly diverse - from chasing mobile prey to scavenging - which may reflect adaptations of their visual systems. To investigate this, we studied the visual field configuration of 15 species of diurnal Accipitriformes that differ in such tactics, first focusing on the binocular field and blind area by using a single-traits approach, and then exploring the shape of the binocular field with a morphometric approach. While the maximum binocular field width did not differ between species with different foraging tactics, the overall shape of their binocular fields did. In particular, raptors chasing terrestrial prey (ground predators) had a more protruding binocular field and a wider blind area above the head than did raptors chasing aerial or aquatic prey and obligate scavengers. Ground predators that forage on mammals from above have a wide but short bill - which increases ingestion rate - and a large suborbital ridge to avoid sun glare. This may explain the protruding binocular field and the wide blind area above the head. By contrast, species from the two other groups have long but narrow bills used to pluck, flake or tear food and may need large visual coverage (and reduced suborbital ridges) to increase their foraging efficiency (e.g. using large visual coverage to follow the escaping prey in three dimensions or detect conspecifics). We propose that binocular field shape is associated with bill and suborbital ridge shape and, ultimately, foraging strategies., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2018. Published by The Company of Biologists Ltd.)

Video filmed by a camera mounted on the head of a Northern Goshawk (Accipiter gentilis) was used to study how the raptor used visual guidance to pursue prey and land on perches. A combination of novel image analysis methods and numerical simulations of mathematical pursuit models was used to determine the goshawk's pursuit strategy. The goshawk flew to intercept targets by fixing the prey at a constant visual angle, using classical pursuit for stationary prey, lures or perches, and usually using constant absolute target direction (CATD) for moving prey. Visual fixation was better maintained along the horizontal than vertical direction. In some cases, we observed oscillations in the visual fix on the prey, suggesting that the goshawk used finite-feedback steering. Video filmed from the ground gave similar results. In most cases, it showed goshawks intercepting prey using a trajectory consistent with CATD, then turning rapidly to attack by classical pursuit; in a few cases, it showed them using curving non-CATD trajectories. Analysis of the prey's evasive tactics indicated that only sharp sideways turns caused the goshawk to lose visual fixation on the prey, supporting a sensory basis for the surprising frequency and effectiveness of this tactic found by previous studies. The dynamics of the prey's looming image also suggested that the goshawk used a tau-based interception strategy. We interpret these results in the context of a concise review of pursuit-evasion in biology, and conjecture that some prey deimatic 'startle' displays may exploit tau-based interception., (© 2015. Published by The Company of Biologists Ltd.)

This study reports on experiments on falcons wearing miniature videocameras mounted on their backs or heads while pursuing flying prey. Videos of hunts by a gyrfalcon (Falco rusticolus), gyrfalcon (F. rusticolus)/Saker falcon (F. cherrug) hybrids and peregrine falcons (F. peregrinus) were analyzed to determine apparent prey positions on their visual fields during pursuits. These video data were then interpreted using computer simulations of pursuit steering laws observed in insects and mammals. A comparison of the empirical and modeling data indicates that falcons use cues due to the apparent motion of prey on the falcon's visual field to track and capture flying prey via a form of motion camouflage. The falcons also were found to maintain their prey's image at visual angles consistent with using their shallow fovea. These results should prove relevant for understanding the co-evolution of pursuit and evasion, as well as the development of computer models of predation and the integration of sensory and locomotion systems in biomimetic robots.

Integrating biomechanics, behavior and ecology requires a mechanistic understanding of the processes producing the movement of animals. This calls for contemporaneous biomechanical, behavioral and environmental data along movement pathways. A recently formulated unifying movement ecology paradigm facilitates the integration of existing biomechanics, optimality, cognitive and random paradigms for studying movement. We focus on the use of tri-axial acceleration (ACC) data to identify behavioral modes of GPS-tracked free-ranging wild animals and demonstrate its application to study the movements of griffon vultures (Gyps fulvus, Hablizl 1783). In particular, we explore a selection of nonlinear and decision tree methods that include support vector machines, classification and regression trees, random forest methods and artificial neural networks and compare them with linear discriminant analysis (LDA) as a baseline for classifying behavioral modes. Using a dataset of 1035 ground-truthed ACC segments, we found that all methods can accurately classify behavior (80-90%) and, as expected, all nonlinear methods outperformed LDA. We also illustrate how ACC-identified behavioral modes provide the means to examine how vulture flight is affected by environmental factors, hence facilitating the integration of behavioral, biomechanical and ecological data. Our analysis of just over three-quarters of a million GPS and ACC measurements obtained from 43 free-ranging vultures across 9783 vulture-days suggests that their annual breeding schedule might be selected primarily in response to seasonal conditions favoring rising-air columns (thermals) and that rare long-range forays of up to 1750 km from the home range are performed despite potentially heavy energetic costs and a low rate of food intake, presumably to explore new breeding, social and long-term resource location opportunities.

Raptors exhibit a diversity of strategies to procure their prey but ultimately kill using their beaks and/or talons. Thus, bite and grip forces are ecologically important variables that have direct survival implications. Whereas hawks rely primarily on their feet for killing prey, falcons tend to employ their beaks. Consequently, falcons are expected to achieve relatively greater bite forces, and hawks are expected to generate relatively greater grip forces. Force estimates predicted from musculoskeletal morphology in a previous study indicated that falcons (Falco spp.) possess greater jaw force capabilities than accipiters (Accipiter spp.) but there were no clear differences in predicted grip-force capacity outside of differences in scaling. The objective of this study was to complement those results with measurements of in vivo forces by inducing captive and wild accipiters and falcons to bite and grasp force transducers. Bite force increased isometrically in both groups whereas grip force tended toward positive allometry. After adjusting for body mass, falcons produced greater bite forces, and accipiters produced greater grip forces. Thus, previous anatomical estimates of forces predicted the expected direction and magnitude of differences in bite forces but the overall greater in vivo grip forces of accipiters deviated from the pattern obtained from biomechanical estimates. Although the scaling relationships were similar between data sets, forces generated by live birds were consistently lower than those predicted from biomechanics. Estimated and in vivo jaw and digital forces were nevertheless correlated, and therefore provide an important link between morphology and killing behavior in these raptors.

Mothers can strongly influence the development of their offspring, and if maternal resources are limited, they may influence optimal reproductive strategies. In birds, maternally deposited carotenoids are a prominent component of egg yolk and are vital for the development of the embryo. However, results of long-lasting fitness consequences of this early nutritional environment have been scarce and inconsistent. In addition, sex-biased sensitivity to different egg components is one of the mechanisms postulated to account for sex-linked environmental vulnerability during early life. However, this important aspect is usually not accounted for when investigating maternal investment in carotenoids. In this study we gave carotenoid (lutein) supplements to female Eurasian kestrels (Falco tinnunculus) before and during egg laying. The experiment increased female plasma carotenoids, but this effect was not apparent in hatchling and fledgling plasma carotenoid concentration. Also, results showed that carotenoid supplementation increased the high density lipoprotein to low density lipoprotein ratio in adult females, suggesting that dietary carotenoids may influence lipid metabolism. Furthermore, the effect of the treatment was manifested in several nestling health state parameters. Nestlings of carotenoid-supplemented females were infested by less intestinal parasite groups, had higher lymphocyte concentrations in blood plasma, and were less stressed (heterophile to lymphocyte ratio) than control nestlings. In addition, an interaction between the experimental treatment and nestling sex was apparent for globulin concentrations, favouring the smaller male nestlings. Thereby, suggesting that males benefited more than females from an increase in maternal carotenoid investment. Our study shows that an increase in carotenoids in the maternal diet during egg laying favours nestling development in kestrels, and may also affect nestlings in a sex-specific way.

It is recognized that carotenoids are useful anti-oxidants in embryo and hatchling avian models. However, recent evidence suggests that the anti-oxidant role of carotenoids in nestling or adult birds may not be as important as previously thought. The aim of the present work was to investigate if supplemental carotenoids decreased the level of oxidative damage (by reactive oxygen metabolites, ROMs) and increased the serum anti-oxidant capacity (OXY) in nestling Eurasian kestrels Falco tinnunculus. Circulating carotenoids in supplemented nestlings increased about 1.5-fold compared to the control and pre-treatment levels at the end of the supplementation period. There was no effect on ROMs, OXY or the level of oxidative stress (ratio between ROMs and OXY), however, or on body mass or body condition of nestlings. ROMs and OXY decreased with age, but this pattern varied across the nests. Our results show that (i) in general, younger nestlings actually have to cope with a high free radical production, and (ii) the ability of wild nestling kestrels to cope with oxidative stress is not affected by carotenoid availability.