Search

Aerodynamic models of bird flight, assuming power minimization, predict a quadratic relationship (i.e. U-shaped curve) between flapping frequency and airspeed. This relationship is supported by experimental bird flight data from wind tunnels, but the degree to which it characterizes natural flight, and the extent to which birds might modify it to accommodate other behaviors, is less known. We hypothesized that the U-shaped relationship would vary or vanish when minimizing power is not a primary consideration. We analyzed videos of wild cliff swallows (Petrochelidon pyrrhonota) engaged in solo and tandem (i.e. following or being followed by a conspecific) flights to collect bird flapping frequencies and airspeeds. Solo birds had a U-shaped flapping frequency to speed relationship. Birds engaged in tandem flights had the opposite pattern; their flapping frequencies varied with speed as an inverse U-shaped curve and were up to 2.1 times higher than solo birds at the same speed., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2025. Published by The Company of Biologists.)

Insects must fly in highly variable natural environments filled with gusts, vortices, and other transient aerodynamic phenomena that challenge flight stability. Furthermore, the aerodynamic forces that support insect flight are produced from rapidly oscillating wings of time-varying orientation and configuration. The instantaneous flight forces produced by these wings are large relative to the average forces supporting body weight. The magnitude of these forces and their time-varying direction add another challenge to flight stability, because even proportionally small asymmetries in timing or magnitude between the left and right wings may be sufficient to produce large changes in body orientation. However, these same large-magnitude oscillating forces also offer an opportunity for unexpected flight stability through nonlinear interactions between body orientation, body oscillation in response to time-varying inertial and aerodynamic forces, and the oscillating wings themselves. Understanding the emergent stability properties of flying insects is a crucial step toward understanding the requirements for evolution of flapping flight and decoding the role of sensory feedback in flight control. Here, we provide a brief review of insect flight stability, with some emphasis on stability effects brought about by oscillating wings, and present some preliminary experimental data probing some aspects of flight stability in free-flying insects., (© The Author(s) 2024. Published by Oxford University Press on behalf of the Society for Integrative and Comparative Biology.)

Birds commonly exploit environmental features such as columns of rising air and vertical windspeed gradients to lower the cost of flight. These environmental subsidies may be especially important for birds that forage via continuous flight, as seen in black skimmers. These birds forage through a unique behavior, called skimming, where they fly above the water surface with their mandible lowered into the water, catching fish on contact. Thus, their foraging flight incurs costs of moving through both air and water. Prior studies of black skimmer flight behavior have focused on reductions in flight cost due to ground effect, but ignored potential beneficial interactions with the surrounding air. We hypothesized a halfpipe skimming strategy for skimmers to reduce the foraging cost by taking advantage of the wind gradient, where the skimmers perform a wind gradient energy extraction maneuver at the end of a skimming bout through a foraging patch. Using video recordings, wind speed and wind direction measurements, we recorded 70 bird tracks over 4 days at two field sites on the North Carolina coast. We found that while ascending, the skimmers flew more upwind and then flew more downwind when descending, a pattern consistent with harvesting energy from the wind gradient. The strength of the wind gradient and flight behavior of the skimmers indicate that the halfpipe skimming strategy could reduce foraging cost by up to 2.5%., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2024. Published by The Company of Biologists Ltd.)

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.)

Many bird species commonly aggregate in flocks for reasons ranging from predator defense to navigation. Available evidence suggests that certain types of flocks-the V and echelon formations of large birds-may provide a benefit that reduces the aerodynamic cost of flight, whereas cluster flocks typical of smaller birds may increase flight costs. However, metabolic flight costs have not been directly measured in any of these group flight contexts [Zhang and Lauder, J. Exp. Biol. 226 , jeb245617 (2023)]. Here, we measured the energetic benefits of flight in small groups of two or three birds and the requirements for realizing those benefits, using metabolic energy expenditure and flight position measurements from European Starlings flying in a wind tunnel. The starlings continuously varied their relative position during flights but adopted a V formation motif on average, with a modal spanwise and streamwise spacing of [0.81, 0.91] wingspans. As measured via CO 2 production, flight costs for follower birds were significantly reduced compared to their individual solo flight benchmarks. However, followers with more positional variability with respect to leaders did less well, even increasing their costs above solo flight. Thus, we directly demonstrate energetic costs and benefits for group flight followers in an experimental context amenable to further investigation of the underlying aerodynamics, wake interactions, and bird characteristics that produce these metabolic effects., Competing Interests: Competing interests statement:The authors declare no competing interest.

The physical principles that govern the function of biological structures also mediate their evolution, but the evolutionary drivers of morphological traits within complex structures can be difficult to predict. Here, we use morphological traits measured from 1096 3-dimensional bird wing scans from 178 species to test the interaction of two frameworks for relating morphology to evolution. We examine whether the evolutionary rate (σ 2 ) and mode is dominated by the modular organization of the wing into handwing and armwing regions, and/or the relationship between trait morphology and functional output (i.e. mechanical sensitivity, driven here by flapping flight aerodynamics). Our results support discretization of the armwing and handwing as morphological modules, but morphological disparity and σ 2 varied continuously with the mechanical sensitivity gradient and were not modular. Thus, mechanical sensitivity should be considered an independent and fundamental driver of evolutionary dynamics in biomechanical traits, distinct from morphological modularity., (© 2023. The Author(s).)

Flight is an efficient way of transport over a unit of distance, but it can be very costly over each unit of time, and reducing flight energy expenditure is a major selective pressure in birds. The common swift (Apus apus) is one of the most aerial bird species, performing most behaviours in flight: foraging, sleeping and also drinking by regularly descending to various waterbodies and skimming over the surface. An energy-saving way to perform such touch-and-go drinking would be to strive to conserve mechanical energy, by transforming potential energy to kinetic energy during the gliding descent, touching water at high speed, and regaining height with minimal muscular work. Using 3D optical tracking, we recorded 163 swift drinking trajectories, over three waterbodies near Rennes, France. Contrary to the energy conservation hypothesis, we show that swifts approaching a waterbody with a higher mechanical energy (higher height and/or speed 5 s before contact) do not reach the water at higher speeds, but do brake, i.e. dissipate mechanical energy to lose both height and speed. Braking seems to be linked with sharp turns and the use of headwind to some extent, but finer turns and postural adjustments, beyond the resolving power of our tracking data, could also be involved. We hypothesize that this surprisingly costly behaviour results from a trade-off between energy expenditure and safety, because approaching a water surface requires fine motor control, and high speed increases the risk of falling into the water, which would have serious energetic and survival costs for a swift., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2023. Published by The Company of Biologists Ltd.)

A new study of flight control in Drosophila using neurogenetic methods and a virtual reality flight arena has revealed a group of descending neurons that fully activate the flight motor and steer the fly by independent regulation of the left and right wings., Competing Interests: Declaration of interests The authors declare no competing interests., (Copyright © 2022 Elsevier Inc. All rights reserved.)

Gliding animals change their body shape and posture while producing and modulating aerodynamic forces during flight. However, the combined effect of these different factors on aerodynamic force production, and ultimately the animal's gliding ability, remains uncertain. Here, we quantified the time-varying morphology and aerodynamics of complete, voluntary glides performed by a population of wild gliding lizards (Draco dussumieri) in a seven-camera motion capture arena constructed in their natural environment. Our findings, in conjunction with previous airfoil models, highlight how three-dimensional (3D) wing shape including camber, planform, and aspect ratio enables gliding flight and effective aerodynamic performance by the lizard up to and over an angle of attack (AoA) of 55° without catastrophic loss of lift. Furthermore, the lizards maintained a near maximal lift-to-drag ratio throughout their mid-glide by changing body pitch to control AoA, while simultaneously modulating airfoil camber to alter the magnitude of aerodynamic forces. This strategy allows an optimal aerodynamic configuration for horizontal transport while ensuring adaptability to real-world flight conditions and behavioral requirements. Overall, we empirically show that the aerodynamics of biological airfoils coupled with the animal's ability to control posture and their 3D wing shape enable efficient gliding and adaptive flight control in the natural habitat., (© 2022. The Author(s).)

Crepuscular mosquitoes, which swarm in low light conditions, exhibit a range of adaptations including large aspect-ratio wings, high flapping frequencies and small stroke amplitudes that taken together, facilitate the generation of wing-tones that are well-suited for acoustic communication. In the current study, we employ computational aeroacoustic modeling to conduct a comparative study of wing-tone and flight efficiency in a mosquito (male Culex ) and a similar sized flying insect: a fruit fly ( Drosophila ). Based on this analysis, we show that pound-for-pound, a mosquito generates wing-tones that are a factor of about 3.4 times more intense than a fruit fly, and the mosquito is more efficient by a factor of about 3.7 in converting mechanical power into acoustic power. The wing-tones for the mosquito are also more tilted in the forward direction, a characteristic that would be more conducive for acoustic signaling during a mate chase. The simulation data also shows that the specific power (mechanical power over mean lift) of the mosquito is nearly equal to that of the fruit fly, indicating that the adaptations that facilitate wing-tone based communication in mosquitoes, do not seem to compromise their flight efficiency., (© 2021 IOP Publishing Ltd.)

The evolution of wing morphology among birds, and its functional consequences, remains an open question, despite much attention. This is in part because the connection between form and function is difficult to test directly. To address this deficit, in prior work, we used computational modeling and sensitivity analysis to interrogate the impact of altering wing aspect ratio (AR), camber, and Reynolds number on aerodynamic performance, revealing the performance landscapes that avian evolution has explored. In the present work, we used a dataset of three-dimensionally scanned bird wings coupled with the performance landscapes to test two hypotheses regarding the evolutionary diversification of wing morphology associated with gliding flight behavior: (1) gliding birds would exhibit higher wing AR and greater chordwise camber than their non-gliding counterparts; and (2) that two strategies for gliding flight exist, with divergent morphological conformations. In support of our first hypothesis, we found evidence of morphological divergence in both wing AR and camber between gliders and non-gliders, suggesting that wing morphology of birds that utilize gliding flight is under different selective pressures than the wings of non-gliding taxa. Furthermore, we found that these morphological differences also yielded differences in coefficient of lift measured both at the maximum lift to drag ratio and at minimum sinking speed, with gliding taxa exhibiting higher coefficient of lift in both cases. Minimum sinking speed was also lower in gliders than non-gliders. However, contrary to our hypothesis, we found that the maximum ratio of the coefficient of lift to the coefficient of drag differed between gliders and non-gliders. This may point to the need for gliders to maintain high lift capability for takeoff and landing independent of gliding performance or could be due to the divergence in flight styles among gliders, as not all gliders are predicted to optimize either quantity. However, direct evidence for the existence of two morphologically defined gliding flight strategies was equivocal, with only slightly stronger support for an evolutionary model positing separate morphological optima for these strategies than an alternative model positing a single peak. The absence of a clear result may be an artifact of low statistical power owing to a relatively small sample size of gliding flyers expected to follow the "aerial search" strategy., (© The Author(s) 2020. Published by Oxford University Press on behalf of the Society for Integrative and Comparative Biology. All rights reserved. For permissions please email: journals.permissions@oup.com.)

The physics of flight influences the morphology of bird wings through natural selection on flight performance. The connection between wing morphology and performance is unclear due to the complex relationships between various parameters of flight. In order to better understand this connection, we present a holistic analysis of gliding flight that preserves complex relationships between parameters. We use a computational model of gliding flight, along with analysis by uncertainty quantification, to (1) create performance landscapes of gliding based on output metrics (maximum lift-to-drag ratio, minimum gliding angle, minimum sinking speed, and lift coefficient at minimum sinking speed) and (2) predict what parameters of flight (chordwise camber, wing aspect ratio [AR], and Reynolds number) would differ between gliding and nongliding species of birds. We also examine performance based on the soaring strategy for possible differences in morphology within gliding birds. Gliding birds likely have greater ARs than non-gliding birds, due to the high sensitivity of AR on most metrics of gliding performance. Furthermore, gliding birds can use two distinct soaring strategies based on performance landscapes. First, maximizing distance traveled (maximizing lift-to-drag ratio and minimizing gliding angle) should result in wings with high ARs and middling-to-low wing chordwise camber. Second, maximizing lift extracted from updrafts should result in wings with middling ARs and high wing chordwise camber. Following studies can test these hypotheses using morphological measurements., (© The Author(s) 2020. Published by Oxford University Press on behalf of the Society for Integrative and Comparative Biology. All rights reserved. For permissions please email: journals.permissions@oup.com.)

It is generally accepted among biology and engineering communities that insects are unstable at hover. However, existing approaches that rely on direct averaging do not fully capture the dynamical features and stability characteristics of insect flight. Here, we reveal a passive stabilization mechanism that insects exploit through their natural wing oscillations: vibrational stabilization. This stabilization technique cannot be captured using the averaging approach commonly used in literature. In contrast, it is elucidated using a special type of calculus: the chronological calculus. Our result is supported through experiments on a real hawkmoth subjected to pitch disturbance from hovering. This finding could be particularly useful to biologists because the vibrational stabilization mechanism may also be exploited by many other creatures. Moreover, our results may inspire more optimal designs for bioinspired flying robots by relaxing the feedback control requirements of flight., (Copyright © 2020 The Authors, some rights reserved; exclusive licensee American Association for the Advancement of Science. No claim to original U.S. Government Works.)

Gliding animals traverse cluttered aerial environments when performing ecologically relevant behaviours. However, it is unknown how gliders execute collision-free flight over varying distances to reach their intended target. We quantified complete glide trajectories amid obstacles in a naturally behaving population of gliding lizards inhabiting a rainforest reserve. In this cluttered habitat, the lizards used glide paths with fewer obstacles than alternatives of similar distance. Their takeoff direction oriented them away from obstacles in their path and they subsequently made mid-air turns with accelerations of up to 0.5 g to reorient towards the target tree. These manoeuvres agreed well with a vision-based steering model which maximized their bearing angle with the obstacle while minimizing it with the target tree. Nonetheless, negotiating obstacles reduced mid-glide shallowing rates, implying greater loss of altitude. Finally, the lizards initiated a pitch-up landing manoeuvre consistent with a visual trigger model, suggesting that the landing decision was based on the optical size and speed of the target. They subsequently followed a controlled-collision approach towards the target, ending with variable impact speeds. Overall, the visually guided path planning strategy that enabled collision-free gliding required continuous changes in the gliding kinematics such that the lizards never attained theoretically ideal steady-state glide dynamics.

The generation of sound from flapping (i.e. wing tones) of mosquito (Culex) wings is investigated using computational modeling. The flow field around the wing is simulated by solving the incompressible Navier-Stokes equations using a sharp-interface immersed boundary method, and the aeroacoustic sound is predicted by the Ffowcs Williams and Hawkings equation using data from the aerodynamic simulations. In addition to the aerodynamics, the characteristics of mosquito's wing tone, spectral directivity patterns, and generation mechanisms are investigated. The effects of wing-beat frequency and stroke amplitude are also studied, and scaling analysis for the mean lift, mechanical power, and overall wing tone sound power are performed to understand the effects of the wing shape and kinematics parameters. The analysis shows that the high wing aspect-ratio, high wing beat frequency, and small stroke amplitude adopted by mosquitoes enable efficient generation of high-intensity wing-tones for acoustic communications. The present findings may also apply to the optimized noise control in the flapping-wing micro air vehicles (FWMAV).

Chimney swifts (Chaetura pelagica) are highly aerial, small, insectivorous birds well known for roosting en masse in chimneys during their autumn migration. These roosting events require hundreds to thousands of birds to enter a small opening (here 0.64 m 2 ) within a short amount of time (15-30 min). Thus, these entry events pose a complex navigational and behavioral challenge as birds identify their entry route, cooperate with other birds present to form an entry flock, and compete with other birds at the time of chimney entry. We used six synchronized cameras to capture and reconstruct the 3D flight trajectories of swifts before and during chimney entry. Navigation into the chimney is consistent with use of a relative retinal expansion velocity cue, which results in an entry/non-entry decision point about 1.5 m above the chimney, or 0.4 s at typical entry speeds. Entries were highly clustered with 91 of 136 entries occurring within 1 s of another entry. We observed both synchronous (entry within 0.2 s) and sequential entry behavior (entry separated by ~0.4 s). Birds entering the chimney flew in close proximity to other birds (median minimum distance 0.51 m; 1.7 wingspans). In cases where two birds appeared to attempt a near-simultaneous entry, the bird either slightly to the rear or with a velocity vector bringing it closer to the current position of the other bird tended to make an avoidance maneuver and abandon its entry attempt. Overall, these results show how groups of animals execute complex landing and collision avoidance maneuvers in a natural setting without central control authority.

Pursuit is a common behavior exhibited by animals chasing prey, competitors and potential mates. Because of their speed and maneuverability, dragonflies are frequently studied as a model system for biological pursuit. Most quantitative studies have focused on prey pursuits in captive environments. To determine whether a different pursuit strategy is used when chasing conspecifics of nearly equal speed and agility, we recorded 3D flight trajectories from nine territorial chases between male Erythemis simplicicollis dragonflies in natural field conditions. During chases, dragonflies used an interception strategy with an unusually high-magnitude gain ( k =-10.03 s -1 horizontal; -8.86 s -1 vertical) and short time delay (τ=50 ms). The product k τ determines how aggressively a pursuer corrects course to achieve interception. Previous studies of prey pursuit have found k τ values close to -1/ e (-0.37), the time-optimal value for achieving pursuit without overshooting. However, we found that dragonflies chasing conspecifics use more negative k τ (-0.50 horizontal; -0.44 vertical), resulting in pursuits with a high degree of overshooting (i.e. moving past the target and alternating position from side to side). We confirmed via simulation that the observed gain and delay produce overshooting. We propose that overshooting is an adaptive feature of conspecific chases that can be achieved with only slight modification of the strategy used for intercepting prey. Overshooting might help avoid potentially damaging collisions while exhibiting the pursuing animal's flight performance and competitive ability. Repeated close approaches might also evoke evasive responses from the other dragonfly, effectively herding the competitor out of the territory., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2019. Published by The Company of Biologists Ltd.)

Animal groups have emergent properties that result from simple interactions among individuals. However, we know little about why animals adopt different interaction rules because of sparse sampling among species. Here, we identify an interaction rule that holds across single and mixed-species flocks of four migratory shorebird species spanning a seven-fold range of body masses. The rule, aligning with a one-wingspan lateral distance to nearest neighbors in the same horizontal plane, scales linearly with wingspan but is independent of nearest neighbor distance and neighbor species. This rule propagates outward to create a global flock structure that we term the compound-V formation. We propose that this formation represents an intermediary between the cluster flocks of starlings and the simple-V formations of geese and other large migratory birds. We explore multiple hypotheses regarding the benefit of this flock structure and how it differs from structures observed in other flocking species., Competing Interests: AC, TH No competing interests declared, (© 2019, Corcoran and Hedrick.)

Although the biomechanics of animal flight have been well studied in laboratory apparatus such as wind tunnels for many years, the applicability of these data to natural flight behaviour has been examined in few instances and mostly in the context of long-distance migration. Here, we used rotational stereo-videography to record the free-flight trajectories of foraging common swifts. We found that, despite their exquisite manoeuvring capabilities, the swifts only rarely performed high-acceleration turns. More surprisingly, we also found that despite feeding on tiny insects probably moving with ambient flow, the birds adjust their air speed to optimize cost of transport over distance. Finally, swifts spent only 25% of their time flapping; the majority of their time (71%) was spent in extended wing gliding, during which the average power expended for changes in speed or elevation was 0.84 W kg -1 and not significantly different from 0. Thus, gliding swifts extracted sufficient environmental energy to pay the cost of flight during foraging., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2018. Published by The Company of Biologists Ltd.)

Flight performance is fundamental to the fitness of flying organisms. Whilst airborne, flying organisms face unavoidable wing wear and wing area loss. Many studies have tried to quantify the consequences of wing area loss to flight performance with varied results, suggesting that not all types of damage are equal and different species may have different means to compensate for some forms of wing damage with little to no cost. Here, we investigated the cost of control during hovering flight with damaged wings, specifically wings with asymmetric and symmetric reductions in area, by measuring maximum load lifting capacity and the metabolic power of hovering flight in hawkmoths ( Manduca sexta ). We found that while asymmetric and symmetric reductions are both costly in terms of maximum load lifting and hovering efficiency, asymmetric reductions are approximately twice as costly in terms of wing area lost. The moths also did not modulate flapping frequency and amplitude as predicted by a hovering flight model, suggesting that the ability to do so, possibly tied to asynchronous versus synchronous flight muscles, underlies the varied responses found in different wing clipping experiments., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2017. Published by The Company of Biologists Ltd.)

Chimney swifts ( Chaetura pelagica ) are highly manoeuvrable birds notable for roosting overnight in chimneys, in groups of hundreds or thousands of birds, before and during their autumn migration. At dusk, birds gather in large numbers from surrounding areas near a roost site. The whole flock then employs an orderly, but dynamic, circling approach pattern before rapidly entering a small aperture en masse We recorded the three-dimensional trajectories of ≈1 800 individual birds during a 30 min period encompassing flock formation, circling, and landing, and used these trajectories to test several hypotheses relating to flock or group behaviour. Specifically, we investigated whether the swifts use local interaction rules based on topological distance (e.g. the n nearest neighbours, regardless of their distance) rather than physical distance (e.g. neighbours within x m, regardless of number) to guide interactions, whether the chimney entry zone is more or less cooperative than the surrounding flock, and whether the characteristic subgroup size is constant or varies with flock density. We found that the swift flock is structured around local rules based on physical distance, that subgroup size increases with density, and that there exist regions of the flock that are less cooperative than others, in particular the chimney entry zone., (© 2017 The Authors.)

The theoretical underpinnings for flight, including animal flight with flapping wings, predict a curvilinear U-shaped or J-shaped relationship between flight speed and the power required to maintain that speed. Experimental data have confirmed this relationship for a variety of bird and bat species but not insects, possibly due to differences in aerodynamics and physiology or experimental difficulties. Here we quantify modulation of the main flight motor muscles (the dorsolongitudinal and dorsoventral) via electromyography in hawkmoths (Manduca sexta) flying freely over a range of speeds in a wind tunnel and show that these insects exhibit a U-shaped speed-power relationship, with a minimum power speed of 2ms -1 , indicating that at least large flying insects achieve sufficiently high flight speeds that drag and power become limiting factors., (Copyright © 2016 Elsevier Ltd. All rights reserved.)

The smallest flying insects commonly possess wings with long bristles. Little quantitative information is available on the morphology of these bristles, and their functional importance remains a mystery. In this study, we (1) collected morphological data on the bristles of 23 species of Mymaridae by analyzing high-resolution photographs and (2) used the immersed boundary method to determine via numerical simulation whether bristled wings reduced the force required to fling the wings apart while still maintaining lift. The effects of Reynolds number, angle of attack, bristle spacing and wing-wing interactions were investigated. In the morphological study, we found that as the body length of Mymaridae decreases, the diameter and gap between bristles decreases and the percentage of the wing area covered by bristles increases. In the numerical study, we found that a bristled wing experiences less force than a solid wing. The decrease in force with increasing gap to diameter ratio is greater at higher angles of attack than at lower angles of attack, suggesting that bristled wings may act more like solid wings at lower angles of attack than they do at higher angles of attack. In wing-wing interactions, bristled wings significantly decrease the drag required to fling two wings apart compared with solid wings, especially at lower Reynolds numbers. These results support the idea that bristles may offer an aerodynamic benefit during clap and fling in tiny insects., (© 2016. Published by The Company of Biologists Ltd.)

The superior manoeuvrability of hummingbirds emerges from complex interactions of specialized neural and physiological processes with the unique flight dynamics of flapping wings. Escape manoeuvring is an ecologically relevant, natural behaviour of hummingbirds, from which we can gain understanding into the functional limits of vertebrate locomotor capacity. Here, we extend our kinematic analysis of escape manoeuvres from a companion paper to assess two potential limiting factors of the manoeuvring performance of hummingbirds: (1) muscle mechanical power output and (2) delays in the neural sensing and control system. We focused on the magnificent hummingbird (Eugenes fulgens, 7.8 g) and the black-chinned hummingbird (Archilochus alexandri, 3.1 g), which represent large and small species, respectively. We first estimated the aerodynamic forces, moments and the mechanical power of escape manoeuvres using measured wing kinematics. Comparing active-manoeuvring and passive-damping aerodynamic moments, we found that pitch dynamics were lightly damped and dominated by the effect of inertia, while roll dynamics were highly damped. To achieve observed closed-loop performance, pitch manoeuvres required faster sensorimotor transduction, as hummingbirds can only tolerate half the delay allowed in roll manoeuvres. Accordingly, our results suggested that pitch control may require a more sophisticated control strategy, such as those based on prediction. For the magnificent hummingbird, we estimated that escape manoeuvres required muscle mass-specific power 4.5 times that during hovering. Therefore, in addition to the limitation imposed by sensorimotor delays, muscle power could also limit the performance of escape manoeuvres., (© 2016. Published by The Company of Biologists Ltd.)

Hummingbirds are nature's masters of aerobatic manoeuvres. Previous research shows that hummingbirds and insects converged evolutionarily upon similar aerodynamic mechanisms and kinematics in hovering. Herein, we use three-dimensional kinematic data to begin to test for similar convergence of kinematics used for escape flight and to explore the effects of body size upon manoeuvring. We studied four hummingbird species in North America including two large species (magnificent hummingbird, Eugenes fulgens, 7.8 g, and blue-throated hummingbird, Lampornis clemenciae, 8.0 g) and two smaller species (broad-billed hummingbird, Cynanthus latirostris, 3.4 g, and black-chinned hummingbirds Archilochus alexandri, 3.1 g). Starting from a steady hover, hummingbirds consistently manoeuvred away from perceived threats using a drastic escape response that featured body pitch and roll rotations coupled with a large linear acceleration. Hummingbirds changed their flapping frequency and wing trajectory in all three degrees of freedom on a stroke-by-stroke basis, likely causing rapid and significant alteration of the magnitude and direction of aerodynamic forces. Thus it appears that the flight control of hummingbirds does not obey the 'helicopter model' that is valid for similar escape manoeuvres in fruit flies. Except for broad-billed hummingbirds, the hummingbirds had faster reaction times than those reported for visual feedback control in insects. The two larger hummingbird species performed pitch rotations and global-yaw turns with considerably larger magnitude than the smaller species, but roll rates and cumulative roll angles were similar among the four species., (© 2016. Published by The Company of Biologists Ltd.)

While prior studies of swallow manoeuvering have focused on slow-speed flight and obstacle avoidance in still air, swallows survive by foraging at high speeds in windy environments. Recent advances in field-portable, high-speed video systems, coupled with precise anemometry, permit measures of high-speed aerial performance of birds in a natural state. We undertook the present study to test: (i) the manner in which barn swallows (Hirundo rustica) may exploit wind dynamics and ground effect while foraging and (ii) the relative importance of flapping versus gliding for accomplishing high-speed manoeuvers. Using multi-camera videography synchronized with wind-velocity measurements, we tracked coursing manoeuvers in pursuit of prey. Wind speed averaged 1.3-2.0 m s(-1) across the atmospheric boundary layer, exhibiting a shear gradient greater than expected, with instantaneous speeds of 0.02-6.1 m s(-1) While barn swallows tended to flap throughout turns, they exhibited reduced wingbeat frequency, relying on glides and partial bounds during maximal manoeuvers. Further, the birds capitalized on the near-earth wind speed gradient to gain kinetic and potential energy during both flapping and gliding turns; providing evidence that such behaviour is not limited to large, fixed-wing soaring seabirds and that exploitation of wind gradients by small aerial insectivores may be a significant aspect of their aeroecology.This article is part of the themed issue 'Moving in a moving medium: new perspectives on flight'., (© 2016 The Author(s).)

We used computational fluid dynamics to determine whether lift- or drag-based mechanisms generate the most vertical force in the flight of the smallest insects. These insects fly at Re on the order of 4-60 where viscous effects are significant. Detailed quantitative data on the wing kinematics of the smallest insects is not available, and as a result both drag- and lift-based strategies have been suggested as the mechanisms by which these insects stay aloft. We used the immersed boundary method to solve the fully-coupled fluid-structure interaction problem of a flexible wing immersed in a two-dimensional viscous fluid to compare three idealized hovering kinematics: a drag-based stroke in the vertical plane, a lift-based stroke in the horizontal plane, and a hybrid stroke on a tilted plane. Our results suggest that at higher Re, a lift-based strategy produces more vertical force than a drag-based strategy. At the Re pertinent to small insect hovering, however, there is little difference in performance between the two strategies. A drag-based mechanism of flight could produce more vertical force than a lift-based mechanism for insects at Re<5; however, we are unaware of active fliers at this scale., (Copyright © 2015 Elsevier Ltd. All rights reserved.)

Despite intense study by physicists and biologists, we do not fully understand the unsteady aerodynamics that relate insect wing morphology and kinematics to lift generation. Here, we formulate a force partitioning method (FPM) and implement it within a computational fluid dynamic model to provide an unambiguous and physically insightful division of aerodynamic force into components associated with wing kinematics, vorticity, and viscosity. Application of the FPM to hawkmoth and fruit fly flight shows that the leading-edge vortex is the dominant mechanism for lift generation for both these insects and contributes between 72-85% of the net lift. However, there is another, previously unidentified mechanism, the centripetal acceleration reaction, which generates up to 17% of the net lift. The centripetal acceleration reaction is similar to the classical inviscid added-mass in that it depends only on the kinematics (i.e. accelerations) of the body, but is different in that it requires the satisfaction of the no-slip condition, and a combination of tangential motion and rotation of the wing surface. Furthermore, the classical added-mass force is identically zero for cyclic motion but this is not true of the centripetal acceleration reaction. Furthermore, unlike the lift due to vorticity, centripetal acceleration reaction lift is insensitive to Reynolds number and to environmental flow perturbations, making it an important contributor to insect flight stability and miniaturization. This force mechanism also has broad implications for flow-induced deformation and vibration, underwater locomotion and flows involving bubbles and droplets.

Hummingbirds are known to defend food resources such as nectar sources from encroachment by competitors (including conspecifics). These competitive intraspecific interactions provide an opportunity to quantify the biomechanics of hummingbird flight performance during ecologically relevant natural behavior. We recorded the three-dimensional flight trajectories of Ruby-throated Hummingbirds defending, being chased from and freely departing from a feeder. These trajectories allowed us to compare natural flight performance to earlier laboratory measurements of maximum flight speed, aerodynamic force generation and power estimates. During field observation, hummingbirds rarely approached the maximal flight speeds previously reported from wind tunnel tests and never did so during level flight. However, the accelerations and rates of change in kinetic and potential energy we recorded indicate that these hummingbirds likely operated near the maximum of their flight force and metabolic power capabilities during these competitive interactions. Furthermore, although birds departing from the feeder while chased did so faster than freely-departing birds, these speed gains were accomplished by modulating kinetic and potential energy gains (or losses) rather than increasing overall power output, essentially trading altitude for speed during their evasive maneuver. Finally, the trajectories of defending birds were directed toward the position of the encroaching bird rather than the feeder.

In hovering flight, hummingbirds reverse the angle of attack of their wings through pitch reversal in order to generate aerodynamic lift during both downstroke and upstroke. In addition, the wings may pitch during translation to further enhance lift production. It is not yet clear whether these pitching motions are caused by the wing inertia or actuated through the musculoskeletal system. Here we perform a computational analysis of the pitching dynamics by incorporating the realistic wing kinematics to determine the inertial effects. The aerodynamic effect is also included using the pressure data from a previous three-dimensional computational fluid dynamics simulation of a hovering hummingbird. The results show that like many insects, pitch reversal of the hummingbird is, to a large degree, caused by the wing inertia. However, actuation power input at the root is needed in the beginning of pronation to initiate a fast pitch reversal and also in mid-downstroke to enable a nose-up pitching motion for lift enhancement. The muscles on the wing may not necessarily be activated for pitching of the distal section. Finally, power analysis of the flapping motion shows that there is no requirement for substantial elastic energy storage or energy absorption at the shoulder joint.

The aerodynamics of flapping flight for the smallest insects such as thrips is often characterized by a 'clap and fling' of the wings at the end of the upstroke and the beginning of the downstroke. These insects fly at Reynolds numbers (Re) of the order of 10 or less where viscous effects are significant. Although this wing motion is known to augment the lift generated during flight, the drag required to fling the wings apart at this scale is an order of magnitude larger than the corresponding force acting on a single wing. As the opposing forces acting normal to each wing nearly cancel during the fling, these large forces do not have a clear aerodynamic benefit. If flight efficiency is defined as the ratio of lift to drag, the clap and fling motion dramatically reduces efficiency relative to the case of wings that do not aerodynamically interact. In this paper, the effect of a bristled wing characteristic of many of these insects was investigated using computational fluid dynamics. We performed 2D numerical simulations using a porous version of the immersed boundary method. Given the computational complexity involved in modeling flow through exact descriptions of bristled wings, the wing was modeled as a homogeneous porous layer as a first approximation. High-speed video recordings of free-flying thrips in take-off flight were captured in the laboratory, and an analysis of the wing kinematics was performed. This information was used for the estimation of input parameters for the simulations. Compared with a solid wing (without bristles), the results of the study show that the porous nature of the wings contributes largely to drag reduction across the Re range explored. The aerodynamic efficiency, calculated as the ratio of lift to drag coefficients, was larger for some porosities when compared with solid wings., (© 2014. Published by The Company of Biologists Ltd.)

A three-dimensional computational fluid dynamics simulation is performed for a ruby-throated hummingbird (Archilochus colubris) in hovering flight. Realistic wing kinematics are adopted in the numerical model by reconstructing the wing motion from high-speed imaging data of the bird. Lift history and the three-dimensional flow pattern around the wing in full stroke cycles are captured in the simulation. Significant asymmetry is observed for lift production within a stroke cycle. In particular, the downstroke generates about 2.5 times as much vertical force as the upstroke, a result that confirms the estimate based on the measurement of the circulation in a previous experimental study. Associated with lift production is the similar power imbalance between the two half strokes. Further analysis shows that in addition to the angle of attack, wing velocity and surface area, drag-based force and wing-wake interaction also contribute significantly to the lift asymmetry. Though the wing-wake interaction could be beneficial for lift enhancement, the isolated stroke simulation shows that this benefit is buried by other opposing effects, e.g. presence of downwash. The leading-edge vortex is stable during the downstroke but may shed during the upstroke. Finally, the full-body simulation result shows that the effects of wing-wing interaction and wing-body interaction are small., (© 2014 The Author(s) Published by the Royal Society. All rights reserved.)

Cliff swallows (Petrochelidon pyrrhonota) are highly maneuverable social birds that often forage and fly in large open spaces. Here we used multi-camera videography to measure the three-dimensional kinematics of their natural flight maneuvers in the field. Specifically, we collected data on tandem flights, defined as two birds maneuvering together. These data permit us to evaluate several hypotheses on the high-speed maneuvering flight performance of birds. We found that high-speed turns are roll-based, but that the magnitude of the centripetal force created in typical maneuvers varied only slightly with flight speed, typically reaching a peak of ~2 body weights. Turning maneuvers typically involved active flapping rather than gliding. In tandem flights the following bird copied the flight path and wingbeat frequency (~12.3 Hz) of the lead bird while maintaining position slightly above the leader. The lead bird turned in a direction away from the lateral position of the following bird 65% of the time on average. Tandem flights vary widely in instantaneous speed (1.0 to 15.6 m s(-1)) and duration (0.72 to 4.71 s), and no single tracking strategy appeared to explain the course taken by the following bird., (© 2014. Published by The Company of Biologists Ltd.)

Stereo videography is a powerful technique for quantifying the kinematics and behavior of animals, but it can be challenging to use in an outdoor field setting. We here present a workflow and associated software for performing calibration of cameras placed in a field setting and estimating the accuracy of the resulting stereoscopic reconstructions. We demonstrate the workflow through example stereoscopic reconstructions of bat and bird flight. We provide software tools for planning experiments and processing the resulting calibrations that other researchers may use to calibrate their own cameras. Our field protocol can be deployed in a single afternoon, requiring only short video clips of light, portable calibration objects., (© 2014. Published by The Company of Biologists Ltd.)

Vertical vortex systems such as tornadoes dramatically affect the flight control and stability of aircraft. However, the control implications of smaller scale vertically oriented vortex systems for small fliers such as animals or micro-air vehicles are unknown. Here we examined the flapping kinematics and body dynamics of hawkmoths performing hovering flights (controls) and maintaining position in three different whirlwind intensities with transverse horizontal velocities of 0.7, 0.9 and 1.2 m s(-1), respectively, generated in a vortex chamber. The average and standard deviation of yaw and pitch were respectively increased and reduced in comparison with hovering flights. Average roll orientation was unchanged in whirlwind flights but was more variable from wingbeat to wingbeat than in hovering. Flapping frequency remained unchanged. Wingbeat amplitude was lower and the average stroke plane angle was higher. Asymmetry was found in the angle of attack between right and left wings during both downstroke and upstroke at medium and high vortex intensities. Thus, hawkmoth flight control in tornado-like vortices is achieved by a suite of asymmetric and symmetric changes to wingbeat amplitude, stroke plane angle and principally angle of attack.

Shedding of vortices is a common phenomenon in the atmosphere over a wide range of spatial and temporal scales. However, it is unclear how these vortices of varying scales affect the flight performance of flying animals. In order to examine these interactions, we trained seven hawkmoths (Manduca sexta) (wingspan ~9 cm) to fly and feed in a wind tunnel under steady flow (controls) and in the von Kármán vortex street of vertically oriented cylinders (two different cylinders with diameters of 10 and 5 cm) at speeds of 0.5, 1 and 2 m s(-1). Cylinders were placed at distances of 5, 25 and 100 cm upstream of the moths. Moths exhibited large amplitude yaw oscillations coupled with modest oscillations in roll and pitch, and slight increases in wingbeat frequency when flying in both the near (recirculating) and middle (vortex dominated) wake regions. Wingbeat amplitude did not vary among treatments, except at 1 m s(-1) for the large cylinder. Yaw and roll oscillations were synchronized with the vortex shedding frequencies in moths flying in the wake of the large cylinder at all speeds. In contrast, yaw and pitch were synchronized with the shedding frequency of small vortices at speeds ≤1 m s(-1). Oscillations in body orientation were also substantially smaller in the small cylinder treatment when compared with the large cylinder, regardless of temporal or non-dimensional spatial scale. Moths flying in steady conditions reached a higher air speed than those flying into cylinder wakes. In general, flight effects produced by the cylinder wakes were qualitatively similar among the recirculating and vortex-dominated wake regions; the magnitude of those effects, however, declined gradually with downstream distance.

Insect wings can undergo significant chordwise (camber) as well as spanwise (twist) deformation during flapping flight but the effect of these deformations is not well understood. The shape and size of butterfly wings leads to particularly large wing deformations, making them an ideal test case for investigation of these effects. Here we use computational models derived from experiments on free-flying butterflies to understand the effect of time-varying twist and camber on the aerodynamic performance of these insects. High-speed videogrammetry is used to capture the wing kinematics, including deformation, of a Painted Lady butterfly (Vanessa cardui) in untethered, forward flight. These experimental results are then analyzed computationally using a high-fidelity, three-dimensional, unsteady Navier-Stokes flow solver. For comparison to this case, a set of non-deforming, flat-plate wing (FPW) models of wing motion are synthesized and subjected to the same analysis along with a wing model that matches the time-varying wing-twist observed for the butterfly, but has no deformation in camber. The simulations show that the observed butterfly wing (OBW) outperforms all the flat-plate wings in terms of usable force production as well as the ratio of lift to power by at least 29% and 46%, respectively. This increase in efficiency of lift production is at least three-fold greater than reported for other insects. Interestingly, we also find that the twist-only-wing (TOW) model recovers much of the performance of the OBW, demonstrating that wing-twist, and not camber is key to forward flight in these insects. The implications of this on the design of flapping wing micro-aerial vehicles are discussed.

Wing damage is common in flying insects and has been studied using a variety of approaches to assess its biomechanical and fitness consequences. Results of these studies range from strong to nil effect among the variety of species, fitness measurements and damage modes studied, suggesting that not all damage modes are equal and that insects may be well adapted to compensate for some types of damage. Here, we examine the biomechanical and neuromuscular means by which flying insects compensate for asymmetric wing damage, which is expected to produce asymmetric flight forces and torques and thus destabilize the animal in addition to reducing its total wing size. We measured the kinematic and neuromuscular responses of hawkmoths (Manduca sexta) hovering in free flight with asymmetrically damaged wings via high-speed videography and extracellular neuromuscular activity recordings. The animals responded to asymmetric wing damage with asymmetric changes to wing stroke amplitude sufficient to restore symmetry in lift production. These asymmetries in stroke amplitude were significantly correlated with bilateral asymmetries in the timing of activation of the dorsal ventral muscle among and within trials. Correspondingly, the magnitude of wing asymmetry was significantly, although non-linearly, correlated with the magnitude of the neuromuscular response among individuals. The strongly non-linear nature of the relationship suggests that active neural compensation for asymmetric wing damage may only be necessary above a threshold (>12% asymmetry in wing second moment of area in this case) below which passive mechanisms may be adequate to maintain flight stability.

Hummingbirds (Trochilidae) are widely known for their insect-like flight strokes characterized by high wing beat frequency, small muscle strains and a highly supinated wing orientation during upstroke that allows for lift production in both halves of the stroke cycle. Here, we show that hummingbirds achieve these functional traits within the limits imposed by a vertebrate endoskeleton and muscle physiology by accentuating a wing inversion mechanism found in other birds and using long-axis rotational movement of the humerus. In hummingbirds, long-axis rotation of the humerus creates additional wing translational movement, supplementing that produced by the humeral elevation and depression movements of a typical avian flight stroke. This adaptation increases the wing-to-muscle-transmission ratio, and is emblematic of a widespread scaling trend among flying animals whereby wing-to-muscle-transmission ratio varies inversely with mass, allowing animals of vastly different sizes to accommodate aerodynamic, biomechanical and physiological constraints on muscle-powered flapping flight.

Flying animals exhibit remarkable degrees of both stability and manoeuvrability. Our understanding of these capabilities has recently been improved by the identification of a source of passive damping specific to flapping flight. Examining how this damping effect scales among different species and how it affects active manoeuvres as well as recovery from perturbations provides general insights into the flight of insects, birds and bats. These new damping models offer a means to predict manoeuvrability and stability for a wide variety of flying animals using prior reports of the morphology and flapping motions of these species. Furthermore, the presence of passive damping is likely to have facilitated the evolution of powered flight in animals by providing a stability benefit associated with flapping.

Insects produce a variety of exquisitely controlled manoeuvres during natural flight behaviour. Here we show how hawkmoths produce and control one such manoeuvre, an avoidance response consisting of rapid pitching up, rearward flight, pitching down (often past the original pitch angle), and then pitching up slowly to equilibrium. We triggered these manoeuvres via a sudden visual stimulus in front of free-flying hawkmoths (Manduca sexta) while recording the animals' body and wing movements via high-speed stereo videography. We then recreated the wing motions in a dynamically scaled model to: (1) associate wing kinematic changes with pitch torque production and (2) extract the open-loop dynamics of an uncontrolled moth. Next, we characterized the closed-loop manoeuvring dynamics from the observed flight behaviour assuming that hawkmoths use feedback control based on translational velocity, pitch angle and angular velocity, and then compared these with the open-loop dynamics to identify the control strategy used by the moth. Our analysis revealed that hawkmoths produce active pitch torque via changes in mean wing spanwise rotation angle. Additionally, body translations produce passive translational damping and pitch torque, both of which are linearly dependent on the translational velocity. Body rotations produce similar passive forces and torques, but of substantially smaller magnitudes. Our comparison of closed-loop and open-loop dynamics showed that hawkmoths rely largely on passive damping to reduce the body translation but use feedback control based on pitch angle and angular velocity to control their orientation. The resulting feedback control system remains stable with sensory delays of more than two wingbeats.

Hummingbirds have the smallest body size and highest wingbeat frequencies of all flying vertebrates, so they represent one endpoint for evaluating the effects of body size on sustained muscle function and flight performance. Other bird species vary neuromuscular recruitment and contractile behavior to accomplish flight over a wide range of speeds, typically exhibiting a U-shaped curve with maxima at the slowest and fastest flight speeds. To test whether the high wingbeat frequencies and aerodynamically active upstroke of hummingbirds lead to different patterns, we flew rufous hummingbirds (Selasphorus rufus, 3 g body mass, 42 Hz wingbeat frequency) in a variable-speed wind tunnel (0-10 m s(-1)). We measured neuromuscular activity in the pectoralis (PECT) and supracoracoideus (SUPRA) muscles using electromyography (EMG, N=4 birds), and we measured changes in PECT length using sonomicrometry (N=1). Differing markedly from the pattern in other birds, PECT deactivation occurred before the start of downstroke and the SUPRA was deactivated before the start of upstroke. The relative amplitude of EMG signal in the PECT and SUPRA varied according to a U-shaped curve with flight speed; additionally, the onset of SUPRA activity became relatively later in the wingbeat at intermediate flight speeds (4 and 6 m s(-1)). Variation in the relative amplitude of EMG was comparable with that observed in other birds but the timing of muscle activity was different. These data indicate the high wingbeat frequency of hummingbirds limits the time available for flight muscle relaxation before the next half stroke of a wingbeat. Unlike in a previous study that reported single-twitch EMG signals in the PECT of hovering hummingbirds, across all flight speeds we observed 2.9+/-0.8 spikes per contraction in the PECT and 3.8+/-0.8 spikes per contraction in the SUPRA. Muscle strain in the PECT was 10.8+/-0.5%, the lowest reported for a flying bird, and average strain rate was 7.4+/-0.2 muscle lengths s(-1). Among species of birds, PECT strain scales proportional to body mass to the 0.2 power (infinityM(b)(0.2)) using species data and infinityM(b)(0.3) using independent contrasts. This positive scaling is probably a physiological response to an adverse scaling of mass-specific power available for flight.

Free-flight body dynamics and wing kinematics were collected from recordings of continuous, low-speed, multi-wingbeat yaw turns in hawkmoths (Manduca sexta) using stereo videography. These data were used to examine the effects of rotational damping arising from interactions between the body rotation and flapping motion (flapping counter-torque, FCT) on continuous turning. The moths were found to accelerate during downstroke, then decelerate during upstroke by an amount consistent with FCT damping. Wing kinematics related to turning were then analysed in a simulation of hawkmoth flight; results were consistent with the observed acceleration-deceleration pattern. However, an alternative wing kinematic which produced more continuous and less damped accelerations was found in the simulation. These findings demonstrate that (i) FCT damping is detectable in the dynamics of continuously turning animals and (ii) FCT-reducing kinematics do exist but were not employed by turning moths, possibly because within-wingbeat damping simplifies control of turning by allowing control systems to target angular velocity rather than acceleration.

Body motions (kinematics) of animals can be dimensionally complex, especially when flexible parts of the body interact with a surrounding fluid. In these systems, tracking motion completely can be difficult, and result in a large number of correlated measurements, with unclear contributions of each parameter to performance. Workers typically get around this by deciding a priori which variables are important (wing camber, stroke amplitude, etc.), and focusing only on those variables, but this constrains the ability of a study to uncover variables of influence. Here, we describe an application of proper orthogonal decomposition (POD) for assigning importances to kinematic variables, using dimensional complexity as a metric. We apply this method to bat flight kinematics, addressing three questions: (1) Does dimensional complexity of motion change with speed? (2) What body markers are optimal for capturing dimensional complexity? (3) What variables should a simplified reconstruction of bat flight include in order to maximally reconstruct actual dimensional complexity? We measured the motions of 17 kinematic markers (20 joint angles) on a bat (Cynopterus brachyotis) flying in a wind tunnel at nine speeds. Dimensional complexity did not change with flight speed, despite changes in the kinematics themselves, suggesting that the relative efficacy of a given number of dimensions for reconstructing kinematics is conserved across speeds. By looking at subsets of the full 17-marker set, we found that using more markers improved resolution of kinematic dimensional complexity, but that the benefit of adding markers diminished as the total number of markers increased. Dimensional complexity was highest when the hindlimb and several points along digits III and IV were tracked. Also, we uncovered three groups of joints that move together during flight by using POD to quantify correlations of motion. These groups describe 14/20 joint angles, and provide a framework for models of bat flight for experimental and modeling purposes.

Hummingbirds are specialized for hovering flight, and substantial research has explored this behavior. Forward flight is also important to hummingbirds, but the manner in which they perform forward flight is not well documented. Previous research suggests that hummingbirds increase flight velocity by simultaneously tilting their body angle and stroke-plane angle of the wings, without varying wingbeat frequency and upstroke: downstroke span ratio. We hypothesized that other wing kinematics besides stroke-plane angle would vary in hummingbirds. To test this, we used synchronized high-speed (500 Hz) video cameras and measured the three-dimensional wing and body kinematics of rufous hummingbirds (Selasphorus rufus, 3 g, N=5) as they flew at velocities of 0-12 m s(-1) in a wind tunnel. Consistent with earlier research, the angles of the body and the stroke plane changed with velocity, and the effect of velocity on wingbeat frequency was not significant. However, hummingbirds significantly altered other wing kinematics including chord angle, angle of attack, anatomical stroke-plane angle relative to their body, percent of wingbeat in downstroke, wingbeat amplitude, angular velocity of the wing, wingspan at mid-downstroke, and span ratio of the wingtips and wrists. This variation in bird-centered kinematics led to significant effects of flight velocity on the angle of attack of the wing and the area and angles of the global stroke planes during downstroke and upstroke. We provide new evidence that the paths of the wingtips and wrists change gradually but consistently with velocity, as in other bird species that possess pointed wings. Although hummingbirds flex their wings slightly at the wrist during upstroke, their average wingtip-span ratio of 93% revealed that they have kinematically ;rigid' wings compared with other avian species.

Maneuvering flight has long been recognized as an important component of the natural behavior of many bird species, but has been the subject of little experimental work. Here we examine the kinematics and neuromuscular control of turning flight in the rose-breasted cockatoo Eolophus roseicapillus (N=6), testing predictions of maneuvering flight and control based on aerodynamic theory and prior kinematic and neuromuscular studies. Six cockatoos were trained to navigate between two perches placed in an L-shaped flight corridor, making a 90 degrees turn midway through each flight. Flights were recorded with three synchronized high-speed video cameras placed outside the corridor, allowing a three-dimensional reconstruction of wing and body kinematics through the turn. We simultaneously collected electromyography recordings from bilateral implants in the pectoralis, supracoracoideus, biceps brachii and extensor metacarpi radialis muscles. The cockatoos maneuvered using flapping, banked turns with an average turn radius of 0.92 m. The mean rate of change in heading during a complete wingbeat varied through the turn and was significantly correlated to roll angle at mid-downstroke. Changes in roll angle were found to include both within-wingbeat and among-wingbeat components that bear no direct relationship to one another. Within-wingbeat changes in roll were dominated by the inertial effects while among-wingbeat changes in roll were likely the result of both inertial and aerodynamic effects.

The reconfigurable, flapping wings of birds allow for both inertial and aerodynamic modes of reorientation. We found evidence that both these modes play important roles in the low speed turning flight of the rose-breasted cockatoo Eolophus roseicapillus. Using three-dimensional kinematics recorded from six cockatoos making a 90 degrees turn in a flight corridor, we developed predictions of inertial and aerodynamic reorientation from estimates of wing moments of inertia and flapping arcs, and a blade-element aerodynamic model. The blade-element model successfully predicted weight support (predicted was 88+/-17% of observed, N=6) and centripetal force (predicted was 79+/-29% of observed, N=6) for the maneuvering cockatoos and provided a reasonable estimate of mechanical power. The estimated torque from the model was a significant predictor of roll acceleration (r(2)=0.55, P<0.00001), but greatly overestimated roll magnitude when applied with no roll damping. Non-dimensional roll damping coefficients of approximately -1.5, 2-6 times greater than those typical of airplane flight dynamics (approximately -0.45), were required to bring our estimates of reorientation due to aerodynamic torque back into conjunction with the measured changes in orientation. Our estimates of inertial reorientation were statistically significant predictors of the measured reorientation within wingbeats (r(2) from 0.2 to 0.37, P<0.0005). Components of both our inertial reorientation and aerodynamic torque estimates correlated, significantly, with asymmetries in the activation profile of four flight muscles: the pectoralis, supracoracoideus, biceps brachii and extensor metacarpi radialis (r(2) from 0.27 to 0.45, P<0.005). Thus, avian flight maneuvers rely on production of asymmetries throughout the flight apparatus rather than in a specific set of control or turning muscles.

The inverse problem of hovering flight, that is, the range of wing movements appropriate for sustained flight at a fixed position and orientation, was examined by developing a simulation of the hawkmoth Manduca sexta. Inverse problems arise when one is seeking the parameters that are required to achieve a specified model outcome. In contrast, forward problems explore the outcomes given a specified set of input parameters. The simulation was coupled to a microgenetic algorithm that found specific sequences of wing and body motions, encoded by ten independent kinematic parameters, capable of generating the fixed body position and orientation characteristic of hovering flight. Additionally, we explored the consequences of restricting the number of free kinematic parameters and used this information to assess the importance to flight control of individual parameters and various combinations of them. Output from the simulated moth was compared to kinematic recordings of hovering flight in real hawkmoths; the real and simulated moths performed similarly with respect to their range of variation in position and orientation. The simulated moth also used average wingbeat kinematics (amplitude, stroke plane orientation, etc) similar to those of the real moths. However, many different subsets of the available kinematic were sufficient for hovering flight and available kinematic data from real moths does not include sufficient detail to assess which, if any, of these was consistent with the real moth. This general result, the multiplicity of possible hovering kinematics, shows that the means by which Manduca sexta actually maintains position and orientation may have considerable freedom and therefore may be influenced by many other factors beyond the physical and aerodynamic requirements of hovering flight.

Regional fascicle strains were recorded in vivo from the pectoralis of carneau pigeons using sonomicrometry during level slow flight, together with regional electromyography (EMG) and deltopectoral crest (DPC) strain measurements of whole muscle force. Fascicle strain measurements were obtained at four sites within the pectoralis: the anterior (Ant), middle (Mid) and posterior (Post) sternobrachium (SB), and the smaller thoracobrachium (TB). Strains were also recorded along the intramuscular aponeurosis of the pectoralis to assess its 'in-series' compliance with respect to strains of Post SB and TB fascicles. In-series segment strains were also obtained along Ant SB and Mid SB fascicles, which insert directly on the DPC without attaching to the intramuscular aponeurosis. In-series segment strains differed from 2% to 17.2%, averaging differences of 6.1% at the Ant SB site and 1.4% at the Mid SB site. Temporal patterns of in-series fascicle segment strain were similar at both sites. Regional fascicle strains also exhibited similar temporal patterns of lengthening and shortening and were most uniform in magnitude at the Ant SB, Mid SB and TB sites (total strain: 33.7%, 35.9% and 33.2% respectively), but were smaller at the Post SB site (24.4%). Strains measured along the aponeurosis tracked the patterns of contractile fascicle strain but were significantly lower in magnitude (19.1%). Fascicle lengthening strains (+25.4%) greatly exceeded net shortening strains (-6.5%) at all sites. Much of the variation in regional fascicle strain patterns resulted from variation of in vivo recording sites among individual animals, despite attempts to define consistent regions for obtaining in vivo recordings. No significant variation in EMG activation onset was found, but deactivation of the Ant SB occurred before the other muscle sites. Even so, the range of variation was small, with all muscle regions being activated midway through lengthening (upstroke) and turned off midway through shortening (downstroke). While subtle differences in the timing and rate of fascicle strain may relate to differing functional roles of the pectoralis, regional patterns of fascicle strain and activation suggest a generally uniform role for the muscle as a whole throughout the wingbeat cycle. Shorter fascicles located in more posterior regions of the muscle underwent generally similar strains as longer fascicles located in more anterior SB regions. The resulting differences in fiber length were accommodated by strain in the intramuscular aponeurosis and rotation of the pectoralis insertion with respect to the origin. As a result, longer Ant and Mid SB fascicles were estimated to contribute substantially more work per unit mass than shorter Post SB and TB fascicles. When the mass fractions of these regions are accounted for, our regional fascicle strain measurements show that the anterior regions of the pectoralis likely contribute 76%, and the posterior regions 24%, of the muscle's total work output. When adjusted for mass fraction and regional fascicle strain, pectoralis work averaged 24.7+/-5.1 J kg(-1) (206.6+/-43.5 W kg(-1)) during level slow (approximately 4-5 m s(-1)) flight.

Differential pressure measurements offer a new approach for studying the aerodynamics of bird flight. Measurements from differential pressure sensors are combined to form a dynamic pressure map for eight sites along and across the wings, and for two sites across the tail, of pigeons flying between two perches. The confounding influence of acceleration on the pressure signals is shown to be small for both wings and tail. The mean differential pressure for the tail during steady, level flight was 25.6 Pa, which, given an angle of attack for the tail of 47.6 degrees , suggests the tail contributes 7.91% of the force required for weight support, and requires a muscle-mass specific power of 19.3 W kg(-1) for flight to overcome its drag at 4.46 m s(-1). Differential pressures during downstroke increase along the wing length, to 300-400 Pa during take-off and landing for distal sites. Taking the signals obtained from five sensors sited along the wing at feather bases as representative of the mean pressure for five spanwise elements at each point in time, and assuming aerodynamic forces act within the x-z plane (i.e. no forces in the direction of travel) and perpendicular to the wing during downstroke, we calculate that 74.5% of the force required to support weight was provided by the wings, and that the aerodynamic muscle-mass specific power required to flap the wings was 272.7 W kg(-1).