ABSTRACT Migration allows animals to use resources that are variable in time and/or space, with different migratory strategies depending on the predictability of resource variation. When food varies seasonally, obligate migrants anticipate and prepare for migration. In contrast, facultative migrants, whose movements are unpredictable in timing and destination, may prepare for either migration or escape when resources are depleted. We propose and test two alternative hypotheses regarding the behavioral and physiological responses of facultative migrants to declining food availability. (1) The prepare hypothesis predicts that facultative migrants prepare for departure by increasing fuel stores in response to declining food availability, and elevations of baseline corticosterone (CORT) facilitate increased activity. (2) The escape hypothesis predicts that facultative migrants do not prepare for departure, body condition declines as food availability declines, and stress-induced levels of CORT induce escape behavior when both energetic condition and food resources are low. We conducted a 16-day experiment, measuring body composition (using quantitative magnetic resonance), activity (using force perches) and baseline CORT in pine siskins ( Spinus pinus ) given ad libitum food or a slow decline, fast decline or randomly changing amount of food. Our results support the escape hypothesis: body condition declined as food declined, decreases in body and fat mass were associated with increases in baseline CORT, and activity increased only when food availability was low. This work suggests that facultative migration in autumn allows birds to escape low-resource areas and that the underlying physiological mechanisms differ from those driving both seasonal, obligate migrations and spring nomadic movements.

ABSTRACT Flexibility in heat generation and dissipation mechanisms provides endotherms the ability to match their thermoregulatory strategy with external demands. However, the degree to which these two mechanisms account for seasonal changes in body temperature regulation is little explored. Here, we present novel data on the regulation of avian body temperature to investigate how birds alter mechanisms of heat production and heat conservation to deal with variation in ambient conditions. We subjected dark-eyed juncos ( Junco hyemalis ) to chronic cold acclimations of varying duration and subsequently quantified their metabolic rates, thermal conductance and ability to maintain normothermia. Cold-acclimated birds adjusted traits related to both heat generation (increased summit metabolic rate) and heat conservation (decreased conductance) to improve their body temperature regulation. Increases in summit metabolic rate occurred rapidly, but plateaued after 1 week of cold exposure. In contrast, changes to conductance occurred only after 9 weeks of cold exposure. Thus, the ability to maintain body temperature continued to improve throughout the experiment, but the mechanisms underlying this improvement changed through time. Our results demonstrate the ability of birds to adjust thermoregulatory strategies in response to thermal cues and reveal that birds may combine multiple responses to meet the specific demands of their environments.

ABSTRACT Alcids, a family of seabirds including murres, guillemots and puffins, exhibit the greatest mass-specific dive depths and durations of any birds or mammals. These impressive diving capabilities have motivated numerous studies on the biomechanics of alcid swimming and diving, with one objective being to compare stroke–acceleration patterns of swimming alcids with those of penguins, where upstroke and downstroke are used for horizontal acceleration. Studies of free-ranging, descending alcids have found that alcids accelerate in the direction of travel during both their upstroke and downstroke, but only at depths <20 m, whereas studies of alcids swimming horizontally report upstroke-based acceleration to be rare (≤16% of upstrokes). We hypothesized that swimming trajectory, via its interaction with buoyancy, determines the magnitude of acceleration produced during the upstroke. Thus, we studied the stroke–acceleration relationships of five species of alcid swimming freely at the Alaska SeaLife Center using videography and kinematic analysis. Contrary to our prediction, we found that upstroke-based acceleration is very common (87% of upstrokes) during both descending and horizontal swimming. We reveal that head-damping – wherein an animal extends and retracts its head to offset periodic accelerations – is common in swimming alcids, underscoring the importance of head stabilization during avian locomotion.

ABSTRACT Many marine organisms and life stages lack specialized respiratory structures, like gills, and rely instead on cutaneous respiration, which they facilitate by having thin integuments. This respiratory mode may limit body size, especially if the integument also functions in support or locomotion. Pycnogonids, or sea spiders, are marine arthropods that lack gills and rely on cutaneous respiration but still grow to large sizes. Their cuticle contains pores, which may play a role in gas exchange. Here, we examined alternative paths of gas exchange in sea spiders: (1) oxygen diffuses across pores in the cuticle, a common mechanism in terrestrial eggshells, (2) oxygen diffuses directly across the cuticle, a common mechanism in small aquatic insects, or (3) oxygen diffuses across both pores and cuticle. We examined these possibilities by modeling diffusive oxygen fluxes across all pores in the body of sea spiders and asking whether those fluxes differed from measured metabolic rates. We estimated fluxes across pores using Fick's law parameterized with measurements of pore morphology and oxygen gradients. Modeled oxygen fluxes through pores closely matched oxygen consumption across a range of body sizes, which means the pores facilitate oxygen diffusion. Furthermore, pore volume scaled hypermetrically with body size, which helps larger species facilitate greater diffusive oxygen fluxes across their cuticle. This likely presents a functional trade-off between gas exchange and structural support, in which the cuticle must be thick enough to prevent buckling due to external forces but porous enough to allow sufficient gas exchange.

ABSTRACT 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.

ABSTRACT 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.

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

ABSTRACT For centuries, naturalists have suggested that the tonal elements of pigeon wing sounds may be sonations (non-vocal acoustic signals) of alarm. However, spurious tonal sounds may be produced passively as a result of aeroelastic flutter in the flight feathers of almost all birds. Using mechanistic criteria emerging from recent work on sonations, we sought to: (1) identify characteristics of rock pigeon flight feathers that might be adapted for sound production rather than flight, and (2) provide evidence that this morphology is necessary for in vivo sound production and is sufficient to replicate in vivo sounds. Pigeons produce tonal sounds (700±50 Hz) during the latter two-thirds of each downstroke during take-off. These tones are produced when a small region of long, curved barbs on the inner vane of the outermost primary feather (P10) aeroelastically flutters. Tones were silenced in live birds when we experimentally increased the stiffness of this region to prevent flutter. Isolated P10 feathers were sufficient to reproduce in vivo sounds when spun at the peak angular velocity of downstroke (53.9–60.3 rad s −1 ), but did not produce tones at average downstroke velocity (31.8 rad s −1 ), whereas P9 and P1 feathers never produced tones. P10 feathers had significantly lower coefficients of resultant aerodynamic force ( C R ) when spun at peak angular velocity than at average angular velocity, revealing that production of tonal sounds incurs an aerodynamic cost. P9 and P1 feathers did not show this difference in C R . These mechanistic results suggest that the tonal sounds produced by P10 feathers are not incidental and may function in communication.

ABSTRACT Slow flight is extremely energetically costly per unit time, yet highly important for takeoff and survival. However, at slow speeds it is presently thought that most birds do not produce beneficial aerodynamic forces during the entire wingbeat: instead they fold or flex their wings during upstroke, prompting the long-standing prediction that the upstroke produces trivial forces. There is increasing evidence that the upstroke contributes to force production, but the aerodynamic and kinematic mechanisms remain unknown. Here, we examined the wingbeat cycle of two species: the diamond dove ( Geopelia cuneata ) and zebra finch ( Taeniopygia guttata ), which exhibit different upstroke styles – a wingtip-reversal and flexed-wing upstroke, respectively. We used a combination of particle image velocimetry and near-wake streamline measures alongside detailed 3D kinematics. We show that during the middle of the wingtip-reversal upstroke, the hand-wing has a high angular velocity (15.3±0.8 deg ms −1 ) and translational speed (8.4±0.6 m s −1 ). The flexed-wing upstroke, in contrast, has low wingtip speed during mid-upstroke. Instead, later in the stroke cycle, during the transition from upstroke to downstroke, it exhibits higher angular velocities (45.5±13.8 deg ms −1 ) and translational speeds (11.0±1.9 m s −1 ). Aerodynamically, the wingtip-reversal upstroke imparts momentum to the wake, with entrained air shed backward (visible as circulation of 14.4±0.09 m 2 s −1 ). In contrast, the flexed-wing upstroke imparts minimal momentum. Clap and peel in the dove enhances the time course for circulation production on the wings, and provides new evidence of convergent evolution on time-varying aerodynamic mechanisms during flapping in insects and birds.

Transitions to and from the air are critical for aerial locomotion and likely shaped the evolution of flying animals. Research on take-off demonstrates that legs generate greater body accelerations compared with wings, and thereby contribute more to initial flight velocity. Here, we explored coordination between wings and legs in two species with different wingbeat styles, and quantified force production of these modules during the final phase of landing. We used the same birds that we had previously studied during take-off: zebra finch ( Taeniopygia guttata, N =4) and diamond dove ( Geopelia cuneata, N =3). We measured kinematics using high-speed video, aerodynamics using particle image velocimetry, and ground-reaction forces using a perch mounted on a force plate. In contrast with the first three wingbeats of take-off, the final four wingbeats during landing featured ~2 times greater force production. Thus, wings contribute proportionally more to changes in velocity during the last phase of landing compared with the initial phase of take-off. The two species touched down at the same velocity (~1 m s −1 ), but they exhibited significant differences in the timing of their final wingbeat relative to touchdown. The ratio of average wing force to peak leg force was greater in diamond doves than in zebra finches. Peak ground reaction forces during landing were ~50% of those during take-off, consistent with the birds being motivated to control landing. Likewise, estimations of mechanical energy flux for both species indicate that wings produce 3–10 times more mechanical work within the final wingbeats of flight compared with the kinetic energy of the body absorbed by legs during ground contact.

SUMMARY Take-off mechanics are fundamental to the ecology and evolution of flying animals. Recent research has revealed that initial take-off velocity in birds is driven mostly by hindlimb forces. However, the contribution of the wings during the transition to air is unknown. To investigate this transition, we integrated measurements of both leg and wing forces during take-off and the first three wingbeats in zebra finch ( Taeniopygia guttata , body mass 15 g, N =7) and diamond dove ( Geopelia cuneata , body mass 50 g, N =3). We measured ground reaction forces produced by the hindlimbs using a perch mounted on a force plate, whole-body and wing kinematics using high-speed video, and aerodynamic forces using particle image velocimetry (PIV). Take-off performance was generally similar between species. When birds were perched, an acceleration peak produced by the legs contributed to 85±1% of the whole-body resultant acceleration in finch and 77±6% in dove. At lift-off, coincident with the start of the first downstroke, the percentage of hindlimb contribution to initial flight velocity was 93.6±0.6% in finch and 95.2±0.4% in dove. In finch, the first wingbeat produced 57.9±3.4% of the lift created during subsequent wingbeats compared with 62.5±2.2% in dove. Advance ratios were <0.5 in both species, even when taking self-convection of shed vortices into account, so it was likely that wing–wake interactions dominated aerodynamics during wingbeats 2 and 3. These results underscore the relatively low contribution of the wings to initial take-off, and reveal a novel transitional role for the first wingbeat in terms of force production.

SUMMARY Wing morphology correlates with flight performance and ecology among adult birds, yet the impact of wing development on aerodynamic capacity is not well understood. Recent work using chukar partridge ( Alectoris chukar ), a precocial flier, indicates that peak coefficients of lift and drag ( C L and C D ) and lift-to-drag ratio ( C L : C D ) increase throughout ontogeny and that these patterns correspond with changes in feather microstructure. To begin to place these results in a comparative context that includes variation in life-history strategy, we used a propeller and force-plate model to study aerodynamic force production across a developmental series of the altricial-flying mallard ( Anas platyrhynchos ). We observed the same trend in mallards as reported for chukar in that coefficients of vertical ( C V ) and horizontal force ( C H ) and C V : C H ratio increased with age, and that measures of gross-wing morphology (aspect ratio, camber and porosity) in mallards did not account for intraspecific trends in force production. Rather, feather microstructure (feather unfurling, rachis width, feather asymmetry and barbule overlap) all were positively correlated with peak C V : C H . Throughout ontogeny, mallard primary feathers became stiffer and less transmissive to air at both macroscale (between individual feathers) and microscale (between barbs/barbules/barbicels) levels. Differences between species were manifest primarily as heterochrony of aerodynamic force development. Chukar wings generated measurable aerodynamic forces early (<8 days), and improved gradually throughout a 100 day ontogenetic period. Mallard wings exhibited delayed aerodynamic force production until just prior to fledging (day 60), and showed dramatic improvement within a condensed 2-week period. These differences in timing may be related to mechanisms of escape used by juveniles, with mallards swimming to safety and chukar flap-running up slopes to take refuge. Future comparative work should test whether the need for early onset of aerodynamic force production in the chukar, compared with delayed, but rapid, change in the mallard wing, leads to a limited repertoire of flight behavior in adult chukar compared with mallards.

SUMMARY Wing-assisted incline running (WAIR) in birds combines the use of the wings and hindlimbs to ascend otherwise insurmountable obstacles. It is a means of escape in precocial birds before they are able to fly, and it is used by a variety of juvenile and adult birds as an alternative to flight for exploiting complex three-dimensional environments at the interface of the ground and air. WAIR and controlled flapping descent (CFD) are the bases of the ontogenetic-transitional wing hypothesis, wherein WAIR and CFD are proposed to be extant biomechanical analogs for incremental adaptive stages in the evolutionary origin of flight. A primary assumption of the hypothesis is that work and power requirements from the primary downstroke muscle, the pectoralis, incrementally increase from shallow- to steep-angled terrestrial locomotion, and between terrestrial and aerial locomotion. To test this assumption, we measured in vivo force, electromyographic (EMG) activity and length change in the pectoralis of pigeons ( Columba livia ) as the birds engaged in shallow and steep WAIR (65 and 85 deg, respectively) and in three modes of slow flight immediately following take-off: ascending at 80 deg, level and descending at –60 deg. Mean EMG amplitude, muscle stress, strain, work and power were minimal during shallow WAIR and increased stepwise from steep WAIR to descending flight and level flight to reach the highest levels during ascending flight. Relative to resting length of the pectoralis, fractional lengthening (maximum muscle strain) was similar among behaviors, but fractional shortening (minimum muscle strain) was absent during WAIR such that the pectoralis did not shorten to less than the resting length. These data dramatically extend the known range of in vivo contractile behavior for the pectoralis in birds. We conclude that WAIR remains a useful extant model for the evolutionary transition from terrestrial to aerial locomotion in birds because work and power requirements from the pectoralis increase incrementally during WAIR and from WAIR to flight.

SUMMARY During slow flight, bird species vary in their upstroke kinematics using either a ‘flexed wing’ or a distally supinated ‘tip-reversal’ upstroke. Two hypotheses have been presented concerning the function of the tip-reversal upstroke. The first is that this behavior is aerodynamically inactive and serves to minimize drag. The second is that the tip-reversal upstroke is capable of producing significant aerodynamic forces. Here, we explored the aerodynamic capabilities of the tip-reversal upstroke using a well-established propeller method. Rock dove ( Columba livia, N =3) wings were spread and dried in postures characteristic of either mid-upstroke or mid-downstroke and spun at in vivo Reynolds numbers to simulate forces experienced during slow flight. We compared 3D wing shape for the propeller and in vivo kinematics, and found reasonable kinematic agreement between methods (mean differences 6.4% of wing length). We found that the wing in the upstroke posture is capable of producing substantial aerodynamic forces. At in vivo angles of attack (66 deg at mid-upstroke, 46 deg at mid-downstroke), the upstroke wings averaged for three birds produced a lift-to-drag ratio of 0.91, and the downstroke wings produced a lift-to-drag ratio of 3.33. Peak lift-to-drag ratio was 2.5 for upstroke and 6.3 for downstroke. Our estimates of total force production during each half-stroke suggest that downstroke produces a force that supports 115% of bodyweight, and during upstroke a forward-directed force (thrust) is produced at 36% of body weight.

SUMMARY The juvenile period is often a crucial interval for selective pressure on locomotor ability. Although flight is central to avian biology, little is known about factors that limit flight performance during development. To improve understanding of flight ontogeny, we used a propeller (revolving wing) model to test how wing shape and feather structure influence aerodynamic performance during development in the precocial chukar partridge ( Alectoris chukar , 4 to >100 days post hatching). We spun wings in mid-downstroke posture and measured lift ( L ) and drag ( D ) using a force plate upon which the propeller assembly was mounted. Our findings demonstrate a clear relationship between feather morphology and aerodynamic performance. Independent of size and velocity, older wings with stiffer and more asymmetrical feathers, high numbers of barbicels and a high degree of overlap between barbules generate greater L and L : D ratios than younger wings with flexible, relatively symmetrical and less cohesive feathers. The gradual transition from immature feathers and drag-based performance to more mature feathers and lift-based performance appears to coincide with ontogenetic transitions in locomotor capacity. Younger birds engage in behaviors that require little aerodynamic force and that allow D to contribute to weight support, whereas older birds may expand their behavioral repertoire by flapping with higher tip velocities and generating greater L . Incipient wings are, therefore, uniquely but immediately functional and provide flight-incapable juveniles with access to three-dimensional environments and refugia. Such access may have conferred selective advantages to theropods with protowings during the evolution of avian flight.

SUMMARY 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 (∞ M b 0.2 ) using species data and ∞ M 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.

SUMMARY The supracoracoideus (SUPRA) is the primary upstroke muscle for avian flight and is the antagonist to the downstroke muscle, the pectoralis (PECT). We studied in vivo contractile properties and mechanical power output of both muscles during take-off, level and landing flight. We measured muscle length change and activation using sonomicrometry and electromyography, and muscle force development using strain recordings on the humerus. Our results support a hypothesis that the primary role of the SUPRA is to supinate the humerus. Antagonistic forces exerted by the SUPRA and PECT overlap during portions of the wingbeat cycle, thereby offering a potential mechanism for enhancing control of the wing. Among flight modes, muscle strain was approximately the same in the SUPRA (33–40%) and the PECT(35–42%), whereas peak muscle stress was higher in the SUPRA(85–126 N m –2 ) than in the PECT (50–58 N m –2 ). The SUPRA mainly shortened relative to resting length and the PECT mainly lengthened. We estimated that elastic energy storage in the tendon of the SUPRA contributed between 28 and 60% of the net work of the SUPRA and 6–10% of the total net mechanical work of both muscles. Mechanical power output in the SUPRA was congruent with the estimated inertial power required for upstroke, but power output from the PECT was only 42–46% of the estimated aerodynamic power requirements for flight. There was a significant effect of flight mode upon aspects of the contractile behavior of both muscles including strain, strain rate, peak stress, work and power.

SUMMARY Power output is a unifying theme for bird flight and considerable progress has been accomplished recently in measuring muscular, metabolic and aerodynamic power in birds. The primary flight muscles of birds, the pectoralis and supracoracoideus, are designed for work and power output, with large stress (force per unit cross-sectional area) and strain (relative length change) per contraction. U-shaped curves describe how mechanical power output varies with flight speed, but the specific shapes and characteristic speeds of these curves differ according to morphology and flight style. New measures of induced, profile and parasite power should help to update existing mathematical models of flight. In turn, these improved models may serve to test behavioral and ecological processes. Unlike terrestrial locomotion that is generally characterized by discrete gaits, changes in wing kinematics and aerodynamics across flight speeds are gradual. Take-off flight performance scales with body size, but fully revealing the mechanisms responsible for this pattern awaits new study. Intermittent flight appears to reduce the power cost for flight, as some species flap–glide at slow speeds and flap–bound at fast speeds. It is vital to test the metabolic costs of intermittent flight to understand why some birds use intermittent bounds during slow flight. Maneuvering and stability are critical for flying birds,and design for maneuvering may impinge upon other aspects of flight performance. The tail contributes to lift and drag; it is also integral to maneuvering and stability. Recent studies have revealed that maneuvers are typically initiated during downstroke and involve bilateral asymmetry of force production in the pectoralis. Future study of maneuvering and stability should measure inertial and aerodynamic forces. It is critical for continued progress into the biomechanics of bird flight that experimental designs are developed in an ecological and evolutionary context.

SUMMARY 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.

SUMMARY Wing-assisted incline running (WAIR) is a form of locomotion in which a bird flaps its wings to aid its hindlimbs in climbing a slope. WAIR is used for escape in ground birds, and the ontogeny of this behavior in precocial birds has been suggested to represent a model analogous to transitional adaptive states during the evolution of powered avian flight. To begin to reveal the aerodynamics of flap-running, we used digital particle image velocimetry (DPIV) and measured air velocity, vorticity, circulation and added mass in the wake of chukar partridge Alectoris chukar as they engaged in WAIR (incline 65–85°; N =7 birds) and ascending flight(85°, N =2). To estimate lift and impulse, we coupled our DPIV data with three-dimensional wing kinematics from a companion study. The ontogeny of lift production was evaluated using three age classes: baby birds incapable of flight [6–8 days post hatching (d.p.h.)] and volant juveniles (25–28 days) and adults (45+ days). All three age classes of birds, including baby birds with partially emerged, symmetrical wing feathers,generated circulation with their wings and exhibited a wake structure that consisted of discrete vortex rings shed once per downstroke. Impulse of the vortex rings during WAIR was directed 45±5° relative to horizontal and 21±4° relative to the substrate. Absolute values of circulation in vortex cores and induced velocity increased with increasing age. Normalized circulation was similar among all ages in WAIR but 67% greater in adults during flight compared with flap-running. Estimated lift during WAIR was 6.6%of body weight in babies and between 63 and 86% of body weight in juveniles and adults. During flight, average lift was 110% of body weight. Our results reveal for the first time that lift from the wings, rather than wing inertia or profile drag, is primarily responsible for accelerating the body toward the substrate during WAIR, and that partially developed wings, not yet capable of flight, can produce useful lift during WAIR. We predict that neuromuscular control or power output, rather than external wing morphology, constrain the onset of flight ability during development in birds.