Turkey vultures tune their airspeed to changing air density.

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.

Musculoskeletal wing-actuation model of hummingbirds predicts diverse effects of primary flight muscles in hovering flight.

Hummingbirds have evolved to hover and manoeuvre with exceptional flight control. This is enabled by their musculoskeletal system that successfully exploits the agile motion of flapping wings. Here, we synthesize existing empirical and modelling data to generate novel hypotheses for principles of hummingbird wing actuation. These may help guide future experimental work and provide insights into the evolution and robotic emulation of hummingbird flight. We develop a functional model of the hummingbird musculoskeletal system, which predicts instantaneous, three-dimensional torque produced by primary (pectoralis and supracoracoideus) and combined secondary muscles. The model also predicts primary muscle contractile behaviour, including stress, strain, elasticity and work. Results suggest that the primary muscles (i.e. the flight 'engine') function as diverse effectors, as they do not simply power the stroke, but also actively deviate and pitch the wing with comparable actuation torque. The results also suggest that the secondary muscles produce controlled-tightening effects by acting against primary muscles in deviation and pitching. The diverse effects of the pectoralis are associated with the evolution of a comparatively enormous bicipital crest on the humerus.

Reconstruction of Flight Parameters of a Bat for Flapping Robots.

The flight of bats is comparatively less documented and understood than birds and insects and may provide novel inspiration for the design of flapping flight robots. This study captured the natural flight of short-nosed fruit bats (Cynopterus sphinx) by an optical motion capture system, "OptiTrack", with pasted markers on the wings and body to reconstruct the flight parameters. Due to the self-occlusion at some moments, points on the membrane wings cannot be captured by any cameras. To draw a smooth trajectory, it is desired to reconstruct all missing data. Therefore, an algorithm is proposed by using numerical techniques, accompanied by modern mathematical and computational tools, to envisage the missing data from the captured flight. The least-square fitted polynomial engendered the parameter equations for x-, y-, and z-coordinates of marked points which were used to reconstruct the trajectory of the flight. The parameter equations of position coordinates were also used to compute the morphological and aerodynamic characteristics of the flight. The most outstanding contribution of the work is that not only the trajectory, velocity, and velocity field but also the morphing areas of the membrane wings were recreated using the reconstructed data. These data and reconstructed curves of trajectory and velocity field will be used for the further aerodynamic analysis and mechanism design of the flapping robot. This method can also be generalized to reconstruct the performance parameters of any other animals for bionic design.

Flow interactions between uncoordinated flapping swimmers give rise to group cohesion.

Many species of fish and birds travel in groups, yet the role of fluid-mediated interactions in schools and flocks is not fully understood. Previous fluid-dynamical models of these collective behaviors assume that all individuals flap identically, whereas animal groups involve variations across members as well as active modifications of wing or fin motions. To study the roles of flapping kinematics and flow interactions, we design a minimal robotic "school" of two hydrofoils swimming in tandem. The flapping kinematics of each foil are independently prescribed and systematically varied, while the forward swimming motions are free and result from the fluid forces. Surprisingly, a pair of uncoordinated foils with dissimilar kinematics can swim together cohesively-without separating or colliding-due to the interaction of the follower with the wake left by the leader. For equal flapping frequencies, the follower experiences stable positions in the leader's wake, with locations that can be controlled by flapping amplitude and phase. Further, a follower with lower flapping speed can defy expectation and keep up with the leader, whereas a faster-flapping follower can be buffered from collision and oscillate in the leader's wake. We formulate a reduced-order model which produces remarkable agreement with all experimentally observed modes by relating the follower's thrust to its flapping speed relative to the wake flow. These results show how flapping kinematics can be used to control locomotion within wakes, and that flow interactions provide a mechanism which promotes group cohesion.

Direct Measurements of the Wing Kinematics of a Bat in Straight Flight.

Bat is the only mammal in the nature that can fly. Compared with birds and insects, bats are quite special in that their wings are formed by an elastic membrane, which renders that the airfoil deforms greatly during downstroke and upstroke. Due to the compliant skin of a bat, the movements of its wings are three-dimensionally complex during diverse flight behaviors. To understand the maneuverability and flight performance, three-dimensional reconstruction of the flight kinematics is essential. This study focuses on the reconstruction of the wing kinematics of the bat and identifies the primary relationship of parameters of aerodynamics in straight flight. With markers pasted on the wings and body of a bat, the motions of these points are recorded by a computerized optical motion capture system. The kinematic analysis shows that the motion of wings is very intricate. The digits of the wing display the sign of coupled motion. A novel approach was developed to measure the angle of attack and flapping angle of the wing. The angle of attack of leading edge differs with the overall angle of attack of the wing. The kinematics of the bat's wing is helpful to interpret the secret of the bat's flight.

Elastic wing deformations mitigate flapping asymmetry during manoeuvres in rose chafers (Protaetia cuprea).

To manoeuvre in air, flying animals produce asymmetric flapping between contralateral wings. Unlike the adjustable vertebrate wings, insect wings lack intrinsic musculature, preventing active control over wing shape during flight. However, the wings elastically deform as a result of aerodynamic and inertial forces generated by the flapping motions. How these elastic deformations vary with flapping kinematics and flight performance in free-flying insects is poorly understood. Using high-speed videography, we measured how contralateral wings elastically deform during free-flight manoeuvring in rose chafer beetles (Protaetia cuprea). We found that asymmetric flapping during aerial turns was associated with contralateral differences in chord-wise wing deformations. The highest instantaneous difference in deformation occurred during stroke reversals, resulting from differences in wing rotation timing. Elastic deformation asymmetry was also evident during mid-strokes, where wing compliance increased the angle of attack of both wings, but reduced the asymmetry in the angle of attack between contralateral wings. A biomechanical model revealed that wing compliance can increase the torques generated by each wing, providing higher potential for manoeuvrability, while concomitantly contributing to flight stability by attenuating steering asymmetry. Such stability may be adaptive for insects such as flower chafers that need to perform delicate low-speed landing manoeuvres among vegetation.

Wind-associated detours promote seasonal migratory connectivity in a flapping flying long-distance avian migrant.

It is essential to gain knowledge about the causes and extent of migratory connectivity between stationary periods of migrants to further the understanding of processes affecting populations, and to allow efficient implementation of conservation efforts throughout the annual cycle. Avian migrants likely use optimal routes with respect to mode of locomotion, orientation and migration strategy, influenced by external factors such as wind and topography. In self-powered flapping flying birds, any increases in fuel loads are associated with added flight costs. Energy-minimizing migrants are therefore predicted to trade-off extended detours against reduced travel across ecological barriers with no or limited foraging opportunities. Here, we quantify the extent of detours taken by different populations of European nightjars Caprimulgus europaeus, to test our predictions that they used routes beneficial according to energetic principles and evaluate the effect of route shape on seasonal migratory connectivity. We combined data on birds tracked from breeding sites along a longitudinal gradient from England to Sweden. We analysed the migratory connectivity between breeding and main non-breeding sites, and en route stopover sites just south of the Sahara desert. We quantified each track's route extension relative to the direct route between breeding and wintering sites, respectively, and contrasted it to the potential detour derived from the barrier reduction along the track while accounting for potential wind effects. Nightjars extended their tracks from the direct route between breeding and main non-breeding sites as they crossed the Mediterranean Sea-Sahara desert, the major ecological barrier in the Palaearctic-African migration system. These clockwise detours were small for birds from eastern sites but increased from east to west breeding longitude. Routes of the tracked birds were associated with partial reduction in the barrier crossing resulting in a trade-off between route extension and barrier reduction, as expected in an energy-minimizing migrant. This study demonstrates how the costs of barrier crossings in prevailing winds can disrupt migratory routes towards slightly different goals, and thereby promote migratory connectivity. This is an important link between individual migration strategies in association with an ecological barrier, and both spatially and demographic population patterns.

Speed control and force-vectoring of bluebottle flies in a magnetically levitated flight mill.

Flies fly at a broad range of speeds and produce sophisticated aerial maneuvers with precisely controlled wing movements. Remarkably, only subtle changes in wing motion are used by flies to produce aerial maneuvers, resulting in little directional tilt of the aerodynamic force vector relative to the body. Therefore, it is often considered that flies fly according to a helicopter model and control speed mainly via force vectoring by body pitch change. Here, we examined the speed control of bluebottle flies using a magnetically levitated (MAGLEV) flight mill, as they fly at different body pitch angles and with different augmented aerodynamic damping. We identified wing kinematic contributors to the changes of estimated aerodynamic force through testing and comparing two force-vectoring models - a constant force-vectoring model and a variable force-vectoring model - while using Akaike's information criterion for selection of the best-approximating model. The results show that the best-approximating variable force-vectoring model, which includes the effects of wing kinematic changes, yields a considerably more accurate prediction of flight speed, particularly in higher velocity range, as compared with those of the constant force-vectoring model. Examination of the variable force-vectoring model reveals that, in the flight mill tethered flight, flies use a collection of wing kinematic variables to control primarily the force magnitude, while the force direction is also modulated, albeit to a smaller extent compared with those due to the changes in body pitch. The roles of these wing kinematic variables are analogous to those of throttle, and collective and cyclic pitch of helicopters.

Aerodynamic characteristics along the wing span of a dragonfly Pantala flavescens.

We investigated the characteristics of interwing aerodynamic interactions across the span of the high aspect ratio, flexible wings of dragonflies under tethered and free-flying conditions. This revealed that the effects of the interactions on the hindwings vary across four spanwise regions. (i) Close to the wing root, a trailing-edge vortex (TEV) is formed by each stroke, while the formation of a leading-edge vortex (LEV) is limited by the short translational distance of the hindwing and suppressed by the forewing-induced flow. (ii) In the region away from the wing root but not quite up to midspan, the formation of the hindwing LEV is influenced by that of the forewing LEV. This vortex synergy can increase the circulation of the hindwing LEV in the corresponding cross-section by 22% versus that of the hindwing in isolation. (iii) In the region about half-way between the wing root and wing tip there is a transition dominated by downwash from the forewing resulting in flow attached to the hindwing. (iv) A LEV is developed in the remaining, outer region of the wing at the end of a stroke when the hindwing captures the vortex shed by the forewing. The interaction effects depend not only on the wing phasing but also on the flapping offset and flight direction. The aerodynamics of the hindwings vary substantially from the wing root to the wing tip. For a given phasing, this spanwise variation in the aerodynamics can be exploited in the design of artificial wings to achieve greater agility and higher efficiency.

Predicting fruit fly's sensing rate with insect flight simulations.

Without sensory feedback, flies cannot fly. Exactly how various feedback controls work in insects is a complex puzzle to solve. What do insects measure to stabilize their flight? How often and how fast must insects adjust their wings to remain stable? To gain insights into algorithms used by insects to control their dynamic instability, we develop a simulation tool to study free flight. To stabilize flight, we construct a control algorithm that modulates wing motion based on discrete measurements of the body-pitch orientation. Our simulations give theoretical bounds on both the sensing rate and the delay time between sensing and actuation. Interpreting our findings together with experimental results on fruit flies' reaction time and sensory motor reflexes, we conjecture that fruit flies sense their kinematic states every wing beat to stabilize their flight. We further propose a candidate for such a control involving the fly's haltere and first basalar motor neuron. Although we focus on fruit flies as a case study, the framework for our simulation and discrete control algorithms is applicable to studies of both natural and man-made fliers.

Flapping wing aerodynamics: from insects to vertebrates.

More than a million insects and approximately 11,000 vertebrates utilize flapping wings to fly. However, flapping flight has only been studied in a few of these species, so many challenges remain in understanding this form of locomotion. Five key aerodynamic mechanisms have been identified for insect flight. Among these is the leading edge vortex, which is a convergent solution to avoid stall for insects, bats and birds. The roles of the other mechanisms - added mass, clap and fling, rotational circulation and wing-wake interactions - have not yet been thoroughly studied in the context of vertebrate flight. Further challenges to understanding bat and bird flight are posed by the complex, dynamic wing morphologies of these species and the more turbulent airflow generated by their wings compared with that observed during insect flight. Nevertheless, three dimensionless numbers that combine key flow, morphological and kinematic parameters - the Reynolds number, Rossby number and advance ratio - govern flapping wing aerodynamics for both insects and vertebrates. These numbers can thus be used to organize an integrative framework for studying and comparing animal flapping flight. Here, we provide a roadmap for developing such a framework, highlighting the aerodynamic mechanisms that remain to be quantified and compared across species. Ultimately, incorporating complex flight maneuvers, environmental effects and developmental stages into this framework will also be essential to advancing our understanding of the biomechanics, movement ecology and evolution of animal flight.

Wing flexibility improves bumblebee flight stability.

Insect wings do not contain intrinsic musculature to change shape, but rather bend and twist passively during flight. Some insect wings feature flexible joints along their veins that contain patches of resilin, a rubber-like protein. Bumblebee wings exhibit a central resilin joint (1m-cu) that has previously been shown to improve vertical force production during hovering flight. In this study, we artificially stiffened bumblebee (Bombus impatiens) wings in vivo by applying a micro-splint to the 1m-cu joint, and measured the consequences for body stability during forward flight in both laminar and turbulent airflow. In laminar flow, bees with stiffened wings exhibited significantly higher mean rotation rates and standard deviation of orientation about the roll axis. Decreasing the wing's flexibility significantly increased its projected surface area relative to the oncoming airflow, likely increasing the drag force it experienced during particular phases of the wing stroke. We hypothesize that higher drag forces on stiffened wings decrease body stability when the left and right wings encounter different flow conditions. Wing splinting also led to a small increase in body rotation rates in turbulent airflow, but this change was not statistically significant, possibly because bees with stiffened wings changed their flight behavior in turbulent flow. Overall, we found that wing flexibility improves flight stability in bumblebees, adding to the growing appreciation that wing flexibility is not merely an inevitable liability in flapping flight, but can enhance flight performance.

Hummingbird flight stability and control in freestream turbulent winds.

Airflow conditions close to the Earth's surface are often complex, posing challenges to flight stability and control for volant taxa. Relatively little is known about how well flying animals can contend with complex, adverse air flows, or about the flight control mechanisms used by animals to mitigate wind disturbances. Several recent studies have examined flight in the unsteady von Kármán vortex streets that form behind cylinders, generating flow disturbances that are predictable in space and time; these structures are relatively rare in nature, because they occur only the immediate, downstream vicinity of an object. In contrast, freestream turbulence is characterized by rapid, unpredictable flow disturbances across a wide range of spatial and temporal scales, and is nearly ubiquitous in natural habitats. Hummingbirds are ideal organisms for studying the influence of freestream turbulence on flight, as they forage in a variety of aerial conditions and are powerful flyers. We filmed ruby-throated hummingbirds (Archilochus colubris) maintaining position at a feeder in laminar and strongly turbulent (intensity ∼15%) airflow environments within a wind tunnel and compared their mean kinematics of the head, body, tail and wing, as well as variability in these parameters. Hummingbirds exhibited remarkably stable head position and orientation in both smooth and turbulent flow while maintaining position at the feeder. However, the hummingbird's body was less stable in turbulent flow and appeared to be most sensitive to disturbances along the mediolateral axis, displaying large lateral accelerations, translations and rolling motions during flight. The hummingbirds mitigated these disturbances by increasing mean wing stroke amplitude and stroke plane angle, and by varying these parameters asymmetrically between the wings and from one stroke to the next. They also actively varied the orientation and fan angle of the tail, maintaining a larger mean fan angle when flying in turbulent flow; this may improve their passive stability, but probably incurs an energetic cost as a result of increased drag. Overall, we observed many of the same kinematic changes noted previously for hummingbirds flying in a von Kármán vortex street, but we also observed kinematic changes associated with high force production, similar to those seen during load-lifting or high-speed flight. These findings suggest that flight may be particularly costly in fully mixed, freestream turbulence, which is the flow condition that hummingbirds are likely to encounter most frequently in natural habitats.

Clap and fling mechanism with interacting porous wings in tiny insect flight.

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.

Aerodynamic response of a red-tailed hawk to discrete transverse gusts.

A limiting factor in the design of smaller size uncrewed aerial vehicles is their inability to navigate through gust-laden environments. As a result, engineers have turned towards bio-inspired engineering approaches for gust mitigation techniques. In this study, the aerodynamics of a red-tailed hawk's response to variable-magnitude discrete transverse gusts was investigated. The hawk was flown in an indoor flight arena instrumented by multiple high-speed cameras to quantify the 3D motion of the bird as it navigated through the gust. The hawk maintained its flapping motion across the gust in all runs; however, it encountered the gust at different points in the flapping pattern depending on the run and gust magnitude. The hawk responded with a downwards pitching motion of the wing, decreasing the wing pitch angle to between -20∘and -5∘, and remained in this configuration until gust exit. The wing pitch data was then applied to a lower-order aerodynamic model that estimated lift coefficients across the wing. In gusts slower than the forward flight velocity (low gust ratio), the lift coefficient increases at a low-rate, to a maximum of around 2-2.5. In gusts faster than the forward flight velocity (high gust ratio), the lift coefficient initially increased rapidly, before increasing at a low-rate to a value around 4-5. In both regimes, the hawk's observed height change due to gust interaction was similar (and small), despite larger estimated lift coefficients over the high gust regime. This suggests another mitigation factor apart from the wing response is present. One potential factor is the tail pitching response observed here, which prior work has shown serves to mitigate pitch disturbances from gusts.

Aerodynamic Effects of Time-Varying Corrugations on Dragonfly Wings in Flapping Flight.

The aerodynamic effects of wing corrugation on insect flight have received widespread attention. However, there has hardly been any specific focus on dynamic changes to corrugation angle in the models. The flexible vein joints containing resilin in the wings of dragonflies and damselflies enable the longitudinal veins to rotate and thereby change the corrugation angles throughout flapping cycles. Therefore, a two-dimensional corrugated airfoil with time-varying corrugation angles is proposed and the aerodynamic performance is evaluated in terms of aerodynamic force, power and efficiency. The results indicate that the airfoil with time-varying corrugations outperforms the rigid one in terms of enhancing thrust and reducing power consumption. The aerodynamic performance of time-varying corrugated airfoils is optimal when the angle varies in a specific range, and an excessively large angle variation may have negative effects. In addition, excessive height or a negative leading edge of the corrugation can lead to a reduction in the thrust. A design concept for the 2D airfoil with time-varying corrugations is provided and the findings are of significance for enhancing the aerodynamic performance of biomimetic flexible flapping-wing vehicles.

Turbulence causes kinematic and behavioural adjustments in a flapping flier.

Turbulence is a widespread phenomenon in the natural world, but its influence on flapping fliers remains little studied. We assessed how freestream turbulence affected the kinematics, flight effort and track properties of homing pigeons (Columba livia), using the fine-scale variations in flight height as a proxy for turbulence levels. Birds showed a small increase in their wingbeat amplitude with increasing turbulence (similar to laboratory studies), but this was accompanied by a reduction in mean wingbeat frequency, such that their flapping wing speed remained the same. Mean kinematic responses to turbulence may therefore enable birds to increase their stability without a reduction in propulsive efficiency. Nonetheless, the most marked response to turbulence was an increase in the variability of wingbeat frequency and amplitude. These stroke-to-stroke changes in kinematics provide instantaneous compensation for turbulence. They will also increase flight costs. Yet pigeons only made small adjustments to their flight altitude, likely resulting in little change in exposure to strong convective turbulence. Responses to turbulence were therefore distinct from responses to wind, with the costs of high turbulence being levied through an increase in the variability of their kinematics and airspeed. This highlights the value of investigating the variability in flight parameters in free-living animals.

Wing deformation improves aerodynamic performance of forward flight of bluebottle flies flying in a flight mill.

Insect wings are flexible structures that exhibit deformations of complex spatiotemporal patterns. Existing studies on wing deformation underscore the indispensable role of wing deformation in enhancing aerodynamic performance. Here, we investigated forward flight in bluebottle flies, flying semi-freely in a magnetic flight mill; we quantified wing surface deformation using high-speed videography and marker-less surface reconstruction and studied the effects on aerodynamic forces, power and efficiency using computational fluid dynamics. The results showed that flies' wings exhibited substantial camber near the wing root and twisted along the wingspan, as they were coupled effects of deflection primarily about the claval flexion line. Such deflection was more substantial for supination during the upstroke when most thrust was produced. Compared with deformed wings, the undeformed wings generated 59-98% of thrust and 54-87% of thrust efficiency (i.e. ratio of thrust and power). Wing twist moved the aerodynamic centre of pressure proximally and posteriorly, likely improving aerodynamic efficiency.

Touchless underwater wall-distance sensing via active proprioception of a robotic flapper.

In this work, we explored a bioinspired method for underwater object sensing based on active proprioception. We investigated whether the fluid flows generated by a robotic flapper, while interacting with an underwater wall, can encode the distance information between the wall and the flapper, and how to decode this information using the proprioception within the flapper. Such touchless wall-distance sensing is enabled by the active motion of a flapping plate, which injects self-generated flow to the fluid environment, thus representing a form of active sensing. Specifically, we trained a long short-term memory (LSTM) neural network to predict the wall distance based on the force and torque measured at the base of the flapping plate. In addition, we varied the Rossby number (Ro, or the aspect ratio of the plate) and the dimensionless flapping amplitude (A∗) to investigate how the rotational effects and unsteadiness of self-generated flow respectively affect the accuracy of the wall-distance prediction. Our results show that the median prediction error is within 5% of the plate length for all the wall-distances investigated (up to 40 cm or approximately 2-3 plate lengths depending on theRo); therefore, confirming that the self-generated flow can enable underwater perception. In addition, we show that stronger rotational effects at lowerRolead to higher prediction accuracy, while flow unsteadiness (A∗) only has moderate effects. Lastly, analysis based on SHapley Additive exPlanations (SHAP) indicate that temporal features that are most prominent at stroke reversals likely promotes the wall-distance prediction.

Force balance in the take-off of a pierid butterfly: relative importance and timing of leg impulsion and aerodynamic forces.

Up to now, the take-off stage has remained an elusive phase of insect flight that was relatively poorly explored compared with other maneuvers. An overall assessment of the different mechanisms involved in force production during take-off has never been explored. Focusing on the first downstroke, we have addressed this problem from a force balance perspective in butterflies taking off from the ground. In order to determine whether the sole aerodynamic wing force could explain the observed motion of the insect, we have firstly compared a simple analytical model of the wing force with the acceleration of the insect's center of mass estimated from video tracking of the wing and body motions. Secondly, wing kinematics were also used for numerical simulations of the aerodynamic flow field. Similar wing aerodynamic forces were obtained by the two methods. However, neither are sufficient, nor is the inclusion of the ground effect, to predict faithfully the body acceleration. We have to resort to the leg forces to obtain a model that best fits the data. We show that the median and hind legs display an active extension responsible for the initiation of the upward motion of the insect's body, occurring before the onset of the wing downstroke. We estimate that legs generate, at various times, an upward force that can be much larger than all other forces applied to the insect's body. The relative timing of leg and wing forces explains the large variability of trajectories observed during the maneuvers.