SUMMARYThe aerodynamic performance of hovering insects is largely explained by the presence of a stably attached leading edge vortex (LEV) on top of their wings. Although LEVs have been visualized on real, physically modeled, and simulated insects, the physical mechanisms responsible for their stability are poorly understood. To gain fundamental insight into LEV stability on flapping fly wings we expressed the Navier-Stokes equations in a rotating frame of reference attached to the wing's surface. Using these equations we show that LEV dynamics on flapping wings are governed by three terms: angular, centripetal and Coriolis acceleration. Our analysis for hovering conditions shows that angular acceleration is proportional to the inverse of dimensionless stroke amplitude, whereas Coriolis and centripetal acceleration are proportional to the inverse of the Rossby number. Using a dynamically scaled robot model of a flapping fruit fly wing to systematically vary these dimensionless numbers, we determined which of the three accelerations mediate LEV stability. Our force measurements and flow visualizations indicate that the LEV is stabilized by the 'quasi-steady' centripetal and Coriolis accelerations that are present at low Rossby number and result from the propeller-like sweep of the wing. In contrast, the unsteady angular acceleration that results from the back and forth motion of a flapping wing does not appear to play a role in the stable attachment of the LEV. Angular acceleration is, however, critical for LEV integrity as we found it can mediate LEV spiral bursting, a high Reynolds number effect. Our analysis and experiments further suggest that the mechanism responsible for LEV stability is not dependent on Reynolds number, at least over the range most relevant for insect flight (100

Gliding birds continually change the shape and size of their wings, presumably to exploit the profound effect of wing morphology on aerodynamic performance. That birds should adjust wing sweep to suit glide speed has been predicted qualitatively by analytical glide models, which extrapolated the wing's performance envelope from aerodynamic theory. Here we describe the aerodynamic and structural performance of actual swift wings, as measured in a wind tunnel, and on this basis build a semi-empirical glide model. By measuring inside and outside swifts' behavioural envelope, we show that choosing the most suitable sweep can halve sink speed or triple turning rate. Extended wings are superior for slow glides and turns; swept wings are superior for fast glides and turns. This superiority is due to better aerodynamic performance-with the exception of fast turns. Swept wings are less effective at generating lift while turning at high speeds, but can bear the extreme loads. Finally, our glide model predicts that cost-effective gliding occurs at speeds of 8-10 m s(-1), whereas agility-related figures of merit peak at 15-25 m s(-1). In fact, swifts spend the night ('roost') in flight at 8-10 m s(-1) (ref. 11), thus our model can explain this choice for a resting behaviour. Morphing not only adjusts birds' wing performance to the task at hand, but could also control the flight of future aircraft.

Helicopter Seed Lift The “helicopter” seeds of maple trees and other similar autorotating seeds detach from their parent tree under windy conditions and gyrate as they are dispersed by the wind. The reproductive success of the tree depends on the flight performance of its seeds. Autorotating seeds are known to generate high lift as they slowly descend through the air, but the means by which they do so is unclear. Lentink et al. (p. 1438 , see the cover) have elucidated the aerodynamic mechanism for high lift in autorotating seeds using a robot model seed and a three-dimensional flow measurement technique. Maple seeds and a hornbeam seed create a prominent leading-edge vortex that is similar to the flow structures that are responsible for the high lift generated by the wings of hovering insects and bats. Thus, both animals and plants have converged on an identical aerodynamic solution for generating lift.

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 mechanismsadded 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.

Hummingbirds are the only birds that can sustain hovering. This unique flight behaviour comes, however, at high energetic cost. Based on helicopter and aeroplane design theory, we expect that hummingbird wing aspect ratio (AR), which ranges from about 3.0 to 4.5, determines aerodynamic efficacy. Previous quasi-steady experiments with a wing spinner set-up provide no support for this prediction. To test this more carefully, we compare the quasi-steady hover performance of 26 wings, from 12 hummingbird taxa. We spun the wings at angular velocities and angles of attack that are representative for every species and measured lift and torque more precisely. The power (aerodynamic torque  angular velocity) required to lift weight depends on aerodynamic efficacy, which is measured by the power factor. Our comparative analysis shows that AR has a modest influence on lift and drag forces, as reported earlier, but interspecific differences in power factor are large. During the downstroke, the power required to hover decreases for larger AR wings at the angles of attack at which hummingbirds flap their wings ( p , 0.05). Quantitative flow visualization demonstrates that variation in hover power among hummingbird wings is driven by similar stable leading edge vortices that delay stall during the down-and upstroke. A side-byside aerodynamic performance comparison of hummingbird wings and an advanced micro helicopter rotor shows that they are remarkably similar.

Dragonfly wings are highly corrugated, which increases the stiffness and strength of the wing significantly, and results in a lightweight structure with good aerodynamic performance. How insect wings carry aerodynamic and inertial loads, and how the resonant frequency of the flapping wings is tuned for carrying these loads, is however not fully understood. To study this we made a three-dimensional scan of a dragonfly (Sympetrum vulgatum) fore-and hindwing with a micro-CT scanner. The scans contain the complete venation pattern including thickness variations throughout both wings. We subsequently approximated the forewing architecture with an efficient three-dimensional beam and shell model. We then determined the wing's natural vibration modes and the wing deformation resulting from analytical estimates of 8 load cases containing aerodynamic and inertial loads (using the finite element solver Abaqus). Based on our computations we find that the inertial loads are 1.5 to 3 times higher than aerodynamic pressure loads. We further find that wing deformation is smaller during the downstroke than during the upstroke, due to structural asymmetry. The natural vibration mode analysis revealed that the structural natural frequency of a dragonfly wing in vacuum is 154 Hz, which is approximately 4.8 times higher than the natural flapping frequency of dragonflies in hovering flight (32.3 Hz). This insight in the structural properties of dragonfly wings could inspire the design of more effective wings for insectsized flapping micro air vehicles: The passive shape of aeroelastically tailored wings inspired by dragonflies can in principle be designed more precisely compared to sail like wings -which can make the dragonfly-like wings more aerodynamically effective.

Airplanes and helicopters use high aspect ratio wings to reduce the power required to fly, but must operate at low angle of attack to prevent flow separation and stall. Animals capable of slow sustained flight, such as hummingbirds, have low aspect ratio wings and flap their wings at high angle of attack without stalling. Instead, they generate an attached vortex along the leading edge of the wing that elevates lift. Previous studies have demonstrated that this vortex and high lift can be reproduced by revolving the animal wing at the same angle of attack. How do flapping and revolving animal wings delay stall and reduce power? It has been hypothesized that stall delay derives from having a short radial distance between the shoulder joint and wing tip, measured in chord lengths. This non-dimensional measure of wing length represents the relative magnitude of inertial forces versus rotational accelerations operating in the boundary layer of revolving and flapping wings. Here we show for a suite of aspect ratios, which represent both animal and aircraft wings, that the attachment of the leading edge vortex on a revolving wing is determined by wing aspect ratio, defined with respect to the centre of revolution. At high angle of attack, the vortex remains attached when the local radius is shorter than four chord lengths and separates outboard on higher aspect ratio wings. This radial stall limit explains why revolving high aspect ratio wings (of helicopters) require less power compared with low aspect ratio wings (of hummingbirds) at low angle of attack and vice versa at high angle of attack.

Since the Wright Flyer, engineers have strived to develop flying machines with morphing wings that can control flight as deftly as birds. Birds morph their wing planform parameters simultaneously—including sweep, span, and area—in a way that has proven to be particularly challenging to embody robotically. Previous solutions have primarily centered around the classical aerospace paradigm of controlling every degree of freedom to ensure predictable performance, but underperform compared with birds. To understand how birds accomplish wing morphing, we measured the kinematics of wing flexion and extension in common pigeons, Columba livia. The skeletal and feather kinematics show that the 20 primary and 20 secondary feathers are coordinated via approximately linear transfer functions controlled by wrist and finger motion. To replicate this control principle in a robot, we developed a biohybrid morphing wing with real feathers to understand the underlying design principles. The outcome, PigeonBot, embodies 42 degrees of freedom that control the position of 40 elastically connected feathers via four servo-actuated wrist and finger joints. Our flight tests demonstrate that the soft feathered wings morph rapidly and robustly under aerodynamic loading. They not only enable wing morphing but also make robot interactions safer, the wing more robust to crashing, and the wing reparable via “preening.” In flight tests, we found that both asymmetric wrist and finger motion can initiate turn maneuvers—evidence that birds may use their fingers to steer in flight.