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Concept: Lift


Small-winged drones can face highly varied aerodynamic requirements, such as high manoeuvrability for flight among obstacles and high wind resistance for constant ground speed against strong headwinds that cannot all be optimally addressed by a single aerodynamic profile. Several bird species solve this problem by changing the shape of their wings to adapt to the different aerodynamic requirements. Here, we describe a novel morphing wing design composed of artificial feathers that can rapidly modify its geometry to fulfil different aerodynamic requirements. We show that a fully deployed configuration enhances manoeuvrability while a folded configuration offers low drag at high speeds and is beneficial in strong headwinds. We also show that asymmetric folding of the wings can be used for roll control of the drone. The aerodynamic performance of the morphing wing is characterized in simulations, in wind tunnel measurements and validated in outdoor flights with a small drone.

Concepts: Aerodynamics, Reynolds number, Drag, Flight, Wing, Lift, Aircraft, Aerospace engineering


A prominent feature of gliding flight in snakes of the genus Chrysopelea is the unique cross-sectional shape of the body, which acts as the lifting surface in the absence of wings. When gliding, the flying snake Chrysopelea paradisi morphs its circular cross-section into a triangular shape by splaying its ribs and flattening its body in the dorsoventral axis, forming a geometry with fore-aft symmetry and a thick profile. Here, we aimed to understand the aerodynamic properties of the snake’s cross-sectional shape to determine its contribution to gliding at low Reynolds numbers. We used a straight physical model in a water tunnel to isolate the effects of 2D shape, analogously to studying the profile of an airfoil of a more typical flyer. Force measurements and time-resolved (TR) digital particle image velocimetry (DPIV) were used to determine lift and drag coefficients, wake dynamics and vortex-shedding characteristics of the shape across a behaviorally relevant range of Reynolds numbers and angles of attack. The snake’s cross-sectional shape produced a maximum lift coefficient of 1.9 and maximum lift-to-drag ratio of 2.7, maintained increases in lift up to 35 deg, and exhibited two distinctly different vortex-shedding modes. Within the measured Reynolds number regime (Re=3000-15,000), this geometry generated significantly larger maximum lift coefficients than many other shapes including bluff bodies, thick airfoils, symmetric airfoils and circular arc airfoils. In addition, the snake’s shape exhibited a gentle stall region that maintained relatively high lift production even up to the highest angle of attack tested (60 deg). Overall, the cross-sectional geometry of the flying snake demonstrated robust aerodynamic behavior by maintaining significant lift production and near-maximum lift-to-drag ratios over a wide range of parameters. These aerodynamic characteristics help to explain how the snake can glide at steep angles and over a wide range of angles of attack, but more complex models that account for 3D effects and the dynamic movements of aerial undulation are required to fully understand the gliding performance of flying snakes.

Concepts: Fluid dynamics, Aerodynamics, Snake, Lift, Airfoil, Angle of attack, Chrysopelea, Lift coefficient


Large ears enhance perception of echolocation and prey generated sounds in bats. However, external ears likely impair aerodynamic performance of bats compared to birds. But large ears may generate lift on their own, mitigating the negative effects. We studied flying brown long-eared bats, using high resolution, time resolved particle image velocimetry, to determine the aerodynamics of flying with large ears. We show that the ears and body generate lift at medium to cruising speeds (3-5 m/s), but at the cost of an interaction with the wing root vortices, likely reducing inner wing performance. We also propose that the bats use a novel wing pitch mechanism at the end of the upstroke generating thrust at low speeds, which should provide effective pitch and yaw control. In addition, the wing tip vortices show a distinct spiraling pattern. The tip vortex of the previous wingbeat remains into the next wingbeat and rotates together with a newly formed tip vortex. Several smaller vortices, related to changes in circulation around the wing also spiral the tip vortex. Our results thus show a new level of complexity in bat wakes and suggest large eared bats are less aerodynamically limited than previous wake studies have suggested.

Concepts: Fluid dynamics, Aerodynamics, Bat, Drag, Spiral, Fixed-wing aircraft, Lift, Wingtip vortices


The theories of aero- and hydrodynamics predict animal movement and device design in air and water through the computation of lift, drag, and thrust forces. Although models of terrestrial legged locomotion have focused on interactions with solid ground, many animals move on substrates that flow in response to intrusion. However, locomotor-ground interaction models on such flowable ground are often unavailable. We developed a force model for arbitrarily-shaped legs and bodies moving freely in granular media, and used this “terradynamics” to predict a small legged robot’s locomotion on granular media using various leg shapes and stride frequencies. Our study reveals a complex but generic dependence of stresses in granular media on intruder depth, orientation, and movement direction and gives insight into the effects of leg morphology and kinematics on movement.

Concepts: Fluid dynamics, Interaction, Aerodynamics, Force, Model, Classical mechanics, Drag, Lift


Understanding how moving organisms generate locomotor forces is fundamental to the analysis of aerodynamic and hydrodynamic flow patterns that are generated during body and appendage oscillation. In the past, this has been accomplished using two-dimensional planar techniques that require reconstruction of three-dimensional flow patterns. We have applied a new, fully three-dimensional, volumetric imaging technique that allows instantaneous capture of wake flow patterns, to a classic problem in functional vertebrate biology: the function of the asymmetrical (heterocercal) tail of swimming sharks to capture the vorticity field within the volume swept by the tail. These data were used to test a previous three-dimensional reconstruction of the shark vortex wake estimated from two-dimensional flow analyses, and show that the volumetric approach reveals a different vortex wake not previously reconstructed from two-dimensional slices. The hydrodynamic wake consists of one set of dual-linked vortex rings produced per half tail beat. In addition, we use a simple passive shark-tail model under robotic control to show that the three-dimensional wake flows of the robotic tail differ from the active tail motion of a live shark, suggesting that active control of kinematics and tail stiffness plays a substantial role in the production of wake vortical patterns.

Concepts: Mathematics, Fluid dynamics, Aerodynamics, Turbulence, Vortex, Vorticity, Lift, Potential flow


To maintain the quality of the feathers, birds regularly undergo moult. It is widely accepted that moult affects flight performance, but the specific aerodynamic consequences have received relatively little attention. Here we measured the components of aerodynamic drag from the wake behind a gliding jackdaw (Corvus monedula) at different stages of its natural wing moult. We found that span efficiency was reduced (lift induced drag increased) and the wing profile drag coefficient was increased. Both effects best correlated with the corresponding reduction in spanwise camber. The negative effects are partially mitigated by adjustments of wing posture to minimize gaps in the wing, and by weight loss to reduce wing loading. By studying the aerodynamic consequences of moult, we can refine our understanding of the emergence of various moulting strategies found among birds.

Concepts: Fluid dynamics, Aerodynamics, Drag, Wing, Flying and gliding animals, Jackdaw, Corvus, Lift


The motion of silica spheres with homeotropic anchoring sedimenting within the nematic liquid crystal 4-cyano-4'-pentylbiphenyl (5CB) has been studied at low Ericksen number. The magnitude of the spheres' velocity depends on the angle θ between the far-field nematic director and the gravitational force, indicating an anisotropic Stokes drag. When the director is oriented at an oblique angle to the gravitational force, the velocity also acquires a component normal to the force, demonstrating the existence of a lift force generated by the fluid. The magnitude and direction of the velocity as functions of θ quantitatively obey theoretically predicted forms.

Concepts: Fluid dynamics, Aerodynamics, Force, Liquid crystal, Drag, Lift, Drag equation, Bernoulli's principle


Mosquitoes exhibit unusual wing kinematics; their long, slender wings flap at remarkably high frequencies for their size (>800 Hz)and with lower stroke amplitudes than any other insect group. This shifts weight support away from the translation-dominated, aerodynamic mechanisms used by most insects, as well as by helicopters and aeroplanes, towards poorly understood rotational mechanisms that occur when pitching at the end of each half-stroke. Here we report free-flight mosquito wing kinematics, solve the full Navier-Stokes equations using computational fluid dynamics with overset grids, and validate our results with in vivo flow measurements. We show that, although mosquitoes use familiar separated flow patterns, much of the aerodynamic force that supports their weight is generated in a manner unlike any previously described for a flying animal. There are three key features: leading-edge vortices (a well-known mechanism that appears to be almost ubiquitous in insect flight), trailing-edge vortices caused by a form of wake capture at stroke reversal, and rotational drag. The two new elements are largely independent of the wing velocity, instead relying on rapid changes in the pitch angle (wing rotation) at the end of each half-stroke, and they are therefore relatively immune to the shallow flapping amplitude. Moreover, these mechanisms are particularly well suited to high aspect ratio mosquito wings.

Concepts: Insect, Fluid dynamics, Aerodynamics, Bat, Computational fluid dynamics, Flight, Wing, Lift


At first sight, echolocating bats face a difficult trade-off. As flying animals, they would benefit from a streamlined geometric shape to reduce aerodynamic drag and increase flight efficiency. However, as echolocating animals, their pinnae generate the acoustic cues necessary for navigation and foraging. Moreover, species emitting sound through their nostrils often feature elaborate noseleaves that help in focussing the emitted echolocation pulses. Both pinnae and noseleaves reduce the streamlined character of a bat’s morphology. It is generally assumed that by compromising the streamlined charactered of the geometry, the head morphology generates substantial drag, thereby reducing flight efficiency. In contrast, it has also been suggested that the pinnae of bats generate lift forces counteracting the detrimental effect of the increased drag. However, very little data exist on the aerodynamic properties of bat pinnae and noseleaves. In this work, the aerodynamic forces generated by the heads of seven species of bats, including noseleaved bats, are measured by testing detailed 3D models in a wind tunnel. Models of Myotis daubentonii, Macrophyllum macrophyllum, Micronycteris microtis, Eptesicus fuscus, Rhinolophus formosae, Rhinolophus rouxi and Phyllostomus discolor are tested. The results confirm that non-streamlined facial morphologies yield considerable drag forces but also generate substantial lift. The net effect is a slight increase in the lift-to-drag ratio. Therefore, there is no evidence of high aerodynamic costs associated with the morphology of bat heads.

Concepts: Fluid dynamics, Aerodynamics, Force, Animal echolocation, Bat, Flight, Wing, Lift


There are three common methods for calculating the lift generated by a flying animal based on the measured airflow in the wake. However, these methods might not be accurate according to computational and robot-based studies of flapping wings. Here we test this hypothesis for the first time for a slowly flying Pacific parrotlet in still air using stereo particle image velocimetry recorded at 1000 Hz. The bird was trained to fly between two perches through a laser sheet wearing laser safety goggles. We found that the wingtip vortices generated during mid-downstroke advected down and broke up quickly, contradicting the frozen turbulence hypothesis typically assumed in animal flight experiments. The quasi-steady lift at mid-downstroke was estimated based on the velocity field by applying the widely used Kutta-Joukowski theorem, vortex ring model, and actuator disk model. The calculated lift was found to be sensitive to the applied model and its different parameters, including vortex span and distance between the bird and laser sheet-rendering these three accepted ways of calculating weight support inconsistent. The three models predict different aerodynamic force values mid-downstroke compared to independent direct measurements with an aerodynamic force platform that we had available for the same species flying over a similar distance. Whereas the lift predictions of the Kutta-Joukowski theorem and the vortex ring model stayed relatively constant despite vortex breakdown, their values were too low. In contrast, the actuator disk model predicted lift reasonably accurately before vortex breakdown, but predicted almost no lift during and after vortex breakdown. Some of these limitations might be better understood, and partially reconciled, if future animal flight studies report lift calculations based on all three quasi-steady lift models instead. This would also enable much needed meta studies of animal flight to derive bioinspired design principles for quasi-steady lift generation with flapping wings.

Concepts: Mathematics, Fluid dynamics, Prediction, Aerodynamics, Wing, Vortex, Lift, Wingtip vortices