Flexible Wing Kinematics of a Free-Flying Beetle (Rhinoceros Beetle Trypoxylus Dichotomus)

Detailed 3-Dimensional (3D) wing kinematics was experimentally presented in free flight of a beetle,Trypoxylus dichotomus,which has a pair of elytra (forewings) and flexible hind wings.The kinematic parameters such as the wing tip trajectory,angle of attack and camber deformation were obtained from a 3D reconstruction technique that involves the use of two synchronized high-speed cameras to digitize various points marked on the wings.Our data showed outstanding characteristics of deformation and flexibility of the beetle's hind wing compared with other measured insects,especially in the chordwise and spanwise directions during flapping motion.The hind wing produced 16％ maximum positive camber deformation during the downstroke.It also experienced twisted shape showing large variation of the angle of attack from the root to the tip during the upstroke.     

Non-Jumping Take off Performance in Beetle Flight （Rhinoceros Beetle Trypoxylus dichotomus）

In recent decades, the take-off mechanisms of flying animals have received much attention in insect flight initiation. Most of previous works have focused on the jumping mechanism, which is the most common take-off mechanism found in flying animals. Here, we presented that the rhinoceros beetle, Trypoxylus dichotomus, takes offwithout jumping. In this study, we used 3-Dimensional （3D） high-speed video techniques to quantitatively analyze the wings and body kinematics during the initiation periods of flight. The details of the flapping angle, angle of attack of the wings and the roll, pitch and yaw angles of the body were investigated to understand the mechanism of take-off in T. dichotomus. The beetle took off gradually with a small velocity and small acceleration. The body kinematic analyses showed that the beetle exhibited stable take-off. To generate high lift force, the beetle modulated its hind wing to control the angle of attack; the angle of attack was large during the upstroke and small during the downstroke. The legs of beetle did not contract and strongly release like other insects. The hind wing could be con- sidered as a main source of lift for heavy beetle.     

Numerical investigation of the aerodynamic characteristics of a hovering Coleopteran insect

The aerodynamic characteristics of the Coleopteran beetle species Epilachna quadricollis, a species with flexible hind wings and stiff elytra (fore wings), are investigated in terms of hovering flight. The flapping wing kinematics of the Coleopteran insect are modeled through experimental observations with a digital high-speed camera and curve fitting from an ideal harmonic kinematics model. This model numerically simulates flight by estimating a cross section of the wing as a two-dimensional elliptical plane. There is currently no detailed study on the role of the elytron or how the elytron-hind wing interaction affects aerodynamic performance. In the case of hovering flight, the relatively small vertical or horizontal forces generated by the elytron suggest that the elytron makes no significant contribution to aerodynamic force.     

Two-and Three-Dimensional Simulations of Beetle Hind Wing Flapping during Free Forward Flight

Aerodynamic characteristic of the beetle, Trypoxylus dichotomus, which has a pair of elytra (forewings) and hind wings, is numerically investigated. Based on the experimental results of wing kinematics, two-dimensional (2D) and three-dimensional (3D) computational fluid dynamic simulations were carried out to reveal aerodynamic performance of the hind wing. The roles of the spiral Leading Edge Vortex (LEV) and the spanwise flow were clarified by comparing 2D and 3D simulations. Mainly due to pitching down of chord line during downstroke in highly inclined stroke plane, relatively high averaged thrust was produced in the free forward flight of the beetle. The effects of the local corrugation and the camber variation were also investigated for the beetle's hind wings. Our results show that the camber variation plays a significant role in improving both lift and thrust in the flapping. On the other hand, the local corrugation pattern has no significant effect on the aerodynamic force due to large angle of attack during flapping.     

Hovering and intermittent flight in birds

Two styles of bird locomotion, hovering and intermittent flight, have great potential to inform future development of autonomous flying vehicles. Hummingbirds are the smallest flying vertebrates, and they are the only birds that can sustain hovering. Their ability to hover is due to their small size, high wingbeat frequency, relatively large margin of mass-specific power available for flight and a suite of anatomical features that include proportionally massive major flight muscles (pectoralis and supracoracoideus) and wing anatomy that enables them to leave their wings extended yet turned over (supinated) during upstroke so that they can generate lift to support their weight. Hummingbirds generate three times more lift during downstroke compared with upstroke, with the disparity due to wing twist during upstroke. Much like insects, hummingbirds exploit unsteady mechanisms during hovering including delayed stall during wing translation that is manifest as a leading-edge vortex (LEV) on the wing and rotational circulation at the end of each half stroke. Intermittent flight is common in small- and medium-sized birds and consists of pauses during which the wings are flexed (bound) or extended (glide). Flap-bounding appears to be an energy-saving style when flying relatively fast, with the production of lift by the body and tail critical to this saving. Flap-gliding is thought to be less costly than continuous flapping during flight at most speeds. Some species are known to shift from flap-gliding at slow speeds to flap-bounding at fast speeds, but there is an upper size limit for the ability to bound (~0.3 kg) and small birds with rounded wings do not use intermittent glides.     

Flow visualization and unsteady aerodynamics in the flight of the hawkmoth,Manduca sexta

The aerodynamic mechanisms employed durng the flight of the hawkmoth, Manduca sexta, have been investigated through smoke visualization studies with tethered moths. Details of the flow around the wings and of the overall wake structure were recorded as stereophotographs and high-speed video sequences. The changes in flow which accompanied increases in flight speed from 0.4 to 5.7 m s^-1 were analysed. The wake consists of an alternating series of horizontal and vertical vortex rings which are generated by successive down- and upstrokes, respectively. The downstroke produces significantly more lift than the upstroke due to a leading-edge vortex which is stabilized by a radia flow moving out towards the wingtip. The leading-edge vortex grew in size with increasing forward flight velocity. Such a phenomenon is proposed as a likely mechanism for lift enhancement in many insect groups. During supination, vorticity is shed from the leading edge as postulated in the ''flex'' mechanism. This vorticity would enhance upstroke lift if it was recaptured diring subsequent translation, but it is not. Instead, the vorticity is left behind and the upstroke circulation builds up slowly. A small jet provides additional thrust as the trailing edges approach at the end of the upstroke. The stereophotographs also suggest that the bound circulation may not be reversed between half strokes at the fastest flight speeds.     

Aerodynamic performance of a hovering hawkmoth with flexible wings: a computational approach

Insect wings are deformable structures that change shape passively and dynamically owing to inertial and aerodynamic forces during flight. It is still unclear how the three-dimensional and passive change of wing kinematics owing to inherent wing flexibility contributes to unsteady aerodynamics and energetics in insect flapping flight. Here, we perform a systematic fluid-structure interaction based analysis on the aerodynamic performance of a hovering hawkmoth, Manduca, with an integrated computational model of a hovering insect with rigid and flexible wings. Aerodynamic performance of flapping wings with passive deformation or prescribed deformation is evaluated in terms of aerodynamic force, power and efficiency. Our results reveal that wing flexibility can increase downwash in wake and hence aerodynamic force: first, a dynamic wing bending is observed, which delays the breakdown of leading edge vortex near the wing tip, responsible for augmenting the aerodynamic force-production; second, a combination of the dynamic change of wing bending and twist favourably modifies the wing kinematics in the distal area, which leads to the aerodynamic force enhancement immediately before stroke reversal. Moreover, an increase in hovering efficiency of the flexible wing is achieved as a result of the wing twist. An extensive study of wing stiffness effect on aerodynamic performance is further conducted through a tuning of Young's modulus and thickness, indicating that insect wing structures may be optimized not only in terms of aerodynamic performance but also dependent on many factors, such as the wing strength, the circulation capability of wing veins and the control of wing movements.     

Effect of outer wing separation on lift and thrust generation in a flapping wing system

We explore the implementation of wing feather separation and lead-lagging motion to a flapping wing. A biomimetic flapping wing system with separated outer wings is designed and demonstrated. The artificial wing feather separation is implemented in the biomimetic wing by dividing the wing into inner and outer wings. The features of flapping, lead-lagging, and outer wing separation of the flapping wing system are captured by a high-speed camera for evaluation. The performance of the flapping wing system with separated outer wings is compared to that of a flapping wing system with closed outer wings in terms of forward force and downward force production. For a low flapping frequency ranging from 2.47 to 3.90 Hz, the proposed biomimetic flapping wing system shows a higher thrust and lift generation capability as demonstrated by a series of experiments. For 1.6 V application (lower frequency operation), the flapping wing system with separated wings could generate about 56% higher forward force and about 61% less downward force compared to that with closed wings, which is enough to demonstrate larger thrust and lift production capability of the separated outer wings. The experiments show that the outer parts of the separated wings are able to deform, resulting in a smaller amount of drag production during the upstroke, while still producing relatively greater lift and thrust during the downstroke.     

Biomechanics and physiology of gait selection in flying birds

Two wing-beat gaits, distinguished by the presence or absence of lift production during the upstroke, are currently used to describe avian flight. Vortex-visualization studies indicate that lift is produced only during the downstroke in the vortex-ring gait and that lift is produced continuously in the continuous-vortex gait. Tip-reversal and feathered upstrokes represent different forms of vortex-ring gait distinguished by wing kinematics. Useful aerodynamic forces may be produced during tip-reversal upstroke in slow flight and during a feathered upstroke in fast flight, but it is probable that downstroke forces are much greater in magnitude. Uncertainty about the function of these types of upstroke may be resolved when more data are available on wake structure in different flight speeds and modes. Inferring from wing kinematics and available data on wake structure, birds with long wings or wings of high aspect ratio use a vortex-ring gait with tip-reversal upstroke at slow speeds, a vortex-ring gait with a feathered upstroke at intermediate speeds, and a continuous-vortex gait at fast speeds. Birds with short wings or wings of low aspect ratio use a vortex-ring gait with a feathered upstroke at all speeds. Regardless of wing shape, species tend to use a vortex-ring gait for acceleration and a continuous-vortex gait for deceleration. Some correlations may exist between gait selection and the function of the muscular and respiratory system. However, overall variation in wing kinematics, muscle activity, and respiratory activity is continuous rather than categorical. To further our understanding of gait selection in flying birds, it is important to test whether upstroke function varies in a similar manner. Transitions between lifting and nonlifting upstrokes may be more subtle and gradual than implied by a binomial scheme of classification.     

Animal flight dynamics II. Longitudinal stability in flapping flight

Stability is essential to flying and is usually assumed to be especially problematic in flapping flight. If so, problems of stability may have presented a particular hurdle to the evolution of flapping flight. In spite of this, the stability of flapping flight has never been properly analysed. Here we use quasi-static and blade element approaches to analyse the stability provided by a flapping wing. By using reduced order approximations to the natural modes of motion, we show that wing beat frequencies are generally high enough compared to the natural frequencies of motion for a quasi-static approach to be valid as a first approximation. Contrary to expectations, we find that there is noting inherently destabilizing about flapping: beating the wings faster simply amplifies any existing stability or instability, and flapping can even enhance stability compared to gliding at the same air speed. This suggests that aerodynamic stability may not have been a particular hurdle in the evolution of flapping flight. Hovering animals, like hovering helicopters, are predicted to possess neutral static stability. Flapping animals, like fixed wing aircraft, are predicted to be stable in forward flight if the mean flight force acts above and/or behind the centre of gravity. In this case, the downstroke will always be stabilizing. The stabilizing contribution may be diminished by an active upstroke with a low advance ratio and more horizontal stroke plane; other forms of the upstroke may make a small positive contribution to stability. An active upstroke could, therefore, be used to lower stability and enhance manoeuvrability. Translatory mechanisms of unsteady lift production are predicted to amplify the stability predicted by a quasi-static analysis. Non-translatory mechanisms will make little or no contribution to stability. This may be one reason why flies, and other animals which rely upon non-translatory aerodynamic mechanisms, often appear inherently unstable.     

Flight Characteristics of Two Plume Moths, Alucita pentadacty/a L. and Orneodes hexadactyla L. (Microlepidoptera)

Norberg, R. Å. (Department of Zoology, University of Gothenburg, Göteborg, Sweden.) Flight characteristics of two plume moths, Alucita pentadactyla L. and Orneodes hexadactyla L. (Microlepidoptera). Zool. Scripta 1 (6): 241–246,1972.–Multiple exposure photographs of up to 100 exposures/sec were taken on two plume moth species in free, unrestrained flight, in order to determine approximate lift/drag ratios and other functional characteristics of their wings, which are of a remarkable structure for insects of this size. In Alucita the forewing is cleft in two fringed lobes, the hind-wing in three, while in Orneodes both forewing and hindwing are deeply cleft in six very narrow, fringed lobes. Wing stroke frequencies are ca. 33 Hz in A. pentadactyla and ca. 40 Hz in O. hexadactyla. During both the downstroke and the upstroke the fringed wing lobes lie edge against edge, thus forming a continuous wing surface. The upstroke seems to contribute no useful forces in A. pentadactyla, possibly some propulsive force in O. hexadactyla. The wings are strongly supinated in the upstroke to minimize drag. From relative wind diagrams, lift/drag ratios of 1.1 and 1.4 (minimum values) can be read for A. pentadactyla and O. hexadactyla, respectively. It is thus clear that these species do not make more use of drag forces than of lift forces. However, in A. pentadactyla the drag force in the downstroke may be almost as large as the lift force. Since drag certainly is small in the upstroke, the drag force probably contributes significantly to useful forces for flight in A. pentadactyla. These plume moths operate at Reynolds numbers of ca. 700. Reynolds numbers are calculated for very small insects. It is obvious that the wings of the smallest insects must be operating at Reynolds numbers of about 1. The fringed wings of small insects are briefly discussed.     

The vortex wake of a 'hovering' model hawkmoth

Visualization experiments with Manduca sexta have revealed the presence of a leading-edge vortex and a highly three-dimensional flow pattern. To further investigate this important discovery, a scaled-up robotic insect was built (the ''flapper'') which could mimic the complex movements of the wings of a hovering hawkmoth. Smoke released from the leading edge of the flapper wing revealed a small but strong leading-edge vortex on the downstroke. This vortex had a high axial flow velocity and was stable, separating from the wing at approximately 75 per cent of the wing length. It connected to a large, tangled tip vortex, extending back to a combining stopping and starting vortex from pronation. At the end of the downstroke, the wake could be approximated as one vortex ring per wing. Based on the size and velocity of the vortex rings, the mean lift force during the downstroke was estimated to be about 1.5 times the body weight of a hawkmoth, confirming that the downstroke is the main provider of lift force.     

The Role of Elytra in Beetle Flight: I. Generation of Quasi-Static Aerodynamic Forces

We conducted a comprehensive study to investigate the aerodynamic characteristics and force generation of the elytra of abeetle,Allomyrina dichotoma.Our analysis included wind tunnel experiments and three-dimensional computational fluiddynamics simulations using ANSYS-CFX software.Our first approach was a quasi-static study that considered the effect ofinduced flapping flow due to the flapping motion of the fore-wings (elytra) at a frequency of around 30 Hz to 40 Hz.The dihedralangle was varied to represent flapping motion during the upstroke and downstroke.We found that an elytron producespositive lift at 0° geometric angle of attack,negative lift during the upstroke,and always produces drag during both the upstrokeand downstroke.We also found that the lift coefficient of an elytron does not drop even at a very high geometric angle of attack.For a beetle with a body weight of 5 g,based on the quasi-static method,the fore-wings (elytra) can produce lift of less than 1%of its body weight.     

Modulation of leading edge vorticity and aerodynamic forces in flexible flapping wings

In diverse biological flight systems, the leading edge vortex has been implicated as a flow feature of key importance in the generation of flight forces. Unlike fixed wings, flapping wings can translate at higher angles of attack without stalling because their leading edge vorticity is more stable than the corresponding fixed wing case. Hence, the leading edge vorticity has often been suggested as the primary determinant of the high forces generated by flapping wings. To test this hypothesis, it is necessary to modulate the size and strength of the leading edge vorticity independently of the gross kinematics while simultaneously monitoring the forces generated by the wing. In a recent study, we observed that forces generated by wings with flexible trailing margins showed a direct dependence on the flexural stiffness of the wing. Based on that study, we hypothesized that trailing edge flexion directly influences leading edge vorticity, and thereby the magnitude of aerodynamic forces on the flexible flapping wings. To test this hypothesis, we visualized the flows on wings of varying flexural stiffness using a custom 2D digital particle image velocimetry system, while simultaneously monitoring the magnitude of the aerodynamic forces. Our data show that as flexion decreases, the magnitude of the leading edge vorticity increases and enhances aerodynamic forces, thus confirming that the leading edge vortex is indeed a key feature for aerodynamic force generation in flapping flight. The data shown here thus support the hypothesis that camber influences instantaneous aerodynamic forces through modulation of the leading edge vorticity.     

Normalized lift: an energy interpretation of the lift coefficient simplifies comparisons of the lifting ability of rotating and flapping surfaces

For a century, researchers have used the standard lift coefficient C(L) to evaluate the lift, L, generated by fixed wings over an area S against dynamic pressure, ?ρv(2), where v is the effective velocity of the wing. Because the lift coefficient was developed initially for fixed wings in steady flow, its application to other lifting systems requires either simplifying assumptions or complex adjustments as is the case for flapping wings and rotating cylinders.This paper interprets the standard lift coefficient of a fixed wing slightly differently, as the work exerted by the wing on the surrounding flow field (L/ρ·S), compared against the total kinetic energy required for generating said lift, ?v(2). This reinterpreted coefficient, the normalized lift, is derived from the work-energy theorem and compares the lifting capabilities of dissimilar lift systems on a similar energy footing. The normalized lift is the same as the standard lift coefficient for fixed wings, but differs for wings with more complex motions; it also accounts for such complex motions explicitly and without complex modifications or adjustments. We compare the normalized lift with the previously-reported values of lift coefficient for a rotating cylinder in Magnus effect, a bat during hovering and forward flight, and a hovering dipteran.The maximum standard lift coefficient for a fixed wing without flaps in steady flow is around 1.5, yet for a rotating cylinder it may exceed 9.0, a value that implies that a rotating cylinder generates nearly 6 times the maximum lift of a wing. The maximum normalized lift for a rotating cylinder is 1.5. We suggest that the normalized lift can be used to evaluate propellers, rotors, flapping wings of animals and micro air vehicles, and underwater thrust-generating fins in the same way the lift coefficient is currently used to evaluate fixed wings.     

ON THE RECONSTRUCTION OF PTEROSAURS AND THEIR MANNER OF FLIGHT, WITH NOTES ON VORTEX WAKES

Classical pterosaur reconstructions are variants on a ‘bat-analogy’, whereby the wing is conceived as a simple membrane with no inherent bending strength, stretched between the arm and leg skeletons. The legs are considered to be splayed out to the sides, as in bats, so that the animal would have to adopt a quadrupedal stance on the ground, supported on its feet and the metacarpo-phalangeal joints. In recent years an alternative ‘bird-analogy’ has come to be generally accepted. This hypothesis, most elements of which are due to Padian (1983 a, b) calls for the animal to stand upright on its legs like a bird. The wings are independent of the legs, as in birds, are stiffened by skeletal fibres in the membrane, and have a very narrow, sharply pointed shape. There are difficulties in reconciling the bird-analogy with the evidence. The long-tailed rhamphorhynchs might conceivably have balanced their weight about their hip joints but this would not have been possible for the short-tailed pterodactyls. The bird pelvis shows modifications which permit bipedal standing in spite of the reduction of the tail, but no equivalent adaptations are seen in pterodactyls. Besides, all known pterosaur pelvises, except that of the giant pterodactyl Pteranodon were open ventrally, which would have precluded the legs from being brought to a parasagittal position, as required for bipedal walking. The notion that the wing was not attached to the legs is based on negative evidence, in that no clear impressions of the inner end of the wing membrane are preserved in the fossils. However one pterodactyl fossil shows a membrane edge approaching the ankle joint. In fossils that are preserved with the wings forward, the legs have been pulled forwards by the ankles. A tendon connecting the ankle to the wing tip is consistent with the evidence. The ‘fibres’ in the wing membranes are actually impressions of surface ridges, with no internal structure, and are better interpreted as surface wrinkles in the skin, caused by contraction of elastic fibres within the membrane. The bird analogy also results in a very unsatisfactory wing from an aerodynamic point of view. The structure of an animal wing is best understood in terms of the type of vortex wake it is adapted to generate. Hummingbirds, and insects capable of economical hovering, have wings that can be inverted on the upstroke, and when hovering, generate a wake consisting of two vortex rings per wingbeat cycle. The span of such wings is fixed, which implies that they create a ‘ladder wake’ in cruising flight, consisting of a pair of undulating wing-tip vortices, joined by a transverse vortex at each transition from downstroke to upstroke and back. Normal birds cannot invert their wings, and so are less efficient in hovering, but they can shorten the wing during the upstroke in cruising flight. This creates a ‘concertina wake’, with no transverse vortices. Hummingbirds show very limited migration performance, compared with normal birds, with the implication that a wing capable of creating a concertina wake is more economical in cruising flight than one creating a ladder wake, and is an essential adaptation for long-distance migration. A revised reconstruction of the pterosaur wing starts from the observations that, contrary to the currently popular bird-analogy, pterosaurs were not bipedal, their wings did not contain stiffening fibres but did contain elastic fibres, and the trailing edge of the membrane was supported by a tendon joining the tip of the wing finger to the ankle. A hypothetical arrangement of elastic fibres, that accounts well for the observed pattern of wrinkles in contracted wings, also allows the planform shape of the wing to be adjusted in much the same way as seen in birds, although using a completely different mechanism. It opens the possibility that pterosaurs could fly with a concertina wake, and thus could have been long-distance migrators like modern birds. Although this hypothetical wing is mechanically somewhat bat-like, it is not a return to the classical bat-analogy. It would not have the high degree of control over profile shape, which gives bats their outstanding manoeuvrability. On the other hand bats do not have the degree of control over their wingspan that is suggested here for pterosaurs, and consequently are not notable for migration performance.     

Design and Demonstration of Insect Mimicking Foldable Artificial Wing Using Four-Bar Linkage Systems

In this work, we develop an artificial foldable wing that mimics the hind wing of a beetle （Allomyrina dichotoma）. In real flight, the beetle unfolds forewings and hind wings, and maintains the unfolded configuration unless it is exhausted. The artificial wing has to be able to maintain a fully unfolded configuration while flapping at a desirable flapping frequency. The artificial foldable hind wing developed in this work is based on two four-bar linkages which adapt the behaviors of the beetle＇s hind wing. The four-bar-linkages are designed to mimic rotational motion of the wing base and the vein folding/unfolding motion of the beetle＇s hind wing. The behavior of the artificial wings, which are installed in a flapping-wing system, is observed using a high-speed camera. The observation shows that the wing could maintain a fully unfolded configuration during flapping motion. A series of thrust measurements are also conducted to estimate the force generated by the flapping-wing system with foldable artificial wings. Although the artificial foldable wings give added burden to the flapping-wing system because of its weight, the thrust measurement results show that the flapping-wing system could still generate reasonable thrust.     

The three-dimensional leading-edge vortex of a 'hovering' model hawkmoth

Recent flow visualisation experiments with the hawkmoth, Manduca sexta, revealed small but clear leading-edge vortex and a pronounced three-dimensional flow. Details of this flow pattern were studied with a scaled-up, robotic insect (''the flapper'') that accurately mimicked the wing movements of a hovering hawkmoth. Smoke released from the leading edge of the flapper wing confirmed the existence of a small, strong and stable leading-edge vortex, increasing in size from wingbase to wingtip. Between 25 and 75 per cent of the wing length, its diameter increased approximately from 10 to 50 per cent of the wing chord. The leading-edge vortex had a strong axial flow veolocity, which stabilized it and reduced its diamater. The vortex separated from the wing at approximately 75 per cent of the wing length and thus fed vorticity into a large, tangled tip vortex. If the circulation of the leading-edge vortex were fully used for lift generation, it could support up to two-thirds of the hawkmoth''s weight during the downstroke. The growth of this circulation with time and spanwise position clearly identify dynamic stall as the unsteady aerodynamic mechanism responsible for high lift production by hovering hawkmoths and possibly also by many other insect species.     

Lift production in the hovering hummingbird

Aerodynamic theory and empirical observations of animals flying at similar Reynolds numbers (Re) predict that airflow over hummingbird wings will be dominated by a stable, attached leading edge vortex (LEV). In insects exhibiting similar kinematics, when the translational movement of the wing ceases (as at the end of the downstroke), the LEV is shed and lift production decreases until the energy of the LEV is re-captured in the subsequent half-cycle translation. We here show that while the hummingbird wing is strongly influenced by similar sharp-leading-edge aerodynamics, leading edge vorticity is inconsistent, varying from 0.7 to 26 per cent (mean 16%) of total lift production, is always generated within 3 mm of the dorsal surface of the wing, showing no retrograde (trailing to leading edge) flow, and does not increase from proximal to distal wing as would be expected with a conical vortex (class III LEV) described for hawkmoths. Further, the bound circulation is not shed as a vortex at the end of translation, but instead remains attached and persists after translation has ceased, augmented by the rotation (pronation, supination) of the wing that occurs between the wing-translation half-cycles. The result is a near-continuous lift production through wing turn-around, previously unknown in vertebrates, able to contribute to weight support as well as stability and control during hovering. Selection for a planform suited to creating this unique flow and nearly-uninterrupted lift production throughout the wingbeat cycle may help explain the relatively narrow hummingbird wing.     

How Could Beetle＇s Elytra Support Their Own Weight during Forward Flight？

The aerodynamic role of the elytra during a beetle＇s flapping motion is not well-elucidated, although it is well-recognized that the evolution of elytra has been a key in the success of coleopteran insects due to their protective function. An experimental study on wing kinematics reveals that for almost concurrent flapping with the hind wings, the flapping angle of the elytra is 5 times smaller than that of the hind wings. Then, we explore the aerodynamic forces on elytra in free forward flight with and without an effect of elytron-hind wing interaction by three-dimensional numerical simulation. The numerical results show that vertical force generated by the elytra without interaction is not sufficient to support even its own weight. However, the elytron-hind wing interaction improves the vertical force on the elytra up to 80%; thus, the total vertical force could fully support its own weight. The interaction slightly increases the vertical force on the hind wind by 6% as well.