Flow Visualization of Rhinoceros Beetle (Trypoxylus dichotomus) in Free Flight

Aerodynamic characteristics of the beetle,Trypoxylus dichotomus,which has a pair of elytra (forewings) and flexible hind wings,are investigated.Visualization experiments were conducted for various flight conditions of a beetle,Trypoxylus dichotomus:free,tethered,hovering,forward and climbing flights.Leading edge,trailing edge and tip vortices on both wings were observed clearly.The leading edge vortex was stable and remained on the top surface of the elytron for a wide interval during the downstroke of free forward flight.Hence,the elytron may have a considerable role in lift force generation of the beetle.In addition,we reveal a suction phenomenon between the gaps of the hind wing and the elytron in upstroke that may improve the positive lift force on the hind wing.We also found the reverse clap-fling mechanism of the T.dichotomus beetle in hovering flight.The hind wings touch together at the beginning of the upstroke.The vortex generation,shedding and interaction give a better understanding of the detailed aerodynamic mechanism of beetle flight.     

A Quasi-Steady Lifting Line Theory for Insect-Like Hovering Flight

A novel lifting line formulation is presented for the quasi-steady aerodynamic evaluation of insect-like wings in hovering flight. The approach allows accurate estimation of aerodynamic forces from geometry and kinematic information alone and provides for the first time quantitative information on the relative contribution of induced and profile drag associated with lift production for insect-like wings in hover. The main adaptation to the existing lifting line theory is the use of an equivalent angle of attack, which enables capture of the steady non-linear aerodynamics at high angles of attack. A simple methodology to include non-ideal induced effects due to wake periodicity and effective actuator disc area within the lifting line theory is included in the model. Low Reynolds number effects as well as the edge velocity correction required to account for different wing planform shapes are incorporated through appropriate modification of the wing section lift curve slope. The model has been successfully validated against measurements from revolving wing experiments and high order computational fluid dynamics simulations. Model predicted mean lift to weight ratio results have an average error of 4% compared to values from computational fluid dynamics for eight different insect cases. Application of an unmodified linear lifting line approach leads on average to a 60% overestimation in the mean lift force required for weight support, with most of the discrepancy due to use of linear aerodynamics. It is shown that on average for the eight insects considered, the induced drag contributes 22% of the total drag based on the mean cycle values and 29% of the total drag based on the mid half-stroke values.     

Soaring and manoeuvring flight of a steppe eagle Aquila nipalensis

We used an onboard inertial measurement unit, together with onboard and ground‐based video cameras, to record the movements of the body, wings and tail of a steppe eagle Aquila nipalensis during wide‐ranging flight. The eagle's flight consisted of a more or less continuous sequence of banked turns, interrupted by occasional wing tucks and roll‐over manoeuvres, and ultimately terminated by a wing‐over manoeuvre leading in to a diving landing approach. The flight configuration of the bird, and its pattern of movement during angular perturbations, together suggest that the eagle is inherently stable in pitch and yaw, and perhaps also in roll. The control inputs used to generate roll moments during banked turns were too subtle to be detected. Control of yaw and pitch during banked turns involved a consistent pattern of tail movement, wherein the tail was spread and depressed immediately before the turn, and then overbanked with respect to the bird during the latter part of the turn. Differential adjustment of wing posture is probably also involved in the control of banked turns, but it was only consistently apparent during more extreme roll manoeuvres. For example, roll‐over and wing‐over manoeuvres were both accomplished by differential changes in the angle of incidence and spread of the wings. In general, however, the bird appeared to maintain positive loading on its wings at all times, except during extreme flight manoeuvres.     

High lift function of the pteroid bone and forewing of pterosaurs

The pteroid bone is a rod-like element found only in pterosaurs, the flying reptiles of the Mesozoic. It articulated at the wrist, and supported a membranous forewing in front of the inner part of the wing spar. The function of this bone, particularly its orientation, has been much debated. It is widely believed that it pointed towards the body, and that the forewing was relatively narrow. An alternative hypothesis states that it was directed forwards during flight, resulting in a much broader forewing that acted as a leading edge flap. We tested scale models in a wind tunnel to determine the aerodynamic consequences of these conflicting hypotheses, and found that performance is greatly improved if the pteroid is directed forwards: the lift: drag ratios are superior and the maximum lift is exceptionally high in comparison with conventional aerofoils. This high lift capability may have enabled even the largest pterosaurs to take off and land without difficulty.     

Comparing aerodynamic efficiency in birds and bats suggests better flight performance in birds

Flight is one of the energetically most costly activities in the animal kingdom, suggesting that natural selection should work to optimize flight performance. The similar size and flight speed of birds and bats may therefore suggest convergent aerodynamic performance; alternatively, flight performance could be restricted by phylogenetic constraints. We test which of these scenarios fit to two measures of aerodynamic flight efficiency in two passerine bird species and two New World leaf-nosed bat species. Using time-resolved particle image velocimetry measurements of the wake of the animals flying in a wind tunnel, we derived the span efficiency, a metric for the efficiency of generating lift, and the lift-to-drag ratio, a metric for mechanical energetic flight efficiency. We show that the birds significantly outperform the bats in both metrics, which we ascribe to variation in aerodynamic function of body and wing upstroke: Bird bodies generated relatively more lift than bat bodies, resulting in a more uniform spanwise lift distribution and higher span efficiency. A likely explanation would be that the bat ears and nose leaf, associated with echolocation, disturb the flow over the body. During the upstroke, the birds retract their wings to make them aerodynamically inactive, while the membranous bat wings generate thrust and negative lift. Despite the differences in performance, the wake morphology of both birds and bats resemble the optimal wake for their respective lift-to-drag ratio regimes. This suggests that evolution has optimized performance relative to the respective conditions of birds and bats, but that maximum performance is possibly limited by phylogenetic constraints. Although ecological differences between birds and bats are subjected to many conspiring variables, the different aerodynamic flight efficiency for the bird and bat species studied here may help explain why birds typically fly faster, migrate more frequently and migrate longer distances than bats.     

Tail effects on yaw stability in birds

Bird tails, which are an aerodynamic surface in the horizontal plane, are treated with regard to their effects on yaw stability. Reference is made to wings of very small aspect ratio similar to the values of bird tails in order to identify features which are significant for the aerodynamic yawing moment characteristics due to sideslip. It is shown that there are yawing moments of considerable magnitude for this aspect ratio region. Furthermore, the lift coefficient, which also exerts an influence, is included in the treatment of yaw stability. To show more concretely the addressed effects for birds, the yawing moment characteristics of the wing-tail combination of a pigeon, which is considered as a representative example, are treated in detail. For this purpose, a sophisticated aerodynamic method capable to deal with the mutual flow interactions between the tail and the wing is used to compute results of high precision. The yawing moment characteristics of the pigeon wing-tail combination with respect to the sideslip angle and the lift coefficient are determined, with emphasis placed on the contribution of the tail. It is shown that there is a significant contribution of the tail to yaw stability. The findings of this paper on the contribution of the tail to the yawing moment characteristics are supported by an evaluation of existing experimental data. Furthermore, the physical mechanisms are considered which are the reasons for the stabilizing role of the tail. These effects concern the contribution of the drag acting at the tail to the yawing moment. In addition, it is shown that extended legs and feet, when exposed to the airflow, can contribute to yaw stability.     

Aerodynamic yawing moment characteristics of bird wings

The aerodynamic yawing moments due to sideslip are considered for wings of birds. Reference is made to the experience with aircraft wings in order to identify features which are significant for the yawing moment characteristics. Thus, it can be shown that wing sweep, aspect ratio and lift coefficient have a great impact. Focus of the paper is on wing sweep which can considerably increase the yawing moment due to sideslip when compared with unswept wings. There are many birds the wings of which employ sweep. To show the effect of sweep for birds, the aerodynamic characteristics of a gull wing which is considered as a representative example are treated in detail. For this purpose, a sophisticated aerodynamic method is used to compute results of high precision. The yawing moments of the gull wing with respect to the sideslip angle and the lift coefficient are determined. They show a significant level of yaw stability which strongly increases with the lift coefficient. It is particularly high in the lift coefficient region of best gliding flight conditions. In order to make the effect of sweep more perspicuous, a modification of the gull wing employing no sweep is considered for comparison. It turns out that the unswept wing yields yawing moments which are substantially smaller than those of the original gull wing with sweep. Another feature significant for the yawing moment characteristics concerns the fact that sweep is at the outer part of bird wings. By considering the underlying physical mechanism, it is shown that this feature is most important for the efficiency of wing sweep. To sum up, wing sweep provides a primary contribution to the yawing moments. It may be concluded that this is an essential reason why there is sweep in bird wings.     

Characteristics of the alula in relation to wing and body size in the Laridae and Sternidae

The alula is a small structure present on the leading edge of bird wings and is known to enhance lift by creating a small vortex at its tip. Alula size vary among birds, but how this variation is associated with the function of the alula remains unclear. In this study, we investigated the relationship between the size and shape of the alula and the features of the wing in the Laridae and Sternidae. Laridae birds have generally longer wings and greater loadings than Sternidae birds. The two families differed in the relationships between body size or wing length and the size or shape of the alula. In the Laridae, the aspect ratio of the alula was smaller in the species that have relatively longer wings, but the pattern was opposite in the Sternidae. The aspect ratio of the alula was greater in the species that are relatively heavier in the Sternidae but not in the Laridae. Combined, these results suggest that the species with high loading potential and long wings exhibit long alula. We hypothesize that heavier species may benefit from having longer alula if they perform flights with higher attack angles than lighter species, as longer alula would better suppress flow separation at higher attack angles. Our results suggest that the size and shape of the alula can be explained in one allometric landscape defined by wing length and loading in these two closely related families of birds with similar wing shapes.     

Particle-Image Velocimetry and Force Measurements of Leading-Edge Serrations on Owl-Based Wing Models

High-resolution Particle-Image Velocimetry （PIV） and time-resolved force measurements were performed to analyze the impact of the comb-like structure on the leading edge of barn owl wings on the flow field and overall aerodynamic performance. The Reynolds number was varied in the range of 40,000 to 120,000 and the range of angle of attack was 0° to 6° for the PIV and -15° to ＋20° for the force measurements to cover the full flight envelope of the owl. As a reference, a wind-tunnel model which possessed a geometry based on the shape of a typical barn owl wing without any owl-specific adaptations was built, and measurements were performed in the aforementioned Reynolds number and angle of attack： range. This clean wing model shows a separation bubble in the distal part of the wing at higher angles of attack. Two types of comb-like structures, i.e., artificial serrations, were manufactured to model the owl＇s leading edge with respect to its length, thickness, and material properties. The artificial structures were able to reduce the size of the separation region and additionally cause a more uniform size of the vortical structures shed by the separation bubble within the Reynolds number range investigated, resulting in stable gliding flight independent of the flight velocity. However, due to increased drag coefficients in conjunction with similar lift coefficients, the overall aerodynamic performance, i.e., lift-to-drag ratio is reduced for the serrated models. Nevertheless, especially at lower Reynolds numbers the stabilizing effect of the uniform vortex size outperforms the lower aerodynamic performance.     

Investigation of Unsteady Aerodynamic Characteristics of a Seagull Wing in Level Flight

Unsteady aerodynamic characteristics of a seagull wing in level flight are investigated using a boundary element method.Anew no-penetration boundary condition is imposed on the surface of the wing by considering its deformation.The geometry andkinematics of the seagull wing are reproduced using the functions and data in the previously published literature.The proposedmethod is validated by comparing the computed results with the published data in the literature.The unsteady aerodynamicscharacteristics of the seagull wing are investigated by changing flapping frequency and advance ratio.It is found that the peakvalues of aerodynamic coefficients increase with the flapping frequency.The thrust and drag generations are complicatedfunctions of frequency and wing stroke motions.The lift is inversely proportional to the advance ratio.The effects of severalflapping modes on the lift and induced drag(or thrust)generation are also investigated.Among three single modes(flapping,folding and lead & lag),flapping generates the largest lift and can produce thrust alone.For three combined modes,both flapping/foldingand flapping/lead & lag can produce lift and thrust larger than the flapping-alone mode can.Folding is shown toincrease thrust when combined with flapping,whereas lead & lag has an effect of increasing the lift when also combined withflapping.When three modes are combined together,the bird can obtain the largest lift among the investigated modes.Eventhough the proposed method is limited to the inviscid flow assumption,it is believed that this method can be used to the designof flapping micro aerial vehicle.     

Effect of slotted wing tips on yawing moment characteristics

The aerodynamic yawing moment characteristics of bird wings with slotted tips are dealt with. Emphasis is placed on the effect of sweep which the separated feathers constituting the wing tips show and which can reach significant values. Reference is made to basic aerodynamic characteristics of wings with sweep which yields a stabilizing yawing moment significantly larger than that of unswept wings. Then, the yawing moment characteristics are determined for a wing, the features of which are considered as representative of bird wings with sweep in their slotted tips. A sophisticated aerodynamic procedure is used for obtaining results of high precision. It is shown that the sweep in the slotted wing tips yields a stabilizing yawing moment of significant magnitude, considerably increasing with the lift coefficient. To make the significance of wing tip sweep for the ability to generate yawing moments more perspicuous, a wing modification the slotted tips of which are unswept is considered for comparison. It turns out that this wing shows yawing moments which are substantially smaller. A physical insight into the effect of slotted wing tip sweep on the aerodynamic yawing moment characteristics is provided by showing the underlying mechanism. From the results presented in this paper it follows that the sweep in slotted wing tips provides a substantial contribution to the aerodynamic yawing moment and, thus, to yaw stability. It may be concluded that this is an essential reason why there is sweep in the slotted tips of bird wings.     

Exploring bird aerodynamics using radio-controlled models

A series of radio-controlled glider models was constructed by duplicating the aerodynamic shape of soaring birds (raven, turkey vulture, seagull and pelican). Controlled tests were conducted to determine the level of longitudinal and lateral-directional static stability, and to identify the characteristics that allowed flight without a vertical tail. The use of tail-tilt for controlling small bank-angle changes, as observed in soaring birds, was verified. Subsequent tests, using wing-tip ailerons, inferred that birds use a three-dimensional flow pattern around the wing tip (wing tip vortices) to control adverse yaw and to create a small amount of forward thrust in gliding flight.     

The avian tail reduces body parasite drag by controlling flow separation and vortex shedding.

The aerodynamic effect of the furled avian tail on the parasite drag of a bird's body was investigated on mounted, frozen European starling Sturnus vulgaris in a wind tunnel at flight speeds between 6 and 14 m s(-1). Removal of tail rectrices and dorsal and ventral covert feathers at the base of the tail increased the total parasite drag of the body and tail by between 25 and 55%. Flow visualization and measurements of dynamic pressure in the tail boundary layer showed that in the intact bird a separation bubble forms on the ventral side of the body, and reattaches to the ventral side of the tail. This bubble is a consequence of the morphology of the body, with a rapid contraction posterior to the pelvis and hind legs. The tail and the covert feathers at its base act as a combined splitter plate and wedge to control vortex shedding and body wake development, and thereby are important to minimize drag. This hitherto unsuspected mechanism is central to understanding the morphology of the avian body, and may have had a significant influence on the evolution of avian tail morphology by pre-adapting the tail for radiation and specialization as an aerodynamic lifting structure and as an organ of communication in sexual selection.     

On the Estimation of Time Dependent Lift of a European Starling (Sturnus vulgaris) during Flapping Flight

We study the role of unsteady lift in the context of flapping wing bird flight. Both aerodynamicists and biologists have attempted to address this subject, yet it seems that the contribution of unsteady lift still holds many open questions. The current study deals with the estimation of unsteady aerodynamic forces on a freely flying bird through analysis of wingbeat kinematics and near wake flow measurements using time resolved particle image velocimetry. The aerodynamic forces are obtained through two approaches, the unsteady thin airfoil theory and using the momentum equation for viscous flows. The unsteady lift is comprised of circulatory and non-circulatory components. Both approaches are presented over the duration of wingbeat cycles. Using long-time sampling data, several wingbeat cycles have been analyzed in order to cover both the downstroke and upstroke phases. It appears that the unsteady lift varies over the wingbeat cycle emphasizing its contribution to the total lift and its role in power estimations. It is suggested that the circulatory lift component cannot assumed to be negligible and should be considered when estimating lift or power of birds in flapping motion.     

Numerical Simulation of the Effect of Bionic Serrated Structures on the Aerodynamic Noise of a Circular Cylinder

Flow control can effectively reduce the aerodynamic noise radiated from a circular cylinder.As one of the flow control methods,a bionic method,inspired by the serrations at the leading edge of owls' wing,was proposed in this paper.The effects of bionic serrated structures arranged on the upper and lower sides of a cylinder on the aerodynamic and aeroacoustic performance of the cylinder were numerically investigated.At a free stream speed of 24.5 m·s-1,corresponding to Reynolds number of 1.58 × 104,the simulation results indicate that the bionic serrated structures can decrease the frequency of the vortex shedding and control the fluctuating aerodynamic force acting on the cylinder,thus reduce the aerodynamic noise.A qualitative-view of the vorticity in the wake of the cylinder suggest that the serrated structures reduce aerodynamic sound by suppressing the unsteady motion of vortices.     

The fish tail motion forms an attached leading edge vortex

The tail (caudal fin) is one of the most prominent characteristics of fishes, and the analysis of the flow pattern it creates is fundamental to understanding how its motion generates locomotor forces. A mechanism that is known to greatly enhance locomotor forces in insect and bird flight is the leading edge vortex (LEV) reattachment, i.e. a vortex (separation bubble) that stays attached at the leading edge of a wing. However, this mechanism has not been reported in fish-like swimming probably owing to the overemphasis on the trailing wake, and the fact that the flow does not separate along the body of undulating swimmers. We provide, to our knowledge, the first evidence of the vortex reattachment at the leading edge of the fish tail using three-dimensional high-resolution numerical simulations of self-propelled virtual swimmers with different tail shapes. We show that at Strouhal numbers (a measure of lateral velocity to the axial velocity) at which most fish swim in nature (approx. 0.25) an attached LEV is formed, whereas at a higher Strouhal number of approximately 0.6 the LEV does not reattach. We show that the evolution of the LEV drastically alters the pressure distribution on the tail and the force it generates. We also show that the tail''s delta shape is not necessary for the LEV reattachment and fish-like kinematics is capable of stabilising the LEV. Our results suggest the need for a paradigm shift in fish-like swimming research to turn the focus from the trailing edge to the leading edge of the tail.     

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.     

Metabolic ‘engines’ of flight drive genome size reduction in birds

The tendency for flying organisms to possess small genomes has been interpreted as evidence of natural selection acting on the physical size of the genome. Nonetheless, the flight–genome link and its mechanistic basis have yet to be well established by comparative studies within a volant clade. Is there a particular functional aspect of flight such as brisk metabolism, lift production or maneuverability that impinges on the physical genome? We measured genome sizes, wing dimensions and heart, flight muscle and body masses from a phylogenetically diverse set of bird species. In phylogenetically controlled analyses, we found that genome size was negatively correlated with relative flight muscle size and heart index (i.e. ratio of heart to body mass), but positively correlated with body mass and wing loading. The proportional masses of the flight muscles and heart were the most important parameters explaining variation in genome size in multivariate models. Hence, the metabolic intensity of powered flight appears to have driven genome size reduction in birds.     

Energy expenditures for flight,aerodynamic quality,and colonization of forest habitats by birds

The power that the birds can use for flight (available power) and the power required for flight according to physical laws (requisite power) grow with an increase in body mass, the exponents of the corresponding functions being different. Small birds can follow different strategies, either improving the aerodynamic quality of the body (thereby saving the excess available power) or sacrifice aerodynamic quality in favor of morphological adaptation to factors other than the demands of flight proper, which provides the possibility of utilizing a wider range of ecological niches. A hypothesis is proposed that the high metabolic rate of passerine birds, compared to representatives of other bird orders, is an adaptation to maneuverable (i.e., relatively low-speed) flight necessary for successful colonization of forest habitats. The speed that birds of such size can develop according to the scaling theory is too high for nesting and foraging in tree crowns, and its reduction is possible in two ways: by increasing air drag or by changing the style of flight (by analogy with airplane vs. helicopter). The first way is feasible, but a high air drag due to morphological modifications (e.g., in the size of the tail or characteristics of the wing) interferes with the possibility of long-distance migration flight, as energy expenditures for it will exceed the energy potential of the bird. This is why migratory nonpasserine birds, which have used this strategy, are practically absent in forests of the temperate zone. Therefore, more promising is the second way involving transition to a new flight style and, in a certain sense, to a new morphophysiological organization. Passerines have achieved this by changing their flight style so that the wing actively generates forces (lift and thrust) only in downstroke. Such a flight requires more energy, and, to provide it in sufficient amounts, passerine birds have increased their basal metabolic rate (BMR). Thus, both their flight energy expenditures and BMR are higher than in nonpasserines. Remarkably, among approximately 8660 extant bird species known today, more than half (about 5100 species) belong to the order Passeriformes. Such a ratio, unknown in any other vertebrate class, is evidence that passerines have gained a considerable biological advantage over all other birds due to their increased BMR.     

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.