Periodic Tail Motion Linked to Wing Motion Affects the Longitudinal Stability of Ornithopter Flight

During slow level flight of a pigeon,a caudal muscle involved in tail movement,the levator caudae pars vertebralis,is activated at a particular phase with the pectoralis wing muscle.Inspired by mechanisms for the control of stability in flying animals,especially the role of the tail in avian flight,we investigated how periodic tail motion linked to motion of the wings affects the longitudinal stability of omithopter flight.This was achieved by using an integrative ornithopter flight simulator that included aeroelastic behaviour of the flexible wings and tail.Trim flight trajectories of the simulated ornithopter model were calculated by time integration of the nonlinear equations of a flexible multi-body dynamics coupled with a semi-empirical flapping-wing and tail aerodynamic models.The unique trim flight characteristics of ornithopter,Limit-Cycle Oscillation,were found under the sets of wingbeat frequency and tail elevation angle,and the appropriate phase angle of tail motion was determined by parameter studies minimizing the amplitude of the oscillations.The numerical simulation results show that tail actuation synchronized with wing motion suppresses the oscillation of body pitch angle over a wide range of wingbeat frequencies.     

Distributed power and control actuation in the thoracic mechanics of a robotic insect

Recent advances in the understanding of biological flight have inspired roboticists to create flapping-wing vehicles on the scale of insects and small birds. While our understanding of the wing kinematics, flight musculature and neuromotor control systems of insects has expanded, in practice it has proven quite difficult to construct an at-scale mechanical device capable of similar flight performance. One of the key challenges is the development of an effective and efficient transmission mechanism to control wing motions. Here we present multiple insect-scale robotic thorax designs capable of producing asymmetric wing kinematics similar to those observed in nature and utilized by dipteran insects to maneuver. Inspired by the thoracic mechanics of dipteran insects, which entail a morphological separation of power and control muscles, these designs show that such distributed actuation can also modulate wing motion in a robotic design.     

Flying at no mechanical energy cost: disclosing the secret of wandering albatrosses

ALBATROSSES DO SOMETHING THAT NO OTHER BIRDS ARE ABLE TO DO: fly thousands of kilometres at no mechanical cost. This is possible because they use dynamic soaring, a flight mode that enables them to gain the energy required for flying from wind. Until now, the physical mechanisms of the energy gain in terms of the energy transfer from the wind to the bird were mostly unknown. Here we show that the energy gain is achieved by a dynamic flight manoeuvre consisting of a continually repeated up-down curve with optimal adjustment to the wind. We determined the energy obtained from the wind by analysing the measured trajectories of free flying birds using a new GPS-signal tracking method yielding a high precision. Our results reveal an evolutionary adaptation to an extreme environment, and may support recent biologically inspired research on robotic aircraft that might utilize albatrosses' flight technique for engineless propulsion.     

Rhythm Patterns Interaction - Synchronization Behavior for Human-Robot Joint Action

Interactive behavior among humans is governed by the dynamics of movement synchronization in a variety of repetitive tasks. This requires the interaction partners to perform for example rhythmic limb swinging or even goal-directed arm movements. Inspired by that essential feature of human interaction, we present a novel concept and design methodology to synthesize goal-directed synchronization behavior for robotic agents in repetitive joint action tasks. The agents’ tasks are described by closed movement trajectories and interpreted as limit cycles, for which instantaneous phase variables are derived based on oscillator theory. Events segmenting the trajectories into multiple primitives are introduced as anchoring points for enhanced synchronization modes. Utilizing both continuous phases and discrete events in a unifying view, we design a continuous dynamical process synchronizing the derived modes. Inverse to the derivation of phases, we also address the generation of goal-directed movements from the behavioral dynamics. The developed concept is implemented to an anthropomorphic robot. For evaluation of the concept an experiment is designed and conducted in which the robot performs a prototypical pick-and-place task jointly with human partners. The effectiveness of the designed behavior is successfully evidenced by objective measures of phase and event synchronization. Feedback gathered from the participants of our exploratory study suggests a subjectively pleasant sense of interaction created by the interactive behavior. The results highlight potential applications of the synchronization concept both in motor coordination among robotic agents and in enhanced social interaction between humanoid agents and humans.     

Ornithopter Flight Simulation Based on Flexible Multi-Body Dynamics

This paper introduces a flight simulation of an ornithopter (flapping-wing air vehicle) based on the flexible multi-body dynamics, a refined flapping-wing aerodynamic model and the fluid-structure interaction approach. A simulated ornithopter was modeled using the multi-body dynamics software, MSC.ADAMS, where the flexible parts can be included by importing a finite element model built in the finite element analysis software, ANSYS. To model the complex aerodynamics of flapping-wing, an improved version of modified strip theory was chosen. The proposed integrative simulation framework of ornithopter was validated by the wind tunnel test data reported in the literature. A magpie-sized model oruithopter was numerically designed and simulated to have the longitudinal trim flight condition. We observed a limit-cycle-oscillation of flight state variables, such as pitch attitude, altitude, flight speed, during the trimmed flight of the model ornithopter. Under the trimmed condition of free flight of the model omithopter, we fixed all the degrees of freedom at the center of gravity to measure the constraint forces and moment. The concept of the "zero moment point" is introduced to explain the physics of ornithopter trimmed longitudinal flight.     

Vortexlet models of flapping flexible wings show tuning for force production and control

Insect wings are compliant structures that experience deformations during flight. Such deformations have recently been shown to substantially affect induced flows, with appreciable consequences to flight forces. However, there are open questions related to the aerodynamic mechanisms underlying the performance benefits of wing deformation, as well as the extent to which such deformations are determined by the boundary conditions governing wing actuation together with mechanical properties of the wing itself. Here we explore aerodynamic performance parameters of compliant wings under periodic oscillations, subject to changes in phase between wing elevation and pitch, and magnitude and spatial pattern of wing flexural stiffness. We use a combination of computational structural mechanics models and a 2D computational fluid dynamics approach to ask how aerodynamic force production and control potential are affected by pitch/elevation phase and variations in wing flexural stiffness. Our results show that lift and thrust forces are highly sensitive to flexural stiffness distributions, with performance optima that lie in different phase regions. These results suggest a control strategy for both flying animals and engineering applications of micro-air vehicles.     

Using Wind Tunnels to Predict Bird Mortality in Wind Farms: The Case of Griffon Vultures

Background

Wind farms have shown a spectacular growth during the last 15 years. Avian mortality through collision with moving rotor blades is well-known as one of the main adverse impacts of wind farms. In Spain, the griffon vulture incurs the highest mortality rates in wind farms.

Methodology/Principal Findings

As far as we know, this study is the first attempt to predict flight trajectories of birds in order to foresee potentially dangerous areas for wind farm development. We analyse topography and wind flows in relation to flight paths of griffon vultures, using a scaled model of the wind farm area in an aerodynamic wind tunnel, and test the difference between the observed flight paths of griffon vultures and the predominant wind flows. Different wind currents for each wind direction in the aerodynamic model were observed. Simulations of wind flows in a wind tunnel were compared with observed flight paths of griffon vultures. No statistical differences were detected between the observed flight trajectories of griffon vultures and the wind passages observed in our wind tunnel model. A significant correlation was found between dead vultures predicted proportion of vultures crossing those cells according to the aerodynamic model.

Conclusions

Griffon vulture flight routes matched the predominant wind flows in the area (i.e. they followed the routes where less flight effort was needed). We suggest using these kinds of simulations to predict flight paths over complex terrains can inform the location of wind turbines and thereby reduce soaring bird mortality.     

AUTOROTATION, SELF-STABILITY, AND STRUCTURE OF SINGLE-WINGED FRUITS AND SEEDS (SAMARAS) WITH COMPARATIVE REMARKS ON ANIMAL FLIGHT

• 1 A samara is a winged fruit or seed that autorotates when falling, thereby reducing the sinking speed of the diaspore and increasing the distance it may be transported by winds. Samaras have evolved independently in a large number of plants.
• 2 Aerodynamical, mechanical, and structural properties crucial for the inherent self-stability are analysed, and formulae for calculation of performance data are given.
• 3 The momentum theorem is applied to samaras to calculate induced air velocities. As a basis for blade element analysis, and for directional stability analysis, various velocity components are put together into resultant relative air velocities normal to the blade's span axis for a samara in vertical autorotation and also in autorotation with side-slip.
• 4 When falling, a samara is free to move in any sense, but in autorotation it possesses static and dynamic stability. Mainly qualitative aspects on static stability are pre sented. Simple experiments on flat plates at Reynolds numbers about 2000 as in samaras, showed that pitch stability prevails when the C. M. (centre of mass) is located 27–35 % of the chord behind the leading edge. The aerodynamic c.p. (centre of pressure) moves forward upon a decrease of the angle of attack, backward upon an increase. In samara blades the c.m. lies ca. one-third chord behind the leading edge, and hence the aerodynamic and centrifugal forces interact so as to give pitch stability, involving stability of the angles of attack and gliding angles.
• 5 Photographs show that the centre of rotation of the samara approximately coincides with its c.m.
• 6 The coning angle (blade angle to tip path plane) taken up by the samara is determined by opposing moments set up by the centrifugal and aerodynamic forces. It is essentially the centrifugal moment (being a tangent function of the coning angle, which is small) that changes upon a change of coning angle, until the centrifugal and aerodynamic moments cancel out at the equilibrium coning angle.
• 7 Directional stability is maintained by keeping the tip path plane horizontal whereby a vertical descent path relative to the ambient air is maintained. Tilting of the tip path plane results in side-slip. Side-slip leads to an increased relative air speed at the blade when advancing, a reduced speed when retreating. The correspondingly fluctuating aerodynamic force and the gyroscopic action of the samara lead to restoring moments that bring the tip path plane back to the horizontal.
• 8 Entrance into autorotation is due to interaction between aerodynamic forces, the force of gravity, and inertial forces (when the blade accelerates towards a trailing position behind the c.m. of the samara).
• 9 The mass distribution must be such that the c.m. lies 0–30 % of the span from one end. In Acer and Plcea samaras the C.M. lies 10–20% from one end, thereby making the disk area swept by the blade large and the sinking speed low.
• 10 The blade plan-form is discussed in relation to aerodynamics. The width is largest far out on the blade where the relative air velocities are large. The large width of the blade contributes to a high Re number and thus probably to a better L/D (lift/drag) ratio and a slower descent.
• 11 The concentration of vascular bundles at the leading edge of the blade and the tapering of the blade thickness towards the trailing edge are essential for a proper chord wise mass distribution.
• 12 Data are given for samaras of Acer and Plcea, and calculations of performance are made by means of the formulae given in the paper. Some figures for an Acer samara are: sinking speed 0.9 m/sec, tip path inclination 15°, average total force coefficient 1.7 (which is discussed), and a L/D ratio of the blade approximately 3.
• 13 The performances of samaras are compared with those of insects, birds, bats, a flat plate, and a parachute. They show the samara to be a relatively very efficient structure in braking the sinking speed of the diaspore.
• 14 In samaras the mass, aerodynamic, and torsion axes coincide, whereas in insect wings the torsicn axis often lies ahead of the other two. Location of the torsion axis in front of the aerodynamic axis in insects tends towards passive wing twisting and passive adjustment of the angles of attack relative to the incident air stream, the direction of which varies along the wing because of wing flapping.
• 15 Location of the mass axis behind the torsion axis may lead to unfavourable

     

Dynamic Modeling, Testing, and Stability Analysis of an Ornithoptic Blimp

In order to study omithopter flight and to improve a dynamic model of flapping propulsion,a series of tests are conducted on a flapping-wing blimp.The blimp is designed and constructed from mylar plastic and balsa wood as a test platform for aerodynamics and flight dynamics.The blimp,2.3 meters long and 420 gram mass,is propelled by its flapping wings.Due to buoyancy the wings have no lift requirement so that the distinction between lift and propulsion can be analyzed in a flight platform at low flight speeds.The blimp is tested using a Vicon motion tracking system and various initial conditions are tested including accelerating flight from standstill,decelerating from an initial speed higher than its steady state,and from its steady-state speed but disturbed in pitch angle.Test results are used to estimate parameters in a coupled quasi-steady aerodynamics/Newtonian flight dynamics model.This model is then analyzed using Floquet theory to determine local dynamic modes and stability.It is concluded that the dynamic model adequately describes the vehicle's nonlinear behavior near the steady-state velocity and that the vehicle's linearized modes are akin to those of a fixed-wing aircraft.     

Directionally compliant legs influence the intrinsic pitch behaviour of a trotting quadruped

Limb design is well conserved among quadrupeds, notably, the knees point forward (i.e. cranial inclination of femora) and the elbows point back (i.e. caudal inclination of humeri). This study was undertaken to examine the effects of joint orientation on individual leg forces and centre of mass dynamics. Steady-speed trotting was simulated in two quadrupedal models. Model I had the knee and elbow orientation of a quadruped and model II had a reversed leg configuration in which knees point back and elbows point forward. The model's legs showed directional compliance determined by the orientation of the knee/elbow. In both models, forward pointing knees/elbows produced a propulsive force bias, while rearward pointing knees/elbows produced a braking force bias. Hence, model I showed the same pattern of hind-leg propulsion and fore-leg braking observed in trotting animals. Simulations revealed minimal pitch oscillations during steady-speed trotting of model I, but substantially greater and more irregular pitch oscillations of model II. The reduced pitch oscillation of model I was a result of fore-leg and hind-leg forces that reduced pitching moments during early and late stance, respectively. This passive mechanism for reducing pitch oscillations was an emergent property of directionally compliant legs with the fore-hind configuration of model I. Such intrinsic stability resulting from mechanical design can simplify control tasks and lead to more robust running machines.     

Functional modes and residue flexibility control the anisotropic response of guanylate kinase to mechanical stress

The coupling between the mechanical properties of enzymes and their biological activity is a well-established feature that has been the object of numerous experimental and theoretical works. In particular, recent experiments show that enzymatic function can be modulated anisotropically by mechanical stress. We study such phenomena using a method for investigating local flexibility on the residue scale that combines a reduced protein representation with Brownian dynamics simulations. We performed calculations on the enzyme guanylate kinase to study its mechanical response when submitted to anisotropic deformations. The resulting modifications of the protein's rigidity profile can be related to the changes in substrate binding affinity observed experimentally. Further analysis of the principal components of motion of the trajectories shows how the application of a mechanical constraint on the protein can disrupt its dynamics, thus leading to a decrease of the enzyme's catalytic rate. Eventually, a systematic probe of the protein surface led to the prediction of potential hotspots where the application of an external constraint would produce a large functional response both from the mechanical and dynamical points of view. Such enzyme-engineering approaches open the possibility to tune catalytic function by varying selected external forces.     

Responses of photosystem II of white elm to UV-B radiation monitored by OJIP fluorescence transients

Photosystem II (PSII) activities in both samara and leaf of white elm (Ulmus pumila L.) were significantly inhibited by enhanced UV-B radiation (UVBR). UVBR disturbed both the donor and acceptor sides of PSII. The plastoquinone (PQ) pool size on the acceptor side, the trapped excited energy for complete reduction of Q_A, and the proportion of closed PSII reaction centers (RCs) increased, with PSII RCs being transformed into dissipative sinks for excitation energy under UVBR. However, samara and leaf responded to UVBR in different ways. A decrease in the F ₀ for leaf induced by UV-B radiation suggests the formation of fluorescence-quenching centers. An increase in the V_I for leaf under UVBR might mean the accumulation of reduced Q_A and PQ. F ₀ and V_I for samara showed opposite change pattern. Leaf has the mechanism of regulation of the amount of light reaching the RC through decreasing the number of light-harvesting chlorophyll molecules under UVBR while samara may be unable to regulate the light-harvesting capacity. PSII in samara was more susceptible to UVBR than that in leaf, with PI_ABS for samara decreasing more rapidly by a factor of 6.4 than that for leaf. Samara can recover more easily from UVBR-induced damage to PSII than the leaf.     

Flight performance of the moth, Manduca sexta, at variable gravity

The tethered and free flight of Manduca sexta were studied during period 1,2, and 0 times normal gravity (g) produced in an aeroplane by flying through parabolic trajectories. Moths in tethered flight did not change their aerodynamic output in response to increases or decreases in gravity. Some moths in free flight at 0 g maintained a position in the box by flying against a surface, or into the angle between two surfaces. In the absence of gravity as an orienting stimulus, the positive dorsophotic response to light was dominant. As the period of 0 g continued, moths were increasingly likely to periodically reduce the amplitude of their wingbeat and/or stop flying, for the equivalent of a few wingbeats. Only at 0 g, moths very occasionally spread their wings and floated freely for a few seconds. At 0 g moths retained control of rolling and yawing movements but stability in pitch was greatly reduced or absent.     

Seed size and dispersal potential of Acer rubrum (Aceraceae) samaras produced by populations in early and late successional environments

In order to determine if red maple dispersal potential or seed size change during secondary succession, samaras were collected from five populations located in early successional environments and five populations located in late successional environments. Wing loading ratios (samara mass—mg/samara area—cm²), which are inversely proportional to dispersal ability, were computed for all samaras, and seeds were excised from each samara and weighed. Samaras from the early successional red maples showed slightly but significantly lower wing loading ratios than those from the late successional environments. This result corresponds with the conclusions reached by several theoretical investigations of seed dispersal evolution that predict that recently founded populations will show greater dispersal abilities than more established populations. The earlier successional populations had slightly heavier seeds than the later successional populations, which suggests that the changes in community composition and dynamics that occur during this successional sequence do not select for heavier seeds in older red maple populations. Coefficients of variation for wing loading and seed size showed no consistent trends with successional stage, which indicates that variation in these characters does not decrease as succession proceeds.     

Wireless Stimulation of Antennal Muscles in Freely Flying Hawkmoths Leads to Flight Path Changes

Insect antennae are sensory organs involved in a variety of behaviors, sensing many different stimulus modalities. As mechanosensors, they are crucial for flight control in the hawkmoth Manduca sexta. One of their roles is to mediate compensatory reflexes of the abdomen in response to rotations of the body in the pitch axis. Abdominal motions, in turn, are a component of the steering mechanism for flying insects. Using a radio controlled, programmable, miniature stimulator, we show that ultra-low-current electrical stimulation of antennal muscles in freely-flying hawkmoths leads to repeatable, transient changes in the animals'' pitch angle, as well as less predictable changes in flight speed and flight altitude. We postulate that by deflecting the antennae we indirectly stimulate mechanoreceptors at the base, which drive compensatory reflexes leading to changes in pitch attitude.     

STUDY OF THE VERTICAL AUTOROTATION OF A SINGLEWINGED SAMARA

1. Autorotation of a single-winged samara is a highly nonlinear phenomenon that represents a delicate equilibrium between gravity, inertia and aerodynamic effects. Therefore, in order to analyse this phenomenon, an accurate detailed model is necessary. Such a model has not been presented in the past. Recently the authors derived a detailed model which is briefly described in the paper. 2. The aerodynamic contributions present the most complicated part of the phenomenon. These contributions are treated using the blade-element/momentuin method, with certain improvements and additions. These improvements are necessary due to inherent differences between samara wings and other rotary wing systems (aircraft propellers, helicopter rotors, etc.). 3. The cross-sectional aerodynamics of the samara is characterized by relatively small Reynolds numbers, high angles of attack and rough surfaces. While these characteristics are different from other rotary wings, they are typical of the wing cross-sections of insects and birds. Therefore the lift and drag coefficients, which are necessary for the analysis, are obtained using available data for insect and bird wings. 4. The results of the theoretical model are compared with experimental results of tlvo kinds. The first kind includes results for a samara of an Acer platanoides that were reported in the literature. In addition, a special experimental model of a samiira was built and tested. This model offers a simple way of varying the configuration in order to study (experimentally) the effect of different geometric parameters on the autorotation. 5. In the light of the uncertainty in the aerodynamic coefficients, it can be concluded that there is quite a good agreement between the theoretical and experimental results. Thus, after LTalidation, the theoretical model is used for a parametric study to find the influence of different parameters on the autorotation. The important results of this study are outlined below. 6. The spanwise flolv component and the tangential component of the induced velocity have a very small influence and thus can be neglected. 7. It is important to include in the analysis the effects of the axial induced velocity, the tip effect, and the drag of the root region. 8. Since chordwise variations of the centre of pressure location, as a function of the angle of attack, were seen in the past (based on over simplified analyses) as the mechanism which is responsible for the samara stability, this effect is also added to the model. While the influence of this effect on the pitch angle is large and small on the sinking rate, it results in an increase in the deviation between the theoretical and experimental results. 9. Autorotation is sensitive to the cross-sectional aerodynamic coefficients. This sensitivity is critical since the available data on these coefficients is, to say the least, unsatisfactory and require significant improvement.     

长白山9种槭树的翅果扩散及种子萌发研究

翅果的风媒传播是槭属植物的主要扩散方式之一,且与种子萌发有着密切关联,但具体机理一直还并不明确。以分布于长白山的9种槭树为对象,探讨翅果的形态特征,测定它们在空气中的垂直沉降速度、不同风速下的水平扩散距离以及在扩散距离上的种子萌发率,进而比较并分析翅果的形态性状与沉降速度、水平扩散距离的相关性以及萌发率在不同扩散距离上的差异性。结果表明：(1) 9种槭树的翅果长、宽和面积与沉降速度、水平扩散距离均呈负相关;尽管如此,翅果形态并不是风传播物种的最佳分类指标,而翅载力能较好地反应物种的风传播能力;(2)翅果垂直沉降速度和水平扩散距离间存在显著负相关,表明沉降速度越小,翅果在空气中停留的时间越长,水平方向上扩散距离越远,且强风有助于提高翅果的扩散能力;(3)沉降速度最慢的花楷槭在不同风速下的水平扩散距离均最远,而沉降速度最快的拧筋槭水平扩散距离最短;(4)种子萌发率随扩散距离的增加呈下降趋势。上述结果不仅为深入理解翅果的风力传播机制以及种子萌发对水平扩散距离的响应机制提供科学依据,还可为种群实生更新方面的理论研究提供参考。     

Intra-flock variability in the body reserve dynamics of meat sheep by analyzing BW and body condition score variations over multiple production cycles

Breeding for resilience requires a better understanding of intra-flock variability and the related mechanisms responsible for robustness traits. Among such traits, the animals’ ability to cope with feed fluctuations by mobilizing or restoring body reserves (BR) is a key mechanism in ruminants. The objective of this work was to characterize individual variability in BR dynamics in productive Romane ewes reared in extensive conditions. The BR dynamics profiles were characterized by combining individual longitudinal measurements of BW and body condition scores (BCS) over several production cycles. Historical data, including up to 2628 records per trait distributed in 1146 ewes, underwent cluster analysis. Two to four trajectories were observed for BW depending on the cycle, while three trajectories were found for BCS, whatever the cycle. Most trajectories suggested that BR dynamics were similar but the level of BR may differ between ewes. Nevertheless, some trajectories suggested that both BR dynamics and levels were different for a proportion of ewes. Clustering on BW and BCS profiles adjusted for individual level trends, resulted in differences only in the level of BW or BCS, rather than differences in trajectories. Thus, the overall shape of trajectories was not changed considering or not the individual level trend across cycles. In addition to individual variability, the ewe’s age at first lambing and litter size contributed to the distribution of the ewes between the trajectories. Regarding the entire productive life, three trajectories were observed for BW and BCS changes over three productive cycles. Increase in BW at each cycle suggested that ewes kept growing up until 3 to 4 years old in our conditions. Similar alternation of BCS gains and losses across cycles suggested BR dynamics might be repeatable. Many individual trajectories remained the same throughout a ewe’s life, whatever the age at first lambing, parity or litter size. Our results demonstrate the relevance of using BW and BCS changes for characterizing the diversity of BR mobilization–accretion profiles in sheep in a long timespan perspective.     

榆属（榆科）一新种——绵竹榆

绵竹榆的花秋季开放,翅果柱头面被毛,其两侧的翅较果核为窄,果核位于翅果上端接近缺口处,与榔榆(Ulmus parvifolia Jacq.)相似,但树皮深灰色,不规则鳞块状浅裂,叶片先端渐尖,花被片裂至基部,宿存,边缘上部生纤毛,翅果狭椭圆形,中部最宽,向两端渐变窄,果梗与花被等长,长约2 mm,果序梗长约1 mm,而明显不同。     

Intraspecific variation in samara morphology and flight behavior in acer saccharinum (Aceraceae)

We studied intraspecific variation in samara morphology and flight behavior within and among parent trees of Acer saccharinum (silver maple), with a particular focus on the effect of samara shape. Samara mass, area, wing loading, and descent rate from a 4.5-m indoor balcony were measured for 50 undamaged mature samaras from each of six parents. We found significant differences among parental types for all morphological variables and descent rate. These differences yielded a 50% range in mean dispersal potential among the six parents. There was a strong linear correlation between descent rate and square root of wing loading when mean values were plotted for each of the six parental types. But there was considerable within-parent variation for all measured variables, including substantial nonallometric variation in wing loading caused in part by poor correlations between wing area and fruit weight. Parents also differed widely in the relationship between square root of wing loading and descent rate (linear r² = 0.150-0.788), with one parental type showing no significant relationship. Fruits from the same parent with similar values of the square root of wing loading showed as much as a 75-100% difference in descent rate. The usefulness of mass : area indices such as wing loading is limited by its exclusion of aerodynamically important factors such as mass distribution and wing shape, which in our case caused the six parents to behave aerodynamically almost as if they were six separate species.