Wing

Abstract: The enhanced aerodynamic performance of insects results from an interaction of three distinct yet interactive mechanisms: delayed stall, rotational circulation, and wake capture. Delayed stall functions during the translational portions of the stroke, when the wings sweep through the air with a large angle of attack. In contrast, rotational circulation and wake capture generate aerodynamic forces during stroke reversals, when the wings rapidly rotate and change direction. In addition to contributing to the lift required to keep an insect aloft, these two rotational mechanisms provide a potent means by which the animal can modulate the direction and magnitude of flight forces during steering maneuvers. A comprehensive theory incorporating both translational and rotational mechanisms may explain the diverse patterns of wing motion displayed by different species of insects.

Abstract: Bat wing morphology is considered in relation to flight performance and flight behaviour to clarify the functional basis for eco-morphological correlations in flying animals. Bivariate correlations are presented between wing dimensions and body mass for a range of bat families and feeding classes, and principal-components analysis is used to measure overall size, wing size and wing shape. The principal components representing wing size and wing shape (as opposed to overall size) are interpreted as being equivalent to wing loading and to aspect ratio. Relative length and area of the hand-wing or wingtip are determined independently of wing size, and are used to derive a wingtip shape index, which measures the degree of roundedness or pointedness of the wingtip. The optimal wing form for bats adapted for different modes of flight is predicted by means of mechanical and aerodynamic models. We identify and model aspects of performance likely to influence flight adaptation significantly; these include selective pressures for economic forward flight (low energy per unit time or per unit distance (equal to cost of transport)), for flight at high or low speeds, for hovering, and for turning. Turning performance is measured by two quantities: manoeuvrability, referring to the minimum space required for a turn at a given speed; and agility, relating to the rate at which a turn can be initiated. High flight speed correlates with high wing loading, good manoeuvrability is favoured by low wing loading, and turning agility should be associated with fast flight and with high wing loading. Other factors influencing wing adaptations, such as migration, flying with a foetus or young or carrying loads in flight (all of which favour large wing area), flight in cluttered environments (short wings) and modes of landing, are identified. The mechanical predictions are cast into a size-independent principal-components form, and are related to the morphology and the observed flight behaviour of different species and families of bats. In this way we provide a broadly based functional interpretation of the selective forces that influence wing morphology in bats. Measured flight speeds in bats permit testing of these predictions. Comparison of open-field free-flight speeds with morphology confirms that speed correlates with mass, wing loading and wingtip proportions as expected; there is no direct relation between speed and aspect ratio. Some adaptive trends in bat wing morphology are clear from this analysis. Insectivores hunt in a range of different ways, which are reflected in their morphology. Bats hawking high-flying insects have small, pointed wings which give good agility, high flight speeds and low cost of transport. Bats hunting for insects among vegetation, and perhaps gleaning, have very short and rounded wingtips, and often relatively short, broad wings, giving good manoeuvrability at low flight speeds. Many insectivorous species forage by `flycatching' (perching while seeking prey) and have somewhat similar morphology to gleaners. Insectivorous species foraging in more open habitats usually have slightly longer wings, and hence lower cost of transport. Piscivores forage over open stretches of water, and have very long wings giving low flight power and cost of transport, and unusually long, rounded tips for control and stability in flight. Carnivores must carry heavy loads, and thus have relatively large wing areas; their foraging strategies consist of perching, hunting and gleaning, and wing structure is similar to that of insectivorous species with similar behaviour. Perching and hovering nectarivores both have a relatively small wing area: this surprising result may result from environmental pressure for a short wingspan or from the advantage of high speed during commuting flights; the large wingtips of these bats are valuable for lift generation in slow flight. The relation between flight morphology (as an indicator of flight behaviour) and echolocation is considered. It is demonstrated that adaptive trends in wing adaptations are predictably and closely paralleled by echolocation call structure, owing to the joint constraints of flying and locating food in different ways. Pressures on flight morphology depend also on size, with most aspects of performance favouring smaller animals. Power rises rapidly as mass increases; in smaller bats the available energy margin is greater than in larger species, and they may have a more generalized repertoire of flight behaviour. Trophic pressures related to feeding strategy and behaviour are also important, and may restrict the size ranges of different feeding classes: insectivores and primary nectarivores must be relatively small, carnivores and frugivores somewhat larger. The relation of these results to bat community ecology is considered, as our predictions may be tested through comparisons between comparable, sympatric species. Our mechanical predictions apply to all bats and to all kinds of bat communities, but other factors (for example echolocation) may also contribute to specialization in feeding or behaviour, and species separation may not be determined solely by wing morphology or flight behaviour. None the less, we believe that our approach, of identifying functional correlates of bat flight behaviour and identifying these with morphological adaptations, clarifies the eco-morphological relationships of bats.

Abstract: INSECTS cannot fly, according to the conventional laws of aerodynamics: during flapping flight, their wings produce more lift than during steady motion at the same velocities and angles of attack1–5. Measured instantaneous lift forces also show qualitative and quantitative disagreement with the forces predicted by conventional aerodynamic theories6–9. The importance of high-life aerodynamic mechanisms is now widely recognized but, except for the specialized fling mechanism used by some insect species1,10–13, the source of extra lift remains unknown. We have now visualized the airflow around the wings of the hawkmoth Manduca sexta and a 'hovering' large mechanical model—the flapper. An intense leading-edge vortex was found on the down-stroke, of sufficient strength to explain the high-lift forces. The vortex is created by dynamic stall, and not by the rotational lift mechanisms that have been postulated for insect flight14–16. The vortex spirals out towards the wingtip with a spanwise velocity comparable to the flapping velocity. The three-dimensional flow is similar to the conical leading-edge vortex found on delta wings, with the spanwise flow stabilizing the vortex.

Abstract: 1. On the assumption that steady-state aerodynamics applies, simple analytical expressions are derived for the average lift coefficient, Reynolds number, the aerodynamic power, the moment of inertia of the wing mass and the dynamic efficiency in animals which perform normal hovering with horizontally beating wings. 2. The majority of hovering animals, including large lamellicorn beetles and sphingid moths, depend mainly on normal aerofoil action. However, in some groups with wing loading less than 10 N m -2 (1 kgf m -2 ), non-steady aerodynamics must play a major role, namely in very small insects at low Reynolds number, in true hover-flies (Syrphinae), in large dragonflies (Odonata) and in many butterflies (Lepidoptera Rhopalocera). 3. The specific aerodynamic power ranges between 1.3 and 4.7 WN -1 (11-40 cal h -1 gf -1 ) but power output does not vary systematically with size, inter alia because the lift/drag ratio deteriorates at low Reynolds number. 4. Comparisons between metabolic rate, aerodynamic power and dynamic efficiency show that the majority of insects require and depend upon an effective elastic system in the thorax which counteracts the bending moments caused by wing inertia. 5. The free flight of a very small chalcid wasp Encarsia formosa has been analysed by means of slow-motion films. At this low Reynolds number (10-20), the high lift co-efficient of 2 or 3 is not possible with steady-state aerodynamics and the wasp must depend almost entirely on non-steady flow patterns. 6. The wings of Encarsia are moved almost horizontally during hovering, the body being vertical, and there are three unusual phases in the wing stroke: the clap , the fling and the flip . In the clap the wings are brought together at the top of the morphological upstroke. In the fling, which is a pronation at the beginning of the morphological downstroke, the opposed wings are flung open like a book, hinging about their posterior margins. In the flip, which is a supination at the beginning of the morphological upstroke, the wings are rapidly twisted through about 180°. 7. The fling is a hitherto undescribed mechanism for creating lift and for setting up the appropriate circulation over the wing in anticipation of the downstroke. In the case of Encarsia the calculated and observed wing velocities at which lift equals body weight are in agreement, and lift is produced almost instantaneously from the beginning of the downstroke and without any Wagner effect. The fling mechanism seems to be involved in the normal flight of butterflies and possibly of Drosophila and other small insects. Dimensional and other considerations show that it could be a useful mechanism in birds and bats during take-off and in emergencies. 8. The flip is also believed to be a means of setting up an appropriate circulation around the wing, which has hitherto escaped attention; but its operation is less well understood. It is not confined to Encarsia but operates in other insects, not only at the beginning of the upstroke (supination) but also at the beginning of the downstroke where a flip (pronation) replaces the clap and fling of Encarsia . A study of freely flying hover-flies strongly indicates that the Syrphinae (and Odonata) depend almost entirely upon the flip mechanism when hovering. In the case of these insects a transient circulation is presumed to be set up before the translation of the wing through the air, by the rapid pronation (or supination) which affects the stiff anterior margin before the soft posterior portions of the wing. In the flip mechanism vortices of opposite sense must be shed, and a Wagner effect must be present. 9. In some hovering insects the wing twistings occur so rapidly that the speed of propagation of the elastic torsional wave from base to tip plays a significant role and appears to introduce beneficial effects. 10. Non-steady periods, particularly flip effects, are present in all flapping animals and they will modify and become superimposed upon the steady-state pattern as described by the mathematical model presented here. However, the accumulated evidence indicates that the majority of hovering animals conform reasonably well with that model. 11. Many new types of analysis are indicated in the text and are now open for future theoretical and experimental research.