Jeremy M. V. Rayner

Bio: Jeremy M. V. Rayner is an academic researcher from University of Bristol. The author has contributed to research in topics: Slow flight & Wing. The author has an hindex of 26, co-authored 31 publications receiving 4365 citations. Previous affiliations of Jeremy M. V. Rayner include University of Leeds & University of Cambridge.

Ecological Morphology and Flight in Bats (Mammalia; Chiroptera): Wing Adaptations, Flight Performance, Foraging Strategy and Echolocation

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.

Form and Function in Avian Flight

Abstract: Flapping flight is a highly effective form of locomotion which has permitted the radiation of birds into a wide range of niches. In this chapter I explore how the mechanics of flapping flight have molded the flight adaptations of birds. The paper has three main threads. First, I describe recent theoretical and experimental studies on flapping flight aerodynamics and demonstrate how the mechanical requirements of locomotion are reflected in wingbeat kinematics, in vortex wake structure, and in the action of the pectoral musculature. Next, I consider how flight performance varies with size; scaling has become a central tool in the analysis of flight in birds and has proved a useful means of predicting how different mechanical, physiological, and ecological parameters change in importance with size, morphology, and behavior. However, scaling is frequently misinterpreted: it is size-dependence of the constraints on adaptation which lead to allometric consistency in avian flight morphology, and many of these constraints can be related directly to flight mechanics. Finally, I use a multivariate analysis of wing morphology to demonstrate how these constraints interact to different degrees in different birds and underlie correlations among flight morphology, ecology, and behavior. These threads are then brought together in a discussion of the conjectural relationships between fitness and the evolution of specializations in flight morphology.

A New Approach to Animal Flight Mechanics

Abstract: The mechanics of lift and thrust generation by flying animals are studied by considering the distribution of vorticity in the wake As wake generation is not continuous, the momentum jet theory, which has previously been used, is not satisfactory, and the vortex theory is a more realistic model The vorticity shed by the wings in the course of each powered stroke deforms into a small-cored vortex ring; the wake is a chain of such rings The momentum of each ring sustains and propels the animal; induced power is calculated as the rate of increase of wake kinetic energy A further advantage of the vortex theory is that lift and induced drag coefficients are not required; estimated instantaneous values of these coefficients are generally too large for steady state aerodynamic theory to be appropriate to natural flapping flight The vortex theory is applied to hovering of insects and to avian forward flight A simple expression for induced power in hovering is found Induced power is always greater than simple momentum jet estimates, and the discrepancy becomes substantial as body mass increases In hovering the wake is composed of a stack of horizontal, coaxial, circular vortex rings In forward flight of birds the rings are elliptic; they are neither horizontal nor coaxial because the momentum of each ring balances the vector sum of parasite and profile drag and the bird9s weight Total power consumption as a function of flight velocity is calculated and compared for several species Power reduction is one of the major factors influencing the choice of flight style A large body of data is used to obtain an approximate scaling between stroke period and the body mass for birds Together with relations between other morphological parameters, this is used to estimate the variation of flight speed and power with body mass for birds, and on this basis deviations from allometric scaling can be related to flight proficiency and to the use of such strategies as the bounding flight of small passerines Note: Present address: Department of Zoology, University of Bristol, Woodland Road, Bristol BS8 IUG, UK

Avian wingtip shape reconsidered: wingtip shape indices and morphological adaptations to migration

Abstract: In this paper we review the existing methods of quantifying avian wingtip shape, and compare their efficiency at detecting morphological adaptations to migration. We use multivariate methods to derive two novel measures of avian wingtip shape, pointedness C2 and convexity C3, based on measurements of primary feather lengths. Size-constrained components analysis, a modified form of principal components analysis, is used to ensure that the measures are independent of isometric size, and have a consistent interpretation in terms of the geometric shape of the wingtip. Our measures of pointedness and convexity can be calculated easily for both live birds and museum skins, and can be applied to any ecomorphological or functional analysis of avian wingtip shape. This approach circumvents many of the interpretational problems associated with previous wingtip shape indices that are often based on less accurate wing measurements. To test the suggested interpretations of previously published wingtip shape indices, we use a comparative interspecific analysis to determine the interrelations of our new shape measures with these published indices, and with aerodynamic parameters which have known functional significance. Published indices do not always measure the quantities that they are claimed to do, and are beset with awkward terminological inconsistencies. We assess the efficacy and utility of these wingtip shape measures with regard to predicted and well-known morphological adaptations in the wings for migration. Once phylogenetic bias and extraneous ecological factors are controlled, the majority of published wingtip shape indices are unable to detect morphological differences between migratory and nonmigratory species. Our measures of pointedness and convexity confirm that migrants have wingtips that are relatively more pointed and more convex; they also have wings of relatively larger aspect ratio. The biomechanical implications of these adaptations for different flight behaviours are discussed. An appendix discusses some of the statistical problems involved in the analysis of size and shape, and introduces the size-constrained components analysis (SCCA) method, which is applicable to any study of morphological variation.

A vortex theory of animal flight. Part 1. The vortex wake of a hovering animal

Abstract: The distribution of vorticity in the wake of a hovering bird or insect is considered. The wake is modelled by a chain of coaxial small-cored circular vortex rings stacked one upon another; each member of the chain is generated by a single wing-stroke. Circulation is determined by the animal's weight and the time for which a single ring must provide lift; ring size is calculated from the circulation distribution on the animal's wing. The theory is equally applicable to birds and insects, although the mechanism of ring formation differs. This approach avoids the use of lift and drag coefficients and is not bound by the constraints of steady-state aerodynamics; it gives a wake configuration in agreement with experimental observations. The classical momentum jet approach has steady momentum flux in the wake, and is difficult to relate to the wing motions of a hovering bird or insect; the vortex wake can be related to the momentum jet, but adjacent vortex elements are disjoint and momentum flux is periodic.The evolution of the wake starting from rest is considered by releasing vortex rings at appropriate time intervals and allowing them to interact in their own velocity fields. The resulting configuration depends on the feathering parameter f (which depends on the animal's morphology); f increases with body size. At the lower end of the wake rings coalesce to form a single large vortex, which breaks away from the rest of the wake at intervals. Wake contraction depends on f; the minimum areal contraction of one-half (as in momentum-jet theory) occurs only in the limit f → 0, but values calculated for smaller insects of just over one-half suggest that the momentum jet may be a good approximation to the wake when f is small.Induced power in hovering is calculated as the limit of the mean rate of increase of wake kinetic energy as time progresses. It can be related to the classical momentum-jet induced power by a simple conversion factor. For an insect or hummingbird the usual momentum-jet estimate may be between 10 and 15% too low, but for a bird it may be as much as 50% too low. This suggests that few, if any, birds are able to sustain aerobic hovering, and that as small a value of f as possible would be necessary if the bird were to hover.Tip losses (energy cost of the vortex-ring wake compared with the equivalent momentum jet) are negligible for insects, but can be in the range 15–20% for birds.

Wing rotation and the aerodynamic basis of insect flight.

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.

Wing rotation and the aerodynamic basis of insect flight

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.

Ecological Morphology and Flight in Bats (Mammalia; Chiroptera): Wing Adaptations, Flight Performance, Foraging Strategy and Echolocation

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.

How Animals Move: An Integrative View

Abstract: Recent advances in integrative studies of locomotion have revealed several general principles. Energy storage and exchange mechanisms discovered in walking and running bipeds apply to multilegged locomotion and even to flying and swimming. Nonpropulsive lateral forces can be sizable, but they may benefit stability, maneuverability, or other criteria that become apparent in natural environments. Locomotor control systems combine rapid mechanical preflexes with multimodal sensory feedback and feedforward commands. Muscles have a surprising variety of functions in locomotion, serving as motors, brakes, springs, and struts. Integrative approaches reveal not only how each component within a locomotor system operates but how they function as a collective whole.

Bats: Important Reservoir Hosts of Emerging Viruses

Abstract: Bats (order Chiroptera, suborders Megachiroptera and Microchiroptera) are abundant, diverse, and geographically widespread. These mammals provide us with resources, but their importance is minimized and many of their populations and species are at risk, even threatened or endangered. Some of their characteristics (food choices, colonial or solitary nature, population structure, ability to fly, seasonal migration and daily movement patterns, torpor and hibernation, life span, roosting behaviors, ability to echolocate, virus susceptibility) make them exquisitely suitable hosts of viruses and other disease agents. Bats of certain species are well recognized as being capable of transmitting rabies virus, but recent observations of outbreaks and epidemics of newly recognized human and livestock diseases caused by viruses transmitted by various megachiropteran and microchiropteran bats have drawn attention anew to these remarkable mammals. This paper summarizes information regarding chiropteran characteristics and information regarding 66 viruses that have been isolated from bats. From these summaries, it is clear that we do not know enough about bat biology, that we are doing too little in terms of bat conservation, and that there remain a multitude of questions regarding the role of bats in disease emergence.