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      Comparing Aerodynamic Efficiency in Birds and Bats Suggests Better Flight Performance in Birds

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          Abstract

          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.

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          Most cited references66

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          Adaptations to migration in birds: behavioural strategies, morphology and scaling effects.

          The annual life cycle of many birds includes breeding, moult and migration. All these processes are time and energy consuming and the extent of investment in any one may compromise the others. The output from breeding is of course the ultimate goal for all birds, while the investment in moult and migration should be selected so that lifetime fitness is maximized. In particular, long-distance migrants breeding at high latitudes face severe time pressures, which is a probable reason why natural selection has evolved efficient behaviours, physiological and morphological adaptations allowing the maximum possible migration speed. Optimal migration theory commonly assumes time minimization as an overall strategy, but the minimization of energy cost and predation risk may also be involved. Based on these assumptions, it is possible to derive adaptive behaviours such as when and at which fuel load a stopover site should be abandoned. I review some core components of optimal migration theory together with some key predictions. A review of accumulated empirical tests of the departure rule indicates that time minimization is an important component of the overall migration strategy, and hence gives support to the assumption about time-selected migration. I also briefly discuss how the optimal policy may be implemented by the bird by applying a set of simple rules. The time constraints on migrants increase with increasing body size. Some consequences of this are discussed.
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            Bat flight generates complex aerodynamic tracks.

            The flapping flight of animals generates an aerodynamic footprint as a time-varying vortex wake in which the rate of momentum change represents the aerodynamic force. We showed that the wakes of a small bat species differ from those of birds in some important respects. In our bats, each wing generated its own vortex loop. Also, at moderate and high flight speeds, the circulation on the outer (hand) wing and the arm wing differed in sign during the upstroke, resulting in negative lift on the hand wing and positive lift on the arm wing. Our interpretations of the unsteady aerodynamic performance and function of membranous-winged, flapping flight should change modeling strategies for the study of equivalent natural and engineered flying devices.
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              A family of vortex wakes generated by a thrush nightingale in free flight in a wind tunnel over its entire natural range of flight speeds.

              In view of the complexity of the wing-beat kinematics and geometry, an important class of theoretical models for analysis and prediction of bird flight performance entirely, or almost entirely, ignores the action of the wing itself and considers only the resulting motions in the air behind the bird. These motions can also be complicated, but some success has previously been recorded in detecting and measuring relatively simple wake structures that can sometimes account for required quantities used to estimate aerodynamic power consumption. To date, all bird wakes, measured or presumed, seem to fall into one of two classes: the closed-loop, discrete vortex model at low flight speeds, and the constant-circulation, continuous vortex model at moderate to high speeds. Here, novel and accurate quantitative measurements of velocity fields in vertical planes aligned with the freestream are used to investigate the wake structure of a thrush nightingale over its entire range of natural flight speeds. At most flight speeds, the wake cannot be categorised as one of the two standard types, but has an intermediate structure, with approximations to the closed-loop and constant-circulation models at the extremes. A careful accounting for all vortical structures revealed with the high-resolution technique permits resolution of the previously unexplained wake momentum paradox. All the measured wake structures have sufficient momentum to provide weight support over the wingbeat. A simple model is formulated and explained that mimics the correct, measured balance of forces in the downstroke- and upstroke-generated wake over the entire range of flight speeds. Pending further work on different bird species, this might form the basis for a generalisable flight model.
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                Author and article information

                Contributors
                Role: Editor
                Journal
                PLoS One
                PLoS ONE
                plos
                plosone
                PLoS ONE
                Public Library of Science (San Francisco, USA )
                1932-6203
                2012
                18 May 2012
                : 7
                : 5
                : e37335
                Affiliations
                [1 ]Department of Biology, Lund University, Lund, Sweden
                [2 ]Department of Biology, University of Washington, Seattle, Washington, United States of America
                [3 ]Department of Natural Sciences, University of Michigan-Dearborn, Dearborn, Michigan, United States of America
                [4 ]Cognitive Neurobiology, Humboldt University and NeuroCure Center of Excellence at the Charité Universitätsmedizin Berlin, Berlin, Germany
                University of Hull, United Kingdom
                Author notes

                Conceived and designed the experiments: FTM LCJ AH. Performed the experiments: FTM MSB. Analyzed the data: FTM MSB. Contributed reagents/materials/analysis tools: FTM LCJ. Wrote the paper: FTM LCJ MSB AH. Provided the experimental bats: YW.

                Article
                PONE-D-11-25065
                10.1371/journal.pone.0037335
                3356262
                22624018
                9c0504fb-8067-466d-9f27-881fcb9d6622
                Muijres et al. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
                History
                : 14 December 2011
                : 20 April 2012
                Page count
                Pages: 9
                Categories
                Research Article
                Biology
                Biophysics
                Biomechanics
                Biological Fluid Mechanics
                Evolutionary Biology
                Organismal Evolution
                Animal Evolution
                Engineering
                Aerospace Engineering
                Fluid Mechanics
                Bioengineering
                Biomimetics
                Physics
                Biophysics
                Biomechanics
                Biological Fluid Mechanics

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                Uncategorized

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