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      Flapping Tail Membrane in Bats Produces Potentially Important Thrust during Horizontal Takeoffs and Very Slow Flight

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      PLoS ONE
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          Abstract

          Historically, studies concerning bat flight have focused primarily on the wings. By analyzing high-speed video taken on 48 individuals of five species of vespertilionid bats, we show that the capacity to flap the tail-membrane (uropatagium) in order to generate thrust and lift during takeoffs and minimal-speed flight (<1 m s−1) was largely underestimated. Indeed, bats flapped the tail-membrane by extensive dorso-ventral fanning motions covering as much as 135 degrees of arc consistent with thrust generation by air displacement. The degree of dorsal extension of the tail-membrane, and thus the potential amount of thrust generated during platform launches, was significantly correlated with body mass (P = 0.02). Adduction of the hind limbs during upstrokes collapsed the tail-membrane thereby reducing its surface area and minimizing negative lift forces. Abduction of the hind limbs during the downstroke fully expanded the tail-membrane as it was swept ventrally. The flapping kinematics of the tail-membrane is thus consistent with expectations for an airfoil. Timing offsets between the wings and tail-membrane during downstrokes was as much as 50%, suggesting that the tail-membrane was providing thrust and perhaps lift when the wings were retracting through the upstoke phase of the wing-beat cycle. The extent to which the tail-membrane was used during takeoffs differed significantly among four vespertilionid species (P = 0.01) and aligned with predictions derived from bat ecomorphology. The extensive fanning motion of the tail membrane by vespertilionid bats has not been reported for other flying vertebrates.

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          Software techniques for two- and three-dimensional kinematic measurements of biological and biomimetic systems.

          Researchers studying aspects of locomotion or movement in biological and biomimetic systems commonly use video or stereo video recordings to quantify the behaviour of the system in question, often with an emphasis on measures of position, velocity and acceleration. However, despite the apparent simplicity of video analysis, it can require substantial investment of time and effort, even when performed with adequate software tools. This paper reviews the underlying principles of video and stereo video analysis as well as its automation and is accompanied by fully functional and freely available software implementation.
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            Transitions from Drag-based to Lift-based Propulsion in Mammalian Swimming

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              Leading-edge vortex improves lift in slow-flying bats.

              Staying aloft when hovering and flying slowly is demanding. According to quasi-steady-state aerodynamic theory, slow-flying vertebrates should not be able to generate enough lift to remain aloft. Therefore, unsteady aerodynamic mechanisms to enhance lift production have been proposed. Using digital particle image velocimetry, we showed that a small nectar-feeding bat is able to increase lift by as much as 40% using attached leading-edge vortices (LEVs) during slow forward flight, resulting in a maximum lift coefficient of 4.8. The airflow passing over the LEV reattaches behind the LEV smoothly to the wing, despite the exceptionally large local angles of attack and wing camber. Our results show that the use of unsteady aerodynamic mechanisms in flapping flight is not limited to insects but is also used by larger and heavier animals.
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                Author and article information

                Contributors
                Role: Editor
                Journal
                PLoS One
                plos
                plosone
                PLoS ONE
                Public Library of Science (San Francisco, USA )
                1932-6203
                2012
                29 February 2012
                : 7
                : 2
                : e32074
                Affiliations
                [1]School of Biological Sciences, University of Northern Colorado, Greeley, Colorado, United States of America
                University of Western Ontario, Canada
                Author notes

                Conceived and designed the experiments: RAA. Performed the experiments: RAA ERS JBS. Analyzed the data: RAA. Contributed reagents/materials/analysis tools: RAA. Wrote the paper: RAA.

                Article
                PONE-D-11-16733
                10.1371/journal.pone.0032074
                3290531
                22393378
                214603eb-a1de-4b27-a8d1-9ee69d4b92e0
                Adams 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
                : 22 August 2011
                : 20 January 2012
                Page count
                Pages: 11
                Categories
                Research Article
                Biology
                Biophysics
                Biomechanics
                Evolutionary Biology
                Zoology
                Veterinary Science
                Animal Types
                Veterinary Anatomy and Physiology

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                Uncategorized

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