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Mechanics and physiology of flight in fossil vertebrates

Published online by Cambridge University Press:  03 November 2011

Jeremy M. V. Rayner
Jeremy M. V. Rayner
Affiliation:
Department of Zoology, University of Bristol, Woodland Road, Bristol BS8 1UG, U.K.
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Abstract

Flight—defined as the ability to produce useful aerodynamic forces by flapping wings—is one of the most demanding adaptations in vertebrates. The mechanical problems of flight ensure considerable external morphological homogeneity and behavioural similarity in extant fliers. Observations of the vortex wakes and wingbeat geometry of modern birds and bats confirm that the two groups are mechanically very similar, despite differences in phylogeny, anatomy and physiology. With this background it is possible to attack two problems: the evolution of flight in vertebrates, and the flight performance of extinct animals such as pterosaurs and Archaeopteryx.

The origin of flight has been surrounded by considerable controversy, due in part to terminological inconsistencies, in part to phylogenetic uncertainty over the relationships of birds, bats and pterosaurs, in part to disagreement over the interpretation of the available fossil evidence, and in part to argument over the relative importance of morphological, mechanical and ecological specialisations. The mechanical changes needed in the course of the evolution of flight favour a gliding origin of tetrapod flight, and on mechanical and ecological grounds the alternative cursorial hypothesis may be discounted. This argument is particularly strong in bats, but has been thought to be weaker in birds owing to apparent inconsistencies with the fossil evidence. However, fossils of the Jurassic theropod dinosaur Archaeopteryx also support a gliding origin for flight, and suggest that this animal was adapted for flapping flight at moderately high speeds associated with gliding; it could fly less well at the slow speeds which would have been required for incipient flight in a running cursor, and at which the wingbeat is aerodynamically and kinematically considerably more complex. Slow flight in birds and bats is the more derived condition, and vertebrate flapping flight apparently evolved through a gliding stage.

The pterosaurs have become the subject of much controversy over the nature of their stance, the wing surface, and the degree of involvement of the leg in the wing membrane. Reconstruction of their wings indicates proficient flying animals, and comparison with birds suggests that most pterosaurs probably occupied marine or coastal/estuarine habitat.

Keywords

aerodynamicsArchaeopteryxbatsbirdsevolutionpalaeoecologypterosaurs
Type
Physiological adaptations in some recent and fossil organisms
Information
Earth and Environmental Science Transactions of The Royal Society of Edinburgh , Volume 80 , Issue 3-4: Environments and Physiology of Fossil Organisms , 1989 , pp. 311 - 320
Copyright
Copyright © Royal Society of Edinburgh 1989

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References

Abel, O. 1912. Grundzüge der Palaeobiologie der Wirbeltiere. Stuttgart: E. Schweizerbart'sche Verlagsbuchhandlung.CrossRefGoogle Scholar
Darwin, C. 1859. The origin of species. London: John Murray.Google Scholar
Desmond, A. J. 1975. The hot-blooded dinosaurs. London: Blond & Briggs.Google Scholar
Feduccia, A. & Tordoff, M. 1979. Feathers of Archaeopteryx: asymmetric vanes indicate aerodynamic function. SCIENCE 203, 1021–1022.Google Scholar
Gauthier, J. 1986. Saurischian monophyly and the origin of birds. In Padian, K. (ed.) The origin of birds and the evolution of flight, 1–55. MEM CALIF ACAD SCI 8.Google Scholar
Hecht, M. K., Ostrom, J. H., Viohl, G. & Wellnhofer, P. (eds) 1985. The beginnings of birds. Eichstätt: JuraMuseum.Google Scholar
Heilmann, G. 1926. The origin of birds. London: Witherby; reprinted New York: Appleton, 1927 and New York: Dover, 1972.Google Scholar
Marsh, O. C. 1880. Odontornithes: a monograph on the extinct toothed birds of North America. PROF PAP ENG DEP U.S. ARMY 18.Google Scholar
Nopsca, F. 1907. Ideas on the origin of flight. PROC ZOOL SOC LONDON, 222–236.Google Scholar
Nopsca, F. 1923. On the origin of flight in birds. PROC ZOOL SOC LONDON, 463–477.Google Scholar
Norberg, R. Å. 1985. Function of vane asymmetry and shaft curvature in bird flight feathers; inference on flight ability in Archaeopteryx. In Hecht, M. K., Ostrom, J. H., Viohl, G. & Wellnhofer, P. (eds) The beginnings of birds, 303–318. Eichstätt: JuraMuseum.Google Scholar
Norberg, U. M. 1985. Evolution of vertebrate flight: an aerodynamic model for the transition from gliding to active flight. AM NAT 126, 303–327.CrossRefGoogle Scholar
Norberg, U. M. & Rayner, J. M. V. 1987. Ecological morphology in bats (Mammalia; Chiroptera): wing adaptations, flight performance, foraging strategy and echolocation. PHILOS TRANS R SOC LONDON B316, 335–427.Google Scholar
Ostrom, J. H. 1974. Archaeopteryx and the origin of flight. Q REV BIOL 49, 27–47.CrossRefGoogle Scholar
Ostrom, J. H. 1979. Bird flight: how did it begin? AM SCI 67, 46–56.Google ScholarPubMed
Padian, K. 1983. A functional analysis of flying and walking in pterosaurs. PALEOBIOLOGY 9, 218–239.Google Scholar
Padian, K. 1985. The origins and aerodynamics of flight in extinct vertebrates. PALAEONTOLOGY 28, 413–433.Google Scholar
Padian, K. (ed.) 1986. The origin of birds and the evolution of flight. MEM CALIF ACAD SCI 8.Google Scholar
Padian, K. 1987. A comparative phylogenetic and functional approach to the origin of vertebrate flight. In Fenton, M. B., Racey, P. A. & Rayner, J.M.V. (eds) Recent advances in the study of bats, 3–19. Cambridge: Cambridge University Press.Google Scholar
Padian, K. 1988. Pterosaurs: were they functional birds or functional bats? In Rayner, J. M. V. (ed.) Biomechanics in evolution. SEMIN SER SOC EXP BIOL. Cambridge: Cambridge University Press, in the press.Google Scholar
Pettigrew, J. D. 1986. Flying primates? Megabats have the advanced pathway from eye to mid-brain. SCIENCE 231, 1304–1306.Google Scholar
Rayner, J. M. V. 1980. Vorticity and animal flight. In Elder, H. Y. & Trueman, E. R., Aspects of animal movement. SEMIN SER SOC EXP BIOL 5, 177–179. Cambridge: Cambridge University Press.Google Scholar
Rayner, J. M. V. 1981. Flight adaptations in vertebrates. SYMP ZOOL SOC LOND 48, 137–172.Google Scholar
Rayner, J. M. V. 1985a. Vorticity and propulsion mechanics in swimming and flying vertebrates. In Riess, J. & Frey, E. (eds) Konstruktionsprinzipien lebender and ausgestorbener Reptilien, 89–118. KONZEPTE SFB230 4, Universität Tübingen.Google Scholar
Rayner, J. M. V. 1985b. Mechanical and ecological constraints on flight evolution. In Hecht, M. K., Ostrom, J. H., Viohl, G. & Wellnhofer, P. (eds) The beginnings of birds, 279–288. Eichstatt: JuraMuseum.Google Scholar
Rayner, J. M. V. 1985c. Cursorial gliding in proto-birds. In Hecht, M. K., Ostrom, J. H., Viohl, G. & Wellnhofer, P. (eds) The beginnings of birds, 289–292. Eichstatt: JuraMuseum.Google Scholar
Rayner, J. M. V. 1986. Vertebrate flapping flight mechanics and aerodynamics, and the evolution of flight in bats. In Nachtigall, W. (ed.) Biona Report 5, Bat flight—Fledermausflug, 27–74. Stuttgart: Gustav Fischer Verlag.Google Scholar
Rayner, J. M. V. 1987. The mechanics of flapping flight in bats. In Fenton, M. B., Racey, P. A. & Rayner, J. M. V. (eds) Recent advances in the study of bats, 23–42. Cambridge: Cambridge University Press.Google Scholar
Rayner, J. M. V. 1988a. Form and function in avian flight. CURR ORNITHOL 5, 1–77.Google Scholar
Rayner, J. M. V. 1988b. The evolution of vertebrate flight. BIOL J LINN SOC 34, 269–287.Google Scholar
Rayner, J. M. V., Jones, G. & Thomas, A. 1986. Vortex flow visualizations reveal change of upstroke function with flight speed in microchiropteran bats. NATURE 321, 162–164.Google Scholar
Sanz, J. I., Bonaparte, J. F. & Lacasa, A. 1988. Unusual Early Cretaceous birds from Spain. NATURE 331, 433–435.CrossRefGoogle Scholar
Scholey, K. D. 1986. The evolution of flight in bats. In Nachtigall, W. (ed.) Biona Report S, Bat flight—Fledermausflug, 1–12. Stuttgart: Gustav Fischer Verlag.Google Scholar
Spedding, G. R. 1986. The wake of a jackdaw (Corvus monedula) in slow flight. J EXP BIOL 125, 287–307.CrossRefGoogle Scholar
Spedding, R. G. 1987. The wake of a kestrel (Falco tinnunculus) in flapping flight. J EXP BIOL 127, 59–78.CrossRefGoogle Scholar
Spedding, G. R., Rayner, J. M. V. & Pennycuick, C. J. 1984. Momentum and energy in the wake of a pigeon (Columbalivia) in slow flight. J EXP BIOL 111, 81–102.Google Scholar
Stephan, B. 1987. Urvögel (3rd edn). Neue Brehm-Bücherei 465. Wittenberg-Lutherstadt: A. Ziemsen Verlag.Google Scholar
Unwin, D. M. 1987. Pterosaur locomotion: joggers or waddlers? NATURE 327, 13–14.CrossRefGoogle Scholar
Unwin, D. M. 1988. New remains of the pterosaur Dimorphodon (Pterosauria: Rhamphortynchoidea) and the terrestrial ability of early pterosaurs. MOD GEOL 13, 57–68.Google Scholar
Wellnhofer, P. 1978. Handbuch der Paläoherpetologie. Teil 19. Pterosauria. Stuttgart: Gustav Fischer Verlag.Google Scholar
Wellnhofer, P. 1980. Flugsaurier-Pterosaurier. Neue Brehm-Bücherei 534. Wittenberg-Lutherstadt: A. Ziemsen Verlag.Google Scholar
Wellnhofer, P. 1985. Neue Pterosaurier aus der Santana-Formation (Apt) der Chapada do Araripe, Brasilien. PALAEONTOGRAPHICA A187, 105–182.Google Scholar
Wellnhofer, P. 1987. Die Flughaut von Pterodactylus (Reptilia, Pterosauria) am Beispiel des Wiener Examplares von Pterodactylus kochi (Wagner). ANN NATHIST MUS WIEN 88A, 149–162.Google Scholar
Wellnhofer, P. 1988a. Terrestrial locomotion in pterosaurs. HIST BIOL 1, 3–16.Google Scholar
Wellnhofer, P. 1988b. A new specimen of Archaeopteryx. SCIENCE 240, 1790–1792.Google Scholar
Wellnhofer, P. 1988c. Ein neues Examplar von Archaeopteryx. ARCHAEOPTERYX 6, 1–30.Google Scholar
Williston, S. W. 1879. Are birds derived from dinosaurs? KANSAS CITY REV SCI 3, 475–460.Google Scholar
Yalden, D. W. 1985. Forelimb function in Archaeopteryx. In Hecht, M. K., Ostrom, J. H., Viohl, G. & Wellnhofer, P. (eds) Thebeginnings of birds, 91–98. Eichstatt: JuraMuseum.Google Scholar