The extent of the pterosaur flight membrane

• Autorzy: Elgin R. , Hone D.W.E. , Frey E.
• Języki publikacji: EN

Abstrakty

EN

The shape and extent of the membranous brachioptagium in pterosaurs remains a controversial topic for those attempting to determine the aerodynamic performance of the first vertebrate fliers. Various arguments in favour of the trailing edge terminating against either the torso or hip, the femur, the ankle, or different locations for various taxa, has resulted in several published reconstructions. Uncertainty over the correct model is detrimental to both aerodynamic and palaeoecological studies that are forced to simultaneously consider multiple and highly variable configurations for individual taxa. A review of relevant pterosaur specimens with preserved soft tissues or impressions of the wing membrane, however, strongly suggests that the trailing edge of the wing extended down to the lower leg or ankle in all specimens where the brachiopatagium is completely preserved. This configuration is seen across a phylogenetically broad range of pterosaurs and is thus likely to have been universally present throughout the Pterosauria. Support for opposing hypotheses where the trailing edge terminates against the body, hip, or knee are based on several specimens where the wing membrane is either incomplete or has undergone post−mortem contraction. An ankle attachment does not rule out a high aspect ratio wing as the curvature of the trailing edge and the ratio of the fore to hind limbs also play a major role in determining the final shape of the membrane.

Słowa kluczowe

EN

paleontology; Reptilia; Pterosauria; flight; wing; Mesozoic; pterosaur; wing membrane; fossil

• Wydawca: -
• Czasopismo: Acta Palaeontologica Polonica
• Rocznik: 2011
• Tom: 56
• Numer: 1
• Opis fizyczny: p.99-111,fig.,ref.

Twórcy

autor

Elgin R.

• Staatliches Museum für Naturkunde Karlsruhe (SMNK), Erbprinzenstraße 13, 76133 Karlsruhe, Germany

autor

Hone D.W.E.

autor

Frey E.

Bibliografia

• Abel, O. 1919. Neue Rekonstruktionen der flugsauriergattungen Pterodactylus and Rhamphorhynchus. Die Naturwissenschaften 37: 661–665. [CrossRef]
• Abel, O. 1925. Geschichte und Methode der Rekonstruktion vorzeitlicher Wirbeltiere. 327 pp. Gustav Fischer Verlag, Stuttgart.
• Andres, B. and Ji, Q. 2008. A new pterosaur from the Liaoning Province of China, the phylogeny of the Pterodactyloidea, and convergence in their cervical vertebrae. Palaeontology 51: 453–469. [CrossRef]
• Andres, B., Clark, J.M., and Xing, X. 2010. A new rhamphorhynchid pterosaur from the Upper Jurassic of Xinjiang, China, and the phylogenetic relationships of basal pterosaurs. Journal of Vertebrate Paleontology 30: 163–187. [CrossRef]
• Bakhurina, N.N. and Unwin, D.M. 2003. Reconstructing the flight apparatus of Eudimorphodon. Rivista del Museo Civico di Scienze Naturali “Enrico Caffi” 22: 5–8.
• Bennett, S.C. 1987. New evidence on the tail of the pterosaur Pteranodon (Archiosauria, Pterosauria). In: P.J. Currie and E.H. Koster (eds.), Fourth Symposium On Mesozoic Terrestrial Ecosystems, Short Papers. Occasional Papers of the Tyrrell Museum of Paleontology 3: 18–23.
• Bennett, S.C. 1990. A pterodactyloid pterosaur pelvis from the Santana Formation of Brazil: Implications for terrestrial locomotion. Journal of Vertebrate Paleontology 10: 80–85. [CrossRef]
• Bennett, S.C. 1997. The arboreal leaping theory of the origin of pterosaur flight. Historical Biology 12: 265–290. [CrossRef]
• Bennett, S.C. 2000. Pterosaur Flight: the role of the actinofibrils in wing function. Historical Biology 14: 255–284. [CrossRef]
• Bennett, S.C. 2001. The osteology and functional morphology of the late cretaceous pterosaur Pteranodon. Part 1. General description of osteology. Palaeontographica A 260: 1–112.
• Bennett, S.C. 2005. Pterosaur science or pterosaur fantasy? Prehistoric Times 70: 21–23, 40–41.
• Bennett, S.C. 2007a. Articulation and function of the pteroid bone in pterosaurs. Journal of Vertebrate Paleontology 27: 881–891. [CrossRef]
• Bennett, S.C. 2007b. A second specimen of the pterosaur Anurognathus ammoni. Paläontologische Zeitschrift 81: 376–398.
• Bramwell, C.D. and Whitfield, G.R. 1974. Biomechanics of Pteranodon. Philosophical Transactions of the Royal Society, London B 267: 503–581. [CrossRef]
• Brower, J.C. 1980. Pterosaurs: How they flew? Episodes 4: 21–24.
• Brower, J.C. 1982. The aerodynamics of an ancient flying reptile. Syracuse Scholar 1982: 45–57.
• Brower, J.C. 1983. The aerodynamics of Pteranodon and Nyctosaurus, two large pterosaurs from the Upper Cretaceous of Kansas. Journal of Vertebrate Paleontology 3: 84–124. [CrossRef]
• Chatterjee, S. and Templin, R.J. 2004. Posture, locomotion, and paleoecology of pterosaurs. Geological Society of America, Special Paper 376: 1–64.
• Clark, J.M., Hopson, J.A., Hernández, R.R., Fastovsky, D.E., and Monotellano, M. 1998. Foot posture in a primitive pterosaur. Nature 391: 886–889. [CrossRef]
• Collini, C. 1784. Sur quelques Zoolithes du Cabinet d’Historie naturelle de S.A.S.E Palatine et du Bavière, à Mannheim. Acta Academiae Theodoro Palatinae, Mannheim, Pars Physica 5: 58–103.
• Cuvier, G. 1801. Reptile volant. Extrait d’un ouvrage sur les espèces de quadrupèdes don’t on a trouvè les ossemens dans l’intérieur de la terre. Journal de Physique, de Chimie et d’Histoire Naturelle 52: 253–267.
• Döderlein, L. 1923. Anurognathus Ammoni, ein neuer Flugsaurier. Sitzungsberichte der Mathematisch−Naturwissenschaftlichen Abteilung der Bayerischen Akademie der Wissenschaften zu München 1923: 306–307.
• Dyke, G.J., Nudds, R.L., and Rayner, J.M. 2006. Limb disparity and wing shape in pterosaurs. Journal of Evolutionary Biology 19: 1339–1342. [CrossRef]
• Faux, C.M. and Padian, K. 2007. The opisthotonic posture of vertebrate skeletons: post−mortem contraction or death throes? Paleobiology 32: 201–226. [CrossRef]
• Frey, E. and Martill, D.M. 1998. Soft tissue preservation in a specimen of Pterodactylus kochi (Wagner) from the Upper Jurassic of Germany. Neues Jahrbuch für Geologie und Paläontologie Abhandlungen 210: 421–441.
• Frey, E., and Tischlinger, H. 2000. Weichteilanatomie der Flugsaurierfüße und Bau der Scheitelkämme: Neue Pterosaurierfunde aus den Solnhofener Schichten (Bayern) und der Crato−Formation. Archaeopteryx 18: 1–16.
• Frey, E., Tischlinger, H., Buchy, M.−C., and Martill, D.M. 2003. New specimens of Pterosauria (Reptilia) with soft parts with implications for pterosaurian anatomy and locomotion. In: E. Buffetaut and J.−M. Mazin (eds.), Evolution and Palaeobiology of Pterosaurs. Geological Society of London, Special Publications 217: 233–266.
• Goldfuss, G.A. 1831. Beiträge zur Kenntnis verschiedener Reptilien der Vorwelt. Nova acta Academiae Caesareae Leopoldino−Carolinae Germanicae Naturae Curiosorum 15: 61–128.
• Hankin, E.H. and Watson, D.S.M. 1914. On the flight of pterodactyls. The Aeronautical Journal 18: 324–335.
• Hazlehurst, G.A. and Rayner, J.M.V. 1992. Flight characteristics of Triassic and Jurassic Pterosauria: an appraisal based on wing shape. Paleobiology 18: 447–463.
• Heptonstall, W.B. 1971. An analysis of the flight of the Cretaceous pterodactyl Pteranodon ingens (Marsh). Scottish Journal of Geology 7: 61–78.
• Jaekel, O. 1910. Rekonstruktion fossiler Tiere. Meyers Großes Konversations Lexikon 22: 1–10.
• Ji, S.−A. and Ji, Q. 1997. Discovery of a new pterosaur from western Liaoning, China. Acta Geologica Sinica 71: 1–6.
• Ji, Q., and Yuan, C.X. 2002. Discovery of two kinds of protofeathered-pterosaurs in the Mesozoic Daohugou Biota in the Ningcheng region and its stratigraphic and biologic significances [in Chinese]. Geological Review 48: 221–224.
• Kellner, A.W.A. 2003. Pterosaur phylogeny and comments on the evolutionary history of the group. In: E. Buffetaut and J.−M. Mazin (eds.), Evolution and Palaeobiology of Pterosaurs. Geological Society of London, Special Publications 217: 105–137.
• Kellner, A.W.A., Wang, X., Tischlinger, H., Campos, D.A., Hone, D.W.E., and Meng, X. 2009.The soft tissue of Jeholopterus(Pterosauria, Anurognathidae, Batrachognathinae) and the structure of the pterosaur wing membrane. Proceedings of the Royal Society B 277 (1679): 321–329. [CrossRef]
• Kripp, D. 1941. Ein Lebenbild von Pteranodon ingens auf flugtechnischer Grundlage. Luftwissen 8: 217–246.
• Lü, J.−C. 2002. Soft tissue in an Early Cretaceous pterosaur from Liaoning province, China. Memoir of the Fukui Prefectural Dinosaur Museum 1: 19–28.
• Lü, J.−C. 2003. A new pterosaur: Beipiaopterus chenianus, gen. et sp. nov. (Reptilia: Pterosauria) from the Western Liaoning Province of China. Memoir of the Fukui Prefectural Dinosaur Museum 2: 153–160.
• Lü, J.−C., Unwin, D.M., Jin, X., Liu, Y., and Ji, Q. 2009. Evidence for modular evolution in a long−tailed pterosaur with a pterodactyloid skull. Proceedings of the Royal Society B 277 (1680): 383–389. [CrossRef]
• MacCready, P. 1985. The great pterodactyl project. Engineering and Science 49: 18–24.
• Marsh, O.C. 1882. The wings of pterodactyls. American Journal of Science 23: 251–256.
• Martill, D.M. and Unwin, D.M. 1989. Exceptionally well preserved pterosaur wing membrane from the Cretaceous of Brazil. Nature 340: 138–140. [CrossRef]
• McGowan, A.J. and Dyke, G. 2009. A morphospace−based test for competitive exclusion amoung flying vertebrates: did birds, bats and pterosaurs get in each other’s space? Trustees of the Natural History Museum 20: 1230–1236.
• Münster, G.G. 1839. Ueber einige neue Versteinerungen in der lithographischen Schiefer von Baiern. Neues Jahrbuch für Mineralogie, Geologie, und Palaeontologie 1839: 676–682.
• Padian, K. 1983. A functional analysis of flying and walking in pterosaurs. Paleobiology 9: 218–239.
• Padian, K. 1985. The origins and aerodynamics of flight in extinct vertebrates. Palaeontology 28: 423–433.
• Padian, K. 1987. The case of the bat−winged pterosaur. Typological taxonomy and the influence of pictorial representation on scientific perception. In: S.J. Czerkas and E.C. Olson (eds.), Dinosaurs Past and Present. Vol. 2, 65–81. Natural History Museum of Los Angeles County and University of Washington Press, Seattle.
• Padian, K. 2003. Pterosaur stance and gait and the interpretation of trackways. Ichnos 10: 115–126. [CrossRef]
• Padian, K. 2008. Were pterosaurs ancestors bipedal or quadrupedal?: Morphometric, functional, and phylogenetic considerations. Zitteliana B 28: 21–33.
• Padian, K. and Rayner, J.M.V. 1993. The wings of pterosaurs. American Journal of Science A 293: 91–166. [CrossRef]
• Palmer, C. and Dyke, G.J. 2009. Biomechanics of the unique pterosaur pteroid. Proceedings of the Royal Society B 277 (1684): 1121–1127. [CrossRef]
• Peters, D. 1995. Wing shape in pterosaurs.Nature 374: 315–316. [CrossRef]
• Peters, D. 2001. A new model for the evolution of the pterosaur wing—with a twist. Historical Biology 15: 277–301. [CrossRef]
• Rayner, J.M.V. 1990. Vertebrate flight and the origins of flying vertebrates. In: K.C. Allen and D.E.G. Briggs (eds.), Evolution and the Fossil Record, 188–217. Belhaven, London.
• Schaller, D. 1985. Wing evolution. In: M.K. Hecht, J.H. Ostrom, G. Viohl, and P. Wellnhofer (eds.), The Beginning of Birds, 333–348. Proceedings of the International Archeaopteryx Conference Eichstätt, Eichstätt: Freundes Jura−Museums, Eichstätt.
• Seeley, H.G. 1901. Dragons of the Air: an Account of Extinct Flying Reptiles. 239 pp. Methuen, London.
• Sharov, A.G. 1971. New flying reptiles from the Mesozoic of Kazakhstan and Kirgizia [in Russian]. Trudy Palaeontologičeskogo Instituta Akademii Nauk SSSR 130: 104–113.
• Sömmerring, S.T. von 1812. Über einen Ornithocephalus oder über das unbekannten Thier der Vorwelt, dessen Fossiles Gerippe Collini im 5. Bande der Actorum Academiae Theodoro−Palatinae nebst einer Abbildung in natürlicher Grösse im Jahre 1784 beschrieb, und welches Gerippe sich gegenwärtig in der Naturalien−Sammlung der königlichen Akademie der Wissenschaften zu München befindet. Denkschriften der königlichen bayerischen Akademie der Wissenschaften, München: mathematisch-physikalische Classe 3: 89–158.
• Stein, R.S. 1975. Dynamic analysis of Pteranodon ingens: a reptilian adaptation to flight. Journal of Paleontology 49: 534–548.
• Stieler, C. 1922. Neuer Rekonstruktionsversuch eines liassischen Flugsauriers. Naturwissenschaftliche Wochenschrift 20: 273–280.
• Strömer, E. 1913. Rekonstruktion des Flugsauriers Rhamphorhynchus gemmingi H. v. M. Neues Jahrbuch für Minerologie, Geologie und Paläontologie 2: 49–68.
• Taquet, P. and Padian, K. 2004. The earliest known restoration of a pterosaur and the philosophical origins of Cuvier’s Ossemen Fossiles. Comptes Rendus Palevol 3: 157–175. [CrossRef]
• Tischlinger, H. and Frey, E. 2002. Ein Rhamphorhynchus (Pterosauria, Reptlia) ungewöhnlicher Flughauterhalrung aus dem Solnhofener Plattenkalk. Archaeopteryx 20: 1–20.
• Unwin, D.M. 1987. Pterosaur Locomotion—joggers or waddlers? Nature 327: 13–14. [CrossRef]
• Unwin, D.M. 1988. A new specimen of Dimorphodon and the terrestrial ability of early pterosaurs. Modern Geology 13: 57–68.
• Unwin, D. 1999. Pterosaurs: back to the traditional model? Trends in Ecology and Evolution 14: 263–268. [CrossRef]
• Unwin, D.M. 2003. On the phylogeny and evolutionary history of pterosaurs. In: E. Buffetaut and J.−M. Mazin (eds.), Evolution and Palaeobiology of Pterosaurs. Geological Society of London, Special Publications 217: 139–190.
• Unwin, D.M. and Bakhurina, N.N. 1994. Sordes pilosus and the nature of the pterosaur flight apparatus. Nature 371: 62–64. [CrossRef]
• Unwin, D.M. and Bakhurina, N.N. 1995. Wing shape in pterosaurs. Nature 374: 315–316. [CrossRef]
• Unwin, D.M and Martill, D.M. 2007. Pterosaurs of the Crato Formation. In: D.M. Martill, G. Bechly, and R.F. Loveridge (eds.), The Crato Fossil Beds of Brazil, 475–524. Cambridge University Press, Cambridge.
• Wang, X., Zhou, Z., Zhang, F., and Xu, X. 2002. A nearly completely articulated rhamphorhynchoid pterosaur with exceptionally well−preserved wing membranes and “hairs” from Inner Mongolia, northeast China. Chinese Science Bulletin 47: 226–230. [CrossRef]
• Wang, X., Kellner, A.W.A., Zhou, Z., and Campos, A. 2005. Pterosaur diversity and faunal turnover in Cretaceous terrestrial ecosystems in China. Nature 437: 875–879. [CrossRef]
• Wang, X., Kellner, A.W.A., Jiang, S., and Meng, X. 2009. An unusual long−tailed pterosaur with elongated neck from western Liaoning of China. Anais da Academia Brasileira de Ciências 89: 793–812.
• Wellnhofer, P. 1970. Die Pterodactyloidea (Pterosauria) der Oberjura-Plattenkalke Süddeutschlands. Abhandlung der Bayerischen Akademie der Wissenschaften 141: 1–133.
• Wellnhofer, P. 1988. Terrestrial locomotion in pterosaurs. Historical Biology 1: 3–16. [CrossRef]
• Wellnhofer, P. 1991. Weitere pterosaurierfunde aus der Santana−Formation (Apt) der Chapada do Araripe, Brasilien. Palaeontographica Abteilung A 215: 43–101.
• Wilkinson, M.T. 2008. Three−dimensional geometry of a pterosaur wing skeleton, and its implications for aerial and terrestrial locomotion. Zoological Journal of the Linnean Society 154: 27–69. [CrossRef]
• Wilkinson, M.T., Unwin, D.M., and Ellington, C.P. 2005. Lift capability of pterosaur wings and the evolution of gigantism. Proceedings of the Royal Society B, Biological Sciences 273: 119–126. [CrossRef]
• Williston, S.W. 1897. Restoration of Ornithostoma (Pteranodon). Kansas University Quarterly 6: 35–51.
• Williston, S.C. 1911. The wing−finger of pterodactyls, with restoration of Nyctosaurus. Journal of Geology 19: 696–705. [CrossRef]
• Wiman, C. 1923. Über Dorygnathus und andere Flugsaurier. Bulletin of the Geological Institute of Upsala 19: 23–55.
• Wiman, C. 1925. Über Pterodactylus westimani und andere Flugsaurier. Bulletin of the Geological Institution of the University of Uppsala 20: 1–38.
• Witton, M.P. 2008. A new approach to determining pterosaur body mass and its implications for pterosaur flight. Zitteliana B 28: 143–158.
• Zhang, F., Zhou, Z., and Hou, L. 2007. Birds. In: M.−M. Chang, P. Chen, Y. Wang, and Y. Wang (eds.), The Jehol Fossils, the Emergence of feathered Dinosaurs, Beaked Birds and Flowering Plants, 128–149. Academic Press, New York.
• Zittel, K.A. Von 1882. Über Flugsaurier aus dem lithographischen Schiefer Bayerns. Paläontographica 29: 47–80.