Structure and function of hindlimb feathers in Archaeopteryx lithographica

Abstract This study examines the morphology and function of hindlimb plumage in Archaeopteryx lithographica. Feathers cover the legs of the Berlin specimen, extending from the cranial surface of the tibia and the caudal margins of both tibia and femur. These feathers exhibit features of flight feathers rather than contour feathers, including vane asymmetry, curved shafts, and a self-stabilizing overlap pattern. Many of these features facilitate lift generation in the wings and tail of birds, suggesting that the hindlimbs acted as airfoils. A new reconstruction of Archaeopteryx is presented, in which the hindlimbs form approximately 12% of total airfoil area. Depending upon their orientation, the hindlimbs could have reduced stall speed by up to 6% and turning radius by up to 12%. Presence of the “four-winged” planform in both Archaeopteryx and basal Dromaeosauridae indicates that their common ancestor used fore- and hindlimbs to generate lift. This finding suggests that arboreal parachuting and gliding preceded the evolution of avian flight.

[1]  Hans-Joachim Gregor,et al.  The beginnings of birds: Proceedings of the International Archaeopteryx Conference, Eichstätt, 1984. M.K. Hecht, J.H. Ostrom, G. Viohl and P. Wellnhofer (Editors). Jura Museum, Eichstätt, 382 pp. DM 90.00 , 1988 .

[2]  Hishida Natsuo,et al.  Technical notes : National Advisory committee for aeronautics , 1942 .

[3]  Jimmy A. McGuire,et al.  Allometric Prediction of Locomotor Performance: An Example from Southeast Asian Flying Lizards , 2003, The American Naturalist.

[4]  F. Novas New information on the systematics and postcranial skeleton of Herrerasaurus ischigualastensis (Ther , 1994 .

[5]  D. Yalden What size was Archaeopteryx , 1984 .

[6]  M. Norell,et al.  The distribution of integumentary structures in a feathered dinosaur , 2001, Nature.

[7]  J. Cracraft The functional morphology of the hind limb of the domestic pigeon, Columba livia. Bulletin of the AMNH ; v. 144, article 3 , 1971 .

[8]  P. Currie,et al.  A new carnosaur (Dinosauria, Theropoda) from the Jurassic of Xinjiang, People's Republic of China , 1993 .

[9]  Colin J Pennycuick,et al.  Mechanical constraints on the evolution of flight , 1986 .

[10]  A. Feduccia,et al.  Feathers of Archaeopteryx: Asymmetric Vanes Indicate Aerodynamic Function , 1979, Science.

[11]  P. Currie,et al.  A new oviraptorosaur [Dinosauria, Theropoda] from Mongolia: the first dinosaur with a pygostyle , 2000 .

[12]  W. J. Bock Ein dritter Archaeopteryx-Fund aus den Solnhofener Plattenkalken von Langenaltheim/Mfr. Florian Heller , 1960 .

[13]  Adrian L. R. Thomas WHY DO BIRDS HAVE TAILS ? THE TAIL AS A DRAG REDUCING FLAP, AND TRIM CONTROL , 1996 .

[14]  Zhonghe Zhou,et al.  Palaeontology: Leg feathers in an Early Cretaceous bird , 2004, Nature.

[15]  S. Ullman,et al.  Neuroscience: Rewiring the adult brain , 2005, Nature.

[16]  J. Anderson,et al.  Fundamentals of Aerodynamics , 1984 .

[17]  L. Chiappe,et al.  The origin of birds and their flight. , 1998, Scientific American.

[18]  鈴木 久仁博 最古の化石鳥類,Archaeopteryx lithographicaの相対生長的計測(地球科学の窓) , 1991 .

[19]  R. V. Mises Theory of Flight , 1959 .

[20]  Ulla M. Norberg,et al.  Evolution of Vertebrate Flight: An Aerodynamic Model for the Transition from Gliding to Active Flight , 1985, The American Naturalist.

[21]  B. Demes,et al.  They seem to glide. Are there aerodynamic effects in leaping prosimian primates? , 1991, Zeitschrift fur Morphologie und Anthropologie.

[22]  Matthew R Evans,et al.  Birds' tails do act like delta wings but delta-wing theory does not always predict the forces they generate , 2003, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[23]  C. Pennycuick Power requirements for horizontal flight in the pigeon Columba livia , 1968 .

[24]  Ilan Kroo,et al.  DRAG DUE TO LIFT: Concepts for Prediction and Reduction , 2001 .

[25]  Bruce A. Young,et al.  On a Flap and a Foot: Aerial Locomotion in the “Flying” Gecko, Ptychozoon kuhli , 2002 .

[26]  A. Feduccia The origin and evolution of birds , 1996 .

[27]  Central Asiatic Expeditions,et al.  On the dinosaurian fauna of the Iren Dabasu Formation. Bulletin of the AMNH ; v. 67, article 2 , 1933 .

[28]  A Tetrapteryx stage in the ancestry of birds , 1915 .

[29]  S. Gatesy,et al.  FROM FROND TO FAN: ARCHAEOPTERYX AND THE EVOLUTION OF SHORT‐TAILED BIRDS , 1996, Evolution; international journal of organic evolution.

[30]  Richard L Essner Three-dimensional launch kinematics in leaping, parachuting and gliding squirrels. , 2002, The Journal of experimental biology.

[31]  M. Norell,et al.  Palaeontology: 'Modern' feathers on a non-avian dinosaur , 2002, Nature.

[32]  O. Abel Grundzüge der Palaeobiologie der Wirbeltiere , 1912 .

[33]  John R. Hutchinson,et al.  Adductors, abductors, and the evolution of archosaur locomotion , 2000, Paleobiology.

[34]  L. Prandtl Induced drag of multiplanes , 1924 .

[35]  M. M. Stewart Arboreal habitat use and parachuting by a subtropical forest frog , 1985 .

[36]  A Hedenström,et al.  Gliding flight in a jackdaw: a wind tunnel study. , 2001, The Journal of experimental biology.

[37]  Russell P. Balda,et al.  The Physics of Leaping Animals and the Evolution of Preflight , 1983, The American Naturalist.

[38]  Fucheng Zhang,et al.  A new maniraptoran dinosaur from China with long feathers on the metatarsus , 2005, Naturwissenschaften.

[39]  C. Pennycuick A wind tunnel study of gliding flight in the pigeon Columba livia , 1968 .

[40]  Adrian L. R. Thomas,et al.  On the origins of birds: the sequence of character acquisition in the evolution of avian flight , 1999, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[41]  J. Speakman,et al.  Flight capabilities of Archaeopteryx , 1994, Nature.

[42]  G. Dyke,et al.  Origin of birds , 2003 .

[43]  James A. Oliver,et al.  "Gliding" in Amphibians and Reptiles, with a Remark on an Arboreal Adaptation in the Lizard, Anolis carolinensis carolinensis Voigt , 1951, The American Naturalist.

[44]  Zhonghe Zhou,et al.  Four-winged dinosaurs from China , 2003, Nature.

[45]  R. Prum Palaeontology: Dinosaurs take to the air , 2003, Nature.

[46]  Origin of flight: Could ‘four-winged’ dinosaurs fly? (Reply) , 2005, Nature.

[47]  Zhonghe Zhou,et al.  Early diversification of birds: Evidence from a new opposite bird , 2001 .

[48]  K. Dial Wing-Assisted Incline Running and the Evolution of Flight , 2003, Science.

[49]  Colin J Pennycuick,et al.  Control of gliding angle in Rüppell's Griffon Vulture Gyps rüppellii , 1971 .

[50]  Ostrom Jh,et al.  Bird flight: how did it begin? , 1979, American scientist.

[51]  Max M Munk,et al.  General Biplane Theory , 1923 .

[52]  R. Åke Norberg,et al.  Delta-wing function of webbed feet gives hydrodynamic lift for swimming propulsion in birds , 2003, Nature.

[53]  N. Pierce Origin of Species , 1914, Nature.

[54]  Adrian L. R. Thomas On the aerodynamics of birds’ tails , 1993 .

[55]  O. C. Marsh,et al.  Odontornithes; a Monograph of the Extinct Toothed Birds of North America , 1880, Nature.

[56]  L. Johansson,et al.  Lift-based paddling in diving grebe. , 2001, The Journal of experimental biology.

[57]  J. H. Ostrom Bird flight: how did it begin? , 1979, American scientist.

[58]  A. Russell The origin of parachuting locomotion in gekkonid lizards (Reptilia: Gekkonidae) , 1979 .

[59]  M. Koehl,et al.  THE INTERACTION OF BEHAVIORAL AND MORPHOLOGICAL CHANGE IN THE EVOLUTION OF A NOVEL LOCOMOTOR TYPE: “FLYING” FROGS , 1990, Evolution; international journal of organic evolution.

[60]  C. A. Walker New subclass of birds from the Cretaceous of South America , 1981, Nature.

[61]  P. Christiansen,et al.  Body plumage in Archaeopteryx: a review, and new evidence from the Berlin specimen , 2004 .

[62]  P. Olsen,et al.  Ratite Footprints and the Stance and Gait of Mesozoic Theropods , 2007 .

[63]  John H. Ostrom,et al.  The Origin of Birds , 1926 .

[64]  A. M. Lucas,et al.  Avian anatomy : integument , 1974 .

[65]  Zhonghe Zhou,et al.  The smallest known non-avian theropod dinosaur , 2000, Nature.

[66]  C. Gans,et al.  Sharovipteryx, a reptilian glider? , 1987, Paleobiology.