Skip to main content
×
×
Home

New heterodontosaurid remains from the Cañadón Asfalto Formation: cursoriality and the functional importance of the pes in small heterodontosaurids

  • Marcos G. Becerra (a1), Diego Pol (a1), Oliver W.M. Rauhut (a2) and Ignacio A. Cerda (a3)
Abstract

New ornithischian remains reported here (MPEF-PV 3826) include two complete metatarsi with associated phalanges and caudal vertebrae, from the late Toarcian levels of the Cañadón Asfalto Formation. We conclude that these fossil remains represent a bipedal heterodontosaurid but lack diagnostic characters to identify them at the species level, although they probably represent remains of Manidens condorensis, known from the same locality. Histological features suggest a subadult ontogenetic stage for the individual. A cluster analysis based on pedal measurements identifies similarities of this specimen with heterodontosaurid taxa and the inclusion of the new material in a phylogenetic analysis with expanded character sampling on pedal remains confirms the described specimen as a heterodontosaurid. Finally, uncommon features of the digits (length proportions among nonungual phalanges of digit III, and claw features) are also quantitatively compared to several ornithischians, theropods, and birds, suggesting that this may represent a bipedal cursorial heterodontosaurid with gracile and grasping feet and long digits. In particular, the elongated non-terminal pedal phalanges and morphology of digit III resemble features present in arboreal birds, a unique condition found so far among ornithischians.

Copyright
References
Hide All
Apaldetti, C., Pol, D., and Yates, A., 2013, The postcranial anatomy of Coloradisaurus brevis (Dinosauria: Sauropodomorpha) from the Late Triassic of Argentina and its phylogenetic implications: Palaeontology, v. 56, p. 277301.
Baumel, J.J., and Witmer, L.M., 1993, Osteologia, in Baumel, J.J., King, A.S., Brazile, J.E., Evans, H.E., and Vanden Berge, J.C., eds., Handbook of Avian Anatomy: Nomina Anatomica Avium, 2d ed.: Cambridge, UK, Nuttall Ornithological Society, p. 45132.
Becerra, M.G., Pol, D., Marsicano, C.A., and Rauhut, O.W.M., 2014, The dentition of Manidens condorensis (Ornithischia; Heterodontosauridae) from the Jurassic Cañadón Asfalto Formation of Patagonia: morphology, heterodonty and the use of statistical methods for identifying isolated teeth: Historical Biology, v. 26, p. 480492.
Birn-Jeffery, A.V., Miller, C.E., Naish, D., Rayfield, E.J., and Hone, D.W.E., 2012, Pedal claw curvature in birds, lizards and Mesozoic dinosaurs – complicated categories and compensating for mass-specific and phylogenetic control: PLoS One, v. 7, p. e50555.
Bonaparte, J.F., 1976, Pisanosaurus mertii Casamiquela and the origin of the Ornithischia: Journal of Paleontology, v. 50, p. 808820.
Bonaparte, J.F., 1978, Coloradia brevis n. g. et n. sp. (Saurischia-Prosauropoda), dinosaurio Plateosauridae de la Formación Los Colorados, Triásico superior de La Rioja, Argentina: Ameghiniana, v. 15, p. 327332.
Boulenger, G.A., 1881, Iguanodon bernissartensis, in van Beneden, P.-J., ed., Sur l’arc pelvien chez les dinosauriens de Bernissart: Bulletins de l´Académie Royale des Sciences, des Lettres et des Beaux-Arts de Belgique, Classe des Sciences, v. 1, p. 600606.
Brochu, C.A., 1996, Closure of neurocentral sutures during crocodilian ontogeny: implications for maturity assessment in fossil archosaurs: Journal of Vertebrate Paleontology, v. 16, p. 4962.
Burnham, D.A., Feduccia, A., Martin, L.D., and Falk, A.R., 2011, Tree climbing-a fundamental avian adaptation: Journal of Systematic Palaeontology, v. 9, p. 103107.
Butler, R.J., 2005, The ‘fabrosaurid’ ornithischian dinosaurs of the Upper Elliot Formation (Lower Jurassic) of South Africa and Lesotho: Zoological Journal of the Linnean Society, v. 145, p. 175218.
Butler, R.J., Smith, R.M.H., and Norman, D.B., 2007, A primitive ornithischian dinosaur from the Late Triassic of South Africa, and the early evolution and diversification of Ornithischia: Proceedings of the Royal Society B, v. 274, p. 20412046.
Butler, R.J., Upchurch, P., and Norman, D.B., 2008, The phylogeny of the ornithischian dinosaurs: Journal of Systematic Palaeontology, v. 6, p. 140.
Butler, R.J., Porro, L.B., Galton, P.M., Chiappe, L.M., Henderson, D.M., and Erickson, G.M., 2010, Lower limits of ornithischian dinosaur body size inferred from a new Upper Jurassic heterodontosaurid from North America: Proceedings of the Royal Society B, v. 277, p. 375381.
Butler, R.J., Liyong, J., Jun, C., and Godefroit, P., 2011, The postcranial osteology and phylogenetic position of the small ornithischian dinosaur Changchunsaurus parvus from the Quantou Formation (Cretaceous: Aptian-Cenomanian) of Jilin province, north-eastern China: Palaeontology, v. 54, p. 667683.
Butler, R.J., Porro, L.B., Galton, P.M., and Chiappe, L.M., 2012, Anatomy and cranial functional morphology of the small-bodied dinosaur Fruitadens haagarorum from the Upper Jurassic of the USA: PLoS One, v. 7, p. e31556.
Cabaleri, N., Volkheimer, W., Armella, C., Gallego, O., Silva Nieto, D., Páez, M., Cagnoni, M., Ramos, A., Panarello, H., and Koukharsky, M., 2010, Estratigrafía, análisis de facies y paleoambientes de la Formación Cañadón Asfalto en el depocentro jurásico Cerro Cóndor, provincia del Chubut: Revista de la Asociación Geológica Argentina, v. 66, p. 349367.
Casamiquela, R.M., 1967, Un nuevo dinosaurio ornitisquio Triásico (Pisanosaurus mertii: Ornithopoda) de la Formación Ischigualasto, Argentina: Ameghiniana, v. 4, p. 4764.
Carpenter, K., 2004, Redescription of Ankylosaurus magniventris Brown 1908 (Ankylosauridae) from the Upper Cretaceous of the Western Interior of North America: Canadian Journal of Earth and Science, v. 41, p. 961986.
Cerda, I.A., and Chinsamy, A., 2012, Biological implications of the bone microstructure of the Late Cretaceous ornithopod dinosaur Gasparinisaura cincosaltensis : Journal of Vertebrate Paleontology, v. 32, p. 355368.
Chinsamy, A., 1995, Ontogenetic changes in the bone histology of the Late Jurassic ornithopod Dryosaurus lettowvorbecki : Journal of Vertebrate Paleontology, v. 15, p 96104.
Chinsamy, A., and Raath, M.A., 1992, Preparation of fossil bone for histological examination: Paleontologia Africana, v. 29, p. 3944.
Chinsamy-Turan, A., 2005, The microstructure of dinosaur bone, p. deciphering biology with fine-scale techniques: Johns Hopkins University Press, Baltimore and London, 195 p.
Clark, J.M., Hopson, J.A., Hernandez, R., Fastovsky, D.E., and Montellano, M., 1998, Foot posture in a primitive pterosaur: Nature, v. 391, p. 886889.
Colbert, E.H., 1981, A primitive ornithischian dinosaur from the Kayenta Formation of Arizona: Museum of Northern Arizona Press, Bulletin Series, v. 53, p. 161.
Coombs, W.P. Jr., 1978, Theoretical aspects of cursorial adaptations in dinosaurs: The Quarterly Review of Biology, v. 53, p. 393418.
Coria, R.A., and Calvo, J.O., 2002, A new iguanodontian ornithopod from Neuquén Basin, Patagonia, Argentina: Journal of Vertebrate Paleontology, v. 22, p. 503509.
Coria, R.A., and Salgado, L., 1996, A basal iguanodontian (Ornithischia: Ornithopoda) from the Late Cretaceous of South America: Journal of Vertebrate Paleontology, v. 16, p. 445457.
Crompton, A.W., and Charig, A.J., 1962, A new ornithischian from the Upper Triassic of South Africa: Nature, v. 196, p. 10741077.
Cuneo, R., Ramezani, J., Scasso, R., Pol, D., Escapa, I., Zavattieri, A.M., and Bowring, S.A., 2013, High-precision U-Pb geochronology and a new chronostratigraphy for the Cañadón Asfalto Basin, Chubut, central Patagonia: Implications for terrestrial faunal and floral evolution in Jurassic: Gondwana Research, v. 24, p. 12671275.
Czerkas, S.A., and Yuan, C., 2002, An arboreal maniraptoran from northeast China, p. 63–95. In S. J. Czerkas (ed.), The Dinosaur Museum Journal 1, Feathered dinosaurs and the origin of flight: Blanding, UT.
Di Rienzo, J.A., Casanoves, F., Balzarini, M.G., Gonzalez, L., Tablada, M., and Robledo, C.W., 2011, InfoStat versión 2011: Grupo InfoStat, FCA, Universidad Nacional de Córdoba, Argentina.
Dodson, P., Forster, C.A., and Sampson, S.D., 2004, Ceratopsidae, in Weishampel, D.B., Dodson, P., and Osmólska, H., eds., The Dinosauria, 2d ed.: Berkeley, University of California Press, p. 494514.
Dong, Z., 1978, A new genus of Pachycephalosauria from Laiyang, Shantung: Vertebrata PalAsiatica, v. 16, p. 225228. [in Chinese]
Dong, Z., 1989, On a small ornithopod (Gongbusaurus wucaiwanensis sp. nov.) from Kelamaili Junggar Basin, Xinjiang, China]: Vertebrata PalAsiatica, v. 27, p. 140146. [in Chinese]
Dong, Z., and Azuma, Y., 1997, On a primitive neoceratopsian from the Early Cretaceous, in Dong, Z.-M., ed: Sino-Japanese Silk Road Dinosaur Expedition, Beijing, China Ocean Press, p. 6889.
Escapa, I.H., Sterli, J., Pol, D., and Nicoli, L., 2008, Jurassic tetrapods and flora of Cañadón Asfalto Formation in Cerro Condor area, Chubut Province: Revista de la Asociación Geológica Argentina, v. 63, p. 613624.
Farris, J.S., 1981, Distance data in phylogenetic analysis, in Funk, V.A., and Brooks, D.R., eds., Advances in Cladistics, vol I: Proceedings of the First Meeting of the Willi Henning Society, New York Botanical Garden, New York. p. 1–23.
Feduccia, A., 1993, Evidence from claw geometry indicating arboreal habits of Archaeopteryx : Science, v. 259, p. 790793.
Fisher, H.I., 1946, Adaptations and comparative anatomy of the locomotor apparatus of New World vultures: American Midland Naturalist, v. 35, p. 545727.
Forster, C.A., Sampson, S.D., Chiappe, L.M., and Krause, D.W., 1998, The theropod ancestry of birds: New evidence from the Late Cretaceous of Madagascar: Science, v. 279, p. 19151919.
Fracillon-Vieillont, H., Debuffrénil, V., Castanet, J., Geraudie, J., Meurier, F.J., Sire, J.Y., Zylberberg, L., and De Ricqlés, A., 1990, Microstructure and mineralization of vertebrate skeletal tissues, in Carter, J.G., ed., Skeletal Biomineralization: Patterns, Processes and Evolutionary Trends, volume 1, New York, Van Nostrand Reinhold, p. 471530.
Galton, P.M., 1974a, Notes on Thescelosaurus, a conservative ornithopod dinosaur from the Upper Cretaceous of North America, with comments on ornithopod classification: Journal of Paleontology, v. 48, p. 10481067.
Galton, P.M., 1974b, The ornithischian dinosaur Hypsilophodon from the Wealden of the Isle of Wight: Bulletin of the British Museum (Natural History) Geology, v. 25, p. 1152.
Galton, P.M., 1978, Fabrosauridae, the basal family of ornithischian dinosaurs (Reptilia: Ornithopoda): Paläontologische Zeitschrift, v. 52, p. 138159.
Galton, P.M., 1981, Dryosaurus, a hypsilophodontid dinosaur from the Upper Jurassic of North America and Africa: postcranial skeleton: Paläontologische Zeitschrift, v. 55, p. 271312.
Galton, P.M., 2006, Teeth of ornithischian dinosaurs (mostly Ornithopoda) from the Morrison Formation (Upper Jurassic) of Western United States, in Carpenter, K., ed., Horns and Beaks, Ceratopsian and Ornithopod Dinosaurs, Bloomington, Indiana University Press, p. 1747.
Galton, P.M., and Jensen, J.A., 1973, Skeleton of a hypsilophodontid dinosaur (Nanosaurus (?) rex) from the Upper Jurassic of Utah: Brigham Young University Geology Studies, v. 20, p. 137157.
Galton, P.M., and Powell, H.P., 1980, The ornithischian dinosaur Camptosaurus prestwichii from the Upper Jurassic of England: Palaeontology, v. 23, p. 411443.
Galton, P.M., and Upchurch, P., 2004a, Prosauropoda, in Weishampel, D.B., Dodson, P., and Osmólska, H., eds., The Dinosauria, 2d ed.: Berkeley, University of California Press, p 393412.
Galton, P.M., and Upchurch, P., 2004b, Stegosauridae, in Weishampel, D.B., Dodson, P., and Osmólska, H., eds., The Dinosauria, 2d ed.: Berkeley, University of California Press, p. 343362.
Gilmore, C.W., 1909, Osteology of the Jurassic reptile Camptosaurus, with a revision of the species of the genus and a description of two new species: Proceedings of the U.S. National Museum, v. 36, p. 197332.
Gilmore, C.W., 1913, A new dinosaur from the Lance Formation of Wyoming: Smithsonian Miscellaneous Collection, v. 61, p. 15.
Gilmore, C.W., 1914, Osteology of the armored dinosaurs in the United States National Museum, with special reference to the genus Stegosaurus : Bulletin of the U.S. National Museum, v. 89, p. 1136.
Gilmore, C.W., 1915, Osteology of Thescelosaurus, an orthopodous dinosaur from the Lance Formation of Wyoming: Proceedings of the U.S. National Museum, v. 49, p. 591616.
Goloboff, P.A., Farris, J.S., and Nixon, K.C., 2008a, TNT, a free program for phylogenetic analysis: Cladistics, v. 24, p. 113.
Goloboff, P.A., Farris, J.S., and Nixon, K.C., 2008b, TNT (Tree analysis using new technology) ver. 1.1.: Published by the authors, Tucumán, Argentina.
Gow, C.E., 1975, A new heterodontosaurid from the Red Beds of South Africa showing clear evidence of tooth replacement: Zoological Journal of the Linnean Society, v. 57, p. 335339.
Hammer, O., Harper, D.A.T., and Ryan, P.D., 2001, PAST: Paleontological Statistics software package for education and data analysis: Palaeontologica Electronica, v. 4, 9 p.
Han, F.L., Barrett, P.M., Butler, R.J., and Xu, X., 2012, Postcranial anatomy of Jeholosaurus shangyuanensis (Dinosauria, Ornithischia) from the Lower Cretaceous Yixian Formation of China: Journal of Vertebrate Paleontology, v. 32, p. 13701395.
Haughton, H., 1924, The fauna and stratigraphy of the Stormberg Series: Annals of the South African Museum, v. 12, p. 323497.
Hayashi, S., Carpenter, K., and Suzuki, D., 2009, Different growth patterns between the skeleton and osteoderms of Stegosaurus (Ornithischia: Thyreophora): Journal of Vertebrate Paleontology, v. 29, p. 123131.
He, X., 1979, A newly discovered ornithopod dinosaur-Yandusaurus from Zigong, China, in Contributions to International Exchange of Geology, Part 2, Stratigraphy and Palaeontology: Beijing, Geological Publishing House, p. 116–123. [in Chinese]
Holtz, T.R. Jr., 1994, The arctometatarsalian pes, an unusual structure of Cretaceous Theropoda (Dinosauria: Saurischia): Journal of Vertebrate Paleontology, v. 14, p. 408519.
Hooley, R.W., 1925, On the skeleton of Iguanodon atherfieldensis sp. nov., from the Wealden Shales of Atherfield (Isle of Wight): Quarterly Journal of the Geological Society, v. 81, p. 161.
Hopson, J.A., 1975, On the generic separation of the ornithischian dinosaurs Lycorhinus and Heterodontosaurus from the Stormberg Series (Upper Triassic) of South Africa: South African Journal of Science, v. 71, p. 302305.
Hopson, J.A., 2001, Ecomorphology of avian and theropod phalangeal proportions, p. implications for the arboreal versus terrestrial origin of bird flight, in Gauthier, J., and Gall, L.F., eds., New Perspectives on the Origin and Early Evolution of Birds: Special Publication of the Peabody Museum of Natural History, Yale University, p. 211235.
Horner, J.R., and Makela, R., 1979, Nest of juveniles provides evidence of family structure among dinosaurs: Nature, v. 282, p. 296298.
Horner, J.R., and Weishampel, D.B., 1988, A comparative embryological study of two ornithischian dinosaurs: Nature, v. 332, p. 256257.
Horner, J.R., De Ricqlés, A., and Padian, K., 2000, Long bone histology of the hadrosaurid dinosaur Maiasaura peeblesorum: growth dynamics and physiology based on an ontogenetic series of skeletal elements: Journal of Vertebrate Paleontology, v. 20, p. 115129.
Horner, J.R., Weishampel, D.B., and Forster, C.A., 2004, Hadrosauridae, in Weishampel, D.B., Dodson, P., and Osmólska, H., eds., The Dinosauria, 2d ed.: Berkeley, University of California Press, p. 438463.
Horner, J.R., De Ricqlés, A., Padian, K., and Scheetz, R.D., 2009, Comparative long bone histology and growth of the ‘hypsilophodontid’ dinosaurs Orodromeus makelai, Dryosaurus altus, and Tenontosaurus tilletii (Ornithischia: Euornithopoda): Journal of Vertebrate Paleontology, v. 29, p. 734747.
Hou, L., 1977, A new primitive Pachycephalosauria from Anhui, China: Vertebrata PalAsiatica, v. 15, p. 198202. [in Chinese]
Hübner, T.R., 2012, Bone histology in Dysalotosaurus lettowvorbecki (Ornithischia: Iguanodontia)-variation, growth, and implications: PLoS One, v. 7, e29958.
Huene, F.V., 1908, Die Dinosaurier der europäischen Triasformation mit Berücksichtigung der aussereuropäischen Vorkommnisse: Geologische und Palaeontologische Abhandlungen, Supplement, v. 1, p. 1419.
Hutchinson, J.R., 2002, The evolution of hindlimb tendons and muscles on the line to crown-group birds: Comparative Biochemistry and Physiology Part A, v. 133, p. 10511086.
Huxley, T.H., 1869, On Hypsilophodon, a new genus of Dinosauria: Abstracts of the Proceedings of the Geological Society of London, v. 204, p. 34.
Irmis, R.B., 2007, Axial skeleton ontogeny in the Parasuchia (Archosauria: Pseudosuchia) and its implications for ontogenetic determination in Archosaurs: Journal of Vertebrate Paleontology, v. 27, p. 350361.
Irmis, R.B., Parker, W.G., Nesbitt, S.J., and Liu, J., 2007, Ornithischian dinosaurs: the Triassic record: Historical Biology, v. 18, p. 322.
Kambic, R.E., 2008, Multivariate analysis of avian and non-avian theropod pedal phalanges, Unpublished Ph.D. dissertation: Bozeman, Montana State University, 90 p.
Langer, M.C., 2004, Basal Saurischia, in Weishampel, D.B., Dodson, P., and Osmólska, H., eds., The Dinosauria, 2d ed.: Berkeley, University of California Press, p 2546.
Longrich, N., 2010, The function of large eyes in Protoceratops: a nocturnal ceratopsian?, in Ryan, M.J., Chinnery-Allgeier, B.J., and Eberth, D.A., eds., New Perspectives on Horned Dinosaurs, The Royal Tyrrell Museum Ceratopsian Symposium, Bloomington and Indianapolis, Indiana University Press, p. 308327.
Makovicky, P.J., and Norell, M.A., 2004, Troodontidae, in Weishampel, D.B., Dodson, P., and Osmólska, H., eds., The Dinosauria, 2d ed.: Berkeley, University of California Press, p. 184195.
Mantell, G.A., 1825, Notice on the Iguanodon, a newly discovered fossil reptile, from the sandstone of Tilgate Forest, in Sussex: Philosophical Transactions of the Royal Society of London, v. 115, p. 179186.
Marsh, O.C., 1878, Principal characters of American Jurassic dinosaurs. Part I: American Journal of Science, v. 16, p. 411416.
Marsh, O.C., 1879, Notice of new Jurassic reptiles: American Journal of Science, v. 18, p. 501505.
Marsh, O.C., 1885, Names of extinct reptiles: American Journal of Science, v. 29, p. 169.
Marsh, O.C., 1894, The typical Ornithopoda of the American Jurassic: American Journal of Science, v. 283, p. 8590.
Maryanska, T., and Osmólska, H., 1974, Pachycephalosauria, a new suborder of ornithischian dinosaurs: Palaeontologia. Polonica, v. 30, p. 45102.
Norell, M.A., and Makovicky, P.J., 2004, Dromaeosauridae, in Weishampel, D.B., Dodson, P., and Osmólska, H., eds., The Dinosauria, 2d ed.: Berkeley, University of California Press, p. 196209.
Norman, D.B., 1980, On the ornithischian dinosaur Iguanodon bernissartensis from Belgium: Mémoire de Institut Royal des Sciences Naturelles de Belgique, v. 178, p. 1103.
Norman, D.B., 2004, Basal Iguanodontia, in Weishampel, D.B., Dodson, P., and Osmólska, H., eds., The Dinosauria, 2d ed.: Berkeley, University of California Press, p. 413437.
Norman, D.B., Sues, H.-D., Witmer, L.M., and Coria, R.A., 2004, Basal Ornithopoda, in Weishampel, D.B., Dodson, P., and Osmólska, H., eds., The Dinosauria, 2d ed.: Berkeley, University of California Press, p. 393412.
Novas, F.E., 1993, New information on the systematics and postcranial skeleton of Herrerasaurus ischigualastensis (Theropoda: Herrerasauridae) from the Ischigualasto Formation (Upper Triassic) of Argentina: Journal of Vertebrate Paleontology, v. 13, p. 400423.
Novas, F.E., Cambiaso, A., and Ambrosio, A., 2004, A new basal iguanodontian (Dinosauria, Ornithischia) from the Upper Cretaceous of Patagonia: Ameghiniana, v. 41, p. 7582.
Perle, A., Maryańska, T., and Osmólska, H., 1982, Goyocephale lattimoeri gen. et sp. n., a new flat-headed pachycephalosaur (Ornithischia, Dinosauria) from the Upper Cretaceous of Mongolia: Acta Palaeontologica Polonica, v. 27, p. 115127.
Osborn, H.F., 1923, Two Lower Cretaceous dinosaurs of Mongolia: American Museum Novitates, v. 95, p. 110.
Osborn, H.F., 1924, Psittacosaurus and Protiguanodon, p. Two Lower Cretaceous iguanodonts from Mongolia: American Museum Novitates, v. 127, p. 116.
Osmólska, H., 1981, Coossified tarsometatarsi in theropod dinosaurs and their bearing on the problem of bird origins: Acta Paleontologica Polonica, v. 42, p. 7995.
Osmólska, H., Currie, P.J., and Barsbold, R., 2004, Oviraptorosauria, in Weishampel, D.B., Dodson, P., and Osmólska, H., eds., The Dinosauria, 2d ed.: Berkeley, University of California Press, p. 165183.
Ostrom, J.H., 1969, Osteology of Deinonychus antirrhopus, an unusual theropod from the Lower Cretaceous of Montana: Bulletin of the Peabody Museum of Natural History, v. 30, p. 1165.
Ostrom, J.H., 1970, Stratigraphy and paleontology of the Cloverly Formation (Lower Cretaceous) of the Bighorn Basin area, Wyoming and Montana: Bulletin of the Peabody Museum of Natural History, v. 35, p. 1234.
Ostrom, J.H., 1974, Archaeopteryx and the origin of flight: The Quarterly Review of Biology, v. 49, p. 2747.
Owen, R., 1861, Monograph of the Fossil Reptilia of the Liassic formations. Part 1. A monograph of a fossil dinosaur (Scelidosaurus harrisonii Owen) of the Lower Lias: Palaeontographical Society Monographs, v. 1, p. 114.
Padian, K., 2004, Basal Avialae, in Weishampel, D.B., Dodson, P., and Osmólska, H., eds., The Dinosauria, 2d ed.: Berkeley, University of California Press, p. 196209.
Parks, W.A., 1926, Thescelosaurus warreni, a new species of ornithopodous dinosaur from the Edmonton Formation of Alberta: University of Toronto Studies (Geological Series), v. 21, p. 142.
Peng, G.-Z., 1990, A new species of small ornithopod from Zigong, Sichuan: Journal of the Zigong Dinosaur Museum, v. 2, p. 1927. [in Chinese]
Peng, G.-Z., 1992, Jurassic ornithopod Agilisaurus louderbacki (Ornithopoda: Fabrosauridae) from Zigong, Sichuan, China: Vertebrata PalAsiatica, v. 30, p. 3951. [in Chinese]
Pike, A.V.L., and Maitland, D.P., 2004, Scaling of bird claws: Journal of Zoology, v. 262, p. 7381.
Pol, D., Rauhut, O.W.M., and Becerra, M.G., 2011, A Middle Jurassic heterodontosaurid dinosaur from Patagonia and the evolution of heterodontosaurids: Naturwissenschaften, v. 98, p. 369379.
Porro, L.B., Butler, R.J., Barrett, P.M., Moore-Fay, S., and Abel, R.L., 2011, New heterodontosaurid specimens from the Lower Jurassic of southern Africa and the early ornithischian dinosaur radiation: Earth and Environmental Transactions of the Royal Society of Edinburgh, v. 101, p. 351366.
Redelstorff, R., Hübner, T.R., Chinsamy, A., and Sander, P.M., 2013, Bone histology of the stegosaur Kentrosaurus aethiopicus (Ornithischia: Thyreophora) from the Upper Jurassic of Tanzania: The Anatomical Record, v. 296, p. 933952.
Reid, R.E.H., 1996, Bone histology of the Cleveland-Lloyd dinosaurs and of dinosaurs in general, part I: introduction to bone tissues: Brigham Young University Geology Studies, v. 41, p. 2572.
Reig, O.A., 1963, La presencia de dinosaurios saurisquios en los “Estratos de Ischigualasto” (Mesotrifisico superior) de las provencias de San Juan y La Rioja (Republica Argentina): Ameghiniana, v. 3, p. 320.
Romer, A.S., 1972, The Chañares (Argentina) Triassic reptile fauna. XV. Further remains of the thecodonts Lagerpeton and Lagosuchus : Breviora, v. 394, p. 17.
Rougier, G.W., Garrido, A., Gaetano, L., Puerta, P., Corbitt, C., and Novacek, M.J., 2007a, A new triconodont from South America: American Museum Novitates, v. 3580, p. 117.
Rougier, G.W., Martinelli, A.G., Forasiepi, A.M., and Novacek, M.J., 2007b, New Jurassic mammals from Patagonia, Argentina: a reappraisal of australosphenidan morphology and interrelationships: American Museum Novitates, v. 3566, p. 154.
Salani, F.M., 2007, Aporte a la edad de la Formación Cañadón Asfalto, Chubut, Argentina: Resúmenes, 30 Simposio Argentino del Jurásico, p 71.
Salgado, L., Coria, R.A., and Heredia, S.E., 1997, New Materials of Gasparinisaura cincosaltensis (Ornithischia, Ornithopoda) from the Upper Cretaceous of Argentina: Journal of Paleontology, v. 71, p. 933940.
Santa Luca, A.P., 1980, The postcranial skeleton of Heterodontosaurus tucki (Reptilia, Ornithischia) from the Stormberg of South Africa: Annals of the South African Museum, v. 79, p. 159211.
Scheetz, R.D., 1999, Osteology of Orodromeus makelai and the phylogeny of basal ornithopod dinosaurs, unpublished Ph.D. dissertation: Bozeman, Montana State University, 186 p.
Sereno, P.C., 1986, Phylogeny of the bird-hipped dinosaurs (Order Ornithischia): National Geographic Research, v. 2, p. 234256.
Sereno, P.C., 1991, Lesothosaurus, “fabrosaurids,” and the early evolution of Ornithischia: Journal of Vertebrate Paleontology, v. 11, p. 168197.
Sereno, P.C., 1997, The origin and evolution of dinosaurs: Annual Review of Earth and Planetary Sciences, v. 25, p. 435489.
Sereno, P.C., 2012, Taxonomy, morphology, masticatory function and phylogeny of heterodontosaurid dinosaurs: ZooKeys, v. 226, p. 1225.
Sereno, P.C., and Arcucci, A.B., 1994, Dinosaurian precursors from the Middle Triassic of Argentina: Marasuchus lilloensis, gen. nov.: Journal of Vertebrate Paleontology, v. 14, p. 5373.
Sereno, P.C., and Rao, C., 1992, Early evolution of avian flight and perching, p. New evidence from the Lower Cretaceous of China: Science, v. 255, p. 845848.
Sternberg, C.M., 1937, Classification of Thescelosaurus: a description of a new species: Proceedings of the Geological Society of America, v. 1936, p. 375.
Thulborn, R.A., 1970, The systematic position of the Triassic ornithischian dinosaur Lycorhinus angustidens : Zoological Journal of the Linnean Society, v. 49, p. 235245.
Tykoski, R.S., and Rowe, T., 2004, Ceratosauria, in Weishampel, D.B., Dodson, P., and Osmólska, H., eds., The Dinosauria, 2d ed.: Berkeley, University of California Press, p. 4770.
Vickaryous, M.K., Maryańska, T., and Weishampel, D.B., 2004, Ankylosauria, in Weishampel, D.B., Dodson, P., and Osmólska, H., eds., The Dinosauria, 2d ed.: Berkeley, University of California Press, p. 363392.
Virchow, H., 1919, Atlas and Epistropheus bei den Schildkröten: Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin, v. 1919, p. 303332.
Volkheimer, W., Gallego, O.F., Cabaleri, N.G., Armella, C., Narváez, P.L., Silva Nieto, D.G., and Páez, M.A., 2009, Stratigraphy, palynology, and conchostracans of a Lower Cretaceous sequence at the Cañadón Calcáreo locality, Extra-Andean central Patagonia: age and palaeoenvironmental significance: Cretaceous Research, v. 30, p. 270282.
Von Meyer, H., 1857, Beiträge zur näheren Kenntnis fossiler Reptilien: Neues Jahrbuch für Mineralogie, Geologie und Paläontolgie, 1857, p. 532–543.
Weishampel, D.B., Dodson, P., and Osmólska, H., 2004, The Dinosauria, 2d ed.: Berkeley, University of California Press, 776 p.
Werning, S., 2012, The ontogenetic osteohistology of Tenontosaurus tilletti : PLoS One, v. 7, p. e33539.
White, M.A., 2009, The subarctometatarsus: intermediate metatarsus architecture demonstrating the evolution of the arctometatarsus and advanced agility in theropod dinosaurs: Alcheringa, v. 33, p. 121.
Woodward, H.N., Rich, T.J., Chinsamy, A., and Vickers-Rich, P., 2011, Growth dynamics of Australia’s Polar dinosaurs: PLoS One, v. 6(8), p. e23339.
Xu, X., Wang, X.-L., and You, H.-L., 2000a, A primitive ornithopod from the Early Cretaceous Yixian Formation of Liaoning: Vertebrata PalAsiatica, v. 38, p. 318325.
Xu, X., Zhou, Z., and Wang, X.-L., 2000b, The smallest known non-avian theropod dinosaur: Nature, v. 408, p. 705708.
Xu, X., Norrel, M.A., Kuang, X., Wang, X., Zhao, Q., and Jia, C., 2004, Basal tyrannosaurids from China and evidence for protofeathers in tyrannosauroids: Nature, v. 431, p. 680684.
Xu, X., Forster, C.A., Clark, J.M., and Mo, J., 2006, A basal ceratopsian with transitional features from the Late Jurassic of northwestern China: Proceedings of the Royal Society B, v. 273, p. 21352140.
Yates, A.M., 2003, The species taxonomy of the sauropodomorph dinosaurs from the Löwenstein Formation (Norian, Late Triassic) of Germany: Palaeontology, v. 46, p. 317337.
You, H.-L., and Dodson, P., 2003, Redescription of neoceratopsian dinosaur Archaeoceratops and early evolution of Neoceratopsia: Acta Paleontologica Polonica, v. 48, p. 261272.
You, H.-L., and Dodson, P., 2004, Basal Ceratopsia, in Weishampel, D.B., Dodson, P., and Osmólska, H., eds., The Dinosauria, 2d ed.: Berkeley, University of California Press, p. 478493.
Zavattieri, A.M., Escapa, I.H., Scasso, R.A., and Olivera, D., 2010, Contribución al conocimiento palinoestratigráfico de la Formación Cañadón Calcáreo en su localidad tipo, provincia del Chubut, Argentina: X Congreso Argentino de Paleontología y Bioestratigrafía-VII Congreso Latinoamericano de Paleontología Resúmenes, La Plata, Argentina, p. 224.
Zhang, F., Zhou, Z., Xu, X., and Wang, X., 2002, A juvenile coelurosaurian theropod from China indicates arboreal habits: Naturwissenschaften, v. 89, p. 394398.
Zheng, X.-T., You, H.-L., Xu, X., and Dong, Z.-M., 2009, An Early Cretaceous heterodontosaurid dinosaur with integumentary structures: Nature, v. 458, p. 333336.
Zhou, Z., and Farlow, J.O., 2001, Flight capability and habits of Confuciusornis , in Gauthier, J., and Gall, L.F., eds., New Perspectives on the Origin and Early Evolution of Birds, New Haven, New Jersey, Special publication of the Peabody Museum of Natural History, Yale University, p. 237254.
Zhou, Z., and Hou, L., 1998, Confuciusornis and the early evolution of birds: Vertebrata PalAsiatica, v. 36, p. 136146.
Recommend this journal

Email your librarian or administrator to recommend adding this journal to your organisation's collection.

Journal of Paleontology
  • ISSN: 0022-3360
  • EISSN: 1937-2337
  • URL: /core/journals/journal-of-paleontology
Please enter your name
Please enter a valid email address
Who would you like to send this to? *
×

Metrics

Altmetric attention score

Full text views

Total number of HTML views: 0
Total number of PDF views: 0 *
Loading metrics...

Abstract views

Total abstract views: 0 *
Loading metrics...

* Views captured on Cambridge Core between <date>. This data will be updated every 24 hours.

Usage data cannot currently be displayed