Hostname: page-component-8448b6f56d-qsmjn Total loading time: 0 Render date: 2024-04-19T15:41:43.482Z Has data issue: false hasContentIssue false

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

Published online by Cambridge University Press:  27 June 2016

Marcos G. Becerra
Affiliation:
CONICET- Museo Palaeontológico Egidio Feruglio, Fontana 140, Trelew, Chubut 9100, Argentina 〈mbecerra@mef.org.ar〉; 〈dpol@mef.org.ar〉
Diego Pol
Affiliation:
CONICET- Museo Palaeontológico Egidio Feruglio, Fontana 140, Trelew, Chubut 9100, Argentina 〈mbecerra@mef.org.ar〉; 〈dpol@mef.org.ar〉
Oliver W.M. Rauhut
Affiliation:
SNSB, Bayerische Staatssammlung für Paläontologie und Geologie and Department of Earth and Environmental Sciences, LMU München, Richard-Wagner-Str. 10, Munich 80333, Germany 〈o.rauhut@lrz.uni-muenchen.de〉
Ignacio A. Cerda
Affiliation:
CONICET- Instituto de Investigación en Paleobiología y Geología, Universidad Nacional de Río Negro, Museo Carlos Ameghino, Belgrano 1700, Paraje Pichi Ruca (predio Marabunta), Cipolletti, Río Negro, Argentina 〈nachocerda6@yahoo.com.ar〉

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.

Type
Articles
Copyright
Copyright © 2016, The Paleontological Society 

Access options

Get access to the full version of this content by using one of the access options below. (Log in options will check for institutional or personal access. Content may require purchase if you do not have access.)

References

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.Google Scholar
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.Google Scholar
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.Google Scholar
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.Google Scholar
Bonaparte, J.F., 1976, Pisanosaurus mertii Casamiquela and the origin of the Ornithischia: Journal of Paleontology, v. 50, p. 808820.Google Scholar
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.Google Scholar
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.Google Scholar
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.Google Scholar
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.Google Scholar
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.Google Scholar
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.Google Scholar
Butler, R.J., Upchurch, P., and Norman, D.B., 2008, The phylogeny of the ornithischian dinosaurs: Journal of Systematic Palaeontology, v. 6, p. 140.Google Scholar
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.Google Scholar
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.CrossRefGoogle Scholar
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.Google Scholar
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.Google Scholar
Casamiquela, R.M., 1967, Un nuevo dinosaurio ornitisquio Triásico (Pisanosaurus mertii: Ornithopoda) de la Formación Ischigualasto, Argentina: Ameghiniana, v. 4, p. 4764.Google Scholar
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.Google Scholar
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.Google Scholar
Chinsamy, A., 1995, Ontogenetic changes in the bone histology of the Late Jurassic ornithopod Dryosaurus lettowvorbecki : Journal of Vertebrate Paleontology, v. 15, p 96104.Google Scholar
Chinsamy, A., and Raath, M.A., 1992, Preparation of fossil bone for histological examination: Paleontologia Africana, v. 29, p. 3944.Google Scholar
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.Google Scholar
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.Google Scholar
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.Google Scholar
Coombs, W.P. Jr., 1978, Theoretical aspects of cursorial adaptations in dinosaurs: The Quarterly Review of Biology, v. 53, p. 393418.Google Scholar
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.Google Scholar
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.Google Scholar
Crompton, A.W., and Charig, A.J., 1962, A new ornithischian from the Upper Triassic of South Africa: Nature, v. 196, p. 10741077.Google Scholar
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.CrossRefGoogle Scholar
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.Google Scholar
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.Google Scholar
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.Google Scholar
Dong, Z., 1978, A new genus of Pachycephalosauria from Laiyang, Shantung: Vertebrata PalAsiatica, v. 16, p. 225228. [in Chinese]Google Scholar
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]Google Scholar
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.Google Scholar
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.Google Scholar
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.Google Scholar
Feduccia, A., 1993, Evidence from claw geometry indicating arboreal habits of Archaeopteryx : Science, v. 259, p. 790793.Google Scholar
Fisher, H.I., 1946, Adaptations and comparative anatomy of the locomotor apparatus of New World vultures: American Midland Naturalist, v. 35, p. 545727.Google Scholar
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.Google Scholar
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.Google Scholar
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.Google Scholar
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.Google Scholar
Galton, P.M., 1978, Fabrosauridae, the basal family of ornithischian dinosaurs (Reptilia: Ornithopoda): Paläontologische Zeitschrift, v. 52, p. 138159.Google Scholar
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.Google Scholar
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.Google Scholar
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.Google Scholar
Galton, P.M., and Powell, H.P., 1980, The ornithischian dinosaur Camptosaurus prestwichii from the Upper Jurassic of England: Palaeontology, v. 23, p. 411443.Google Scholar
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.Google Scholar
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.Google Scholar
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.Google Scholar
Gilmore, C.W., 1913, A new dinosaur from the Lance Formation of Wyoming: Smithsonian Miscellaneous Collection, v. 61, p. 15.Google Scholar
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.Google Scholar
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.CrossRefGoogle Scholar
Goloboff, P.A., Farris, J.S., and Nixon, K.C., 2008a, TNT, a free program for phylogenetic analysis: Cladistics, v. 24, p. 113.CrossRefGoogle Scholar
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.Google Scholar
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.Google Scholar
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.Google Scholar
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.Google Scholar
Haughton, H., 1924, The fauna and stratigraphy of the Stormberg Series: Annals of the South African Museum, v. 12, p. 323497.Google Scholar
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.Google Scholar
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]Google Scholar
Holtz, T.R. Jr., 1994, The arctometatarsalian pes, an unusual structure of Cretaceous Theropoda (Dinosauria: Saurischia): Journal of Vertebrate Paleontology, v. 14, p. 408519.Google Scholar
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.CrossRefGoogle Scholar
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.Google Scholar
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.Google Scholar
Horner, J.R., and Makela, R., 1979, Nest of juveniles provides evidence of family structure among dinosaurs: Nature, v. 282, p. 296298.Google Scholar
Horner, J.R., and Weishampel, D.B., 1988, A comparative embryological study of two ornithischian dinosaurs: Nature, v. 332, p. 256257.Google Scholar
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.Google Scholar
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.Google Scholar
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.Google Scholar
Hou, L., 1977, A new primitive Pachycephalosauria from Anhui, China: Vertebrata PalAsiatica, v. 15, p. 198202. [in Chinese]Google Scholar
Hübner, T.R., 2012, Bone histology in Dysalotosaurus lettowvorbecki (Ornithischia: Iguanodontia)-variation, growth, and implications: PLoS One, v. 7, e29958.Google Scholar
Huene, F.V., 1908, Die Dinosaurier der europäischen Triasformation mit Berücksichtigung der aussereuropäischen Vorkommnisse: Geologische und Palaeontologische Abhandlungen, Supplement, v. 1, p. 1419.Google Scholar
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.CrossRefGoogle ScholarPubMed
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.Google Scholar
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.Google Scholar
Irmis, R.B., Parker, W.G., Nesbitt, S.J., and Liu, J., 2007, Ornithischian dinosaurs: the Triassic record: Historical Biology, v. 18, p. 322.CrossRefGoogle Scholar
Kambic, R.E., 2008, Multivariate analysis of avian and non-avian theropod pedal phalanges, Unpublished Ph.D. dissertation: Bozeman, Montana State University, 90 p.Google Scholar
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.Google Scholar
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.Google Scholar
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.Google Scholar
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.Google Scholar
Marsh, O.C., 1878, Principal characters of American Jurassic dinosaurs. Part I: American Journal of Science, v. 16, p. 411416.CrossRefGoogle Scholar
Marsh, O.C., 1879, Notice of new Jurassic reptiles: American Journal of Science, v. 18, p. 501505.Google Scholar
Marsh, O.C., 1885, Names of extinct reptiles: American Journal of Science, v. 29, p. 169.Google Scholar
Marsh, O.C., 1894, The typical Ornithopoda of the American Jurassic: American Journal of Science, v. 283, p. 8590.Google Scholar
Maryanska, T., and Osmólska, H., 1974, Pachycephalosauria, a new suborder of ornithischian dinosaurs: Palaeontologia. Polonica, v. 30, p. 45102.Google Scholar
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.Google Scholar
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.Google Scholar
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.Google Scholar
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.Google Scholar
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.Google Scholar
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.Google Scholar
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.Google Scholar
Osborn, H.F., 1923, Two Lower Cretaceous dinosaurs of Mongolia: American Museum Novitates, v. 95, p. 110.Google Scholar
Osborn, H.F., 1924, Psittacosaurus and Protiguanodon, p. Two Lower Cretaceous iguanodonts from Mongolia: American Museum Novitates, v. 127, p. 116.Google Scholar
Osmólska, H., 1981, Coossified tarsometatarsi in theropod dinosaurs and their bearing on the problem of bird origins: Acta Paleontologica Polonica, v. 42, p. 7995.Google Scholar
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.Google Scholar
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.Google Scholar
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.Google Scholar
Ostrom, J.H., 1974, Archaeopteryx and the origin of flight: The Quarterly Review of Biology, v. 49, p. 2747.Google Scholar
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.Google Scholar
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.Google Scholar
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.Google Scholar
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]Google Scholar
Peng, G.-Z., 1992, Jurassic ornithopod Agilisaurus louderbacki (Ornithopoda: Fabrosauridae) from Zigong, Sichuan, China: Vertebrata PalAsiatica, v. 30, p. 3951. [in Chinese]Google Scholar
Pike, A.V.L., and Maitland, D.P., 2004, Scaling of bird claws: Journal of Zoology, v. 262, p. 7381.Google Scholar
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.Google Scholar
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.Google Scholar
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.Google Scholar
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.Google Scholar
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.Google Scholar
Romer, A.S., 1972, The Chañares (Argentina) Triassic reptile fauna. XV. Further remains of the thecodonts Lagerpeton and Lagosuchus : Breviora, v. 394, p. 17.Google Scholar
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.Google Scholar
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.Google Scholar
Salani, F.M., 2007, Aporte a la edad de la Formación Cañadón Asfalto, Chubut, Argentina: Resúmenes, 30 Simposio Argentino del Jurásico, p 71.Google Scholar
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.Google Scholar
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.Google Scholar
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.Google Scholar
Sereno, P.C., 1986, Phylogeny of the bird-hipped dinosaurs (Order Ornithischia): National Geographic Research, v. 2, p. 234256.Google Scholar
Sereno, P.C., 1991, Lesothosaurus, “fabrosaurids,” and the early evolution of Ornithischia: Journal of Vertebrate Paleontology, v. 11, p. 168197.Google Scholar
Sereno, P.C., 1997, The origin and evolution of dinosaurs: Annual Review of Earth and Planetary Sciences, v. 25, p. 435489.Google Scholar
Sereno, P.C., 2012, Taxonomy, morphology, masticatory function and phylogeny of heterodontosaurid dinosaurs: ZooKeys, v. 226, p. 1225.Google Scholar
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.Google Scholar
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.Google Scholar
Sternberg, C.M., 1937, Classification of Thescelosaurus: a description of a new species: Proceedings of the Geological Society of America, v. 1936, p. 375.Google Scholar
Thulborn, R.A., 1970, The systematic position of the Triassic ornithischian dinosaur Lycorhinus angustidens : Zoological Journal of the Linnean Society, v. 49, p. 235245.Google Scholar
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.Google Scholar
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.Google Scholar
Virchow, H., 1919, Atlas and Epistropheus bei den Schildkröten: Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin, v. 1919, p. 303332.Google Scholar
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.Google Scholar
Von Meyer, H., 1857, Beiträge zur näheren Kenntnis fossiler Reptilien: Neues Jahrbuch für Mineralogie, Geologie und Paläontolgie, 1857, p. 532–543.Google Scholar
Weishampel, D.B., Dodson, P., and Osmólska, H., 2004, The Dinosauria, 2d ed.: Berkeley, University of California Press, 776 p.Google Scholar
Werning, S., 2012, The ontogenetic osteohistology of Tenontosaurus tilletti : PLoS One, v. 7, p. e33539.Google Scholar
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.Google Scholar
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.Google Scholar
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.Google Scholar
Xu, X., Zhou, Z., and Wang, X.-L., 2000b, The smallest known non-avian theropod dinosaur: Nature, v. 408, p. 705708.Google Scholar
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.Google Scholar
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.Google Scholar
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.Google Scholar
You, H.-L., and Dodson, P., 2003, Redescription of neoceratopsian dinosaur Archaeoceratops and early evolution of Neoceratopsia: Acta Paleontologica Polonica, v. 48, p. 261272.Google Scholar
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.Google Scholar
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.Google Scholar
Zhang, F., Zhou, Z., Xu, X., and Wang, X., 2002, A juvenile coelurosaurian theropod from China indicates arboreal habits: Naturwissenschaften, v. 89, p. 394398.Google Scholar
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.Google Scholar
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.Google Scholar
Zhou, Z., and Hou, L., 1998, Confuciusornis and the early evolution of birds: Vertebrata PalAsiatica, v. 36, p. 136146.Google Scholar