First Record of Gregarious Behavior in Robust Medium-Sized Jurassic Ornithopods: Evidence from the Kimmeridgian Trackways of Asturias (N. Spain) and Some General Considerations on Other Medium-Large Ornithopod Tracks in the Mesozoic Record

References

• Agirrezabala, L. M., Torres, J. A., and Viera, L. I. 1985. El Weald de Igea (Cameros-La Rioja). Sedimentología, Bioestratigrafía y Paleoicnología de grandes reptiles (Dinosaurios). Munibe, 37: 111–138.
   Google Scholar
• Alexander, R. M. (1976): Estimates of speeds of dinosaurs. Nature, 261: 129–130.
   Web of Science ®Google Scholar
• Aramburu, C., and Bastida, F. (eds.). 1995. Geología de Asturias. Ediciones Trea, Gijón, 314 p.
   Google Scholar
• Barco, J. L., Ardévol, L., and Canudo, J. I. 2001. Descripción de los primeros rastros asignados a Hadrosauridae (Ornithopoda, Dinosauria) del Maastricthtiense de la Península Ibérica (Areny, Huesca). Geogaceta, 30: 235–238.
   Google Scholar
• Carpenter, K. 1992. Behavior of hadrosaurs as interpreted from footprints in the “Mesaverde” Group (Campanian) of Colorado, Utah, and Wyoming. Contributions to Geology, University of Wyoming, 29: 81–96.
   Google Scholar
• Casanovas Cladellas, M. L., Ezquerra Miguel, R., Fernández Ortega, A., Pérez-Lorente, F., and Santafe Llopis, J. V., Torcida Fernández, F. 1993. Tracks of a herd of webbed ornithopods and other footprints found in the same site (Igea, La Rioja, Spain). Revue de Paléobiologie. Vol., spec. 7: 29–36.
   Google Scholar
• Castanera, D., Vila, B., Razzolini, N. L., Falkingham, P. L., and Canudo, J. I. 2013a. Manus track preservation bias as a key factor for assessing trackmaker identity and quadrupedalism in sasal ornithopods. PLoS ONE, 8: e54177.
   PubMed Web of Science ®Google Scholar
• Castanera, D., Pascual, C., Razzolini, N. L., Vila, B., Barco, J. L., and Canudo, J. I. 2013b. Discriminating between medium-sized tridactyl trackmakers: Tracking ornithopod tracks in the base of the Cretaceous (Berriasian, Spain). PLoS ONE, 8: e81830.
   PubMed Web of Science ®Google Scholar
• Castanera, D., Vila, B., Razzolini, N. L., Santos, V. F., Pascual, C., and Canudo, J. I. 2014. Sauropod trackways of the Iberian Peninsula: palaeoetological and palaeoenvironmental implications. Journal of Iberian Geology, 40: 49–59.
   Web of Science ®Google Scholar
• Cobos, A., and Gascó, F. 2012. Presencia del icnogénero Iguanodontipus en el Cretácico Inferior de la provincia de Teruel (España). Geogaceta, 52: 185–188.
   Google Scholar
• Cotton, W. D., Cotton, J. E., and Hunt, A. P. 1998. Evidence for social behavior in ornithopod dinosaurs from the Dakota Group of northeastern New Mexico, U.S.A. Ichnos, 6: 141–149.
   Google Scholar
• Currie, P. J. 1983. Hadrosaur trackways from the Lower Cretaceous of Canada. Acta Palaeontologica Polonica, 28: 63–73.
   Google Scholar
• Currie, P. J. 1989. Dinosaur footprints of western Canada. In Gillette, D. D. and Lockley, M. G. (eds.). Dinosaur Tracks and Traces. Cambridge University Press, Cambridge, 293–300.
   Google Scholar
• Currie, P. J. 1995. Ornithopod trackways from the Lower Cretaceous of Canada. In Sarjeant, W. A. S. (ed.). Vertebrate Fossils and the Evolution of Scientific Concepts. Gordon and Breach, Newark, NJ, 431–442.
   Google Scholar
• Currie, P. J. and Sarjeant, W. A. S. 1979. Lower Cretaceous dinosaur footprints from the Peace River Canyon, British Columbia, Canada. Palaeogeography, Palaeoclimatology, Palaeoecology, 28: 103–115.
   Web of Science ®Google Scholar
• Currie, P. J., Badamgarav, D., and Koppelhus, E. B. 2003. The first Late Cretaceous footprints from the Nemegt locality in the Gobi of Mongolia. Ichnos, 10: 1–13.
   Google Scholar
• Currie, P. J., Nadon, G. C., and Lockley, M. G. 1991. Dinosaur footprints with skin impressions from the Cretaceous of Alberta and Colorado. Canadian Journal of Earth Sciences, 28: 102–115.
   Web of Science ®Google Scholar
• Dalla Vecchia, F. M. 1998. Remains of Sauropoda (Reptilia, Saurischia) in the Lower Cretaceous (Upper Hauterivian/Lower Barremian) limestones of SW Istria (Croatia). Geologica Croatica, 51: 105–134.
   Google Scholar
• Dalla Vecchia, F. M., Vlahović, I., Posocco, L., Tarlao, A., and Tentor, M. 2002. Late Barremian and Late Albian (Early Cretaceous) dinosaur track sites in the Main Brioni/Brijun Island (SW Istria, Croatia). Natura Nascosta, 25: 1–36.
   Google Scholar
• Delair, J. B. 1989. A history of dinosaur footprint discoveries in the British Weald. In Gillette, D. D. and Lockley, M. G. (eds.). Dinosaur Tracks and Traces. Cambridge University Press, Cambridge, 19–25.
   Google Scholar
• Delair, J. B., and Lander, A. B. 1973. A short history of the discovery of reptilian footprints in the Purbeck Beds of Dorset, with notes on their distribution. Proceedings of the Dorset Natural History and Archaeological Society, 94: 17–20.
   Google Scholar
• Díaz-Martínez, I., Pereda-Suberbiola, X., Pérez-Lorente, F., and Canudo, J. I. 2015. Ichnotaxonomic review of large ornithopod dinosaur tracks: Temporal and geographic implications. PLoS ONE, 10: e0115477.
   PubMed Web of Science ®Google Scholar
• Diedrich, C. 2004. New important iguanodontid and theropod trackways of the tracksite Obernkirchen in the Berriasian of NW Germany and megatracksite concept of Central Europe. Ichnos, 11: 215–228.
   Google Scholar
• Diedrich, C. 2011. Upper Jurassic tidal flat megatracksites of Germany: Coastal dinosaur migration highways between European islands, and a review of the dinosaur footprints. Palaeobiodiversity and Palaeoenvironments, 91: 129–155.
   Google Scholar
• Dollo, L. 1906. Les allures des Iguanodons, d´après les empreintes des pieds et de la queue. Bulletin Scientifique de la France et de la Belgique, series 5, 9: 1–12.
   Google Scholar
• Dong, Z. M., Zhou, Z. L., and Wu, S. Y. 2003. Note on a hadrosaur footprint from Heilongjiang River area of China. Vertebrata PalAsiatica, 41: 324–326. (In Chinese with an English abstract).
   Google Scholar
• Farlow, J. O., Langston, W. Jr., Deschner, E. E., Solis, R., Ward, W., Kirkland, B. L., Hovorka, S., Reece, T. L., and Whitcraft, J. 2006. Texas giants: dinosaurs of the heritage museum on the Texas Hill Country. The Heritage Museum of the Texas Hill Country, Canyon Lake, Texas, 105 p.
   Google Scholar
• Fanti, F., Bell, P. R., and Sissons, R. L. 2013. A diverse, high-latitude ichnofauna from the Late Cretaceous Wapiti Formation, Alberta, Canada. Cretaceous Research, 41: 256–269.
   Web of Science ®Google Scholar
• Fiorillo, A. R., and Parrish J. T. 2004. The first record of a Cretaceous dinosaur from southwestern Alaska. Cretaceous Research, 25: 453–458.
   Web of Science ®Google Scholar
• Fiorillo, A. R., Hasiotis, S. T., and Kobayashi, Y. 2014. Herd structure in Late Cretaceous polar dinosaurs: A remarkable new dinosaur tracksite, Denali National Park, Alaska, USA. Geology, 42: 719–722.
   Web of Science ®Google Scholar
• Fischer, R. 1998. Die Saurierfährten im Naturdenkmal Münchehagen. Mitteilungen aus dem Institut für Geologie und Paläontologie der Univestität Hannover, 37: 3–59.
   Google Scholar
• Fürsich, F. T., Werner, W., Delvene, G., García-Ramos, J. C., Bermúdez-Rochas, D. D., and Piñuela, L. 2012. Taphonomy and palaeoecology of high-stress benthic associations from the Upper Jurassic of Asturias, northern Spain. Palaeogeography, Palaeoclimatology, Palaeoecology, 358–360: 1–18.
   Web of Science ®Google Scholar
• Galton, P. M., and Upchurch, P. 2004. Stegosauria. In Weishampel, D. B., Dodson, P., and Osmólska, H. (eds.). The Dinosauria (second edition). University of California Press, Berkeley, 343–362.
   Google Scholar
• Gangloff, R. A., May, K. C., and Storer, J. E. 2004. An early Late Cretaceous dinosaur tracksite in central Yukon Territory, Canada. Ichnos, 11: 299–309.
   Google Scholar
• García-Ortiz, E., and Pérez-Lorente, F. 2014. Palaeoecological inferences about dinosaur gregarious behaviour based on the study of tracksites from La Rioja area in the Cameros Basin (Lower Cretaceous, Spain). Journal of Iberian Geology, 40: 113–127.
   Web of Science ®Google Scholar
• García-Ramos, J. C. 1977. Hallazgo de huellas de dinosaurio (Theropoda y Ornithopoda) en la costa asturiana. Asturnatura, 3: 171–172.
   Google Scholar
• García-Ramos, J. C., Piñuela, L., and Aramburu, C. 2010a. Introducción al Jurásico asturiano. In García-Ramos, J. C. and Aramburu, C. (cords.). Las sucesiones litorales y marinas restringidas del Jurásico Superior. Acantilados de Tereñes (Ribadesella) y de la playa de La Griega (Colunga). V Congreso del Jurásico de España, Guía de campo (excursión B), Museo del Jurásico de Asturias, Colunga, 5–10.
   Google Scholar
• García-Ramos, J. C., Piñuela, L., and Aramburu, C. 2010b. La Formación Tereñes en su localidad tipo. In García-Ramos, J. C. and Aramburu, C. (eds.). Las sucesiones litorales y marinas restringidas del Jurásico Superior. Acantilados de Tereñes (Ribadesella) y de la playa de La Griega (Colunga). V Congreso del Jurásico de España, Guía de campo (excursión B), Museo del Jurásico de Asturias, Colunga, 15–39.
   Google Scholar
• García-Ramos, J. C., Piñuela, L., and Lires, J. 2006. Atlas del Jurásico de Asturias. Ediciones Nobel, Oviedo, 225 p.
   Google Scholar
• García-Ramos, J. C., Piñuela, L., and Rodríguez-Tovar, F. J. 2011. Post-workshop field trip guide of the XI International Ichnofabric Workshop. Museo del Jurásico de Asturias, Colunga, 89 p.
   Google Scholar
• García-Ramos, J. C., Piñuela, L., Avanzini, M., and Ruiz-Omeñaca J. I. (2009). Deep theropod undertracks look like ornithopod tracks. A conclusion from a three-dimensional study of dinosaur footprints. In Delgado Buscalioni, A. and Fregenal Martínez, M. (eds.). Tenth International Symposium on Mesozoic Terrestrial Ecosystems and Biota, Teruel. Universidad Autónoma de Madrid, 279–281.
   Google Scholar
• García-Ramos, J. C., Piñuela L., Ruiz-Omeñaca, J. I., and Pereda Suberbiola, X. 2008. Costas jurásicas frecuentadas por estegosaurios. In Ruiz-Omeñaca, J. I., Piñuela, L., and García-Ramos, J. C. (eds.). Libro de resúmenes XXIV Jornadas de la Sociedad Española de Paleontología. Museo del Jurásico de Asturias (MUJA), Colunga, 15–18 de octubre de 2008, 33–34.
   Google Scholar
• Gierliński, G. D., and Sabath, K. 2008. Stegosaurian footprints from the Morrison Formation of Utah and their implications for interpreting other ornithischian tracks. Oryctos, 8: 29–46.
   Google Scholar
• Gierliński, G. D., Ploch, I., Gawor-Biedowa, E., and Niedźwiedzki, G. 2008. The first evidence of dinosaur tracks in the Upper Cretaceous of Poland. Oryctos, 8: 107–113.
   Google Scholar
• Hamblin, A. H., and Foster, J. R. 2000. Ancient animal footprints and traces in the Grand Staircase-Escalante National Monument, south-central Utah. In Sprinkel, D. A., Chidsey, T. C. Jr., and Anderson, P. B. (eds.). Geology of Utah's Parks and Monuments, Utah Geological Association Publication, 28: 1–12.
   Google Scholar
• Hitchcock, E. 1848. An attempt to discriminate and describe the animals that made the fossil footmarks of the United States, and especially of New England. Memoirs of the American Academy of Arts and Sciences, new series, 3: 129–256.
   Google Scholar
• Hornung, J. J., Böhme, A., van der Lubbe, T., Reich, M., and Richter, A. 2012. Vertebrate tracksites in the Obernkirchen Sandstone (late Berriasian, Early Cretaceous) of northwest Germany–their stratigraphical, palaeogeographical, palaeoecological, and historical context. Paläontologische Zeitschrift, 86: 231–267.
   Web of Science ®Google Scholar
• Huh, M., Paik, I. S., Chung, C. H., Park, J. B., and Kim, B. S. 2001. Dinosaur tracks from islands in Yeosu, Cheollanam-do Korea. Journal of the Geological Society of Korea, 37: 653–658. (In Korean with an English abstract).
   Google Scholar
• Hunt, A. P., and Lucas, S. G. 2003. A new hadrosaur track from the Upper Cretaceous Fruitland Formation of Northwestern New Mexico. New Mexico Geological Society Guidebook, 54th Conference, Geology of the Zuni Plateau, 379–381.
   Google Scholar
• Ishigaki, S., Watabe, M., Tsogtbaatar, Kh., and Saneyoshi, M. 2009. Dinosaur footprints from the Upper Cretaceous of Mongolia. Geological Quarterly, 53: 449–460.
   Web of Science ®Google Scholar
• Jaillard, E., Cappeta, H., Ellenberger, P., Feist, M., Grambast-Fessard, N., Lefranc, J. P., and Sigé, B. 1993. Sedimentology, palaeontology, biostratigraphy and correlation of the Late Cretaceous Vilquechico Group of southern Peru. Cretaceous Research, 14: 623–661.
   Web of Science ®Google Scholar
• Kim, J. Y., Lockley, M. G., Kim, H. M., Lim, J.-D., and Kim, K. S. 2009. New dinosaur tracks from Korea, Ornithopodichnus masanensis ichnogen. et ichnosp. nov. (Jindong Formation, Lower Cretaceous): implications for polarities in ornithopod foot morphology. Cretaceous Research, 30: 1387–1397.
   Web of Science ®Google Scholar
• Lapparent, A. F. de 1962. Footprints of dinosaur in the Lower Cretaceous of Vestspitsbergen-Svalbard. Norsk Polarinstitutt Årbok, 1960: 14–21.
   Google Scholar
• Lee, Y.-N., Yang, S.-Y., Seo, S.-J., Baek, K.-S., Yi, M.-S., Lee, D.-J., Park, E.-J., and Han, S.-W. 2000. Distribution and paleobiological significance of dinosaur tracks from the Jindong Formation (Albian) in Kosong County, Korea. Paleontolological Society of Korea, Special Publication, 4: 1–12.
   Google Scholar
• Leonardi, G. 1979. Nota preliminar sobre seis pistas de dinossauros Ornithischia da Bacia do Rio do Peixe, em Soussa, Paraíba, Brasil. Anais da Academia Brasileira de Ciências, 51: 501–516.
   Google Scholar
• Leonardi, G. 1984. Le impronte fossili di dinosauri. In Bonaparte J. F. et al. (eds.). Sulle Orme dei Dinosauri. Erizzo Editrice, Venezia, 163–186.
   Google Scholar
• Lim, S.-Y., Yang, S.-Y., and Lockley, M. G. 1989. Large dinosaur footprint assemblages from the Cretaceous Jindong Formation of southern Korea. In Gillette, D. D. and Lockley, M. G. (eds.). Dinosaur Tracks and Traces. Cambridge University Press, Cambridge, 333–336.
   Google Scholar
• Lim, S.-K., Lockley, M. G., Yang, S.-Y., Fleming, R. F., and Houck, K. 1995. A preliminary report on sauropod tracksites from the Cretaceous of Korea. In Lockley, M. G., Santos, V. F., Meyer, C. A., and Hunt, A. P. (eds.). Aspects of Sauropod Paleobiology. Gaia, 10: 109–117.
   Google Scholar
• Llompart, C. 2006. Presencia de icnitas de dinosaurios en el Cretácico Superior del margen sur del sinclinal de Tremp (prov. de Lleida, Cataluña, España). Revista Española de Paleontología, 21: 1–14.
   Google Scholar
• Lockley, M. G. 1987. Dinosaur footprints from the Dakota Group of eastern Colorado. The Mountain Geologist, 24: 107–122.
   Google Scholar
• Lockley, M. G. 1989. Tracks and traces: New perspectives on dinosaurian behavior, ecology and biogeography. In Padian, K. and Chure, D. J. (eds.). The Age of Dinosaurs, Paleontological Society, Knoxville, Tennessee. Short courses in Paleontology, 2, 135–145.
   Google Scholar
• Lockley, M. G. 1995. Track records. Natural History, 104: 46–51.
   Web of Science ®Google Scholar
• Lockley, M. G. 1997. The paleoecological and paleoenvironmental utility of dinosaur tracks. In Farlow, J. O. and Brett-Surman, M. K. (eds.). The Complete Dinosaur. Indiana University Press, Bloomington, Indiana, 554–578.
   Google Scholar
• Lockley, M. G. 2009. New perspectives on morphological variation in tridactyl footprints: Clues to widespread convergence in developmental dynamics. Geological Quarterly, 53, 415–432.
   Web of Science ®Google Scholar
• Lockley, M. G., and Hunt, A. P. 1995. Dinosaur tracks and other fossil footprints of the Western United States. Columbia University Press, New York, 338 p.
   Google Scholar
• Lockley, M. G., and Hunt, A. P. 1998. A probable stegosaur track from the Morrison Formation of Utah. Modern Geology, 23: 331–342.
   Google Scholar
• Lockley, M. G., and Matsukawa, M. 1999. Some observations on trackway evidence for gregarious behavior among small bipedal dinosaurs. Palaeogeography, Palaeoclimatology, Palaeoecology, 150: 25–31.
   Web of Science ®Google Scholar
• Lockley, M. G., Huh, M., and Kim, B. S. 2012b. Ornithopodichnus and pes-only sauropod trackways from the Hwasun tracksite, Cretaceous of Korea. Ichnos, 19: 93–100.
   Web of Science ®Google Scholar
• Lockley, M. G., Lucas, S. G., and Hunt, A. P. 2000. Dinosaur tracksites in New Mexico: A review. In Lucas, S. G. and Heckert, A. B. (eds.). New Mexico Museum of Natural History and Science Bulletin, 17: 9–16.
   Google Scholar
• Lockley, M. G., Nadon, G., and Currie, P. J. 2003. A diverse dinosaur-bird footprint assemblage from the Lance Formation, Upper Cretaceous, Eastern Wyoming: implications for ichnotaxonomy. Ichnos, 11: 229–249.
   Google Scholar
• Lockley, M. G., Young, B. H., and Carpenter, K. 1983. Hadrosaur locomotion and herding behavior: evidence from footprints in the Mesaverde Formation Grand Mesa coal field, Colorado. The Mountain Geologist, 20: 5–14.
   Google Scholar
• Lockley, M., García-Ramos, J. C., Piñuela, L., and Avanzini, M. 2008. A review of vertebrate track assemblages from the Late Jurassic of Asturias, Spain with comparative notes on coeval ichnofaunas from the western USA: Implications for faunal diversity in siliciclastic facies assemblages. Oryctos, 8: 53–70.
   Google Scholar
• Lockley, M., Santos, V. F., Meyer, C., and Hunt, A. 1998a. A new dinosaur tracksite in the Morrison Formation, Boundary Butte, Southeastern Utah. In Carpenter, K., Chure, D., and Kirkland, J. (eds.). The Upper Jurassic Morrison Formation: An Interdisciplinary Study. Modern Geology, 23: 317–330.
   Google Scholar
• Lockley, M. G., Xing, L., Lockwood, J. A. F., and Pond, S. 2014. A review of large Cretaceous ornithopod tracks, with special reference to their ichnotaxonomy. Biological Journal of the Linnean Society, 113: 721–736.
   Web of Science ®Google Scholar
• Lockley, M. G., Holbrook, J., Hunt, A. P., Matsukawa, M., and Meyer, C. 1992. The dinosaur freeway: a preliminary report on the Cretaceous megatracksite, Dakota Group, Rocky Mountain Front Range and Highplains; Colorado, Oklahoma and New Mexico. In Flores, R. (ed.). Mesozoic of the Western Interior. SEPM, Tulsa, OK, Midyear Mtg. Fieldtrip Guidebook, 39–54.
   Google Scholar
• Lockley, M. G., Houck, K., Yang, S.-Y., Matsukawa, M., and Lim, S.-K. 2006. Dinosaur-dominated footprint assemblages from the Cretaceous Jindong Formation, Hallyo Haesang National Park, Goseong County, South Korea: Evidence and implications. Cretaceous Research, 27: 70–101.
   Web of Science ®Google Scholar
• Lockley, M. G., Huh, M., Gwak, S.-G., Hwang, K. G., and Paik, I. S. 2012a. Multiple tracksites with parallel trackways from the Cretaceous of the Yeosu City area, Korea: Implications for gregarious behavior in ornithopod and sauropod dinosaurs. Ichnos, 19: 105–114.
   Web of Science ®Google Scholar
• Lockley, M. G., Hunt, A. P., Meyer, C., Rainforth, E. C., and Schultz, R. J. 1998b. A survey of fossil footprint sites at Glen Canyon National Recreation Area (western USA): A case study in documentation of trace fossil resources at a National Preserve. Ichnos, 5: 177–211.
   Google Scholar
• Lockwood, J. A. F., Lockley, M. G., and Pond, S. 2014. A review of footprints from the Wessex Formation (Wealden Group, Lower Cretaceous) at Hanover Point, the Isle of Wight, southern England. Biological Journal of the Linnean Society, 113: 707–720.
   Web of Science ®Google Scholar
• Lucas, S. G., Sullivan, R. M., Jasinski, S. E., and Ford, T. L. 2011. Hadrosaur footprints from the Upper Cretaceous Fruitland Formation, San Juan basin, New Mexico, and the ichnotaxonomy of large ornithopod footprints. In Sullivan, R. M. et al. (eds.). New Mexico Museum of Natural History and Science Bulletin, 53: 357–362.
   Google Scholar
• Maidment, S. C. R., and Barrett, P. M. 2014. Osteological correlates for quadrupedality in ornithischian dinosaurs. Acta Palaeontologica Polonica, 59: 53–70.
   Web of Science ®Google Scholar
• Maidment, S. C. R., Bates, K. T., Falkingham, P. L., VanBuren, C., Arbour, V., and Barrett, P. M. 2014. Locomotion in ornithischian dinosaurs: An assessment using three-dimensional computational modelling. Biological Reviews, 89: 588–617.
   PubMed Web of Science ®Google Scholar
• Marty, D. 2008. Sedimentology, taphonomy, and ichnology of Late Jurassic dinosaur tracks from the Jura carbonate platform (Chevenez-Combe Ronde tracksite, NW Switzerland): Insights into the tidal-flat palaeoenvironment and dinosaur diversity, locomotion, and palaeoecology. GeoFocus, 21: 1–278.
   Google Scholar
• Mateus, O., and Antunes, T. 2001. Draconyx loureiroi, a new camptosauridae (Dinosauria, Ornithopoda) from the Late Jurassic of Lourinhã, Portugal. Annales de Paléontologie, 87: 61–73.
   Google Scholar
• Mateus, O., and Milan, J. 2008. Ichnological evidence for giant ornithopod dinosaurs in the Upper Jurassic Lourinhá Formation, Portugal. Oryctos, 8: 47–52.
   Google Scholar
• Matsukawa, M., Hamuro, T., Mizukami, T., and Fujii, S. 1997. First trackway evidence of gregarious dinosaurs from the Lower Cretaceous Tetori Group of eastern Toyama prefecture, central Japan. Cretaceous Research, 18: 603–619.
   Web of Science ®Google Scholar
• Matsukawa, M., Matsui, T., and Lockley, M. G. 2001. Trackway evidence of herd structure among ornithopod dinosaurs from the Cretaceous Dakota Group of Northeastern New Mexico, USA. Ichnos, 8: 197–206.
   Google Scholar
• McDonald, A. T. 2011. The taxonomy of species assigned to Camptosaurus (Dinosauria: Ornithopoda). Zootaxa, 2783: 52–68.
   Web of Science ®Google Scholar
• Meyer, C. A., and Thüring, B. 2003. The first iguanodontid dinosaur tracks from the Swiss Alps (Schrattenkalk Formation, Aptian). Ichnos, 10: 221–228.
   Google Scholar
• Moratalla, J. J., Sanz, J. L., and Jiménez, S. 1988. Nueva evidencia icnológica de dinosaurios en el Cretácico Inferior de La Rioja (España). Estudios Geológicos, 44: 119–131.
   Google Scholar
• Moratalla, J. J., Sanz, J. L., and Jiménez, S. 1994. Dinosaur tracks from the Lower Cretaceous of Regumiel de la Sierra (province of Burgos, Spain): Inferences on a new quadrupedal ornithopod trackway. Ichnos, 3: 89–97.
   Google Scholar
• Moratalla, J. J., Sanz, J. L., Jiménez, S., and Lockley, M. G. 1992. A quadrupedal ornithopod trackway from the Lower Cretaceous of La Rioja (Spain): Inferences on gait and hand structure. Journal of Vertebrate Paleontology, 12: 150–157.
   Google Scholar
• Norman, D. B. 1980. On the ornithischian dinosaur Iguanodon bernissartensis of Bernissart (Belgium). Mémoirs de l´Institut Royal des Sciences Naturelle de Belgique, 178: 1–105.
   Google Scholar
• Norman, D. B. 2004. Basal Iguanodontia. In Weishampel, D. B., Dodson, P., and Osmólska, H. (eds.). The Dinosauria. Second Edition. University of California Press, Berkeley, Los Angeles, CA, 413–437.
   Google Scholar
• Olsen, P. E., and Rainforth, E. C. 2003. The Early Jurassic ornithischian dinosaurian ichnogenus Anomoepus. In Le Tourneau, P. M., and Olsen, P. E. (eds.). The Great Rift Valleys of Pangea in Eastern North America, Volume 2: Sedimentology and Paleontology. Columbia University Press, New York, 314–368.
   Google Scholar
• Ostrom, J. H. 1972. Were some dinosaurs gregarious? Palaeogeography, Palaeoclimatology, Palaeoecology, 11: 287–301.
   Web of Science ®Google Scholar
• Paik, I. S., Huh, M., Park, K. H., Hwang, K. G., Kim, K. S., and Kim, H. J. 2006. Yeosu dinosaur track sites of Korea: The youngest dinosaur track records in Asia. Journal of Asian Earth Sciences, 28: 457–468.
   Web of Science ®Google Scholar
• Parker, L. R., and Rowley, R. L. Jr. 1989. Dinosaur footprints from a coal mine in east-central Utah. In Gillette, D. D. and Lockley, M. G. (eds.). Dinosaur Tracks and Traces, Cambridge University Press, Cambridge, 361–366.
   Google Scholar
• Pascual-Arribas, C., Hernández-Medrano, N., Latorre-Macarrón, P., and Sanz-Pérez, E. 2009. El icnogénero Iguanodontipus en el yacimiento de “Las Cuestas I” (Santa Cruz de Yanguas, Soria, Espanña). Studia Geologica Salmanticensia, 45: 105–128.
   Google Scholar
• Peabody, F. E. 1955. Taxonomy and the footprints of tetrapods. Palaeontology, 29, 915–918.
   Google Scholar
• Perez-Lorente, F., Cuenca-Besos, G., Aurell, M., Canudo, J. I., Soria, A. R., and Ruiz-Omeñaca, J. I. 1997. Las Cerradicas tracksite (Berriasian, Galve, Spain): Growing evidence for quadrupedal ornithopods. Ichnos, 5: 109–120.
   Google Scholar
• Piñuela, L., Farlow, J. O., García-Ramos, J. C., Ruiz-Omeñaca, J. I., Avanzini, M., and Lockley, M. 2008. Probable Anomoepus from the Late Jurassic of Asturias, Spain. The Second International Congress on Ichnology, Abstract book and the intra-Congress field trip guide book. Cracow, Poland, 106–107.
   Google Scholar
• Piñuela, L., García-Ramos, J. C., and Ruiz-Omeñaca, J. I. 2007. El yacimiento de icnitas de dinosaurios de Tereñes (Ribadesella, Asturias). In Braga, J. C., Checa, A., and Company, M. Libro de resúmenes, XXIV Jornadas de la Sociedad Española de Paleontología, Caravaca de la Cruz 3–6 de octubre, 185–186.
   Google Scholar
• Piñuela, L., García-Ramos, J. C., Fernández, L. A., and Ruiz-Omeñaca, J. I. 2014. Nuevos rastros de estegosaurios en el Jurásico Superior de Asturias. In Royo-Torres, R., Verdú, F. J., and Alcalá, L. (eds.). XXX Jornadas de Paleontología de la Sociedad Española de Paleontología. Fundamental, 24: 187–190.
   Google Scholar
• Piñuela Suárez, L. 2000. Icnitas de dinosaurios bípedos del Jurásico de Asturias. Morfometría, morfología e interpretación. MSc, University of Oviedo, 63 p. (Unpublished).
   Google Scholar
• Piñuela Suárez, L. 2015. Huellas de dinosaurios y de otros reptiles del Jurásico Superior de Asturias. PhD thesis, University of Oviedo, 326 p.
   Google Scholar
• Piñuela, L. 2012. Dinosaur true tracks and undertracks. Recognition criteria and nomenclature problems. The Asturian case (Spain). Abstract Book of Qijiang International Dinosaur Tracks Symposium, Chongqing, China, 91–95.
   Google Scholar
• Razzolini, N. L., Vila, B., Castanera, D., Falkingham, P. L., Barco, J. L., Canudo, J. I, Manning, P. L., and Galobart, A. 2014. Intra-trackway morphological variations due to substrate consistency: The El Frontal dinosaur tracksite (Lower Cretaceous, Spain). PLoS ONE, 9: e93708.
   PubMed Web of Science ®Google Scholar
• Rodríguez-de la Rosa, R. A., Aguillón-Martínez, M. C., López-Espinoza, J., and Eberth, D. A. 2004. The fossil record of vertebrate tracks in Mexico. Ichnos, 11: 27–37.
   Google Scholar
• Romilio, A., and Salisbury, S. W. 2011. A reassessment of large theropod dinosaur tracks from the mid-Cretaceous (late Albian-Cenomanian) Winton Formation of Lark Quarry, central-western Queensland, Australia: A case for mistaken identity. Cretaceous Research, 32: 135–142.
   Web of Science ®Google Scholar
• Ruiz-Omeñaca J. I., Piñuela, L., and García-Ramos, J. C. 2008. Nuevos restos de ornitópodo Ornithischia: Ankylopollexia) del Jurásico Superior de Tazones, Asturias (Formación Tereñes). Geogaceta, 45: 59–62.
   Google Scholar
• Ruiz-Omeñaca J. I., Piñuela, L., and García-Ramos, J. C. 2010. El primer diente de ornitópodo del Jurásico Superior de España (Asturias). Geogaceta, 48: 83–86.
   Google Scholar
• Ruiz-Omeñaca J. I., Piñuela, L., and García-Ramos, J. C. 2012. New ornithopod remains from the Upper Jurassic of Asturias (North Spain). Fundamental, 20: 219–222.
   Google Scholar
• Santos, V. F., Callapez, P. M., and Rodrigues, N. P. C. 2013. Dinosaur footprints from the Lower Cretaceous of the Algarve Basin (Portugal): New data on the ornithopod palaeoecology and palaeobiogeography of the Iberian Peninsula. Cretaceous Research, 40: 158–169.
   Web of Science ®Google Scholar
• Sanz, E., Pascual, C., and Bárez, S. 1999. Icnitas de dinosaurios en las facies Weald de “El Royo·” (Soria, España). Boletín de la Real Sociedad Española de Historia Natural (Sección Geología), 95: 101–115.
   Google Scholar
• Sarjeant, W. A. S. 1988. Fossil vertebrate footprints. Geology Today, 4: 125–130.
   Google Scholar
• Sarjeant, W. A. S. 1989. ‘Ten paleoichnological commandments’: A standardized procedure for the description of fossil vertebrate footprints. In Gillette, D. D. and Lockley, M. G. (eds.). Dinosaur Tracks and Traces, Cambridge University Press, Cambridge, 369–370.
   Google Scholar
• Sarjeant, W. A. S., Delair, J. B., and Lockley, M. G. 1998. The footprints of Iguanodon: A history and taxonomic study. Ichnos, 6: 183–202.
   Google Scholar
• Schulp, A. S., and Al-Wosabi, M. 2012. Telling apart ornithopod and theropod trackways: A closer look at a large, Late Jurassic tridactyl dinosaur trackway at Serwah, Republic of Yemen. Ichnos, 19: 194–198.
   Web of Science ®Google Scholar
• Sternberg, C. M. 1932. Dinosaur tracks from Peace River, British Columbia. National Museum of Canada, Annual Report, 1930: 59–85.
   Google Scholar
• Thulborn, R. A. 1990. Dinosaur Tracks. Chapman and Hall, London, 410 p.
   Google Scholar
• Valenzuela, M. 1988. Estratigrafía, sedimentología y paleogeografía del Jurásico de Asturias. PhD thesis, University of Oviedo, 748 p.
   Google Scholar
• Vila, B., Oms, O., Fondevilla, V., Gaete, R., Galobart, A., Riera, V., and Canudo, J. I. 2013. The latest succession of dinosaur tracksites in Europe: Hadrosaur ichnology, track production and palaeoenvironments. PLoS ONE, 8: e72579.
   PubMed Web of Science ®Google Scholar
• Weems, R. E., and Bachman, J. M. 1997. Cretaceous anuran and dinosaur footprints from the Patuxent Formation of Virginia. Proceedings of the Biological Society of Washington, 110: 1–17.
   Web of Science ®Google Scholar
• Whyte, M. A., and Romano, M. 1994. Probable sauropod footprints from the Middle Jurassic of Yorkshire, England. Gaia, 10: 15–26.
   Google Scholar
• Whyte, M. A., and Romano, M. 2001. Probably stegosaurian dinosaur tracks from the Saltwick Formation (Middle Jurassic) of Yorkshire, England. Proceedings of the Geologists´ Association, 112: 45–54.
   Web of Science ®Google Scholar
• Woodhams, K. E., and Hines, J. S. 1989. Dinosaur footprints from the Lower Cretaceous of East Sussex, England. In Gillette, D. D. and Lockley, M. G. (eds.). Dinosaur Tracks and Traces. Cambridge University Press, Cambridge, 301–307.
   Google Scholar
• Wright, J. L. 1999. Ichnological evidence for the use of the forelimb in iguanodontoid locomotion. Special Papers in Palaeontology, 60: 209–219.
   Google Scholar
• Xing, L. D., Wang, F. P., Pan, S. G., and Chen, W. 2007. The discovery of dinosaur footprints from the Middle Cretaceous Jiaguan Formation of Qijiang County, Chongqing City. Acta Geologica Sinica, 81: 1591–1602. (In Chinese with an English abstract).
   Google Scholar
• Xing, L. D., Harris, J. D., Wang, K. B., Chen, S. Q., Zhao, C. J., and Li, R. H. 2010. An Early Cretaceous non-avian dinosaur and bird footprint assemblage from the Laiyang Group in the Zhucheng basin, Shandong Province, China. Geological Bulletin of China, 29: 1105–1112.
   Google Scholar
• Xing, L. D., Harris, J. D., Dong, Z. M., Lin, Y. L., Chen, W., Guo, S. B., and Ji, Q. 2009. Ornithopod (Dinosauria: Ornitischia) tracks from the Upper Cretaceous Zhutian Formation in the Nanxiong basin, Guangdong, China and general observations on large Chinese ornithopod footprints. Geological Bulletin of China, 28: 829–843.
   Google Scholar
• You, H., and Azuma, Y. 1995. Early Cretaceous dinosaur footprints form Luanping, Hebei province, China. In Sun, A. and Wang, Y. (eds.). Sixth Symposium of Mesozoic Terrestrial Ecosystems and Biota. China Ocean Press, Beijing, China, 151–156.
   Google Scholar
• Zhang, J. P., Li, D. Q., Li, M., Lockley, M. G., and Bai, Z. 2006. Diverse dinosaur-, pterosaur-, and bird-track assemblages from the Hakou Formation, Lower Cretaceous of Gansu Province, northwest China. Cretaceous Research, 27: 44–55.
   Web of Science ®Google Scholar