Abstract
The El Castellar Formation (upper Hauterivian-lowermost Barremian, Lower Cretaceous) in the southwest of the Maestrat Basin (Spain) has yielded diverse dinosaur remains, mostly fragmentary bones. This study describes, morphometrically analyses, and compares several isolated postcranial fossils of large ornithopods from three new different sites in the municipality of Cabra de Mora (Peñagolosa sub-basin, Teruel Province). The fossils are attributed to styracosternan ornithopods, and some of them are related to the species Iguanodon galvensis, representing the oldest known record related to this taxon in the Iberian Peninsula. Moreover, this study verifies the hypothesis of the coexistence of at least two different styracosternans—a medium-sized form and a large-sized one—during the late Hauterivian-early Barremian in the Peñagolosa sub-basin. In addition, we formally describe the first dinosaur tracksite of the El Castellar Formation in this sub-basin. All ichnites are preserved as natural casts, and their trackmakers are likely to have been related to large styracosternans. Hence, the evidence provided here by both bones and tracks supports the idea that large ornithopods were the predominant dinosaurs in the wetlands of an extensive coastal plain.
Resumen
La Formación El Castellar (Hauteriviense superior-Barremiense basal) en el suroeste de la Cuenca del Maestrazgo (España) ha proporcionado diversos fósiles de dinosaurios bastante fragmentarios. Aquí describimos, analizamos morfométricamente y comparamos varios restos postcraneales aislados de grandes ornitópodos procedentes de tres yacimientos en el municipio de Cabra de Mora (subcuenca de Peñagolosa, provincia de Teruel). Los fósiles son atribuidos sistemáticamente a grandes ornitópodos estiracosternos, y algunos de ellos son relacionados con Iguanodon galvensis, lo que implica que estos restos pueden representar el registro más antiguo relacionado con dicho taxón en la Península Ibérica. Además, este estudio comprueba la hipótesis de la presencia de al menos dos formas de estiracosternos durante el Hauteriviense tardío-Barremiense temprano en la subcuenca de Peñagolosa, uno de talla grande y otro de talla mediana. Adicionalmente, se describe formalmente el primer yacimiento de icnitas de dinosaurios en la Formación El Castellar de esta subcuenca. Todas las icnitas están preservadas como hiporrelieves convexos y sus productores podrían estar relacionados también con grandes estiracosternos. Por lo tanto, la evidencia aquí proporcionada tanto por los restos directos como por las icnitas respaldan la idea de que los grandes ornitópodos eran los dinosaurios predominantes en los diferentes humedales desarrollados en una extensa llanura costera.
Similar content being viewed by others
Availability of data and material
The remains studied here are deposited in a public institution (Museo Aragonés de Paleontología).
Code availability
Not applicable for that section.
References
Alcalá, L., Cobos, A., & Royo-Torres, R. (2018). Dinosaurios de la Península Ibérica. Revista PH, 94, 116–153. https://doi.org/10.33349/2018.0.4145
Alexander, R. M. (1976). Estimates of speeds of dinosaurs. Nature, 261(5556), 129–130. https://doi.org/10.1038/261129a0
Arribas, I., Buscalioni, A. D., Royo-Torres, R., Espílez, E., Mampel, L., & Alcalá, L. (2019). A new goniopholidid crocodyliform, Hulkepholis rori sp. Nov. from the Camarillas Formation (early Barremian) in Galve, Spain. PeerJ, 7, e7911. https://doi.org/10.7717/peerj.7911
Aurell, M., Bádenas, B., Gascá, J. M., Canudo, J. I., Liesa, C. L., Soria, A. R., et al. (2016). Stratigraphy and evolution of the Galve sub-basin (Spain) in the middle Tithonian-early Barremian: Implications for the setting and age of some dinosaur fossil sites. Cretaceous Research, 65, 138–162.
Barrett, P. M. (2016). A new specimen of Valdosaurus canaliculatus (Ornithopoda: Dryosauridae) from the Lower Cretaceous of the Isle of Wight, England. Memoirs of Museum Victoria, 74, 29–48. https://doi.org/10.24199/j.mmv.2016.74.04
Benito, M. I., Cobos, A., Mas, J. R., Alcalá, L., Campos-Soto, S., & Fernández-Donado, L., et al. (2015). “Walking on tides”: Sedimentary controls on dinosaur tracks formation and preservation in the Barremian coastal-alluvial to tidal-flat deposits of SE Spain (W Penyagolosa Sub-basin, Maestrat Basin). In: The 9th International Conference on Tidal Sedimentology (Ed.), Abstracts Tidalites (pp. 117–120). Puerto Madryn, Patagonia, Argentina: Asociación Argentina de Sedimentología y CONICET/CENPAT.
Bertozzo, F., Dalla-Vecchia, F. M., & Fabbri, M. (2017). The Venice specimen of Ouranosaurus nigeriensis (Dinosauria, Ornithopoda). PeerJ, 5, e3403. https://doi.org/10.7717/peerj.3403
Brochu, C. (1996). Closure of neurocentral sutures during crocodilian ontogeny: Implications for maturity assessment in fossil archosaurs. Journal of Vertebrate Paleontology, 16(1), 49–62.
Campos-Soto, S., Benito, M. I., Cobos, A., Caus, E., Quijada, I. E., Suarez-González, P., et al. (2019). Revisiting the age and palaeoenvironments of the Upper Jurassic-Lower Cretaceous? Dinosaur-bearing sedimentary record of eastern Spain: Implications for Iberian palaeogeography. Journal of Iberian Geology, 45, 471–510.
Campos-Soto, S., Cobos, A., Caus, E., Benito, M. I., Fernández-LabradorSuarez-González, L. P., et al. (2017). Jurassic Coastal Park: A great diversity of palaeoenvironments for the dinosaurs of the Villar del Arzobispo Formation (Teruel, eastern Spain). Palaeogeography, Palaeoclimatology, Palaeoecology, 485, 154–177. https://doi.org/10.1016/j.palaeo.2017.06.010
Carpenter, K., & Wilson, Y. (2008). A new species of Camptosaurus (Ornithopoda: Dinosauria) from the Morrison Formation (Upper Jurassic) of Dinosaur National Monument, Utah, and a biomechanical analysis of its forelimb. Annals of Carnegie Museum, 76(4), 227–263.
Cobos, A., & Gascó, F. (2012). Presencia del género Iguanodontipus en el Cretácico Inferior de la provincia de Teruel (España). Geogaceta, 52, 185–188.
Cobos, A., Gascó, F., Royo-Torres, R., & Alcalá, L. (2013). “Dinoichnofacies” barremienses en Teruel (España). In C. Álvarez-Vázquez & I. López-Rodríguez (Eds.), Libro de Resúmenes de las XXIX Jornadas de la Sociedad Española de Paleontología (pp. 65–66). Real Jardín Botánico de Córdoba.
Cobos, A., Gascó, F., Royo-Torres, R., Lockley, M. G., & Alcalá, L. (2016). Dinosaur tracks as “four-dimensional phenomena” reveal how different species moved. In P. L. Falkingham, D. Marty, & A. Richter (Eds.), Dinosaur tracks: The next steps (pp. 244–256). Indiana University Press.
Cobos, A., Royo-Torres, R., & Alcalá, L. (2012). Restos postcraneales de ornitópodo en el Cretácico Inferior de El Castellar (Subcuenca de Peñagolosa, Teruel). Geogaceta, 52, 181–184.
Cuesta, E., Vidal, D., Ortega, F., Shibata, M., & Sanz, J. L. (2021). Pelecanimimus (Theropoda: Ornithomimosauria) postcranial anatomy and the evolution of the specialized manus in ornithomimosaurs and sternum in maniraptoriforms. Zoological Journal of the Linnean Society. https://doi.org/10.1093/zoolinnean/zlab013
Díaz-Martínez, I., Pereda-Suberbiola, X., Pérez-Lorente, F., & Canudo, J. I. (2015). Ichnotaxonomic review of large ornithopod dinosaur tracks: Temporal and geographic implications. PLoS ONE, 10(2), e0115477. https://doi.org/10.1371/journal.pone.0115477
Dieudonné, P. E., Tortosa, T., Fernández-Baldor, F. T., Canudo, J. I., & Díaz-Martínez, I. (2016). An unexpected early rhabdodontid from Europe (Lower Cretaceous of Salas de los Infantes, Burgos Province, Spain) and a re-examination of basal iguanodontian relationships. PLoS ONE, 11(6), e0156251.
Dollo, L. (1888). Iguanodontidae et Camptonotidae. Comptes Rendus De Academie Des Sciences, Paris, 106, 775–777.
Fernández-Baldor, F. T., Canudo, J. I., Huerta, P., Montero, D., Suberbiola, X. P., & Salgado, L. (2011). Demandasaurus darwini, a new rebbachisaurid sauropod from the Early Cretaceous of the Iberian Peninsula. Acta Palaeontologica Polonica, 56(3), 535–552.
Figueiredo, S. D., Carvalho, I. S., Pereda-Suberbiola, X., Cunha, P. P., Antunes, V., & Díaz-Martínez, I. (2022). New ornithopod footprints from the Areia do Mastro Formation (Lower Cretaceous), Espichel Cape (Portugal, Western Iberia) and their context in the Iberian ichnological ornithopod record. Cretaceous Research. https://doi.org/10.1016/j.cretres.2021.105069
Fuentes-Vidarte, C., Meijide-Calvo, M., Meijide-Fuentes-Vidarte, F., & Meijide-Fuentes-Vidarte, M. (2016). Un nuevo dinosaurio estiracosterno (Ornithopoda: Ankylopollexia) del Cretácico Inferior de España. Spanish Journal of Paleontology, 31(2), 407–446.
Galton, P. M. (1974). The ornithischian dinosaur Hypsilophodon from the Wealden of the Isle of Wight. Bulletin of the British Museum (Natural History). Geology, 25, 1–152.
Galton, P. M. (2009). Notes on Neocomian (Lower Cretaceous) ornithopod dinosaurs from England—Hypsilophodon, Valdosaurus, “Camptosaurus”, “Iguanodon”—and referred specimens from Romania and elsewhere. Revue De Paléobiologie, 28, 211–273.
Galton, P. M. (2012). Hypsilophodon foxii and other smaller bipedal ornithischian dinosaurs from the Lower Cretaceous of Southern England. In P. Godefroit (Ed.), Bernissart dinosaurs and Early Cretaceous terrestrial ecosystems (pp. 225–281). Indiana University Press.
Gasca, J. M. (2011). Descripción de una tibia de dinosaurio ornitópodo de la Formación El Castellar (Cretácico Inferior, Teruel). In J. Marigó, M. Pérez de los Ríos, D. de Miguel, & A. Bolet (Eds.), Libro de Resúmenes de las XXVII Jornadas de la Sociedad Española de Paleontología (pp. 179–183). Sabadell: Institut Català de Paleontologia Miquel Crusafont.
Gasca, J. M., Canudo, J. I., & Moreno-Azanza, M. (2014a). On the diversity of Iberian iguanodont dinosaurs: New fossils from the lower Barremian, Teruel province, Spain. Cretaceous Research, 50, 264–272. https://doi.org/10.1016/j.cretres.2014.05.009
Gasca, J. M., Canudo, J. I., & Moreno-Azanza, M. (2014b). A large-bodied theropod (Tetanurae: Carcharodontosauria) from the Mirambel Formation (Barremian) of Spain. Neues Jahrbuch Für Geologie Und Paläontologie-Abhandlungen, 273(1), 13–23.
Gasca, J. M., Díaz-Martínez, I., Moreno-Azanza, M., Canudo, J. I., & Alonso, A. (2018). A hypertrophied ungual phalanx from the lower Barremian of Spain: Implications for the diversity and palaeoecology of Spinosauridae (Theropoda) in Iberia. Cretaceous Research, 84, 141–152.
Gasca, J. M., Moreno-Azanza, M., & Canudo, J. I. (2008). Dientes de dinosaurios terópodos espinosáuridos de la Formación El Castellar (Cretácico Inferior, Teruel). Paleontologica Nova, SEPAZ, 8, 233–234.
Gasulla, J. M., Escaso, F., Narváez, I., Ortega, F., & Sanz, J. L. (2015). A new sail-backed styracosternan (Dinosauria: Ornithopoda) from the Early Cretaceous of Morella, Spain. PLoS ONE, 10(12), e0144167. https://doi.org/10.1371/journal.pone.0144167
Gasulla, J. M., Escaso, F., Ortega, F., & Sanz, J. L. (2014). New hadrosauriform cranial remains from the Arcillas de Morella Formation (lower Aptian) of Morella, Spain. Cretaceous Research, 47, 19–24. https://doi.org/10.1016/j.cretres.2013.10.004
Gasulla, J. M., Ortega, F., Pereda-Suberbiola, X., Escaso, F., & Sanz, J. L. (2011). Elementos de la armadura dérmica del dinosaurio anquilosaurio Polacanthus Owen, 1865, en el Cretácico Inferior de Morella (Castellón, España). Ameghiniana, 48(4), 508–519.
Gilmore, C. W. (1909). Osteology of the Jurassic reptile Camptosaurus, with a revision of the species of the genus, and descriptions of two new species. Proceedings of the United States National Museum, 36, 197–332.
Godefroit, P., Dong, Z. M., Bultynck, P., Li, H., & Feng, L. (1998). New Bactrosaurus (Dinosauria: Hadrosauroidea) material from Iren Dabasu (Inner Mongolia, PR China). Bulletin De L’institut Royal Des Sciences Naturelles De Belgique, Sciences De La Terra, 68(Supplement), 3–70.
Griffin, C. T., Stocker, M. R., Colleary, C., Stefanic, C. M., Lessner, E. J., Riegler, M., et al. (2020). Assessing ontogenetic maturity in extinct saurian reptiles. Biological Reviews. https://doi.org/10.1111/brv.12666
Guerrero, A., & Cobos, A. (2017). Nuevos fósiles de dinosaurios cretácicos del yacimiento La Balsa 2 (El Castellar, Teruel, España). In S. Barrios de Pedro, C. Blanco-Moreno, A. de Celis, J. Colmenar, E. Cuesta, D. García-Martínez, et al. (Eds.), A Glimpse of the Past (pp. 15–19). Lisboa: Abstract book of the XV Encuentro de Jóvenes Investigadores en Paleontología.
Hammer, Ø., Harper, D. A. T., & Ryan, P. D. (2001). PAST: Paleontological statistics software package for education and data analysis. Palaeontologia Electronica, 4(1), 1–9.
Horner, J. R., Weishampel, D. B., & Forster, C. A. (2004). Hadrosauridae. In D. B. Weishampel, P. Donson, & H. Osmólska (Eds.), The Dinosauria (2nd ed., pp. 438–463). University of California Press.
Hübner, T. (2018). The postcranial ontogeny of Dysalotosaurus lettowvorbecki (Ornithischia: Iguanodontia) and implications for the evolution of ornithopod dinosaurs. Palaeontographica Abteilung A, 310, 43–120. https://doi.org/10.1127/pala/2018/0072
Ikejiri, T. (2012). Histology-based morphology of the neurocentral synchondrosis in Alligator mississippiensis (Archosauria, Crocodylia). The Anatomical Record: Advances in Integrative Anatomy and Evolutionary Biology, 295(1), 18–31.
Ikejiri, T. (2015). Modes of ontogenetic allometric shifts in crocodilian vertebrae. Biological Journal of the Linnean Society, 116, 649–670.
Isasmendi, E., Sáez-Benito, P., Torices, A., Navarro-Lorbés, P., & Pereda-Suberbiola, X. (2020). New insights about theropod palaeobiodiversity in the Iberian Peninsula and Europe: Spinosaurid teeth (Theropoda, Megalosauroidea) from the Lower Cretaceous of La Rioja (Spain). Cretaceous Research, 116, 104600. https://doi.org/10.1016/j.cretres.2020.104600
Knoll, F. (2009). A large iguanodont from the upper Barremian of the Paris Basin. Geobios, 42, 755–764. https://doi.org/10.1016/j.geobios.2009.06.002
Leonardi, G., et al. (1984). Le impronte fossili dei dinosauri. In J. F. Bonaparte, E. H. Colbert, P. H. Currie, A. de Ricqles, Z. Kielan-Jaworowska, & G. Leonardi (Eds.), Sulle Orme dei dinosauri (pp. 161–186). Erizzo Editrice.
Llandres, M., Vullo, R., Marugán-Lobón, J., Ortega, F., & Buscalioni, A. D. (2013). An articulated hindlimb of a basal iguanodont (Dinosauria, Ornithopoda) from the Early Cretaceous Las Hoyas Lagerstätte (Spain). Geological Magazine, 150(3), 572–576. https://doi.org/10.1017/S0016756813000095
Lockley, M. G., Xing, L., Lockwood, J. A. F., & 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.
Lockley, M. G., Young, B. H., & Carpenter, K. (1983). Hadrosaur locomotion and herding behavior: Evidence from footprints in the Mesaverde Formation, Grand Mesa Coal Field, Colorado. The Mountain Geologist, 20(4), 5–14.
Lockwood, J. A. F., Martill, D. M., & Maidment, C. R. (2021). A new hadrosauriform dinosaur from the Wessex Formation, Wealden Group (Early Cretaceous), of the Isle of Wight, southern England. Journal of Systematic Palaeontology. https://doi.org/10.1080/14772019.2021.1978005
Luque, L., Espílez, E., Cobos, A., Alcalá, L., Royo-Torre, R., Aberasturi, A., et al. (2006). Iguanodóntidos y otros vertebrados cretácicos de Miravete de la Sierra (Teruel). Teruel, 91(1), 57–87.
Madzia, D., Arbour, V. M., Boyd, C. A., Farke, A. A., Cruzado-Caballero, P., & Evans, D. C. (2021). The phylogenetic nomenclature of ornithischian dinosaurs. PeerJ, 9, e12362. https://doi.org/10.7717/peerj.12362
Malafaia, E., Gasulla, J. M., Escaso, F., Narvaéz, I., & Ortega, F. (2020a). An update of the spinosaurid (Dinosauria: Theropoda) fossil record from the Lower Cretaceous of the Iberian Peninsula: Distribution, diversity, and evolutionary history. Journal of Iberian Geology, 46(4), 431–444.
Malafaia, E., Gasulla, J. M., Escaso, F., Narváez, I., Sanz, J. L., & Ortega, F. (2020b). A new spinosaurid theropod (Dinosauria: Megalosauroidea) from the upper Barremian of Vallibona, Spain: Implications for spinosaurid diversity in the Early Cretaceous of the Iberian Peninsula. Cretaceous Research, 106, 104221. https://doi.org/10.1016/j.cretres.2019.104221
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, 115, 179–186. https://doi.org/10.1098/rstl.1825.0010
Marsh, O. (1881). Classification of the Dinosauria. American Journal of Sciences, 3, 81–86.
Martín-Closas, C. (1989). Els caròfits del Cretaci Inferior de les conques perifèriques del Bloc de l’Ebre. PhD Thesis, Universidad de Barcelona, pp. 1–581.
McDonald, A. T. (2011). The taxonomy of species assigned to Camptosaurus (Dinosauria: Ornithopoda). Zootaxa, 2783(1), 52–68. https://doi.org/10.11646/zootaxa.2783.1.4
McDonald, A. T., Bird, J., Kirkland, J. I., & Dodson, P. (2012). Osteology of the basal hadrosauroid Eolambia caroljonesa (Dinosauria: Ornithopoda) from the Cedar Mountain Formation of Utah. PLoS ONE, 7, e45712.
McDonald, A. T., Kirkland, J. I., DeBlieux, D. D., Madsen, S. K., Cavin, J., et al. (2010). New basal iguanodonts from the Cedar Mountain Formation of Utah and the evolution of thumb-spiked dinosaurs. PLoS ONE, 5(11), e14075. https://doi.org/10.1371/journal.pone.0014075
McDonald, A. T., Maidment, S. C., Barrett, P. M., You, H. L., & Dodson, P. (2014). Osteology of the basal hadrosauroid Equijubus normani (Dinosauria, Ornithopoda) from the Early Cretaceous of China. In D. A. Eberth & D. C. Evans (Eds.), Hadrosaurs (pp. 3–34). Indiana University Press.
Meyer, C. A., & Thüring, B. (2003). The first iguanodontid dinosaur tracks from the Swiss Alps (Schrattenkalk Formation, Aptian). Ichnos, 10, 221–228.
Norman, D. B. (1980). On the ornithischian dinosaurs Iguanodon bernissartensis from the Lower Cretaceous of Bernissart (Belgium). Memoires De L’institut Royal Des Sciences Naturelles De Belgique, 178, 1–103.
Norman, D. B. (1986). On the anatomy of Iguanodon atherfieldensis (Ornithischia: Ornithopoda). Bulletin De L’institut Royal Des Sciences Naturelles De Belgique, 56, 281–372.
Norman, D. B. (2002). On Asian ornithopods (Dinosauria: Ornithischia). Probactrosaurus Rozhdestvensky, 1966. Zoological Journal of the Linnean Society, 136, 113–144.
Norman, D. B. (2004). Basal iguanodontia. In D. B. Weishampel, P. Donson, & H. Osmólska (Eds.), The Dinosauria (pp. 413–437). University of California Press.
Norman, D. B. (2011). On the osteology of the lower Wealden (Valanginian) iguanodontian Barilium dawsoni (Iguanodontia: Styracosterna). Special Papers in Palaeontology, Palaeontological Association, 86, 165–198.
Norman, D. B. (2012). Iguanodontian taxa (Dinosauria: Ornithischia) from the Lower Cretaceous of Britain and Belgium. In P. Godefroit (Ed.), Bernissart dinosaurs and Early Cretaceous terrestrial ecosystems (pp. 174–212). Indiana University Press.
Norman, D. B. (2014). Iguanodonts from the Wealden of England: Do they contribute to the discussion concerning hadrosaur origins? In D. A. Eberth & D. C. Evans (Eds.), Hadrosaurs (life of the past) (pp. 10–43). Indiana University Press.
Norman, D. B. (2015). On the history, osteology, and systematic position of the Wealden (Hastings group) dinosaur Hypselospinus fittoni (Iguanodontia: Styracosterna). Zoological Journal of the Linnean Society, 173(1), 92–189.
Ortega, F., Escaso, F., & Sanz, J. L. (2010). A bizarre, humped Carcharodontosauria (Theropoda) from the Lower Cretaceous of Spain. Nature, 467, 203–206.
Ösi, A., Prondvai, E., Butler, R., & Weishampel, D. B. (2012). Phylogeny, histology and inferred body size evolution in a new rhabdodontid dinosaur from the Late Cretaceous of Hungary. PLoS ONE, 7(9), e44318. https://doi.org/10.1371/journal.pone.0044318
Owen, R. (1842). Report on British fossil reptiles, Part II. In: Report of the Eleventh Meeting of the British Association Advancement of the Science Vol. 1841 (pp. 60–204).
Paul, G. S. (2008). A revised taxonomy of the iguanodont dinosaur genera and species. Cretaceous Research, 29, 192–216. https://doi.org/10.1016/j.cretres.2007.04.009
Pereda-Suberbiola, X., Fuentes, C., Meijide, M., Meijide-Fuentes, F., & Meijide-Fuentes, M., Jr. (2007). New remains of the ankylosaurian dinosaur Polacanthus from the Lower Cretaceous of Soria, Spain. Cretaceous Research, 28(4), 583–596.
Pereda-Suberbiola, X., Ruiz-Omeñaca, J. I., Canudo, J. I., Torcida, F., & Sanz, J. L. (2012). Dinosaur faunas from the Early Cretaceous (Valanginian-Albian) of Spain. In P. Godefroit (Ed.), Bernissart dinosaurs and early cretaceous terrestrial ecosystems (pp. 379–407). Indiana University Press.
Pereda-Suberbiola, X., Ruiz-Omeñaca, J. I., Fernández-Baldor, F. T., Maisch, M. W., Huerta, P., Contreras, R., et al. (2011). A tall-spined ornithopod dinosaur from the Early Cretaceous of Salas de los Infantes (Burgos, Spain). Comptes Rendus Palevol, 10, 551–558. https://doi.org/10.1016/j.crpv.2011.04.003
Pérez-García, A., Cobos, A., & Royo-Torres, R. (2017). The oldest evidence of a dortokid turtle (stem Pleurodira) from the uppermost Hauterivian-basal Barremian El Castellar Formation (Teruel, Spain). Journal of Iberian Geology, 43(2), 139–146.
Pérez-García, A., Scheyer, T. M., & Murelaga, X. (2013). The turtles from the uppermost Jurassic and Early Cretaceous of Galve (Iberian Range, Spain): Anatomical, systematic, biostratigraphic and palaeobiogeographical implications. Cretaceous Research, 44, 64–82.
Pérez-Moreno, B. P., Sanz, J. L., Buscalioni, Á. D., Moratalla, J. J., Ortega, F., & Rasskin-Gutman, D. (1994). A unique multitoothed ornithomimosaur dinosaur from the Lower Cretaceous of Spain. Nature, 370, 363–367.
Prieto-Márquez, A., & Norell, M. A. (2010). Anatomy and relationships of Gilmoreosaurus mongoliensis (Dinosauria: Hadrosauroidea) from the Late Cretaceous of Central Asia. American Museum Novitates, 3694, 1–49.
Royo-Torres, R., & Cobos, A. (2007). Teeth of Oplosaurus armatus (Sauropoda) from El Castellar (Teruel, Spain). In: J. Le Loeuf, (Ed.), Abstract book of European Association of Vertebrate Paleontologists 5th anual meeting (pp. 52–55). Francia: Carcassonne.
Royo-Torres, R., Fuentes, C., Meijide, M., Meijide-Fuentes, F., & Meijide-Fuentes, M. (2017). A new brachiosauridae sauropod dinosaur from the Lower Cretaceous of Europe (Soria province, Spain). Cretaceous Research, 80, 38–55. https://doi.org/10.1016/j.cretres.2017.08.012
Ruiz-Omeñaca, J. I., Canudo, J. I., Cuenca-Bescós, G., Cruzado-Caballero, P. L., Gasca, J. M., & Moreno-Azanza, M. (2012). A new basal ornithopod dinosaur from the Barremian of Galve, Spain. Comptes Rendus Palevol, 11(6), 435–444. https://doi.org/10.1016/j.crpv.2012.06.001
Salas, R. (1987). El Malm i el Cretaci inferior entre el Massif de Garraf i la Serra d´Espadà. Anàlisi de conca. PhD Thesis, Universidad de Barcelona, pp. 1–365.
Salas, R., & Guimerà, J. (1996). Rasgos estructurales principales de la cuenca Cretácica Inferior del Maestrazgo (Cordillera Ibérica oriental). Geogaceta, 20, 1704–1706.
Salas, R., Guimerà, J., Mas, R., Martín-Closas, C., Meléndez, A., & Alonso, A. (2001). Peri-Tethys Memoir 6: Peri-Tethyan Rift/Wrench Basins and passive margins. In P. A. Ziegler, W. Cavazza, A. F. H. Robertson, & S. Crasquin-Soleau (Eds.), Mémoires du Muséum national d‘Histoire naturelle (pp. 145–185). Editions du Muséum.
Sanguino, F., & Buscalioni, Á. D. (2018). The Iguanodon locality of Barranco Pata la Mona (upper Barremian, Buenache de la Sierra, Cuenca) revisited. In E. Amayuelas, P. Bilbao-Lasa, O. Bonilla, M. del Val, J. Errandonea-Martin, I. Garate-Olave, et al. (Eds.), Life finds a way (pp. 103–106). Gasteiz: Abstract book of the XVI Encuentro de Jóvenes Investigadores en Paleontología.
Santos-Cubedo, A., de Santisteban, C., Poza, B., & Meseguer, S. (2021). A new styracosternan hadrosauroid (Dinosauria: Ornithischia) from the Early Cretaceous of Portell, Spain. PLoS ONE, 16(7), e0253599. https://doi.org/10.1371/journal.pone.0253599
Sanz, J. L., Casanovas, M. I., & Santafé, J. V. (1982). Paleontología. In J. V. Santafé, M. L. Casanovas, J. L. Sanz, & S. Calzada (Eds.), Geología y paleontología (dinosaurios) de las capas rojas de Morella (Castellón, España) (pp. 69–169). Diputación Provincial de Castellón y Diputación de Barcelona.
Schudack, U., & Schudack, M. (2009). Ostracod biostratigraphy in the Lower Cretaceous of the Iberian chain (eastern Spain). Journal of Iberian Geology, 35, 141–168.
Seeley, H. G. (1887). On the classification of the fossil animals commonly named Dinosauria. Proceedings of the Royal Society of London, 43, 165–171. https://doi.org/10.1098/rspl.1887.0117
Sereno, P. C. (1986). Phylogeny of the bird-hipped dinosaurs (Order Ornithischia). National Geographic Research, 2, 234–256.
Sereno, P. C. (1997). The origin and evolution of dinosaurs. Annual Review of Earth and Planetary Sciences, 25(1), 435–489.
Sigovini, M., Keppel, E., & Tagliapietra, D. (2016). Open Nomenclature in the biodiversity era. Methods in Ecology and Evolution, 7(10), 1217–1225.
Taquet, P. (1976). Geologie et paleontologie du gisement de Gadoufaoua (Aptien du Niger), Cahiers Paléontologie (pp. 1–191). es: Centre National de la Recherche Scientifiqu.
Torrente, M. Á., Verdú, F. J., Cobos, A., & Alcalá, L. (2018). Diente de un coelurosaurio (Theropoda, Dinosauria) del Barremiense (Cretácico Inferior) de El Castellar (Teruel, España). Geogaceta, 63, 79–82.
Verdú, F. J. (2017). Sistemática, filogenia y paleobiología de Iguanodon galvensis (Ornithopoda, Dinosauria) del Barremiense inferior (Cretácico Inferior) de Teruel (España). PhD Thesis, Universitat de València, pp. 1–445.
Verdú, F. J., Cobos, A., Royo-Torres, R., & Alcalá, L. (2019). Diversity of large ornithopod dinosaurs in the Upper Hauterivian-Lower Barremian (Lower Cretaceous) of Teruel, Spain: A morphometric approach. Spanish Journal of Palaeontology, 34(2), 269–288.
Verdú, F. J., Godefroit, P., Royo-Torres, R., Cobos, A., & Alcalá, L. (2017). Individual variation in the postcranial skeleton of the Early Cretaceous Iguanodon bernissartensis (Dinosauria: Ornithopoda). Cretaceous Research, 74, 65–86. https://doi.org/10.1016/j.cretres.2017.02.006
Verdú, F. J., Royo-Torres, R., Cobos, A., & Alcalá, L. (2015). Perinates of a new species of Iguanodon (Ornithischia: Ornithopoda) from the lower Barremian of Galve (Teruel, Spain). Cretaceous Research, 56, 250–264.
Verdú, F. J., Royo-Torres, R., Cobos, A., & Alcalá, L. (2018). New systematic and phylogenetic data about the early Barremian Iguanodon galvensis (Ornithopoda: Iguanodontoidea) from Spain. Historical Biology, 30, 437–474. https://doi.org/10.1080/08912963.2017.1287179
Verdú, F. J., Royo-Torres, R., Cobos, A., & Alcalá, L. (2020). Systematics and paleobiology of a new articulated axial specimen referred to Iguanodon cf. galvensis (Ornithopoda, Iguanodontoidea). Journal of Vertebrate Paleontology, 40(6), 8202. https://doi.org/10.1080/02724634.2021.1878202
Vialov, O. S. (1988). On the classification of dinosaurian traces. Ezhegodnik Vsesoyuznogo Paleontologicheskogo Obshchestva, 31, 322–325.
Villanueva-Amadoz, U., Sender, L. M., Royo-Torres, R., Verdú, F. J., Pons, D., Alcalá, L., et al. (2015). Palaeobotanical remains associated with dinosaur fossils from the Camarillas Formation (Barremian) of Galve (Teruel, Spain). Historical Biology, 27, 374–388.
Weishampel, D. B., Jianu, C. M., Cziki, Z., & Norman, D. B. (2003). Osteology and phylogeny of Zalmoxes (n. g.), an unusual ornithopod dinosaur from the latest Cretaceous of Romania. Journal of Systematic Paleontology, 1, 123–143. https://doi.org/10.1017/S1477201903001032
Acknowledgements
This research was funded by the Departamento de Educación, Cultura y Deporte (Gobierno de Aragón), Research Group E04_20R FOCONTUR financed by Departamento de Ciencia, Universidad y Sociedad del Conocimiento (Gobierno de Aragón), the Instituto Aragonés de Fomento, the Ministerio de Ciencia e Innovación (MCI), Agencia Estatal de Investigación (AEI), and Fondo Europeo de Desarrollo Regional (FEDER) from Spanish Government (PGC2018-094034-B-C22 and Unidad de Paleontología de Dinosaurios de Teruel). The authors thank Pascal Godefroit, Annelise Folie and all the RBINS (Royal Belgian Institute of Natural Sciences) staff for providing access to the Bernissart iguanodontian collection on 2016. We appreciate useful comments and suggestions made by the associate editor Laura Domingo, and the two reviewers Fernando Escaso and anonymous. Finally, we would like to express our gratitude to Mr. Gabriel Fuentes Bellido, neighbour of Cabra de Mora, for donating part of the material studied here to the Museo Aragonés de Paleontología (MAP).
Funding
This work was supported by the Departamento de Educación, Cultura y Deporte (Gobierno de Aragón), Research Group E04_20R FOCONTUR financed by Departamento de Ciencia, Universidad y Sociedad del Conocimiento (Gobierno de Aragón), the Instituto Aragonés de Fomento, the Ministerio de Ciencia e Innovación (MCI), Agencia Estatal de Investigación (AEI), and Fondo Europeo de Desarrollo Regional (FEDER) from Spanish Government (PGC2018-094034-B-C22 and Unidad de Paleontología de Dinosaurios de Teruel).
Author information
Authors and Affiliations
Contributions
Not applicable for that section.
Corresponding author
Ethics declarations
Conflicts of interest
The authors declare that they have no conflict of interest.
Ethics approval
Not applicable for that section.
Consent to participate
Informed consent was obtained from all individual participants included in the study. All authors consent to the publication of the study.
Supplementary Information
Below is the link to the electronic supplementary material.
Rights and permissions
About this article
Cite this article
García-Cobeña, J., Verdú, F.J. & Cobos, A. Abundance of large ornithopod dinosaurs in the El Castellar Formation (Hauterivian-Barremian, Lower Cretaceous) of the Peñagolosa sub-basin (Teruel, Spain). J Iber Geol 48, 107–127 (2022). https://doi.org/10.1007/s41513-021-00185-w
Received:
Accepted:
Published:
Issue Date:
DOI: https://doi.org/10.1007/s41513-021-00185-w