Journal of Iberian Geology

, Volume 44, Issue 1, pp 7–23 | Cite as

A new contribution to our knowledge of the large-bodied theropods from the Barremian of the Iberian Peninsula: the “Barranco del Hocino” site (Spain)

  • A. Alonso
  • J. M. Gasca
  • P. Navarro-Lorbés
  • C. Rubio
  • J. I. Canudo
Research paper



Barranco del Hocino-1 is a new fossil site located near Estercuel, Teruel province, Spain. The fossil site is located geologically within the Oliete sub-basin, in the Blesa Formation (Barremian in age). Barranco del Hocino-1 shows a diverse assemblage of tetrapod vertebrates similar to other sites in the Blesa Formation.

Materials and methods

Six isolated teeth belonging to Theropoda have been found. A study of their qualitative and quantitative characters, along with statistical (DFA) and cladistic analyses, enable us to identify four different dental morphotypes.


These morphotypes belong to separate tetanuran theropod taxa. One is related to Spinosauridae. The other morphotypes show affinities with non-spinosaurid tetanurans, probably related to Carcharodontosauria.


The results are congruent with the known theropod record of the Iberian Peninsula and western Europe. This work is a new contribution to what is known of the palaeobiodiversity and distribution of large-bodied theropods from the Barremian of the Iberian Peninsula.


Lower Cretaceous Blesa Formation Theropoda Dinosaur teeth Spain 



Barranco del Hocino-1 es un nuevo yacimiento localizado en el entorno de Estercuel, provincial de Teruel (España). Geológicamente se sitúa en la Formación Blesa (subcuenca de Oliete) de edad Barremiense. El yacimiento presenta una asociación diversa de vertebrados similar a otros de la misma formación.

Materiales y métodos

Se han encontrado seis dientes aislados de dinosaurios terópodos. Mediante el estudio de los caracteres cualitativos y cuantitativos junto con el uso de análisis estadístico multivariante (DFA) y análisis cladístico se han podido identificar cuatro morfotipos diferentes.


Los morfotipos identificados pertenecen a diferentes grupos de tetanuros basales. Uno de los morfotipos está relacionado con Spinosauridae, mientras que el resto presentan afinidades con tetanuros no espinosáuridos, posiblemente relacionados con el clado Carcharodontosauria.


Los resultados son coherentes con el registro de terópodos conocido tanto en la península ibérica y Europa Occidental durante el Cretácico Inferior. El hallazgo supone una nueva contribución al conocimiento de la paleobiodiversidad y distribución de grandes terópodos del Barremiense de la península ibérica.

Palabras clave

Cretácico Inferior Formación Blesa Theropoda Dientes de dinosaurio España 



This paper forms part of the project CGL2014-53548 and is subsidized by the Spanish Ministry of Economy and Competitiveness, the European Regional Development Fund, and the Government of Aragón (Grupos Consolidados). Antonio Alonso is the recipient of a Ph.D. Grant from the DGA (Diputación General de Aragón). José Manuel Gasca is supported by the Ministerio de Ciencia, Tecnología e Innovación Productiva, Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET) of Argentina (Postdoctoral Fellowship). Pablo Navarro-Lorbés is supported by a Ph.D. fellowship of the Government of La Rioja. The authors thank the Town Council of Estercuel, the Caja Rural de Teruel and the Provincial Deputation of Teruel for the funding provided, the General Direction of Cultural Patrimony of the Government of Aragón for authorizing the fieldwork, and Javier Rubio of Paleoymás for the photographs of the theropod teeth. Miguel Moreno-Azanza helped with cladistic analyses and Carmen Núñez-Lahuerta provided help with photographs. We especially thank Juan Rubio and Australair SL for their help, funding and participation in the fieldwork. Finally, we thank Christophe Hendrickx, Julio Company and the anonymous reviewers for the useful comments that resulted in substantial improvements to the manuscript, and Rupert Glasgow, who edited the text in English.

Supplementary material

41513_2018_51_MOESM1_ESM.xlsx (445 kb)
Supplementary material 1 (XLSX 444 kb)
41513_2018_51_MOESM2_ESM.docx (61 kb)
Supplementary material 2 (DOCX 60 kb)


  1. Allain, R. (2005). The enigmatic theropod dinosaur Erectopus superbus (Sauvage 1882) from the Lower Albian of Louppy-le-Chateau. In Carpenter, K. (Ed.), The carnivorous dinosaurs (pp. 72–86). Bloomington, Indiana: Indiana University Press.Google Scholar
  2. Alonso, A., & Canudo, J. I. (2016). On the spinosaurid theropod teeth from the early Barremian (Early Cretaceous) Blesa Formation (Spain). Historical Biology, 28(6), 823–834.CrossRefGoogle Scholar
  3. Alonso, A., Gasca, J. M., Navarro-Lorbés, P., Núñez-Lahuerta, C., Galán, J., Parrilla-Bel, J., et al. (2016). La asociación faunística de Barranco del Hocino 1, un nuevo yacimiento de vertebrados del Barremiense (Cretácico Inferior) de Teruel. Cuadernos del Museo Geominero, 20, 303–307.Google Scholar
  4. Badiola, A., Canudo, J. I., & Cuenca-Bescós, C. (2008). New multituberculate mammals from the Hauterivian/Barremian transition of Europe (Iberian Peninsula). Palaeontology, 51(6), 1455–1469.CrossRefGoogle Scholar
  5. Benson, R. B., Brusatte, S. L., Hutt, S., & Naish, D. (2009). A new large basal tetanuran (Dinosauria: Theropoda) from the Wessex Formation (Barremian) of the isle of Wight, England. Journal of Vertebrate Paleontology, 29(2), 612–615.CrossRefGoogle Scholar
  6. Brusatte, S. L., Benson, R. B., Carr, T. D., Williamson, T. E., & Sereno, P. C. (2007). The systematic utility of theropod enamel wrinkles. Journal of Vertebrate Paleontology, 27, 1052–1056.CrossRefGoogle Scholar
  7. Brusatte, S. L., Carr, T. D., & Norell, M. A. (2012). The osteology of Alioramus, a gracile and long-snouted tyrannosaurid (Dinosauria: Theropoda) from the Late Cretaceous of Mongolia. Bulletin of the American Museum of Natural History, 366, 1–197.CrossRefGoogle Scholar
  8. Brusatte, S. L., Norell, M. A., Carr, T. D., Erickson, G. M., Hutchinson, J. R., Balanoff, A. M., et al. (2010). Tyrannosaur paleobiology: New research on ancient exemplar organisms. Science, 329, 1481–1485.CrossRefGoogle Scholar
  9. Brusatte, S. L., & Sereno, P. C. (2007). A new species of Carcharodontosaurus (Dinosauria: Theropoda) from the Cenomanian of Niger and a revision of the genus. Journal of Vertebrate Paleontology, 27(4), 902–916.CrossRefGoogle Scholar
  10. Brusatte, S. L. & Sereno, P. C. (2008). Phylogeny of Allosauroidea (Dinosauria: Theropoda): Comparative analysis and resolution. Journal of Systematic Palaeontology, 6(2), 155–182.CrossRefGoogle Scholar
  11. Buffetaut, E. (2007). The spinosaurid dinosaur Baryonyx (Saurischia, Theropoda) in the Early Cretaceous of Portugal. Geological Magazine, 144(6), 1021–1025.CrossRefGoogle Scholar
  12. Buckley, L. G. (2009). Individual and ontogenetic variation in theropod dinosaur teeth: A case study of Coelophysis bauri (Theropoda: Coelophysoidea) and implications for identifying isolated theropod teeth (pp. 109). MSc. Dissertation, University of Alberta, Edmonton, Alberta, Canada.Google Scholar
  13. Canudo, J. I., Gasca, J. M., Aurell, M., Badiola, A., Blain, H.-A., Cruzado-Caballero, P., et al. (2010). La Cantalera: An exceptional window onto the vertebrate biodiversity of the Hauterivian–Barremian transition in the Iberian Peninsula. Journal of Iberian Geology, 36(2), 205–224.CrossRefGoogle Scholar
  14. Canudo, J. I., Gasca, J. M., Moreno-Azanza, M., & Aurell, M. (2012). New information about the stratigraphic position and age of the sauropod Aragosaurus ischiaticus from the Early Cretaceous of the Iberian Peninsula. Geological Magazine, 149(2), 252–263.CrossRefGoogle Scholar
  15. Canudo, J. I., Gasulla, J. M., Gómez-Fernández, D., Ortega, F., Sanz, J. L., & Yagüe, P. (2008a). Primera evidencia de dientes aislados atribuidos a Spinosauridae (Theropoda) en el Aptiano inferior (Cretácico Inferior) de Europa: Formación Arcillas de Morella (España). Ameghiniana, 45(3), 649–652.Google Scholar
  16. Canudo, J. I., Royo-Torres, R., & Cuenca-Bescós, G. (2008b). A new sauropod: Tastavinsaurus sanzi gen. et sp. nov. from the Early Cretaceous (Aptian) of Spain. Journal of Vertebrate Paleontology, 28(3), 712–731.CrossRefGoogle Scholar
  17. Charig, A. J., & Milner, A. C. (1997). Baryonyx walkeri, a fish-eating dinosaur from the Wealden of Surrey. Bulletin of the Natural History Museum of London, 53, 11–70.Google Scholar
  18. Csiki-Sava, Z., Brusatte, S. L., & Vasile, S. (2016). “Megalosaurus cf. superbus” from southeastern Romania: The oldest known Cretaceous carcharodontotosaurid (Dinosauria: Theropoda) and its implications for earliest Cretaceous Europe-Gondwana connections. Cretaceous Research, 60, 221–238.CrossRefGoogle Scholar
  19. Currie, P. J., Rigby, J. K. J., & Sloan, R. E. (1990). Theropod teeth from the Judith River Formation of southern Alberta, Canada. In K. Carpenter & P. J. Currie (Eds.), Dinosaur systematics: Approaches and perspectives (pp. 107–125). New York: Cambridge University Press.CrossRefGoogle Scholar
  20. Fanti, F., Cau, A., Martinelli, A., & Contessi, M. (2014). Integrating palaeoecology and morphology in theropod diversity estimation: A case from the Aptian-Albian of Tunisia. Palaeogeography, Palaeoclimatology, Palaeoecology, 410, 39–57.CrossRefGoogle Scholar
  21. Gasca, J. M., Canudo, J. I., & Moreno-Azanza, M. (2014a). On the Iberian iguanodont dinosaur diversity: New fossils from the lower Barremian, Teruel province, Spain. Cretaceous Research, 50, 264–272.CrossRefGoogle Scholar
  22. 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.CrossRefGoogle Scholar
  23. Gasca, J. M., Moreno-Azanza, M., & Canudo, J. I. (2008). Dientes de dinosaurios terópodos espinosáuridos de la Formación El Castellar (Cretácico Inferior, Teruel). Palaeontologica Nova SEPAZ, 8, 233–234.Google Scholar
  24. 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.CrossRefGoogle Scholar
  25. Gauthier, J. A. (1986). Saurischian monophyly and the origin of birds. In K. Padian (Ed.), The origin of birds and the evolution of flight. Memoirs of the California Academy of Sciences (Vol. 8, pp. 1–55). San Francisco: California Academy of Sciences.Google Scholar
  26. Gerke, O., & Wings, O. (2016). Multivariate and cladistic analyses of isolated teeth reveal sympatry of theropod dinosaurs in the Late Jurassic of northern Germany. PLoS One, 11(7), e0158334.CrossRefGoogle Scholar
  27. Goloboff, P. A., & Catalano, S. A. (2016). TNT versión 1.5, including a full implementation of phylogenetic morphometrix. Cladistics, 32, 221–238. Scholar
  28. Hammer, Ø., & Harper, D. A. T. (2006). Paleontological data analysis (p. 351). Oxford: Blackwell.Google Scholar
  29. 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.Google Scholar
  30. Harris, J. D. (1998). A reanalysis of Acrocanthosaurus atokensis, its phylogenetic status, and paleobiogeographic implications, based on a new specimen from Texas. New Mexico Museum of natural History and Science Bulletin, 13, 1–75.Google Scholar
  31. Hendrickx, C. (2015). Evolution of Teeth and Quadrate in Non-avian Theropoda (Dinosauria: Saurischia), with the Description of Torvosaurus Remains from Portugal. Ph.D. Dissertation. Universidade Nova de Lisboa, p. 646.Google Scholar
  32. Hendrickx, C., & Mateus, O. (2014a). Torvosaurus gurneyi n. sp., the largest terrestrial predator from Europe, and a proposed terminology of the maxilla anatomy in nonavian theropods. PLoS One, 9(3), e88905. Scholar
  33. Hendrickx, C., & Mateus, O. (2014b). Abelisauridae (Dinosauria: Theropoda) from the Late Jurassic of Portugal and dentition-based phylogeny as a contribution for the identification of isolated theropod teeth. Zootaxa, 3759(1), 1–74.CrossRefGoogle Scholar
  34. Hendrickx, C., Mateus, O., & Araújo, R. (2015a). A proposed terminology of theropod teeth (Dinosauria, Saurischia). Journal of Vertebrate Paleontology, 35(5), e982797.CrossRefGoogle Scholar
  35. Hendrickx, C., Mateus, O., & Araújo, R. (2015b). The dentition of megalosaurid theropods. Acta Palaeontologica Polonica, 60(3), 627–642.Google Scholar
  36. Hocknull, S. A., White, M. A., Tischler, T. R., Cook, A. G., Calleja, N. D., Sloan, T., et al. (2009). New mid-Cretaceous (Latest Albian) dinosaurs from Winton, Queensland, Australia. PLoS One, 4(7), e6190.CrossRefGoogle Scholar
  37. Hutt, S., Martill, D. M., & Barker, M. J. (1996). The first European allosauroid dinosaur (Lower Cretaceous, Wealden Group, England. Neues Jahrbuch für Geologie und Paläontologie Monatshefte, 10, 635–644.Google Scholar
  38. Infante, P., Canudo, J. I., & Ruiz-Omeñaca, J. I. (2005). First evidence of theropod dinosaurs from the Mirambel Formation (Lower Barremian, Lower Cretaceous) from Castellote, Teruel. Geogaceta, 38, 31–34.Google Scholar
  39. Kellner, A. W. A., & Campos, D. A. (1996). First early Cretaceous theropod dinosaur from Brazil with comments on Spinosauridae. Neues Jahrbuch für Geologie und Paläontologie Abhandlungen, 199(2), 151–166.Google Scholar
  40. Mateus, O., Araújo, R., Natário, C., & Castanhinha, R. (2011). A new specimen of the theropod dinosaur Baryonyx from the early Cretaceous of Portugal and taxonomic validity of Suchosaurus. Zootaxa, 2827, 54–68.Google Scholar
  41. Moreno-Azanza, M., Canudo, J. I., & Gasca, J. M. (2014). Unusual theropod eggshells from the Early Cretaceous Blesa Formation of the Iberian Range, Spain. Acta Paleontologica Polonica, 59(4), 843–854.Google Scholar
  42. Norell, M. A., Makovicky, P. J., Bever, G. S., Balanoff, A. M., Clark, J. M., Barsbold, R., et al. (2009). A review of the Mongolian Cretaceous dinosaur Saurornithoides (Troodontidae: Theropoda). American Museum Novitates, 3654, 1–63.CrossRefGoogle Scholar
  43. Novas, F. E., de Valais, S., Vickers-Rich, P. A., & Rich, T. H. (2005). A large Cretaceous theropod from Patagonia, Argentina, and the evolution of carcharodontosaurids. Naturwissenschaften, 92, 226–230.CrossRefGoogle Scholar
  44. Ortega, F., Escaso, F., & Sanz, J. L. (2010). A bizarre, humped Carcharodontosauria (Theropoda) from the lower Cretaceous of Spain. Nature, 467(7312), 203–206.CrossRefGoogle Scholar
  45. Parrilla-Bel, J., & Canudo, J. I. (2015). On the presence of plesiosaurs in the Blesa Formation (Barremian) in Teruel (Spain). Neues Jahrbuch für Geologie und Paläontologie Abhandlungen, 278(2), 213–227.CrossRefGoogle Scholar
  46. 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 ecosystem (pp. 379–407). Bloomington: Indiana University Press.Google Scholar
  47. Pérez-Moreno, B. P., Sanz, J. L., Buscalioni, A. 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.CrossRefGoogle Scholar
  48. Puértolas-Pascual, E., Rabal-Garcés, R., & Canudo, J. I. (2015). Exceptional crocodylomorph biodiversity of “La Cantalera” site (lower Barremian; Lower Cretaceous) in Teruel, Spain. Palaeontologia Electronica, 18(2), 1–16.Google Scholar
  49. Rauhut, O. W. M. (2002). Dinosaur teeth from the Barremian of Uña, Province of Cuenca, Spain. Cretaceous Research, 23(2), 255–263.CrossRefGoogle Scholar
  50. Rauhut, O. W. M., & López-Arbarello, A. (2009). Considerations on the age of the Tiouaren Formation (Iullemmeden Basin, Niger, Africa): Implications for Gondwanan Mesozoic terrestrial vertebrate faunas. Palaeogeography, Palaeoclimatology, Palaeoecology, 271, 259–267.CrossRefGoogle Scholar
  51. Rauhut, O. W. M., Milner, A. C., & Moore-Fay, S. (2010). Cranial osteology and phylogenetic position of the theropod dinosaur Proceratosaurus bradleyi (Woolward, 1910) from the Middle Jurassic of England. Zoological Journal of the Linnean Society, 158, 155–195.CrossRefGoogle Scholar
  52. Rauhut, O. W. M., & Werner, C. (1995). First record of the family Dromaeosauridae (Dinosauria: Theropoda) in the Cretaceous of Gondwana (Wadi Milk Formation, northern Sudan). Paläontologische Zeitshrift, 69, 475–489.CrossRefGoogle Scholar
  53. Riveline, J., Berger, J. P., Feist, M., Martín-Closas, C., Schudack, M., & Soulié-Märsche, I. (1996). European Mesozoic-Cenozoic charophyte biozonation. Bulletin de la Société géologique de France, 167(3), 453–468.Google Scholar
  54. Ruiz-Omeñaca, J. I. (2011). Delapparentia turolensis nov. gen et sp., un nuevo dinosaurio iguanodontoideo (Ornithischia: Ornithopoda) en el Cretácico Inferior de Galve. Estudios Geológicos, 67(1), 83–110.CrossRefGoogle Scholar
  55. Ruiz-Omeñaca, J. I., Canudo, J. I., & Cuenca-Bescós, G. (1996). Dientes de dinosaurios (Ornitischia y Saurischia) del Barremiense superior (Cretácico inferior) de Vallipón (Castellote, Teruel). Mas de las Matas, 15, 59–103.Google Scholar
  56. Ruiz-Omeñaca, J. I., Canudo, J. I., & Cuenca-Bescós, G. (1998). Primera cita de dinosaurios barionícidos (Saurischia: Theropoda) en el Barremiense superior (Cretácico Inferior) de Vallipón (Castellote, Teruel). Mas de las Matas, 17, 201–223.Google Scholar
  57. 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 Paleovol, 11(6), 435–444.CrossRefGoogle Scholar
  58. Ruiz-Omeñaca, J. I., Cruzado-Caballero, P., Infante, P., & Moreno-Azanza, M. (2005). Baryonychine teeth (Theropoda: Spinosauridae) from the lower Cretaceous of La Cantalera (Josa, NE Spain). Kaupia, 14, 59–63.Google Scholar
  59. Salas, R., Guimerà, J., Más, R., Martín-Closas, C., Meléndez, A., & Alonso, A. (2001). Evolution of the Mesozoic central Iberian Rift System and its Cainozoic inversion (Iberian Chain). Mémoires du Muséum Nationale de l’Histoire Naturelle, Paris, 186, 145–185.Google Scholar
  60. Samman, T., Powell, G. L., Currie, P. J., & Hills, L. V. (2005). Morphometry of the teeth of western North American tyrannosaurids and its applicability to quantitative classification. Acta Palaeontologica Polonica, 50, 757–776.Google Scholar
  61. Sánchez-Hernández, B. R., & Benton, M. (2014). Filling the ceratosaur gap: A new ceratosaurian theropod from the Early Cretaceous of Spain. Acta Palaeontologica Polonica, 59(3), 581–600.Google Scholar
  62. Sánchez-Hernández, B., Benton, M. J., & Naish, D. (2007). Dinosaurs and other fossil vertebrates from the Late Jurassic and Early Cretaceous of the Galve area, NE Spain. Palaeogeography, Palaeoclimatology, Palaeoecology, 249, 180–215.CrossRefGoogle Scholar
  63. Sanz, J. L., Bonaparte, J. F., & Lacasa, A. (1988). Unusual Early Cretaceous birds from Spain. Nature, 331, 433–435.CrossRefGoogle Scholar
  64. Sanz, J.L., Buscalioni, A.D., Casanovas, M.L., Santafé, J.V. (1987). Dinosaurios del Cretácico Inferior de Galve (Teruel, España). Estudios geológicos. Volumen extraordinario Galve-Tremp, pp. 45–64.Google Scholar
  65. Sereno, P. C., Beck, A. L., Dutheil, D. B., Larsson, H. C. E., Lyon, G. H., Marcot, J. D., et al. (1998). A long-snouted predatory dinosaur from Africa and the evolution of spinosaurids. Science, 282(5392), 1298–1302.CrossRefGoogle Scholar
  66. Sereno, P. C., & Brusatte, S. L. (2008). Basal abelisaurid and carcharodontosaurid theropods from the lower Cretaceous Elrhaz Formation of Niger. Acta Palaeontologica Polonica, 53(1), 15–46.CrossRefGoogle Scholar
  67. Sereno, P. C., Dutheil, D. B., Iarochene, M., Larsson, H. C. E., Lyon, G. H., Magwene, P. M., et al. (1996). Predatory dinosaurs from the Sahara and Late Cretaceous faunal differentiation. Science, 272, 986–991.CrossRefGoogle Scholar
  68. Serrano-Martínez, A., Vidal, D., Sciscio, L., & Ortega, F. (2016). Isolated theropod teeth from the Middle Jurassic of Niger and the early dental evolution of Spìnosauridae. Acta Palaeontologica Polonica, 61(2), 403–415.Google Scholar
  69. Smith, J. B., & Dodston, P. (2003). A proposal for a standard terminology of anatomical notation and orientation in fossil vertebrate dentitions. Journal of Vertebrate Paleontology, 23(1), 1–12.CrossRefGoogle Scholar
  70. Smith, J. B., Vann, D. R., & Dodson, P. (2005). Dental morphology and variation in theropod dinosaurs: Implications for the taxonomic identification of isolated teeth. The Anatomical Record, 285A, 699–736.CrossRefGoogle Scholar
  71. Stromer, E. (1915). Ergebnisse der Forschungsreisen Prof. E. Stromers in den Wüsten Ägyptens. II. Wirbeltier-Reste der Baharîje-Stufe (unterstes Cenoman). 3. Das Original des Theropoden Spinosaurus aegyptiacus nov. gen., nov. spec. Abhandlungen der Königlich Bayerischen Akademie der Wissenschaften, Mathemtatisch-physikalische Klasse, 28(3), 1–32.Google Scholar
  72. Sues, H. D., Frey, E., Martill, D. M., & Scott, D. M. (2002). Irritator challengeri, a spinosaurid (Dinosauria: Theropoda) from the Lower Cretaceous of Brazil. Journal of Vertebrate Paleontology, 22(3), 535–547.CrossRefGoogle Scholar
  73. Torices, A., Currie, P., Canudo, J. I., & Pereda Suberbiola, X. (2015). Theropod dinosaurs from the Upper Cretaceous of the South Pyrenees Basin of Spain. Acta Paleontologica Polonica, 60(3), 611–626.Google Scholar
  74. 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.CrossRefGoogle Scholar
  75. White, M. A., Bell, P. R., Cook, A. G., Poropat, S. F., & Elliott, D. A. (2015). The dentary of Australovenator wintonensis (Theropoda, Megaraptoridae); implications for megaraptorid dentition. PeerJ, 3, e1512.CrossRefGoogle Scholar

Copyright information

© Springer International Publishing AG, part of Springer Nature 2018

Authors and Affiliations

  1. 1.Grupo Aragosaurus-IUCA, Área de Paleontología, Facultad de CienciasUniversidad de ZaragozaZaragozaSpain
  2. 2.CONICET-Museo OlsacherZapalaArgentina
  3. 3.Cátedra Extraordinaria de Paleontología, Departamento de Ciencias HumanasUniversidad de La RiojaLogroñoSpain
  4. 4.Paleoymás S.L. Pol. EmpresariumLa Cartuja BajaSpain
  5. 5.Museo de Ciencias Naturales de la Universidad de ZaragozaZaragozaSpain

Personalised recommendations