An update of the spinosaurid, Dinosauria, Theropoda, fossil record from the Lower Cretaceous of the Iberian Peninsuladistribution, diversity, and evolutionary history
- E. Malafaia 1
- J. M. Gasulla 2
- F. Escaso 2
- I. Narvaéz 2
- F. Ortega 2
-
1
Universidade de Lisboa
info
-
2
Universidad Nacional de Educación a Distancia
info
ISSN: 1886-7995, 1698-6180
Année de publication: 2020
Titre de la publication: Dinosaurs and other Biota: from Bones to Environments
Volumen: 46
Número: 4
Pages: 431-444
Type: Article
D'autres publications dans: Journal of iberian geology: an international publication of earth sciences
Résumé
The fossil record of spinosaurid theropods from the Lower Cretaceous of the Iberian Peninsula is relatively abundant, but not very informative. This record, as generally occurs in other geographic areas, is mainly represented by isolated teeth, probably due to the distinctive features of the dentition of these theropods. However, cranial and postcranial elements attributed to spinosaurids have been described and reported from different Iberian fossil sites. The currently known record of these theropods spans from the upper Hauterivian to the lower Aptian of the Cameros and Maestrazgo basins in Eastern Spain whereas it is early Barremian in age in the Lusitanian Basin of Western Portugal. Spinosaurid remains from the Iberian fossil record were traditionally attributed to Baryonyx, firstly described in the upper Barremian–lower Aptian of United Kingdom. However, the recent description of a specimen collected in the locality of Santa Águeda (Vallibona, Castellón), from sediments of the Morella Formation, implies the presence of a new taxon with a combination of derived features more closely related with some Gondwanan spinosaurines than with Baryonyx. Therefore, the currently known Iberian spinosaurid record would be made up of at least three taxa: Baryonyx (or a closely related baryonychine), the spinosaurine Vallibonavenatrix cani, and possibly Camarillasaurus cirugedae, which was recently re-interpreted as belonging to a megalosauroid tentatively related with spinosaurids. This faunal diversity is also compatible with the assortment of spinosaurid tooth morphotypes (distinguished mostly by the presence or absence of serrated mesial carina) that has been described in the Lower Cretaceous of the Iberian Peninsula.
Références bibliographiques
- Allain, R., Xaisanavong, T., Richir, P., & Khentavong, B. (2012). The first definitive Asian spinosaurid (Dinosauria:Theropoda) from the Early Cretaceous of Laos. Naturwissenschaften, 99, 369–377.
- 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.
- Alonso, A., Canudo, J. I., Fernández-Baldor, F. T., & Huerta, P. (2017). Isolated theropod teeth associated with sauropod remains from El Oterillo II (Early Cretaceous) site of Salas de los Infantes (Burgos, Spain). Journal of Iberian Geology, 43, 193–215.
- Alonso, A., Gasca, J. M., Navarro-Lorbés, P., Rubio, C., & Canudo, J. I. (2018). A new contribution to our knowledge of the largebodied theropods from the Barremian of the Iberian Peninsula: the “Barranco del Hocino” site (Spain). Journal of Iberian Geology, 44(1), 7–23.
- Arden, T. M. S., Klein, C. G., Zouhri, S., & Longrich, N. R. (2019). Aquatic adaptation in the skull of carnivorous dinosaurs (Theropoda: Spinosauridae) and the evolution of aquatic habits in spinosaurs. Cretaceous Research, 93, 275–284.
- Aurell, M., Soria, A. R., Bádenas, B., Liesa, C. L., Canudo, J. I., Gasca, J. M., et al. (2018). Barremian synrift sedimentation in the Oliete sub-basin (Iberian Basin, Spain): Palaeogeographical evolution and distribution of vertebrate remains. Journal of Iberian Geology, 44, 285–308.
- Barrett, P. M., Benson, R. B. J., Rich, T. H., & Vickers-Rich, P. (2011). First spinosaurid dinosaur from Australia and the cosmopolitanism of Cretaceous dinosaur faunas. Biology Letters, 7, 933–936.
- Benson, R. B. J., & Xing, X. (2008). The anatomy and systematic position of the theropod dinosaur Chilantaisaurus tashuikouensis Hu, 1964 from the Early Cretaceous of Alanshan, People’s Republic of China. Geological Magazine, 145(6), 778–789.
- Bertin, T. (2010). A catalogue of material and review of the Spinosauridae. PalArch’s Journal of Vertebrate Palaeontology, 7, 1–39.
- Bittencourt, J. D. S., & Kellner, A. W. A. (2004). On a sequence of sacrocaudal theropod dinosaur vertebrae from the Lower Cretaceous Santana Formation, northeastern Brazil. Arquivos do Museu Nacional, Rio de Janeiro, 62(3), 309–320.
- Bover-Arnal, T., Moreno-Bedmar, J. A., Frijia, G., Pascual-Cebrian, E., & Salas, R. (2016). Chronostratigraphy of the Barremiane Early Albian of the Maestrat Basin (E Iberian Peninsula): Integrating strontium-isotope stratigraphy and ammonoid biostratigraphy. Newsletters on Stratigraphy, 49(1), 41–68.
- Brusatte, S. L., Benson, R. B. J., & Xu, X. (2010). The evolution of large-bodied theropod dinosaurs during the Mesozoic in Asia. Journal of Iberian Geology, 36, 275–296.
- Buffetaut, E. (2007). The spinosaurid dinosaur Baryonyx (Saurischia, Theropoda) in the Early Cretaceous of Portugal. Geological Magazine, 144, 1021–1025.
- Buffetaut, E. (2008). Spinosaurids before Stromer. Early discoveries of spinosaurid theropod and their interpretation. In R. Moody, E. Buffetaut, D.M. Martill, & D. Naish (Ed.), Dinosaurs and other extinct saurians. A historical perspective(pp. 20–21). London: History of Geology Group.
- Buffetaut, E., & Ingavat, R. (1986). Unusual theropod dinosaur teeth from the upper Jurassic of Phu Wiang, northeastern Thailand. Revue de Paléobiologie, 5, 217–220.
- Buffetaut, E., & Ouaja, M. (2002). A new specimen of Spinosaurus (Dinosauria, Theropoda) from the Lower Cretaceous of Tunisia, with remarks on the evolutionary history of the Spinosauridae. Bulletin de la Société géologique de France, 173, 415–421.
- Buffetaut, E., Suteethorn, V., Tong, H., & Amiot, R. (2008). An Early Cretaceous spinosaurid theropod from southern China. Geological Magazine, 145, 745–748.
- Canudo, J. I., & Ruiz-Omeñaca, J. I. (2003). Los restos directos de dinosaurios terópodos (excluyendo aves) en España. Dinosaurios Yotros Reptiles Mesozoicos de España, 26, 347–373.
- Canudo, J. I., Gasulla, J. M., Gómez-Fernández, D., Ortega, F., Sanz, J. L., & Yagüe, P. (2008). 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.
- Canudo, J. I., Barco, J. L., Pereda-Suberbiola, X., Ruiz-Omeñaca, J. I., Salgado, L., Fernández-Baldor, F. T., et al. (2009). What Iberian dinosaurs reveal about the bridge said to exist between Gondwana and Laurasia in the Early Cretaceous. Bulletin de la Société Géologique de France, 180(1), 5–11.
- 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.
- 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.
- Carrano, M. T., & Sampson, S. D. (2008). The Phylogeny of Ceratosauria (Dinosauria: Theropoda). Journal of Systematic Palaeontology, 6(2), 183–236.
- Carrano, M. T., Benson, R. B. J., & Sampson, S. D. (2012). The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology, 10(2), 211–300.
- Charig, A. J., & Milner, A. C. (1986). Baryonyx, a remarkable new theropod dinosaur. Nature, 324, 359–361.
- Charig, A. J., & Milner, A. C. (1997). Baryonyx walkeri, a fish eating dinosaur from the Wealden of Surrey. Bulletin of the Natural History Museum, London, 53, 11–70.
- Clemente, P. (2010). Review of the Upper Jurassic-Lower Cretaceous stratigraphy in Western Cameros basin, Northern Spain. Revista de la Sociedad Geológica de España, 23(3–4), 101–143.
- Dal Sasso, C., Maganuco, S., & Buffetaut, E. (2005). New information on the skull of the enigmatic theropod Spinosaurus, with remarks on its size and affinities. Journal of Vertebrate Paleontology, 25, 888–896.
- Evers, S. W., Rauhut, O. W. M., Milner, A. C., McFeeters, B., & Allain, R. (2015). A reappraisal of the morphology and systematic position of the theropod dinosaur Sigilmassasaurus from the “middle” Cretaceous of Morocco. PeerJ, 3, e1323.
- Figueiredo, S. D., Rosina, P., & Figuti, L. (2015). Dinosaurs and other vertebrates from the Papo-Seco Formation (Lower Cretaceous) of southern Portugal. Journal of Iberian Geology, 41(3), 301–314.
- Fowler, D. W. (2007). Recently rediscovered baryonychine teeth (Dinosauria: Theropoda): New morphologic data, range extension & similarity to Ceratosaurus. Journal of Vertebrate Paleontology, 27(3), 76A.
- Fuentes Vidarte, C., Calvo, M.M., Izquierdo, L.A., Montero, D., Pérez, G., Torcida, F., Urién, V., Fuentes, F.M., & Fuentes, M.M. (2001). Restos fósiles de Baryonyx (Dinosauria, Theropoda) en el Cretácico Inferior de Salas de los Infantes (Burgos, España). In Colectivo Arqueológico-Paleontológico de Salas (Ed.), Actas de las I Jornadas Internacionales sobre Dinosaurios y su entorno (pp. 349–359).Burgos: Colectivo Arqueológico y Paleontológico de Salas.
- Gasca, J. M., Moreno-Azanza, M., & Canudo, J. I. (2008). Dientes de dinosaurios terópodos espinosáuridos de la Formación El Castellar (Cretácico Inferior, Teruel). Publicaciones del Seminario de Paleontología de Zaragoza (SEPAZ), 8, 233–234.
- Gasca, J.M., Canudo, J.I., & Moreno-Azanza, M. (2009). Dientes aislados de dinosaurio de la Formación El Castellar en Miravete de la Sierra (Cretácico Inferior, Teruel, España). In Colectivo Arqueológico y Paleontológico de Salas (Ed.). Actas de las IV Jornadas Internacionales sobre Paleontología de Dinosaurios y su Entorno (pp. 221–234). Salas de los Infantes: Colectivo Arqueológico y Paleontológico de Salas.
- Gasca, J.M., Moreno-Azanza, M., & Canudo, J.I. (2011). Dientes de dinosaurios en el Barremiense de Allepuz, Teruel. In A. Pérez- García, F. Gascó, J.M. Gasulla, F. Escaso & I. Narváez (Ed.). Libro de resúmenes del IX Encuentro de Jóvenes Investigadores en Paleontologia (pp. 145–155). Morella.
- 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.
- Gasulla, J.M., Ortega, F., Escaso, F., & Sanz, J.L. (2006). Diversidad de terópodos del Cretácico Inferior (Fm. Arcillas de Morella, Aptiense) en los yacimientos del Mas de la Parreta (Morella, Castellón). In E. Fernández-Martínez (Ed.). Libro de resúmenes de las 22ª Jornadas de la Sociedad Española de Paleontología(pp. 117–118). León: Universidad de León.
- Gasulla, J.M., Ortega, F., Sanz, J.L., Escaso, F., & Pérez Garcia, A. (2009). A spinosaurid cervical vertebra from the Morella Formation (lower Aptian) of Morella, Spain. In D. Schwarz-Wings, O. Wings & F. Sattler (Ed.), Abstract Book 7th Annual Meeting of the European Association of Vertebrate Paleontologists(pp. 31). Berlin: Shaker Verlag.
- Gómez-Fernández, D., Canudo, J.I., & Cano-Llop, V. 2007. Cintura pelviana de un nuevo dinosaurio terópodo de la Formación Morella (Aptiense inferior) en Vallibona (Castellón). Libro de Resúmenes XXIII Jornadas de la Sociedad Española de Paleontología(pp. 96–97). Caravaca de la Cruz: Sociedad Española de Paleontología.
- Hasegawa, Y., Buffetaut, E., Manabe, M., & Takakuwa, Y. (2003). A possible spinosaurid tooth from the Sebayashi formation (Lower Cretaceous), Gunma, Japan. Bulletin of Gunma Museum of Natural History, 7, 1–5.
- Hendrickx, C., & Mateus, O. (2014). 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.
- Holtz, T. R., Jr., Chapman, R. E., & Lamanna, M. C. (2004). Mesozoic Biogeography of Dinosauria. In D. B. Weishampel, P. Dodson, & H. Osmólska (Eds.), The Dinosauria (2nd ed., pp. 627–642). Berkeley: University of California Press.
- Hone, D. W. E., & Holtz, T. R. J. (2017). A century of spinosaurs e a review and revision of the Spinosauridae with comments on their ecology. Acta Geologica Sinica, 91, 1120–1132.
- Hone, D. W. E., Xu, X., & Wang, D. A. (2010). A probable baryonychine tooth from the Late Cretaceous of Henan Province, China. Vertebrata Palasiatica, 48, 19–26.
- Ibrahim, N., Sereno, P. C., Dal Sasso, C., Maganuco, S., Fabbri, M., Martill, D. M., et al. (2014). Semiaquatic adaptations in a giant predatory dinosaur. Science, 345, 1613–1616.
- Ibrahim, N., Sereno, P. C., Varricchio, D. J., Martill, D. M., Dutheil, D. B., Unwin, D. M., et al. (2020a). Geology and paleontology of the Upper Cretaceous Kem Kem Group of eastern Mococco. ZooKeys, 928, 1–216.
- Ibrahim, N., Maganuco, S., Dal Sasso, C., Fabbri, M., Auditore, M., Bindellini, G., et al. (2020b). Tail-propelled aquatic locomotion in a theropod dinosaur. Nature, 581, 67–70.
- Infante, P., Canudo, J. I., & Ruiz-Omeñaca, J. I. (2005). Primera evidencia de dinosaurios terópodos en la Formación Mirambel (Barremiense inferior, Cretácico Inferior) en Castellote, Teruel. Geogaceta, 38, 31–34.
- Kellner, A. W. A., & Campos, D. A. (1996). First Early Cretaceous theropod dinosaur from Brazil with comments on Spinosauridae. Neues Jahrbuch für Geologie und Paleaontologie Abhandlungen, 199(2), 151–166.
- Kellner, A. W., Azevedo, S. A. K., Machado, E. B., de Carvalho, L. B., & Henriques, D. D. R. (2011). A new dinosaur (Theropoda, Spinosauridae) from the Cretaceous (Cenomanian) Alcântara Formation, Cajual Island, Brazil. Anais da Academia Brasileira de Ciências, 83, 99–108.
- Kubota, K., Takakuwa, Y., & Hasegawa, H. (2017). Second discovery of a spinosaurid tooth from the Sebayashi Formation (Lower Cretaceous), Kanna Town, Gunma Prefecture, Japan. Bulletin Gunma Museum of Natural History, 21, 1–6.
- Kullberg, J. C., Terrinha, P., Pais, J., Reis, R. P., & Legoinha, P. (2006). Arrábida e Sintra: Dois Exemplos de Tectónica Pós-Rifting da Bacia Lusitaniana. In R. Dias, A. Araújo, P. Terrinha, & J. C. Kullberg (Eds.), Geologia de Portugal no contexto da Ibéria (pp. 369–396). Évora: Universidade de Évora.
- Lakin, R. J., & Longrich, N. R. (2019). Juvenile spinosaurs (Theropoda: Spinosauridae) from the middle Cretaceous of Morocco and implications for spinosaur ecology. Cretaceous Research, 93, 129–142.
- Machado, E. B., & Kellner, A. W. A. (2005). Notas sobre Spinosauridae (Theropoda, Dinosauria). Anuário do Instituto de Geociências- UFR J, 28(1), 158–173.
- Maganuco, S., & Dal Sasso, C. (2018). The smallest biggest theropod dinosaur: a tiny pedal ungual of a juvenile Spinosaurus from the Cretaceous of Morocco. PeerJ, 6, e4785.
- Malafaia, E., Ortega, F., Escaso, F., & Mocho, P. (2013). Rediscovery of a lost portion of the holotype of Suchosaurus girardi (Sauvage, 1897–98), now related to the spinosaurid theropod Baryonyx. In F. T. Fernández-Baldor & P. Huerta (Eds.), Actas de las VI Jornadas Internacionales sobre Dinosaurios y su entorno (pp. 82–84). Burgos: Colectivo Arqueológico-Paleontológico de Salas.
- Malafaia, E., Gasulla, J. M., Escaso, F., Narváez, I., Sanz, J. L., & Ortega, F. (2018). New spinosaurid (Theropoda, Megalosauroidea) remains from the Arcillas de Morella Formation (upper Barremian) of Morella, Spain. Cretaceous Research, 92, 174–183.
- Malafaia, E., Gasulla, J. M., Escaso, F., Narváez, I., Sanz, J. L., & Ortega, F. (2020). 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.cretr es.2019.10422 1.
- Manuppella, G., Pais, J., Legoinha, P., & Rey, J. (1994). Carta Geológica de Portugal à escala 1:50 000, Folha 38-B. Setúbal. 2ª Edição. Instituto Geológico e Mineiro, Portugal.
- Martill, D. M., & Hutt, S. (1996). Possible baryonychid dinosaur teeth from the Wessex Formation (Lower Cretaceous, Barremian) of the Isle of Wight, England. Proceedings of the Geologists’ Association, 107, 81–84.
- Martín-Chivelet, J., López-Gómez, J., Aguado, R., Arias, C., Arribas, J., Arribas, M. E., et al. (2019). The Late Jurassic-Early Cretaceous Rifting. In C. Quesada & J. Oliveira (Eds.), The Geology of Iberia: A Geodynamic Approach (pp. 169–249). Cham: Springer.
- 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.
- McFeeters, B., Ryan, M. J., Hinic-Frlog, S., Schrӧder-Adams, C., & Sues, H. (2013). A reevaluation of Sigilmassasaurus brevicollis (Dinosauria) from the Cretaceous of Morocco. Canadian Journal of Earth Sciences, 50, 636–649.
- Milner, A.C. (2003). Fish-Eating Theropods: A Short Review of the Systematics, Biology and Palaeobiogeography of Spinosaurs. In Colectivo Arqueológico-Paleontológico de Salas (Ed.), Actas de las II Jornadas Internacionales sobre Paleontología de dinosaurios y su entorno (pp. 129–138). Burgos: Colectivo Arqueológico- Paleontológico de Salas.
- Milner, A., Buffetaut, E., & Suteethorn, V. (2007). A tall-spined spinosaurid theropod from Thailand and the biogeography of spinosaurs. Journal of Vertebrate Paleontology, 27, 118A.
- Novas, F. E., Agnolín, F. L., Ezcurra, M. D., Porfiri, J., & Canale, J. I. (2013). Evolution of the carnivorous dinosaurs during the Cretaceous: The evidence from Patagonia. Cretaceous Research, 45, 174–215.
- Ortega, F., Escaso, F., Gasulla, J. M., Dantas, P., & Sanz, J. L. (2006). Dinosaurios de la Península Ibérica. Estudios Geológicos, 62(1), 219–240.
- 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). Bloomington: Indiana University Press.
- Pereira, D. I., Pereira, P., Brilha, J., & Cunha, P. P. (2015). The IBerian Massif Landscape and Fluvial Network in Portugal: a geoheritage inventory based on the scientific value. Proceedings of the Geologists’ Association, 126, 252–265.
- Rauhut, O. W. M. (2003). The Interrelationships and Evolution of Basal Theropod Dinosaurs. Special Papers in Palaeontology, 69, 1–213.
- Rauhut, O. W. M. (2011). Theropod dinosaurs from the Late Jurassic of Tendaguru (Tanzania). Special Papers in Palaeontology, 86, 195–239.
- Rauhut, O. W. M., Hübner, T. R., & Lanser, K.-P. (2016). A new megalosaurid theropod dinosaur from the late Middle Jurassic (Callovian) of north-western Germany: Implications for theropod evolution and faunal turnover in the Jurassic. Palaeontologia Electronica, 19.2.26A, 1–65.
- Rauhut, O.W.M., Canudo, J.I., & Castanera, D. (2019). A reappraisal of the Early Cretaceous theropod dinosaur Camarillasaurus from Spain. In European Association of Vertebrate Paleontologists (EAVP) (Ed.), Program and Abstracts XVII Conference of the EAVP(pp. 96), Brussels: EAVP.
- Ruiz-Omeñaca, J.I., Canudo, J.I., & Cuenca-Bescós, G. (1995). Dientes de dinosaurios (Ornitischia y Saurischia) del Barremiense superior (Cretácico inferior) de Vallipón (Castellote, Teruel). Mas de las Matas, pp. 59–103.
- Ruiz-Omeñaca, J. I., Canudo, J. I., & Cuenca-Bescós, G. (1997). Primera cita de dinosaurios barionícidos (Saurischia: Theropoda) en el Barremiense superior (Cretácico Inferior) de Vallipón (Castellote, Teruel). Becas del Museo de Mas de las Matas, 17, 201–223.
- Ruiz-Omeñaca, J. I., Canudo, J. I., Aurell, M., Bádenas, B., Barco, J. L., Cuenca-Bescós, G., et al. (2004). Estado de las investigaciones sobre los vertebrados del Jurásico Superior y Cretácico Inferior de Galve (Teruel). Estudios Geológicos, 60, 179–202.
- Ruiz-Omeñaca, J. I., Canudo, 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.
- Russell, D. A. (1996). Isolated dinosaur bones from the Middle Cretaceous of the Tafilalt, Morocco. Bulletin du Muséum National d’Histoire Naturelle, Paris, 4(18), 349–402.
- Sales, M. A. F., & Schultz, C. L. (2017). Spinosaur taxonomy and evolution of craniodental feautures: Evidence from Brazil. PLoS ONE, 12(11), e0187070.
- Salgado, L., Canudo, J. I., Garrido, A. C., Ruiz-Omeñaca, J. I., García, R. A., de la Fuente, M. S., et al. (2009). Upper Cretaceous vertebrates from El Anfiteatro area, Río Negro, Patagonia, Argentina. Cretaceous Research, 30, 767–784.
- Sánchez-Hernández, B., & Benton, M. J. (2014). Filling the ceratosaur gap: A new ceratosaurian theropod from the Early Cretaceous of Spain. Acta Palaeontologica Polonica, 59(3), 581–600.
- 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.
- Sauvage, H.E. (1897–98). Vertébrés fossiles du Portugal. Contribution à l’étude des poissons et des reptiles du Jurassique et du Crétacé. Direction Travaux. Géologique du Portugal, pp. 1–47.
- Sereno, P. C., Beck, A. L., Dutheuil, D. B., Gado, B., Larsson, H. C. E., Lyon, G. H., et al. (1998). A long-snouted predatory dinosaur from Africa and the evolution of spinosaurids. Science, 282, 1298–1302.
- Serrano-Martínez, A., Ortega, F., Sciscio, L., Tent-Manclús, J. E., Fierro Bandera, I., & Knoll, F. (2015). New theropod remains from the Tiouraren Formation (Middle Jurassic, Niger) and their bearing on the dental evolution in basal tetanurans. Proceedings of the Geologists’ Association, 126, 107–118.
- Smith, J. B., Lamanna, M. C., Mayr, H., & Lacovara, K. J. (2006). New information regarding the holotype of Spinosaurus aegyptiacus Stromer, 1915. Journal of Vertebrate Paleontology, 80(2), 400–406.
- Smyth, R. S. H., Ibrahim, N., & Martill, D. M. (2020). Sigilmassasaurus is Spinosaurus: a reappraisal of African spinosaurines. Cretaceous Research. https ://doi.org/10.1016/j.cretr es.2020.104520.
- Stromer, E. (1915). Ergebnisse der Forschungsreisen Prof. E. Stromers in den Wüsten Agyptens. II. Wirbeltier-Reste der Baharîje- Stufe (unterstes Cenoman). 3. Das Original des Theropoden Spinosaurus aegyptiacus nov. gen., nov. spec. Abhandlungen der Kӧniglich Bayerischen Akademie der Wissenschaften. Mathematisch- physikalische Klasse Abhandlung, 28, 1–31.
- 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, 535–547.
- Taquet, P., & Russell, D. A. (1998). New data on spinosaurid dinosaurs from the Early Cretaceous of the Sahara. Compte Rendus de l’Académie des Sciences, Sciences de la terre et des planetes, 327, 347–353.
- Torcida Fernández-Baldor, F. (2006). Restos directos de dinosaurios en Burgos (Sistema Ibérico): un balance provisional. In Colectivo Arqueológico-Paleontológico Salense (Ed.), Actas de las III Jornadas sobre Dinosaurios y su Entorno (pp. 105–128). Burgos: Colectivo Arqueológico-Paleontológico Salense.
- Torcida Fernández-Baldor, F., Fuentes, C., Izquierdo, L. A., Montero, D., & Urién, V. (1997). Dientes de dinosaurios terópodos (cf. Baryonyx) en el Weald de Burgos. Stvdia Geologica Salmanticensis, 33, 59–65.
- Torcida Fernández-Baldor, F., Izquierdo, M.L.A., Huerta, H.P., Montero, H.D., & Pérez M.G. (2003). Dientes de dinosaurios (Theropoda, Sauropoda), en el Cretácico Inferior de Burgos (España). In F. Pérez-Lorente (Ed.), Dinosaurios y otros Reptiles Mesozoicos en España(pp. 335–346). Logroño: Fundación Patrimonio Paleontológico de La Rioja, Instituto de Estudios Riojanos, Universidad de La Rioja, p. 26.
- Viera, L. I., & Torres, J. A. (1995). Presencia de Baryonyx walkeri (Saurischia, Theropoda) en el Weald de La Rioja (España). Nota previa. Munibe. Sociedad de Ciencias Naturales Aranzadi (San Sebastian), 47, 57–61.
- Vullo, R., Abit, D., Ballèvre, M., Billon-Bruyat, J.-P., Bourgeais, R., Buffetaut, E., et al. (2014). Palaeontology of the Purbeck-type (Tithonian, Late Jurassic) bonebeds of Chassiron (Oléron Island, western France). Comptes Rendus Palevol, 13, 421–441.
- Wilson, J. A. (1999). A nomenclature for vertebral laminae in sauropods and other saurischian dinosaurs. Journal of Vertebrate Paleontology, 19(4), 639–653.
- Wilson, J. A., D’Emic, M. D., Ikejiri, T., Moacdieh, E. M., & Whitlock, J. A. (2011). A Nomenclature for Vertebral Fossae in Sauropods and other Saurischian Dinosaurs. PLoS ONE, 6(2), e17114.