Journal of Iberian Geology

, Volume 43, Issue 2, pp 257–291 | Cite as

Analysis of diversity, stratigraphic and geographical distribution of isolated theropod teeth from the Upper Jurassic of the Lusitanian Basin, Portugal

  • Elisabete MalafaiaEmail author
  • Fernando Escaso
  • Pedro Mocho
  • Alejandro Serrano-Martínez
  • Angelica Torices
  • Mário Cachão
  • Francisco Ortega
Research Article



Isolated theropod teeth are abundant in the Upper Jurassic of the Lusitanian Basin and are an important source to reconstruct the diversity of this group as well as its geographic and stratigraphic distribution. However, reliably identification of isolated teeth is complex, especially for those morphotypes related to poorly represented groups. Herein a set of isolated theropod teeth collected in different sites from the Upper Jurassic of the Lusitanian Basin ranging from the late Kimmeridgian to late Tithonian in age are described and discussed.


These teeth were grouped in seventeen distinct morphotypes based first on morphology and comparative anatomy. Multivariate statistical analyses were performed in order to assign each morphotype to a certain taxon.


The current analysis shows the presence of several groups of theropods such as Ceratosaurus, Torvosaurus, and Allosaurus beside morphotypes identified as belonging to indeterminate Megalosauroidea and Allosauroidea and morphotypes tentatively assigned to Tyrannosauroidea, Dromaeosauridae, and Richardoestesia. This faunal composition, namely the presence of a non-megalosaurid megalosauroid possibly related to the piatnitzkysaurid Marshosaurus, indicates a higher diversity of theropods in the Late Jurassic of the Lusitanian Basin than previously known, based on more complete specimens. Results obtained from this analysis partially agree with previous studies of other collections with isolated theropod teeth from the Upper Jurassic of Portugal such as those of the Guimarota coal mine. However, the presence of velociraptorine dromaeosaurids, compsognathids, and troodontids reported from this site could not be confirmed in the sample herein analyzed. This analysis also indicates a great similarity of the theropod faunas from the Late Jurassic of the Lusitanian Basin and other European chronocorrelative localities such as those from Spain and Germany.


Multivariate analysis Ceratosauria Megalosauroidea Allosauroidea Coelurosauria 



Los dientes aislados de dinosaurios terópodos son un registro abundante en el Jurásico Superior de la cuenca lusitánica pudiendo llegar ser una importante contribución para comprender la diversidad y la distribución geográfica y estratigráfica de estas faunas. Sin embargo, la identificación de dientes aislados y su asignación a un determinado taxón es compleja, especialmente en el caso de morfotipos relacionados con grupos poco conocidos en el mismo registro. En este estudio se presenta el resultado del análisis de un conjunto de dientes aislados de terópodos procedentes de diferentes localidades de la cuenca lusitánica datadas en el Jurásico Superior, concretamente entre el Kimmeridgiense superior y el Tithoniense superior.


Estos dientes se han agrupado en diecisiete morfotipos a partir del estudio morfológicco y de la comparación anatómica. Se realizaron análisis estadístico multivariante para comprobar la identificación de cada morfotipo.


El resultado de este análisis ha revelado una gran diversidad de grupos de terópodos que incluye Ceratosaurus, Torvosaurus y Allosaurus además de morfotipos identificados como pertenecientes a Megalosauroidea indet. y Allosauroidea indet. Además, se han reconocido también algunos morfotipos preliminarmente asignados a Tyrannosauroidea, Dromaeosauridae y Richardoestesia. Esta composición faunística, tal como la presencia de non-megalosauridos megalosauroides posiblemente relacionado al piatnitzkysaurido Marshosaurus, sugiere una mayor diversidad de terópodos de la que se conoce actualmente a partir de ejemplares más completos. Los resultados obtenidos soportan, en parte, algunos estudios previos de otras colecciones con dientes aislados del Jurásico Superior de Portugal, como por ejemplo los de la mina de Guimarota. No obstante, la presencia de terópodos velociraptorinos, compsognathidos y troodontidos, citados en Guimarota, no se ha podido confirmar en la muestra estudiada. Este análisis indica también una grande semejanza de las faunas de terópodos del Jurásico Superior de la cuenca lusitánica y de otras localidades sincrónicas europeas como por ejemplo de España y Alemania.

Palabras clave

Análisis multivariante Ceratosauria Megalosauroidea Allosauroidea Coelurosauria 



This work was supported by SFRH/BD/84746/2012 PhD scholarship, financed by the “Fundação para a Ciência e Tecnologia” (Portugal). Individual Grants to E.M. visits for review collections were financed by the Jurassic Foundation, Fundação Luso-Americana para o Desenvolvimento [Grant Number L07-V-22/2010] and Synthesys [Grant Number GB-TAF-2160 and FR-TAF-4911]. The study was also supported by a protocol between CMTV and SHN. We thank to the reviewers D. Weishampel, C. Hendrickx and O. Gerke for the comments and suggestions to the paper. We also thank S. Pereira for photographs of some elements, J. J. Santos and N. Pimentel for field assistance, and for allow accessing specimens to B. C. Silva (SHN, Portugal), R. Castanhinha and C. Tomás (ML, Portugal), V. Santos (MUHNAC, Portugal), E. Espilez and R. Royo-Torres (Fundación Conjunto Paleontológico de Teruel-Dinópolis, Spain), E. D. Berenguer and J. I. Canudo (Museo Paleontológico de Zaragoza, Spain), R. Allain (MNHN, France), L. Chiappe (NHMLAC, USA), L. Ivy and K. Carpenter (DMNH, USA), R. Scheetz and B. Britt (BYU, USA), M. Getty, M. Loewen, and R. Irmis (NHMU, USA), D. Chure (DINO, USA), S. Chapman (NHMUK, UK), P. Jeffery (OUMNH, UK), and T. Schossleitner and D. Schwarz (MfN).

Supplementary material

41513_2017_21_MOESM1_ESM.pdf (29 kb)
Supplementary material 1 (PDF 29 kb) Table 1S Geographic and stratigraphic distribution of the specimens studied in this work
41513_2017_21_MOESM2_ESM.pdf (39 kb)
Supplementary material 2 (PDF 38 kb) Table 2S Morphometric variables of the isolated teeth studied in this work. All measurements are in millimeters. * Estimated measurement
41513_2017_21_MOESM3_ESM.pdf (29 kb)
Supplementary material 3 (PDF 29 kb) Table 3S Results of the DFA analysis based on the complete dataset of Gerke and Wings (2016). Prob, probability; Res, resolution
41513_2017_21_MOESM4_ESM.pdf (33 kb)
Supplementary material 4 (PDF 33 kb) Table 4S Results of the DFA analysis based on the reduced dataset of Gerke and Wings (2016) and on the dataset of Hendrickx et al. (2015). The green lines mark the specimens identified to the same taxon on both analyses. Inc spec, incomplete specimens; Morph, morphotype; Prob, probability; Res, resolution; Spec, specimen


  1. Allain, R. (2005). The enigmatic theropod dinosaur Erectopus superbus (Sauvage 1882) from the Lower Albian of Louppy-le-Château (Meuse, France). In K. Carpenter (Ed.), The Carnivorous Dinosaurs (pp. 72–86). Bloomington: University of Indiana Press.Google Scholar
  2. Allain, R., & Taquet, P. (2000). A new genus of Dromaeosauridae (Dinosauria, Theropoda) from the Upper Cretaceous of France. Journal of Vertebrate Paleontology, 20, 404–407. doi:10.1671/0272-4634(2000)020[0404:ANGODD]2.0.CO;2.Google Scholar
  3. Benson, R. B. J. (2010). A description of Megalosaurus bucklandii (Dinosauria: Theropoda) from the Bathonian of the UK and the relationships of Middle Jurassic theropods. Zoological Journal of the Linnean Society, 158(4), 882–935. doi: 10.1111/j.1096-3642.2009.00569.x.CrossRefGoogle Scholar
  4. Bouaziz, S., Buffetaut, E., Ghanmi, M., Jaeger, J.-J., Martin, M., Mazin, J.-M., et al. (1988). Nouvelles découvertes de vertébrés fossils dans l’Albien du Sud tunisien. Bulletin de la Société Géologique de France, 8(4), 335–339.Google Scholar
  5. Britt, B. (1991). Theropods of Dry Mesa Quarry (Morrison Formation, Late Jurassic), Colorado, with emphasis on the osteology of Torvosaurus tanneri. In B. J. Kowallis & K. Seeley (Eds.), Young University Geology Studies (Vol. 37, pp. 1–72).Google Scholar
  6. Brochu, C. A. (2003). Osteology of Tyrannosaurus rex: Insights from a nearly complete skeleton and high-resolution computed tomographic analysis of the skull. Journal of Vertebrate Paleontology, Memoir, 7, 1–138.CrossRefGoogle Scholar
  7. Brusatte, S. L., Carr, T. D., Erickson, G. M., Bever, G. S., & Norell, M. A. (2009). A long-snouted, multihorned tyrannosaurid from the Late Cretaceous of Mongolia. Proceedings of the National academy of Sciences of the United States of America, 41, 17261–17266. doi: 10.1073/pnas.0906911106.CrossRefGoogle Scholar
  8. Buckley, L. (2009). Determining ontogenetic and individual variation in Coelophysis bauri (Theropoda: Coelophysoidea) using multivariate analyses and implications for identifying isolated theropod teeth. Journal of Vertebrate Paleontology, 29(Supplement to Number 3), 72A.Google Scholar
  9. Buffetaut, E. (2007). The spinosaurid dinosaur Baryonyx (Saurischia, Theropoda) in the Early Cretaceous of Portugal. Geological Magazine, 144(6), 1021–1025. doi: 10.1017/S0016756807003883.CrossRefGoogle Scholar
  10. Buffetaut, E. (2011). An early spinosaurid dinosaur from the Late Jurassic of Tendaguru (Tanzania) and the evolution of the spinosaurid dentition. Oryctos, 10, 1–8.Google Scholar
  11. Canudo, J. I., Gasulla, J. M., Gómez-Fernández, D., Ortega, F., Sanz, J. L., & Yagüe, P. (2008). First evidence of isolated teeth referred to Spinosauridae (Theropoda) from the lower Aptian (Lower Cretaceous) from Europe: Arcillas de Morella Formation (Spain). [In Spanish, with English abstract]. Ameghiniana, Revista de la Asociación Paleontogica Argentina. Buenos Aires, 45(4), 649–662.Google Scholar
  12. Canudo, J. I., Ruiz-Omeñaca, J. I., Aurell, M., Barco, J. L., & Cuenca-Bescós, G. (2006). A megatheropod tooth from the late Tithonian-middle Berriasian (Jurassic-Cretaceous transition). Neues Jahrbuch für Geologie und Paläontologie, 239, 77–99.Google Scholar
  13. Carrano, M. T., Sampson, S. D., & Forster, C. A. (2002). The osteology of Masiakasaurus knopfleri, a small abelisauroid (Dinosauria: Theropoda) from the Late Cretaceous of Madagascar. Journal of Vertebrate Paleontology, 22(3), 510–534. doi:10.1671/0272-4634(2002)022[0510:TOOMKA]2.0.CO;2.Google Scholar
  14. 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
  15. Cobos, A., Lockley, M. G., Gascó, F., Royo-Torres, R., & Alcalá, L. (2014). Megatheropods as apex predators in the typically Jurassic ecosystems of the Villar del Arzobispo Formation (Iberian Range, Spain). Palaeogeography, Palaeoclimatology, Palaeoecology, 399, 31–41. doi: 10.1016/j.palaeo.2014.02.008.CrossRefGoogle Scholar
  16. Codrea, V., Smith, T., Dica, P., Folie, A., Garcia, G., Godefroit, P., et al. (2002). Dinosaur egg nests, mammals and other vertebrates from a new Maastrichtian site of the Hateg Basin (Romania). Comptes Rendus Palevol, 1, 173–180.CrossRefGoogle Scholar
  17. Currie, P. J., & Carpenter, K. (2000). A new specimen of Acrocanthosaurus atokensis (Theropoda, Dinosauria) from the Lower Cretaceous Antlers Formation (Lower Cretaceous, Aptian) of Oklahoma, USA. Geodiversitas, 22(2), 207–246.Google Scholar
  18. Currie, P. J., & Chen, P.-J. (2001). Anatomy of Sinosauropteryx prima from Liaoning, northeastern China. Canadian Journal of Earth Sciences, 38(1), 705–727. doi: 10.1139/cjes-38-12-1705.Google Scholar
  19. Currie, P. J., Rigby, J. K. Jr., & 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) Cambridge: Cambridge University Press. doi: 10.1017/cbo9780511608377.011.
  20. Currie, P. J., & Zhao, X.-J. (1993). A new carnosaur (Dinosauria, Theropoda) from the Jurassic of Xinjiang, People’s Republic of China. Canadian Journal of Earth Sciences, 30(10–11), 2037–2081.CrossRefGoogle Scholar
  21. Currie, P. J., & Zhiming, D. (2001). New information on Cretaceous troodontids (Dinosauria, Theropoda) from the People’s Republic of China. Canadian Journal of Earth Sciences, 38, 1753–1766.CrossRefGoogle Scholar
  22. Dal Sasso, C., & Maganuco, S. (2011). Scipionyx samniticus (Theropoda: Compsognathidae) from the Lower Cretaceous of Italy. Osteology, ontogenetic assessment, phylogeny, soft tissue anatomy, taphonomy and palaeobiology. Memoire della Società Italiana di Scienze Naturali e del Museo Civico do Storia Naturale di Milano. Volume XXXVII (Fascilolo I).Google Scholar
  23. Dal Sasso, C., Maganuco, S., Buffetaut, E., & Mendez, M. A. (2005). New information on the skull of the enigmatic theropod Spinosaurus, with remarks on its size and affinities. Journal of Vertebrate Paleontology, 25(4), 888–896. doi:10.1671/0272-4634(2005)025[0888:NIOTSO]2.0.CO;2.Google Scholar
  24. 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.Google Scholar
  25. Fürsich, F. T. (1981). Salinity-controlled benthic associations from the Upper Jurassic of Portugal. Lethaia, 14, 203–223. doi: 10.1111/j.1502-3931.1981.tb01690.x.CrossRefGoogle Scholar
  26. Gascó, F., Cobos, A., Royo-Torres, R., Alcalá, L., & Mampel, L. (2012). Theropod teeth diversity from Villar del Arzobispo Formation (Tithonian–Berriasian) at Riodeva (Teruel, Spain). Palaeobiodiversity and Palaeoenvironments, 92(2), 273–286.CrossRefGoogle Scholar
  27. 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. doi: 10.1371/journal.pone.0158334.CrossRefGoogle Scholar
  28. Gianechini, F. A., Makovicky, P. J., & Apesteguía, S. (2011). The teeth of the unenlagiine theropod Buitreraptor from the Cretaceous of Patagonia, Argentina, and the unusual dentition of the Gondwanan dromaeosaurids. Acta Palaeontologica Polonica, 56(2), 279–290. doi: 10.4202/app.2009.0127.CrossRefGoogle Scholar
  29. Gilmore, C. W. (1920). Osteology of the carnivorous Dinosauria in the United States National Museum, with special reference to the genera Antrodemus (Allosaurus) and Ceratosaurus. Bulletin United States National Museum, 110, 1–159.Google Scholar
  30. Godefroit, P., Currie, P. J., Hong, L., Yong, S. C., & Zhi-Ming, D. (2008). A new species of Velociraptor (Dinosauria: Dromaeosauridae) from the Upper Cretaceous of Northern China. Journal of Vertebrate Paleontology, 28(2), 432–438. doi:10.1671/0272-4634(2008)28[432:ANSOVD]2.0.CO;2.Google Scholar
  31. Göhlich, U. B., & Chiappe, L. M. (2006). A new carnivorous dinosaur from the Late Jurassic Solnhofen archipelago. Nature, 440, 329–332. doi: 10.1038/nature04579.CrossRefGoogle Scholar
  32. Hammer, Ø., Harper, D. A. T., Ryan, P. D. (2001). PAST: Paleontological statistics software package for education and data analysis. Palaeontologia Electronica, 4(1), 9.Google Scholar
  33. Han, F., Clark, J. M., Xu, X., Sullivan, C., Choiniere, J., & Hone, D. W. E. (2011). Theropod teeth from the Middle-Upper Jurassic Shishugou Formation of northwest Xinjiang, China. Journal of Vertebrate Paleontology, 31(1), 111–126. doi: 10.1080/02724634.2011.546291.CrossRefGoogle Scholar
  34. 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 (Vol. 13).Google Scholar
  35. 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. doi: 10.1371/journal.pone.0088905.CrossRefGoogle Scholar
  36. 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. doi: 10.11646/zootaxa.3759.1.1.CrossRefGoogle Scholar
  37. Hendrickx, C., Mateus, O., & Araújo, R. (2015). The dentition of megalosaurid theropods. Acta Palaeontologica Polonica, 60(3), 627–642. doi: 10.4202/app.00056.2013.Google Scholar
  38. Hendrickx, C., Mateus, O., & Araújo, R. (2016). A proposed terminology of theropod Teeth (Dinosauria, Saurischia). Journal of Vertebrate Paleontology. doi: 10.1080/02724634.2015.982797.
  39. Hill, G. (1988). The sedimentology and lithostratigraphy of the Upper Jurassic Lourinhã Formation, Lusitanian Basin, Portugal. Unpublished PhD Thesis, Open University.Google Scholar
  40. Hill, G. (1989). Distal alluvial fan sediments from the Upper Jurassic of Portugal: Controls on their cyclicity and channel formation. Journal of the Geological Society of London, 146, 539–555. doi: 10.1144/gsjgs.146.3.0539.CrossRefGoogle Scholar
  41. Huene, F. (1926). The carnivorous Saurischia in the Jura and Cretaceous Formations, principally in Europe. Revista del Museo de La Plata 29, 35–167.Google Scholar
  42. Janensch, W. (1920). Ueber Elaphrosaurus bambergi und die Megalosaurier aus den Tendaguru-Schichten Deutsch-Ostafrikas. In: Sitzungsberichte der Gesellschaft naturforschender Freunde zu Berlin (pp. 225–235).Google Scholar
  43. Kirkland, J. I., Zanno, L. E., Sampson, S. D., Clark, J. M., & DeBlieux, D. D. (2005). A primitive therizinosauroid dinosaur from the Early Cretaceous of Utah. Nature, 7038, 84–87. doi: 10.1038/nature03468.CrossRefGoogle Scholar
  44. Kullberg, J. C., Rocha, R. B., Soares, A. F., Rey, J., Terrinha, P., Azerêdo, A. C., Callapez, P., Duarte, L. V., Kullberg, M. C., Martins, L., Miranda, R., Alves, C., Mata, J., Madeira, J., Mateus, O., Moreira, M., & Nogueira. C. R. (2013). A Bacia Lusitaniana: Estratigrafia, Paleogeografia e Tectónica. In R. Dias, A. Araújo, P. Terrinha & J. C. Kullberg (Eds.), Geologia de Portugal, Volume II—Geologia Meso-cenozóica de Portugal (pp. 195–347).Google Scholar
  45. Larson, D. W. (2008). Diversity and variation of theropod dinosaur teeth from the uppermost Santonian Milk River Formation (Upper Cretaceous), Alberta: A quantitative method supporting identification of the oldest dinosaur tooth assemblage in Canada. Canadian Journal of Earth Sciences, 45, 1455–1468.CrossRefGoogle Scholar
  46. Larson, D. W., & Currie, P. J. (2013). Multivariate analyses of small theropod dinosaur teeth and implications for paleoecological turnover through time. PLoS One, 8(1), e54329. doi: 10.1371/journal.pone.0054329.CrossRefGoogle Scholar
  47. Louchart, A. & Pouech, J. (2017). A tooth of Archaeopterygidae (Aves) from the Lower Cretaceous of France extends the spatial and temporal occurrence of the earliest birds. Cretaceous Research, 73, 40–46. doi: 10.1016/j.cretres.2017.01.004.
  48. Madsen, J. H., Jr. (1976). A second new theropod dinosaur from the Late Jurassic of east central Utah. Utah Geology, 3(1), 51–60.Google Scholar
  49. Malafaia, E., Ortega, F., & Escaso, F. (2014). New post-cranial elements assigned to coelurosaurian theropods from the Late Jurassic of Lusitanian Basin, Portugal. Fundamental, 20, 123–126.Google Scholar
  50. Malafaia, E., Ortega, F., Escaso, F., Dantas, P., Pimentel, N., Gasulla, J. M., et al. (2010). Vertebrate fauna at the Allosaurus fossil-site of Andrés (Upper Jurassic), Pombal, Portugal. Journal of Iberian Geology, 36(2), 193–204. doi: 10.5209/rev_JIGE.2010.v36.n2.7.CrossRefGoogle Scholar
  51. Malafaia, E., Ortega, F., Escaso, F., & Silva, B. (2015). New evidence of Ceratosaurus (Dinosauria: Theropoda) from the Late Jurassic of the Lusitanian Basin, Portugal. Historical Biology, 27(7), 938–946. doi: 10.1080/08912963.2014.915820.CrossRefGoogle Scholar
  52. Manuppella, G., Antunes, M. T., Pais, J., Ramalho, M. M., & Rey, J. (1999). Notícia explicativa da Folha 30-A, Lourinhã. Lisboa: Departamento de Geologia do Instituto Geológico e Mineiro.Google Scholar
  53. Mateus, O. (1998). Lourinhanosaurus antunesi, a new upper Jurassic allosauroid (Dinosauria: Theropoda) from Lourinhã, Portugal. Memórias da Academia de Ciências de Lisboa, 37, 111–124.Google Scholar
  54. 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
  55. Mateus, O., Dinis, J, & Cunha, P.P. (2013). Upper Jurassic to Lowermost Cretaceous of the Lusitanian Basin, Portugal—Landscapes where dinosaurs walked. Ciências da Terra, número especial VIII.Google Scholar
  56. Mateus, O., Walen, A., & Antunes, M. T. (2006). The large theropod fauna of the Lourinhã Formation (Portugal) and its similarity to the Morrison Formation, with a description of a new species of Allosaurus. In J. R. Foster & S. G. Lucas (Eds.), Paleontology and Geology of the Upper Jurassic Morrison Formation (Vol. 36, pp. 123–129). New Mexico Museum of Natural History and Science, Bulletin.Google Scholar
  57. Meyer, C. A., & Thuring, B. (2003). Dinosaurs of Switzerland. Comptes Rendus Palevol, 2, 103–117. doi: 10.1016/S1631-0683(03)00005-8.CrossRefGoogle Scholar
  58. Norell, M. A., Makovicky, O. P., & Clark, J. M. (2000). A new troodontid theropod from Ukhaa Tolgod, Mongolia. Journal of Vertebrate Paleontology, 20(1), 7–11.CrossRefGoogle Scholar
  59. Novas, F. E., Ezcurra, M. D., & Lecuona, A. (2008a). Orkoraptor burkei nov. gen. et sp., a large theropod from the Maastrichtian Pari Aike Formation, Southern Patagonia,Argentina. Cretaceous Research, 29, 468–480. doi: 10.1016/j.cretres.2008.01.001.CrossRefGoogle Scholar
  60. Novas, F. E., Pol, D., Canale, J. I., Porfiri, J. D., & Calvo, J. O. (2008b). A bizarre Cretaceous theropod dinosaur from Patagonia and the evolution of Gondwanan dromaeosaurids. Proceedings of the Royal Society B, 276, 1101–1107. doi: 10.1098/rspb.2008.1554.CrossRefGoogle Scholar
  61. Oliveira, J. T., Pereira, H., Ramalho, M., & Antunes, M. T. (1992). Carta Geológica de Portugal, na escala 1:50 0000. Serviços Geológicos de Portugal.Google Scholar
  62. Osmólska, H., Currie, P. J., & Barsbold, R. (2004). Oviraptorosauria. In D.B. Weishampel, P. Dodson & H. Osmólska (Eds.), The Dinosauria, Second edition (pp. 517–606). Berkeley, Los Angeles, and London: University of California Press. doi: 10.1017/s001675680624305x.
  63. Pérez-Moreno, B. P., Chure, D. J., Pires, C., da Silva, C. M., Santos, V., Dantas, P., et al. (1999). On the presence of Allosaurus fragilis (Theropoda: Carnosauria) in the Upper Jurassic of Portugal: First evidence of an intercontinental dinosaur species. Journal of the Geological Society, 156, 449–452.CrossRefGoogle Scholar
  64. 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. doi: 10.1038/370363a0.CrossRefGoogle Scholar
  65. Perle, A., Norell, M. A., Chiappe, L. M., & Clark, J. M. (1993). Flightless bird from the Cretaceous of Mongolia. Nature, 362, 623–626. doi: 10.1038/362623a0.CrossRefGoogle Scholar
  66. Peyer, K. (2006). A reconsideration of Compsognathus from the upper Tithonian of Canjuers, southeastern France. Journal of Vertebrate Paleontology, 26(4), 879–896. doi:10.1671/0272-4634(2006)26[879:AROCFT]2.0.CO;2.Google Scholar
  67. Rauhut, O. W. M. (2002). Dinosaur teeth from the Barremian of Uña, Province of Cuenca, Spain. Cretaceous Research, 23, 255–263. doi: 10.1006/cres.2002.1003.CrossRefGoogle Scholar
  68. Rauhut, O. W. M. (2003). The interrelationships and evolution of basal theropod dinosaurs. Special Papers in Palaeontology, 69, 1–213.Google Scholar
  69. Rauhut, O. W. M. (2004). Provenance and anatomy of Genyodectes serus, a large-toothed ceratosaur (Dinosauria: Theropoda) from Patagonia. Journal of Vertebrate Paleontology, 24(4), 894–902. doi:10.1671/0272-4634(2004)024[0894:PAAOGS]2.0.CO;2.Google Scholar
  70. Rauhut, O. W. M. (2011). Theropod dinosaurs from the Late Jurassic of Tendaguru (Tanzania). Special Papers in Palaeontology, 86, 195–239.Google Scholar
  71. Rauhut, O. W. M. (2012). A reappraisal of a putative record of abelisauroid theropod dinosaur from the Middle Jurassic of England. Proceedings of the Geologists’ Association, 123, 779–786. doi: 10.1016/j.pgeola.2012.05.008.CrossRefGoogle Scholar
  72. Rauhut, O. W. M., & Fechner, R. (2005). Early development of the facial region in a non-avian theropod dinosaur. Proceedings of the Royal Society B: Biological Sciences, 272, 1179–1183. doi: 10.1098/rspb.2005.3071.CrossRefGoogle Scholar
  73. Rauhut, O. W. M., Milner, A. C., & Moore-Fay, S. (2010). Cranial osteology and phylogenetic position of the theropod dinosaur Proceratosaurus bradleyi (Woodward, 1910) from the Middle Jurassic of England. Zoological Journal of the Linnean Society, 158, 155–195. doi: 10.1111/j.1096-3642.2009.00591.x.CrossRefGoogle Scholar
  74. Rauhut, O. W. M., & Werner, C. H. (1995). First record of the family Dromaeosauridae (Dinosauria: Theropoda) in the Cretaceous of Gondwana (Wadi Milk Formation, northern Sudan). Paläontologische Zeitschrift, 69(3/4), 475–489. doi: 10.1007/BF02987808.CrossRefGoogle Scholar
  75. Rocha, R. B, Marques, B. L., Kullberg, J. C., Caetano, P. C., Lopes, C., Soares, A. F., Duarte, L.V., Marques, J. F., & Gomes, C. R. (1996). The 1st and 2nd rifting phases of the Lusitanian Basin: Stratigraphy, sequence analysis and sedimentary evolution. Final Report C.E.C. Proj. MILUPOBAS 4 vol. Lisboa.Google Scholar
  76. Royo-Torres, R., Cobos, A., & Alcalá, L. (2009). Tooth of a large-sized theropod dinosaur (Allosauroidea) from the Villar del Arzobispo Formation (Tithonian–Berresian) of Riodeva (Spain). [In Spanish, with English abstract]. Estudios Geológicos, 65(1), 91–99.CrossRefGoogle Scholar
  77. Schneider, S., Fürsich, F. T., & Werner, W. (2009). Sr-isotope of the Upper Jurassic of central Portugal (Lusitanian Basin) based on oyster shells. International Journal of Earth Sciences, Geologische Rundschau, 98, 1949–1970. doi: 10.1007/s00531-008-0359-3.CrossRefGoogle Scholar
  78. Sereno, P. C., Beck, A. L., Dutheil, 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.CrossRefGoogle Scholar
  79. Serrano-Martínez, A., Ortega, F., Sciscio, L., Tent-Manclús, J. E., Bandera, I. F., & Knoll, F. (2015). New theropod remains from the Tiourarén Formation (?Middle Jurassic, Niger) and their bearing on the dental evolution in basal tetanurans. Proceedings of the Geologists’ Association, 126(1), 107–118. doi: 10.1016/j.pgeola.2014.10.005.CrossRefGoogle Scholar
  80. Serrano-Martínez, A., Vidal, D., Sciscio, L., Ortega, F., & Knoll, F. (2016). Isolated theropod teeth from the Middle Jurassic of Niger and the early dental evolution of Spinosauridae. Acta Palaeontologica Polonica, 61(2), 403–415. doi: 10.4202/app.00101.2014.Google Scholar
  81. Smith, J. B. (2005). Heterodonty in Tyrannosaurus rex: Implications for the taxonomic and systematic utility of theropod dentitions. Journal of Vertebrate Paleontology, 25(4), 865–887. doi:10.1671/0272-4634(2005)025[0865:HITRIF]2.0.CO;2.Google Scholar
  82. 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 Part A, 285, 699–736. doi: 10.1002/ar.a.20206.CrossRefGoogle Scholar
  83. Soto, M., & Perea, D. (2008). A ceratosaurid (Dinosauria, Theropoda) from the Late Jurassic-Early Cretaceous of Uruguay. Journal of Vertebrate Paleontology, 28(2), 439–444. doi:10.1671/0272-4634(2008)28[439:ACDTFT]2.0.CO;2.Google Scholar
  84. 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. doi:10.1671/0272-4634(2002)022[0535:ICASDT]2.0.CO;2.Google Scholar
  85. Sullivan, R. M. (2006). Saurornitholestes robustus, n. sp. (Theropoda: Dromaeosauridae) from the Upper Cretaceous Kirtland Formation (De-Na-Zin Member), San Juan Basin, New Mexico. In S. G. Lucas, & R. M. Sullivan (Eds.), Late Cretaceous vertebrates from the Western Interior (Vol. 35, pp. 253–256). New Mexico Museum of Natural History and Science Bulletin.Google Scholar
  86. Suñer, M., de Santisteban, C., & Galobart, A. (2005). New Upper Jurassic—Lower Cretaceous Theropoda remains from 'Los Serranosʼ region (Valencia). [In Spanish, with English abstract] Revista Española de Paleontología, N.E. X, 10, 93–99.Google Scholar
  87. Taylor, A. M., Gowland, S., Leary, S., Keogh, K. J., & Martinius, A. W. (2014). Stratigraphical correlation of the Late Jurassic Lourinhã Formation in the Consolação Sub-basin (Lusitanian Basin), Portugal. Geological Journal, 49(2), 143–162. doi: 10.1002/gj.2505.CrossRefGoogle Scholar
  88. Torices, A., Currie, P. J., Canudo, J. I., & Pereda-Suberbiola, X. (2015). Theropod dinosaurs from the Upper Cretaceous of the South Pyrenees Basin of Spain. Acta Palaeontologica Polonica, 60(3), 611–626. doi: 10.4202/app.2012.0121.Google Scholar
  89. Weigert, A. (1995). Isolierte Zähne von cf. Archaeopteryx sp. aus dem Oberen Jura der Kohlengrube Guimarota (Portugal). Neues Jahrbuch für Geologie und Paläontologie, 9, 562–576.Google Scholar
  90. Weishampel, D. A., Csiki, Z., Benton, M. J., Grigorescu, D., & Codrea, V. (2010). Palaeobiogeographic relationships of the Hateg biota—Between isolation and innovation. Palaeogeography, Palaeoclimatology, Palaeoecology, 293, 419–437.CrossRefGoogle Scholar
  91. Madsen, J. H. Jr., & Welles, S. P. (2000). Ceratosaurus (Dinosauria, Theropoda) a revised osteology. Utah Geological Survey. Miscellaneous Publication (Vol. 00-2, pp. 1–80).Google Scholar
  92. Werner, W. (1986). Palökologische und biofazielle Analyse des Kimmeridge (Oberjura) von Consolacao, Mittelportugal [Paleoecological and biofacies analysis of the Kimmeridgian (Upper Jurassic) of Consolação, central Portugal]. Zitteliana, 13, 3–109.Google Scholar
  93. Williamson, T. E., & Brusatte, S. L. (2014). Small theropod teeth from the Late Cretaceous of the San Juan Basin, northwestern New Mexico and their implications for understanding latest Cretaceous dinosaur evolution. PLoS One, 9(4), e93190.CrossRefGoogle Scholar
  94. Xu, X., Zhou, Z.-H., & Wang, X.-L. (2000). The smallest known non-avian theropod dinosaur. Nature, 408, 705–708.CrossRefGoogle Scholar
  95. Zinke, J. (1998). Small theropod teeth from the Upper Jurassic coal mine of Guimarota (Portugal). Palaontologische Zeitschrift, 72, 179–189.CrossRefGoogle Scholar
  96. Zinke, J., & Rauhut, O. W. M. (1994). Small theropods (Dinosauria, Saurischia) from the Upper Jurassic and Lower Cretaceous of the Iberian Peninsula. Berliner geowiss. Abh., E13, 163–177.Google Scholar

Copyright information

© Springer International Publishing AG 2017

Authors and Affiliations

  • Elisabete Malafaia
    • 1
    • 2
    • 3
    Email author
  • Fernando Escaso
    • 2
    • 4
  • Pedro Mocho
    • 2
    • 4
    • 5
  • Alejandro Serrano-Martínez
    • 4
  • Angelica Torices
    • 6
  • Mário Cachão
    • 1
  • Francisco Ortega
    • 2
    • 4
  1. 1.Faculdade de Ciências and Instituto Dom Luiz, Universidade de LisboaLisbonPortugal
  2. 2.Laboratório de Paleontologia e PaleoecologiaSociedade de História NaturalTorres VedrasPortugal
  3. 3.Museu Nacional de História Natural e da CiênciaUniversidade de LisboaLisbonPortugal
  4. 4.Grupo de Biología EvolutivaUniversidad Nacional de Educación a DistanciaMadridSpain
  5. 5.The Dinosaur Institute, Natural History Museum of Los Angeles CountyLos AngelesUSA
  6. 6.Departamento de Ciencias HumanasUniversidad de La RiojaLogroñoSpain

Personalised recommendations