Skip to main content

Theropod teeth from the basalmost Cretaceous of Anoual (Morocco) and their palaeobiogeographical significance

  • F. KNOLL (a1) and J. I. RUIZ-OMEÑACA (a2)

The theropod teeth from the Berriasian (Early Cretaceous) site of Anoual (N Morocco) are described. The assemblage is important in that it comes from one of the very few dinosaur sites of this age globally and the only one for the whole of Gondwana. The theropod teeth from Anoual are morphologically diverse. Most of the material possibly belongs to the clade Dromaeosauridae, which would be an early occurrence for this taxon. The palaeogeographic position of Anoual enables it to provide data on the dispersal events that affected terrestrial faunas during Mesozoic times. A Laurasian influence is evidenced by the presence of Velociraptorinae and, on the whole, the theropod fauna from Anoual provides support for the existence of a trans-Tethyan passage allowing terrestrial faunal interchanges during Late Jurassic and/or earliest Cretaceous times. Additionally, Anoual records the existence of diminutive theropods. However, it cannot yet be determined whether the small size of the specimens is genetic or ontogenetic.

Corresponding author
Author for correspondence:
Hide All
Alifanov, V. R. & Sennikov, A. G. 2001. [On the discovery of dinosaur remains in Moscow region]. Doklady Akademii nauk 376, 73–5 (in Russian).
Antunes, M. T. & Sigogneau, D. 1992. La faune de petits dinosaures du Crétacé terminal portugais. Comunicações dos Serviços Geológicos de Portugal 78, 4962.
Antunes, M. T. & Sigogneau-Russell, D. 1996. Le Crétacé terminal portugais et son apport au problème de l'extinction des dinosaures. Bulletin du Muséum national d'Histoire naturelle, Section C 4, 595606.
Averianov, A. O., Martin, T. & Bakirov, A. A. 2005. Pterosaur and dinosaur remains from the Middle Jurassic Balabansai Svita in the northern Fergana Depression, Kyrgyzstan (Central Asia). Palaeontology 48, 135–55.
Bakker, R. T., Williams, M. & Currie, P. J. 1988. Nanotyrannus, a new genus of pygmy tyrannosaur, from the latest Cretaceous of Montana. Hunteria 1, 126.
Barco, J. L. & Ruiz-Omeñaca, J. I. 2001 a. Primeros dientes de terópodos (Dinosauria, Saurischia) en la Formación Villar del Arzobispo (Tithónico-Berriasiense): yacimientos Cuesta Lonsal y Las Cerradicas 2 (Galve, Teruel). In XVII Jornadas de la Sociedad Española de Paleontología: los Fósiles y la Paleogeografía, y Simposios de los Proyectos del Programa Internacional de Correlación Geológica (UNESCO/IUGS) N° 410 y 421 (eds Meléndez, G., Herrera, Z., Delvene, G. & Azanza, B.), pp. 239–46. Zaragoza: Seminario de Paleontología de Zaragoza.
Barco, J. L. & Ruiz-Omeñaca, J. I. 2001 b. Primeros restos postcraneales de terópodo (Dinosauria, Saurischia) en la Formación Villar del Arzobispo (Tithónico-Berriasiense): un centro vertebral caudal del yacimiento Carretera (Galve, Teruel). In XVII Jornadas de la Sociedad Española de Paleontología: los Fósiles y la Paleogeografía, y Simposios de los Proyectos del Programa Internacional de Correlación Geológica (UNESCO/IUGS) N° 410 y 421 (eds Meléndez, G., Herrera, Z., Delvene, G. & Azanza, B.), pp. 247–54. Zaragoza: Seminario de Paleontología de Zaragoza.
Barco, J. L., Ruiz-Omeñaca, J. I., Canudo, J. I. & Cuenca-Bescós, G. 2004. Dinosaurios. In Guía del Parque Paleontológico de Galve (ed. Barco, J. L.), pp. 6488. Zaragoza: Paleoymás.
Barsbold, R. 1983. [Carnivorous dinosaurs from the Cretaceous of Mongolia]. Trudy–Sovmestnaya Sovetsko-Mongol'skaya Paleontologicheskaya Ekspeditsiya 19, 5119 (in Russian, with English summary).
Barsbold, R. & Osmólska, H. 1999. The skull of Velociraptor (Theropoda) from the Late Cretaceous of Mongolia. Acta Palaeontologica Polonica 44, 189219.
Baszio, S. 1997. Systematic palaeontology of isolated dinosaur teeth from the latest Cretaceous of south Alberta, Canada. Courier Forschunginstitut Senckenberg 196, 3377.
Brinkman, D. L., Cifelli, R. L. & Czaplewski, N. J. 1998. First occurrence of Deinonychus antirrhopus (Dinosauria: Theropoda) from the Antlers Formation (Lower Cretaceous: Aptian–Albian) of Oklahoma. Oklahoma Geological Survey Bulletin 146, 127.
Burnham, D. A., Derstler, K. L., Currie, P. J., Bakker, R. T., Zhou, Z. & Ostrom, J. H. 2000. Remarkable new birdlike dinosaur (Theropoda: Maniraptora) from the Upper Cretaceous of Montana. The University of Kansas Paleontological Contributions 13, 114.
Canudo, J. I., Cuenca-Bescós, G. & Ruiz-Omeñaca, J. I. 1997. Dinosaurios dromeosaúridos [sic] (Saurischia: Theropoda) en el Barremiense Superior (Cretácico Inferior) de Castellote, Teruel. Geogaceta 22, 3942.
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) of Galve (Aragón, NE Spain). Neues Jahrbuch für Geologie und Paläontologie Abhandlungen 239, 7799.
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, 510–34.
Codrea, V., Smith, T., Dica, P., Folie, A., Garcia, G., Godefroit, P. & Van Itterbeeck, J. 2002. Dinosaur egg nests, mammals and other vertebrates from a new Maastrichtian site of the Haţeg Basin (Romania). Comptes Rendus Palevol 1, 173–80.
Company, J., Pereda Suberbiola, X., Ruiz-Omeñaca, J. I. & Buscalioni, A. D. 2005. A new species of Doratodon (Crocodyliformes: Ziphosuchia) from the Late Cretaceous of Spain. Journal of Vertebrate Paleontology 25, 343–53.
Currie, P. J. 1995. New information on the anatomy and relationships of Dromaeosaurus albertensis (Dinosauria: Theropoda). Journal of Vertebrate Paleontology 15, 576–91.
Currie, P. J. & Azuma, Y. 2006. New specimens, including a growth series, of Fukuiraptor (Dinosauria, Theropoda) from the Lower Cretaceous Kitadani Quarry of Japan. Journal of the Paleontological Society of Korea 22, 173–93.
Currie, P. J. & Chen, P. J. 2001. Anatomy of Sinosauropteryx prima from Liaoning, northeastern China. Canadian Journal of Earth Sciences 38, 1705–27.
Currie, P. J., Rigby, K. Jr & Sloan, R. E. 1990. Theropod teeth from the Judith River Formation of southern Alberta, Canada. In Dinosaur Systematics: Approaches and Perspectives (eds Carpenter, K. & Currie, P. J.), pp. 107–25. New York: Cambridge University Press.
Currie, P. J. & Varricchio, D. J. 2004. A new dromaeosaurid from the Horseshoe Canyon Formation (Upper Cretaceous) of Alberta, Canada. In Feathered Dragons (eds Currie, P. J., Koppelhus, E. B., Shugar, M. A. & Wright, J. L.), pp. 112–32. Bloomington: Indiana University Press.
Dal Sasso, C. & Signore, M. 1998. Exceptional soft-tissue preservation in a theropod dinosaur from Italy. Nature 392, 383–7.
Dercourt, J., Guetani, M., Vrielynk, B., Barrier, E., Biju-Duval, B., Brunet, M.-F., Cadet, J.-P., Crasquin, S. & Sandulescu, M. (eds) 2000. Peri-Tethys palaeogeographical Atlas-Explanatory Notes. Paris: Gauthier-Villars, 268 pp., 24 maps.
Dercourt, J., Ricou, L.-E. & Vrielynck, B. (eds) 1993. Atlas Tethys-Palaeoenvironmental Maps-Explanatory Notes. Paris: Gauthier-Villars, 307 pp., 14 maps.
Evans, S. E. & Milner, A. R. 1994. Middle Jurassic microvertebrate assemblages from the British Isles. In In the Shadow of the Dinosaurs: Early Mesozoic Tetrapods (eds Fraser, N. and Sues, H.-D..), pp. 303–21. Cambridge: Cambridge University Press.
Evans, S. E. & Sigogneau-Russell, D. 2001. A stem-group caecilian (Lissamphibia: Gymnophiona) from the Lower Cretaceous of North Africa. Palaeontology 44, 259–73.
Fanti, F. & Therrien, F. 2007. Theropod tooth assemblages from the Late Cretaceous Maevarano Formation and the possible presence of dromaeosaurids in Madagascar. Acta Palaeontologica Polonica 52, 155–66.
Farlow, J. O., Brinkman, D. L., Abler, W. L. & Currie, P. J. 1991. Size, shape, and serration density of theropod dinosaur lateral teeth. Modern Geology 16, 161–98.
Galton, P. M. 1996. Notes on Dinosauria from the Upper Cretaceous of Portugal. Neues Jahrbuch für Geologie und Paläontologie Monatshefte 1996, 8390.
Garcia, G., Pincemaille, M., Vianey-Liaud, M., Marandat, B., Lorenz, E., Cheylan, G., Cappetta, H., Michaux, J. & Sudre, J. 1999. Découverte du premier squelette presque complet de Rhabdodon priscus (Dinosauria, Ornithopoda) du Maastrichtien inférieur de Provence. Comptes Rendus de l'Académie des Sciences de Paris, Sciences de la Terre et des Planètes 328, 415–21.
Gardner, J. D., Evans, S. E. & Sigogneau-Russell, D. 2003. New albanerpetontid amphibians from the Early Cretaceous of Morocco and Middle Jurassic of England. Acta Palaeontologica Polonica 48, 301–19.
Gauthier, J. A. 1986. Saurischian monophyly and the origin of birds. Memoirs of the California Academy of Sciences 8, 155.
Gheerbrant, E. & Rage, J.-C. 2006. Paleobiogeography of Africa: How distinct from Gondwana and Laurasia? Palaeogeography, Palaeoclimatology, Palaeoecology 241, 224–46.
Göhlich, U. B. & Chiappe, L. M. 2006. A new carnivorous dinosaur from the Late Jurassic Solnhofen archipelago. Nature 440, 329–32.
Goodwin, M. B., Clemens, W. A., Hutchison, J. H., Wood, C. B., Zavada, M. S., Kemp, A., Duffin, C. J. & Schaff, C. R. 1999. Mesozoic continental vertebrates with associated palynostratigraphic dates from the northwestern Ethiopian plateau. Journal of Vertebrate Paleontology 19, 728–41.
Grigorescu, D. 1984. New paleontological data on the dinosaur beds from the Hateg Basin. In 75 years of the Laboratory of Paleontology (eds Petrescu, J. & Dragastan, O..), pp. 111–18. Bucureşti: Universitatea din-Bucureşti.
Haddoumi, H., Alméras, Y., Bodergat, A. M., Charrière, A., Mangold, C. & Benshili, K. 1998. Ages et environnements des couches rouges d'Anoual (Jurassique moyen et Crétacé inférieur, Haut-Atlas oriental, Maroc). Comptes Rendus de l'Académie des Sciences de Paris, Sciences de la Terre et des Planètes 327, 127–33.
Hahn, G. & Hahn, R. 2003. New multituberculate teeth from the Early Cretaceous of Morocco. Acta Palaeontologica Polonica 48, 349–56.
Hartman, S., Lovelace, D. & Wahl, W. 2005. Phylogenetic assessment of a maniraptoran from the Morrison Formation. Journal of Vertebrate Paleontology 25, 67A68A.
Holtz, T. R. Jr. 1995. A new phylogeny of the Theropoda. Journal of Vertebrate Paleontology 15, 35A.
Holtz, T. R. Jr, Molnar, R. E. & Currie, P. J. 2004. Basal Tetanurae. In The Dinosauria, Second Edition (eds Weishampel, D. B., Dodson, P. & Osmólska, H..), pp. 71110. Berkeley: University of California Press.
Holtz, T. R. Jr & Osmólska, H. 2004. Saurischia. In The Dinosauria, Second Edition (eds Weishampel, D. B., Dodson, P. & Osmólska, H..), pp. 21–4. Berkeley: University of California Press.
von Huene, F. 1914. Das natürliche System der Saurischia. Zentralblatt für Mineralogie, Geologie und Paläontologie, Abteilung B 1914, 154–8.
Hwang, S. H., Norell, M. A., Ji, Q. & Gao, K. 2002. New specimens of Microraptor zhaoianius (Theropoda: Dromaeosauridae) from northeastern China. American Museum Novitates 3381, 144.
Hwang, S. H., Norell, M. A., Ji, Q. & Gao, K. 2004. A large compsognathid from the Early Cretaceous Yixian Formation of China. Journal of Systematic Palaeontology 2, 1330.
Jones, M. E. H., Evans, S. E. & Sigogneau-Russell, D. 2003. Early Cretaceous frogs from Morocco. Annals of Carnegie Museum 72, 6597.
Knoll, F. 2000. Pterosaurs from the Lower Cretaceous (?Berriasian) of Anoual, Morocco. Annales de Paléontologie 86, 157–64.
Laurent, Y., Bilotte, M. & Le Loeuff, J. 2002. Late Maastrichtian continental vertebrates from southwestern France: correlation with marine fauna. Palaeogeography, Palaeoclimatology, Palaeoecology 187, 121–35.
Leonardi, G. & Carvalho, I. de S. 2002. Icnofósseis da Bacia do Rio do Peixe, PB: o mais marcante registro de pegadas de dinossauros do Brasil. In Sítios geológicos e paleontológicos do Brasil (eds Schobbenhaus, C., Campos, D. de A., Queiroz, E. T., Winge, M. & Berbert-Born, M..), pp. 101–11. Brasilia: SIGEP.
Leonardi, G. & Carvalho, I. de S. 2007. Dinosaur ichnocoenosis from Sousa and Uiraúna-Brejo das Freiras Basins, northeast Brazil. In Paleontologia: Cenários de Vida, vol. 1 (eds Carvalho, I. de S., Cassab, R. de C. T., Schwanke, C., Carvalho, M. de A., Fernandes, A. C. S., Rodrigues, M. A. da C., Carvalho, M. S. S. de, Arai, M. & Oliveira, M. E. Q..), pp. 355–69. Rio de Janeiro: Interciência.
Lindgren, J., Currie, P. J., Rees, J., Siverson, M., Lindström, S., Alwmark, C. 2008. Theropod dinosaur teeth from the lowermost Cretaceous Rabekke Formation on Bornholm, Denmark. Geobios 41, 253–62.
Maganuco, S., Cau, A. & Pasini, G. 2005. First description of theropod remains from the Middle Jurassic (Bathonian) of Madagascar. Atti della Società Italiana di Scienze Naturali e del Museo civico di Storia Naturale di Milano 146, 165202.
Makovicky, P. J., Apesteguía, S. & Agnolín, F. L. 2005. The earliest dromaeosaurid theropod from South America. Nature 437, 1007–11.
Makovicky, P. & Norell, M. A. 2004. Troodontidae. In The Dinosauria, Second Edition (eds Weishampel, D. B., Dodson, P. and Osmólska, H..), pp. 184–95. Berkeley: University of California Press.
Marsh, O. C. 1881. Principal characters of American Jurassic dinosaurs; Part V. American Journal of Science 21, 417–23.
Mateer, N. J. 1987. A new report of a theropod dinosaur from South Africa. Palaeontology 30, 141–5.
Matthew, W. D. & Brown, B. 1922. The family Deinodontidae, with notice of a new genus from the Cretaceous of Alberta. Bulletin of the American Museum of Natural History 46, 367–85.
Milner, A. 2002. Theropod dinosaurs of the Purbeck Limestone Group, Southern England. Special Papers in Palaeontology 68, 191201.
Mojon, P. O., Haddoumi, H. & Charrière, A. 2005. Nouvelles données sur les Charophytes et Ostracodes du Jurassique–Crétacé de l'Atlas marocain. In Recherches sur la Pangée mésozoïque (ed. Mojon, P. O..), pp. 447. Lausanne: Digit Presse.
Norell, M. A., Clark, J. M., Dashzeveg, D., Barsbold, R., Chiappe, L. M., Davidson, A. R., McKenna, M. C., Perle, A. & Novacek, M. J. 1994. A theropod dinosaur embryo and the affinities of the Flaming Cliffs dinosaur eggs. Science 266, 779–82.
Norell, M. A., Clark, J. M., Turner, A. H., Makovicky, P. J., Barsbold, R. & Rowe, T. 2006. A new dromaeosaurid theropod from Ukhaa Tolgod (Ömnögov, Mongolia). American Museum Novitates 3545, 151.
Novas, F. E. & Pol, D. 2005. New evidence on deinonychosaurian dinosaurs from the Late Cretaceous of Patagonia. Nature 433, 858–61.
Osborn, H. F. 1924. Three new Theropoda, Protoceratops zone, central Mongolia. American Museum Novitates 144, 112.
Ostrom, J. H. 1969. Osteology of Deinonychus antirrhopus, an unusual theropod from the Lower Cretaceous of Montana. Peabody Museum of Natural History Bulletin 30, 1165.
Ostrom, J. H. 1978. The osteology of Compsognathus longipes Wagner. Zitteliana 4, 73118.
Ostrom, J. H. 1990. Dromaeosauridae. In The Dinosauria (eds Weishampel, D. B., Dodson, P. & Osmólska, H..), pp. 269–79. Berkeley: University of California Press.
Owen, R. 1842. Report on British fossil reptiles; Part II. Report of the British Association for the Advancement of Science 1841, 60204.
Paul, G. S. 1988 a. The small predatory dinosaurs of the mid-Mesozoic: the horned theropods of the Morrison and Great Oolite – Ornitholestes and Proceratosaurus – and the sickle-claw theropods of the Cloverly, Djadokhta and Judith River – Deinonychus, Velociraptor and Saurornitholestes. Hunteria 2, 19.
Paul, G. S. 1988 b. Predatory Dinosaurs of the World. New York: Simon and Schuster, 464 pp.
Pereda Suberbiola, X. 1999. Las faunas finicretácicas de dinosaurios ibéricos. Zubía 17, 259–79.
Pol, C., Buscalioni, A. D., Carballeira, J., Francés, V., López Martinez, N., Marandat, B., Moratalla, J. J., Sanz, J. L., Sigé, B. & Villatte, J. 1992. Reptiles and mammals from the Late Cretaceous new locality Quintanilla del Coco (Burgos Province, Spain). Neues Jahrbuch für Geologie und Paläontologie Abhandlungen 184, 279314.
Prieto-Márquez, A., Gaete, R., Galobart, A. & Ardèvol, L. 2000. A Richardoestesia-like theropod tooth from the Late Cretaceous foredeep, south-central Pyrenees, Spain. Eclogae Geologicae Helvetiae 93, 497501.
Rauhut, O. W. M. 2000. The dinosaur fauna from the Guimarota mine. In Guimarota – A Jurassic Ecosystem (eds Martin, T. & Krebs, B..), pp. 7582. München: Dr Friedrich Pfeil.
Rauhut, O. W. M. 2001. Herbivorous dinosaurs from the Late Jurassic (Kimmeridgian) of Guimarota, Portugal. Proceedings of the Geologists' Association 112, 275–83.
Rauhut, O. W. M. 2002. Dinosaur teeth from the Barremian of Uña, Province of Cuenca, Spain. Cretaceous Research 23, 255–63.
Rauhut, O. W. M. 2004. Provenance and anatomy of Genyodectes serus, a large-toothed ceratosaur (Dinosauria: Theropoda) from Patagonia. Journal of Vertebrate Paleontology 24, 894902.
Rauhut, O. W. M. 2005. Osteology and relationships of a new theropod dinosaur from the Middle Jurassic of Patagonia. Palaeontology 48, 87110.
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 Zeitschrift 69, 475–89.
Rauhut, O. W. M. & Zinke, J. 1995. A description of the Barremian dinosaur fauna from Uña with a comparison to that of Las Hoyas. In II International Symposium on Lithographic Limestones, Lleida-Cuenca (Spain), 9th–16th July 1995 – Extended Abstracts (ed. Meléndez, N..), pp. 123–6. Cantoblanco: Universidad Autónoma de Madrid.
Royo-Torres, R., Cobos, A., Alcalá, L. & Bello, Y. 2003. Primeros restos de dinosaurio en el Cretácico Inferior de Riodeva (Teruel). In XIX Jornadas de la Sociedad Española de Paleontología, Morella, 16/18 Octubre 2003 – Libro de Resúmenes (eds Alonso, M. V. Pardo & Gozalo, R..), p. 147. Morella: Ajuntament de Morella.
Ruiz-Omeñaca, J. I. & Canudo, J. I. 2001. Dos yacimientos excepcionales con vertebrados continentales del Barremiense (Cretácico Inferior) de Teruel: Vallipón y La Cantalera. Naturaleza aragonesa 8, 817.
Ruiz-Omeñaca, J. I. & Canudo, J. I. 2003. Un nuevo dinosaurio terópodo (“Prodeinodon” sp.) en el Cretácico Inferior de la Cantalera (Teruel). Geogaceta 34, 111–14.
Ruiz-Omeñaca, J. I., Canudo, J. I. & Cuenca-Bescós, G. 1996. Dientes de dinosaurios (Ornithischia y Saurischia) del Barremiense superior (Cretácico inferior) de Vallipón (Castellote, Teruel). Mas de las Matas 15, 59103.
Ruiz-Omeñaca, J. I., Canudo, J. I., Cuenca-Bescós, G. & Amo, O. 1998. Theropod teeth from the Lower Cretaceous of Galve (Teruel, Spain). In Third European Workshop of Vertebrate Palaeontology: Programme and Abstracts, Field Guide (eds Jagt, J. W. M., Lambers, P. H., Mulder, E. W. A. & Schulp, A. S..), pp. 62–3. Maastricht: Natuurhistorisch Museum Maastricht.
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, 757–76.
Sankey, J. T. 2008. Diversity of Latest Cretaceous (Late Maastrichtian) small theropods and birds: teeth from the Lance and Hell Creek Formations, USA. In Vertebrate Microfossil Assemblages: their role in paleoecology and paleobiogeography (eds by Sankey, J. T. & Baszio, S..), pp. 117–34. Bloomington: Indiana University Press.
Sankey, J. T., Brinkman, D. B., Guenther, M. & Currie, P. J. 2002. Small theropod and bird teeth from the Late Cretaceous (Late Campanian) Judith River Group, Alberta. Journal of Paleontology 76, 751–63.
Sankey, J. T., Standhardt, B. R. & Schiebout, J. A. 2005, Theropod teeth from the Upper Cretaceous (Campanian–Maastrichtian), Big Bend National Park, Texas. In The Carnivorous Dinosaurs (ed. Carpenter, K..), pp. 127–52. Bloomington: Indiana University Press.
Seeley, H. G. 1888. On the classification of the fossil animals commonly named Dinosauria. Proceedings of the Royal Society of London 43, 165–71.
Sereno, P. C. & Brusatte, S. L. 2008. Basal abelisaurid and carcharodontosaurid theropods from the Lower Cretaceous Elrhaz Formation of Niger. Acta Palaeontologica Polonica 53, 1546.
Sereno, P. C., Dutheil, D. B., Iarochene, M., Larsson, H. C. E., Lyon, G. H., Magwene, P. M., Sidor, C. A., Varricchio, D. J. & Wilson, J. A. 1996. Predatory dinosaurs from the Sahara and Late Cretaceous faunal differentiation. Science 272, 986–91.
Sigogneau-Russell, D. 1999 a. Réévaluation des Peramura (Mammalia, Cladotheria) sur la base de nouveaux spécimens du Crétacé inférieur d'Angleterre et du Maroc. Geodiversitas 21, 93127.
Sigogneau-Russell, D. 1999 b. Le site d'Anoual et sa riche faune de microvertébrés. In Maroc, Mémoire de la Terre (ed. Blandin, P..), pp. 148–51. Paris: Muséum national d'Histoire naturelle.
Sigogneau-Russell, D. 2003. Diversity of triconodont mammals from the Early Cretaceous of North Africa – affinities of the amphilestids. Palaeovertebrata 32, 2755.
Sigogneau-Russell, D., Evans, S. E., Levine, J. F. & Russell, D. A. 1998. The Early Cretaceous microvertebrate locality of Anoual, Morocco: a glimpse at the small vertebrate assemblages of Africa. New Mexico Museum of Natural History and Science Bulletin 14, 177–81.
Sigogneau-Russell, D., Monbaron, M. & Kaenel, de E. 1990. Nouvelles données sur le gisement à Mammifères mésozoïques du Haut–Atlas marocain. Geobios 23, 461–83.
Smith, J. B. 2005. Heterodonty in Tyrannosaurus rex: implications for the taxonomic and systematic utility of theropod dentitions. Journal of Vertebrate Paleontology 25, 865–87.
Smith, J. B. 2007. Dental morphology and variation in Majungasaurus crenatissimus (Theropoda: Abelisauridae) from the Late Cretaceous of Madagascar. Society of Vertebrate Paleontology Memoir 8, 103–26.
Smith, J. B. & Dodson, P. 2003. A proposal for a standard terminology of anatomical notation and orientation in fossil vertebrate dentitions. Journal of Vertebrate Paleontology 23, 114.
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: Discoveries in Molecular, Cellular, and Evolutionary Biology 285A, 699736.
Stromer, E. 1934. Die Zähne des Compsognathus und Bemerkungen über dass Gebiss der Theropoda. Centralblatt für Mineralogie, Geologie und Paläontologie Abteilung B 1934, 7485.
Sues, H.-D. 1978. A new small theropod dinosaur from the Judith River Formation (Campanian) of Alberta, Canada. Zoological Journal of the Linnean Society 62, 381400.
Suñer, M., Santisteban, C. de & Galobart, A. 2005. Nuevos restos de Theropoda del Jurásico Superior–Cretácico Inferior de la Comarca de los Serranos (Valencia). Revista Española de Paleontología, Número extraordinario 10, 93–9.
Sweetman, S. C. 2004. The first record of velociraptorine dinosaurs (Saurischia, Theropoda) from the Wealden (Early Cretaceous, Barremian) of southern England. Cretaceous Research 25, 353–64.
Torices Hernández, A. 2002. Los dinosaurios terópodos del Cretácico superior de la Cuenca de Tremp (Pirineos Sur-Centrales, Lleida). Coloquios de Paleontología 53, 139–46.
Torices Hernández, A. 2003. Estudio preliminar de dientes aislados de terópodos en el Cretácico superior de la Cuenca de Tremp (Pirineos Sur-Centrales, Lleida). In II Jornadas internacionales sobre Paleontología de Dinosaurios y su Entorno (eds Vidarte, C. Fuentes & Fernández-Baldor, F. Torcida.), pp. 213–20. Salas de los Infantes: Colectivo Arqueológico y Paleontológico de Salas.
Torices, A., Ruiz-Omeñaca, J. I., Canudo, J. I. & López-Martínez, N. 2004. Nuevos datos sobre los dinosaurios terópodos (Saurischia: Theropoda) del Cretácico superior de los Pirineos Sur-Centrales (Huesca y Lleida). Geo-Temas 6, 71–4.
Turner, A. H., Hwang, S. H. & Norell, M. A. 2007. A small derived theropod from Öösh, Early Cretaceous, Baykhangor Mongolia. American Museum Novitates 3557, 127.
Turner, A. H., Pol, D., Clarke, J. A., Erickson, G. M. &. Norell, M. A. 2007. A basal dromaeosaurid and size evolution preceding avian flight. Science 317, 1378–81.
Valença, L. M. M., Neumann, V. H. & Mabesoone, J. M. 2003. An overview on Callovian–Cenomanian intracratonic basins of Northeast Brazil: onshore stratrigraphic record of the opening of the southern Atlantic. Geologica Acta 1, 261–75.
Wang, S. C. & Dodson, P. 2006. Estimating the diversity of dinosaurs. Proceedings of the National Academy of Sciences of the United States of America 103, 13601–5.
Weishampel, D. B., Barrett, P. M., Coria, R. A., Le Loeuff, J., Xu, X., Zhao, X.-J., Sahni, A., Gomani, E. M. P. & Noto, C. R. 2004. Dinosaur distribution. In The Dinosauria, Second Edition (eds Weishampel, D. B., Dodson, P. & Osmólska, H..), pp. 517606. Berkeley: University of California Press.
Werner, C. 1995. Neue Funde von mesozoischen Wirbeltieren in Äthiopien. Berliner Geowissenschaftliche Abhandlungen 16, 377–83.
Xu, X. & Wang, X. 2004. A new dromaeosaur (Dinosauria: Theropoda) from the Early Cretaceous Yixian Formation of western Liaoning. Vertebrata PalAsiatica 42, 111–19.
Xu, X. & Wu, X. 2001. Cranial morphology of Sinornithosaurus millenii Xu et al. 1999 (Dinosauria: Theropoda: Dromaeosauridae) from the Yixian Formation of Liaoning, China. Canadian Journal of Earth Sciences 38, 1739–52.
Xu, X., Zhou, Z.-H. & Wang, X.-L. 2000. The smallest known non-avian theropod dinosaur. Nature 408, 705–8.
Zinke, J. 1998. Small theropod teeth from the Upper Jurassic coal mine of Guimarota (Portugal). Paläontologische Zeitschrift 72, 179–89.
Recommend this journal

Email your librarian or administrator to recommend adding this journal to your organisation's collection.

Geological Magazine
  • ISSN: 0016-7568
  • EISSN: 1469-5081
  • URL: /core/journals/geological-magazine
Please enter your name
Please enter a valid email address
Who would you like to send this to? *



Altmetric attention score

Full text views

Total number of HTML views: 0
Total number of PDF views: 0 *
Loading metrics...

Abstract views

Total abstract views: 0 *
Loading metrics...

* Views captured on Cambridge Core between <date>. This data will be updated every 24 hours.

Usage data cannot currently be displayed