Abstract
Thirty-seven well-preserved, isolated theropod teeth from the Early Cenomanian Kem Kem beds, Morocco, are identified by using morphometric data and direct comparison with teeth previously described in the literature. Direct comparison reveals that four different morphotypes (MT 1–4) are present in the sample. The teeth of MT 1 are characterised by unserrated carinae and belong to spinosaurid dinosaurs. The teeth of MT 2–4 have serrated carinae, and our data analysis indicates they are of carcharodontosaurid, dromaeosaurid, and abelisaurid origin. Three types of crown enamel ornamentation are present among the teeth of MT 1, which implies that, apart from Spinosaurus aegyptiacus STROMER 1915, more than one species of spinosaurine theropods may be present in the Early Cenomanian of Northern Africa. Our results also confirm the occurrence of abelisaurids, dromaeosaurids, and carcharodontosaurids in Morocco.
Kurzfassung
Mit Hilfe morphometrischer Daten und durch einen direkten Vergleich mit Zähnen aus der Literatur können 37 isolierte Zähne von Theropoden aus den Kem Kem Schichten (Unteres Cenomanium) von Marokko 4 verschiedenen Morphotypen (MT) zugeordnet werden. Die Zähne des MT 1 zeichnen sich durch ein Fehlen der Serrationen der Carinae aus und gehören deshalb in die Gruppe der spinosauriden Dinosaurier. Die Zähne der MT 2–4 sind durch serrierte Carinae charakterisiert und die morphometrische Datenanalyse zeigt, dass es sich um Zähne von caracharodontosauriden, dromaeosauriden und abelisauriden Dinosauriern handelt. An Zähnen des MT 1 sind drei verschiedene Ornamentierungstypen des Schmelzes zu beobachten, was darauf hindeutet, dass es neben Spinosaurus aegyptiacus STROMER 1915 noch mehr als nur eine Art von spinosaurinen Theropoden zur Zeit des Unteren Cenomanium in Nord-Afrika gegeben haben könnte. Die Ergebnisse unserer Studie zeigen außerdem, dass abelisauride, dromaeosauride und carcharodontosauride Theropoden zu dieser Zeit in Nord-Afrika gelebt haben müssen.
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References
Amiot, R., E. Buffetaut, H. Tong, L. Boudad, and L. Kabiri. 2004. Isolated theropod teeth from the Cenomanian of Morocco and their palaeobiogeographical significance. Revue de Paleobiologie Vol. spec 9: 143–149.
Bertin, T. 2010. A catalogue of material and review of the Spinosauridae. PalArch’s Journal of Vertebrate Palaeontology 7(4): 1–39.
Brusatte, S.L., R.B.J. Benson, T.D. Carr, T.E. Williamson, and P.C. Sereno. 2007. The systematic utility of theropod enamel wrinkles. Journal of Vertebrate Paleontology 27(4): 1052–1056.
Buffetaut, E. 1989. New remains of the enigmatic dinosaur Spinosaurus from the Cretaceous of Morocco and the affinities between Spinosaurus and Baryonyx. Neues Jahrbuch Geologische und Paläontologische Mitteilungen Monatshefte H 2: 79–87.
Buffetaut, E., F. Escuillié, and B. Pohl. 2005. First theropod dinosaur from the Maastrichtian phosphates of Morocco. Kaupia 14: 3–8.
Buffetaut, E., V. Suteethorn, H. Tong, and R. Amiot. 2008. An Early Cretaceous spinosaurid theropod from Southern China. Geological Magazine 145(5): 745–748.
Bouaziz, S., E. Buffetaut, M. Ghanmi, J.J. Jaeger, M. Martin, J.M. Mazin, and H. Tong. 1988. Nouvelles découvertes des vertébrés fossiles dans l’Albien du Sud tunisien. Bulletin de la Société géologique de France Série 8, 4(2): 335–339.
Carrano, M.T., S.D. Sampson, and C.A. Foster. 2002. The osteology of Masiakasaurus knopfleri, a small abelisauroid (Dinosauria: Theropoda) from the Late Cretaceous of Madagascar. Journal of Vertebrate Paleontology 22: 510–534.
Cavin, L., L. Boudad, S. Duffaud, L. Kabiri, J. LeLoeuff, I. Rouget, and H. Tong. 2001. L’évolution paléoenvironnementale des faunes des poissons du Crétacé supérieur du basin de Tafilalt et des régions avoisinantes (Sud-Est du Maroc); implications paléobiogeographiques. Comptes Rendus de l’Académie des Science—Series IIA—Earth and Planetary Science 333(10): 677–683.
Cavin, L., H. Tong, L. Boudad, C. Meister, A. Piuz, J. Tabouelle, M. Aarab, R. Amiot, E. Buffetaut, G. Dyke, S. Hua, and J. Le Loeuff. 2010. Vertebrate assemblages from the early Late Cretaceous of southeastern Morocco: An overview. Journal of African Earth Sciences 57: 391–412.
Charig, A.J., and A.C. Milner. 1997. Baryonyx walkeri: A fish-eating dinosaur from the Wealden of Surrey. Bulletin of the Natural History Museum, Geology series 53: 11–70.
Currie, P.J., J.K. Rigby Jr, and R.E. Sloan. 1990. Theropod teeth from the Judith River Formation of southern Alberta, Canada. In Dinosaur systematics: Perspectives and approaches, ed. K. Carpenter, and P.J. Currie, 107–125. Cambridge: Cambridge University Press.
Dutheil, D.B. 1999. An overview of the freshwater fish fauna from the Kem Kem beds (Late Cretaceous: Cenomanian) of southeastern Morocco. In Mesozoic fishes 2—systematics and fossil record, ed. G. Arratia, and H.P. Schultze, 553–563. München: Verlag Friedrich Pfeil.
Hammer, Ø., D.A.T. Harper, and P.D. Ryan. 2001. PAST: Paleontological statistics software package for education and data analysis. Palaeontologia Electronica 4(1). http://palaeo-electronica.org/2001_1/past/issue1_01.htm.
Hammer, Ø., and D.A.T. Harper. 2005. Paleontological data analysis. Malden: Wiley-Blackwell Publishing Ltd.
Hasegawa, Y., G. Tanaka, Y. Takakuwa, and S. Koike. 2010. Fine sculptures on a tooth of Spinosaurus (Dinosauria: Theropoda) from Morocco. Bulletin of the Gunma Museum of Natural History 14: 11–20.
Hwang, S. H., M. A. Norell, J. Qiang, and G. Geqin. 2002. New specimens of Microraptor zhaoianus (Theropoda: Dromaeosauridae) from Northeastern China. American Museum Novitates 3381.
Joly, F. 1962. Etudes sur le relief du sud-est marocain. Géographie Physique: Traveaux de l’Institut de Sciences Chérifien Série Géologie. 10.
Knoll, F., and J.I. Ruiz-Omeñaca. 2009. Theropod teeth from the basalmost Cretaceous of Anoual (Morocco) and their palaeobiogeographical significance. Geological Magazine 146(4): 602–616.
Lavocat, R. 1954. Reconnaissance géologique dans les hammadas des confines algero-marocains du sud. Notes Mémoires du Service géologique (Maroc) 116.
Mahler, L. 2005. Record of Abelisauridae (Dinosauria, Theropoda) from the Cenomanian of Morocco. Journal of Vertebrate Paleontology 25: 236–239.
Medeiros, M.A. 2006. Large theropod teeth from the Eocenomanian of northeastern Brazil and the occurrence of Spinosauridae. Revista Brasileira de Paleontologia 9(3): 333–338.
Norell, M.A., and P.J. Makovicky. 2004. Dinosaur systematics, chapter 10: Dromaeosauridae. In The Dinosauria, ed. D.B. Weishampel, P. Dodson, and H. Osmolska, 196–209. Berkley: University of California Press.
Novas, F.E., F. Dalla Vecchia, and D.F. Pais. 2005. Theropod pedal unguals from the Late Cretaceous (Cenomanian) of Morocco, Africa. Revista del Museo Argentino de Ciencias Naturales 7(2): 167–175.
Rauhut, O.W.M., and C. Werner. 1995. First record of the family Dromaeosauridae (Dinosauria: Theropoda) in the Cretaceous of Gondwana (Wadi Milk Formation, northern Sudan). Paläontologische Zeitschrift 69: 475–489.
Rauhut, O.W.M. 2003. The interrelationships and evolution of basal theropod dinosaurs. Special Papers in Palaeontology 69: 1–213.
Rauhut, O.W.M. 2004. Provenance and anatomy of Genyodectes serus, a large-toothed ceratosaur (Dinosauria: Theropoda) from Patagonia. Journal of Vertebrate Paleontology 24: 894–902.
Russell, D. 1996. Isolated dinosaur bones from the Middle Cretaceous of the Tafilalt, Morocco. Bulletin du Muséum national d’Histoire naturelle, Série 4, 18, Section C:349–402.
Sankey, J.T., D.B. Brinkman, M. Guenther, and P.J. Currie. 2002. Small theropod and bird teeth from the Late Cretaceous (Late Campanian) Judith River Group, Alberta. Journal of Paleontology 76(4): 751–763.
Sereno, P.C., D.B. Dutheil, M. Larochene, H.C.E. Larsson, G.H. Lyon, P.M. Magwene, C.A. Sidor, D.J. Varricchio, and J.A. Wilson. 1996. Predatory dinosaurs from the Sahara and Late Cretaceous faunal distribution. Science 272: 986–991.
Sereno, P.C., A.L. Beck, D.B. Dutheil, B. Gado, H.C.E. Larsson, G.H. Lyon, J.D. Marcot, O.W.M. Rauhut, R.W. Sadleir, C.A. Sidor, D.D. Varricchio, G.P. Wilson, and J.A. Wilson. 1998. A long-snouted predatory dinosaur from Africa and the evolution of Spinosaurids. Science 282: 1298–1302.
Sereno, P.C., and S.L. Brusatte. 2008. Basal abelisaurid and carcharodontosaurid theropods from the Lower Cretaceous Elrhaz Formation of Niger. Acta Palaeontologica Polonica 53(1): 15–46.
Smith, J.B. 2002. An examination of dental morphology and variation in theropod dinosaurs: Implications for the identification of isolated teeth. Ph.D. dissertation, University of Pennsylvania, Philadelphia.
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.
Smith, J.B. 2007. Dental morphology and variation in Majungasaurus crenatissimus (Theropoda: Abelisauridae) from the Late Cretaceous of Madagascar. Journal of Vertebrate Paleontology 27(2): 103–126.
Smith, J.B., D.R. Vann, and P. Dodson. 2005. Dental morphology and variation in theropod dinosaurs: implications for the taxonomic identification of isolated teeth. The Anatomical Record Part A 285: 699–736.
Smith, J.B., and M.C. Lamanna. 2006. An abelisaurid from the Late Cretaceous of Egypt: implications for theropod biogeography. Naturwissenschaften 93: 242–245.
Smith, J.B., and F.M. Dalla Vecchia. 2006. An abelisaurid (Dinosauria: Theropoda) tooth from the Lower Cretaceous Chicla formation of Libya. Journal of African Earth Sciences 46: 240–244.
Sokal, R.R., and F.J. Rohlf. 1995. Biometry: The principles and practice of statistics in biological research, 3rd edition. vi–xix, 1–887. New York: W. H. Freeman and Company.
Stromer, F. 1915. Ergebnisse der Forschungsreisen Prof. E. Stromers in den Wüsten Ägyptens 11. Wirbeltier-Reste der Baharijestufe (unterstes Cenoman).
3. Das Original des Theropoden Spinosaurus aegyptiacus nov. gen., nov. spec. Abhandlungen der Königlich Bayerischen Akademie der Wissenschaften, Mathematisch-physikalische Klasse 28(3):1–32.
Stromer, F. 1931. Ergebnisse der Forschungsreisen Prof. E. Stromers in den Wüsten Ägyptens 11. Wirbeltier-Reste der Baharijestufe (unterstes Cenoman).
10. Ein Skelett-Rest von Carcharodontosaurus nov. gen. Abhandlungen der Bayerischen Akademie der Wissenschaften, Mathematisch-naturwissenschaftliche Abteilung Neue Folge 9:1–23.
Tong, H., and E. Buffetaut. 1996. A new genus and species of pleurodiran turtle from the Cretaceous of southern Morocco. Neues Jahrbuch für Geologie und Paläontologie Abhandlungen 199(1):133–150.
Wellnhofer, P., and E. Buffetaut. 1999. Pterosaur remains from the Cretaceous of Morocco. Paläontologische Zeitschrift 73: 133–142.
Acknowledgments
We thank Dr. Mike Reich, Institute of Geobiology, Museum des Geowissenschaftlichen Zentrums, Georg August University Göttingen, for the loan of the theropod teeth specimens from the Alberti Collection of Morocco and Prof. Dr. Ulrich Joger for the loan of the theropod teeth specimens from the collection of the Naturhistorischen Museum Braunschweig.
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Richter, U., Mudroch, A. & Buckley, L.G. Isolated theropod teeth from the Kem Kem Beds (Early Cenomanian) near Taouz, Morocco. Paläontol Z 87, 291–309 (2013). https://doi.org/10.1007/s12542-012-0153-1
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DOI: https://doi.org/10.1007/s12542-012-0153-1
Keywords
- Theropoda
- Spinosaurinae
- Dromaeosauridae
- Carcharodontosauridae
- Abelisauridae
- Morocco
- Kem Kem beds
Schlüsselwörter
- Theropoda
- Spinosaurinae
- Dromaeosauridae
- Carcharodontosauridae
- Abelisauridae
- Marokko