A combined approach to identify isolated theropod teeth from the Cenomanian Kem Kem Group of Morocco: cladistic, discriminant, and machine learning analyses

LITERATURE CITED

• Alonso, A., Gasca, J. M., Navarro-Lorbés, P., Rubio, C., & Canudo, J. I. (2018). A new contribution to our knowledge of the large-bodied theropods from the Barremian of the Iberian Peninsula: The “Barranco del Hocino” site (Spain). Journal of Iberian Geology, 44(1), 7–23. https://doi.org/10.1007/s41513-018-0051-9
   Web of Science ®Google Scholar
• Alonso, A., Canudo, J. I, Navarro-Lorbés, P., & Cuenca-Bescós, G. (2015). Morfolometría geométrica aplicada al estudio de dientes aislados de dinosaurios terópodos. XIII EJIP Conference Proceedings, 19–21.
   Google Scholar
• Ameghino, F. (1899). Nota preliminar sobre el Loncosaurus argentinus, un representante de la familia de los Megalosauridae en la República Argentina. Anales de La Sociedad Científica Argentina, 47, 61–62.
   Google Scholar
• Amiot, R., Buffetaut, E., Tong, H., Boudad, L., & Kabiri, L. (2004). Isolated theropod teeth from the Cenomanian of Morocco and their palaeobiogeographical significance. Revue de Paléobiologie, 9, 143–149.
   Google Scholar
• Antunes, M. T., & Sigogneau-Russell, D. (1991). Nouvelles données sur les dinosaures du Crétacé supérieur du Portugal. Comptes rendus de l’Académie des sciences. Série 2, Mécanique, Physique, Chimie, Sciences de l’univers, Sciences de la Terre, 313(1), 113–119.
   Google Scholar
• Apesteguía, S., Smith, N. D., Valieri, R. J., & Makovicky, P. J. (2016). An unusual new theropod with a didactyl manus from the Upper Cretaceous of Patagonia, Argentina. PLoS ONE, 11(7), e0157793. https://doi.org/10.1371/journal.pone.0157793
   PubMed Web of Science ®Google Scholar
• 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 spinosaurids. Cretaceous Research, 93, 275–284. https://doi.org/10.1016/j.cretres.2018.06.013
   Web of Science ®Google Scholar
• Argast, S., Farlow, J. O., Gabet, R. M., & Brinkman, D. L. (1987). Transport-induced abrasion of fossil reptilian teeth: Implications for the existence of Tertiary dinosaurs in the Hell Creek Formation, Montana. Geology, 15(10), 927–930. https://doi.org/10.1130/0091-7613(1987)15<927:TAOFRT>2.0.CO;2
   Web of Science ®Google Scholar
• Averianov, A. O., Ivantsov, S. V., & Skutschas, P. P. (2019). Theropod teeth from the Lower Cretaceous Ilek Formation of Western Siberia, Russia. Proceedings of the Zoological Institute RAS, 323(2), 65–84.
   Google Scholar
• Beevor, T., Quigley, A., Smith, R. E., Smyth, R. S. H., Ibrahim, N., Zouhri, S., & Martill, D. M. (2021). Taphonomic evidence supports an aquatic lifestyle for Spinosaurus. Cretaceous Research, 117, 104627. https://doi.org/10.1016/j.cretres.2020.104627
   Web of Science ®Google Scholar
• Belvedere, M., Jalil, N.-E., Breda, A., Gattolin, G., Bourget, H., Khaldoune, F., & Dyke, G. J. (2013). Vertebrate footprints from the Kem Kem beds (Morocco): A novel ichnological approach to faunal reconstruction. Palaeogeography, Palaeoclimatology, Palaeoecology, 383–384, 52–58. https://doi.org/10.1016/j.palaeo.2013.04.026
   Web of Science ®Google Scholar
• Benson, R., Carrano, M., & Brusatte, S. (2010). A new clade of archaic large-bodied predatory dinosaurs (Theropoda: Allosauroidea) that survived to the latest Mesozoic. Naturwissenschaften, 97(1), 71–78. https://doi.org/10.1007/s00114-009-0614-x
   PubMedGoogle Scholar
• Benyoucef, M., Läng, E., Cavin, L., Mebarki, K., Adaci, M., & Bensalah, M. (2015). Overabundance of piscivorous dinosaurs (Theropoda: Spinosauridae) in the mid-Cretaceous of North Africa: The Algerian dilemma. Cretaceous Research, 55, 44–55. https://doi.org/10.1016/j.cretres.2015.02.002
   Web of Science ®Google Scholar
• Bonaparte, J. F., & Novas, F. E. (1985). Abelisaurus comahuensis, n.g., Carnosauria del Crétacico Tardio de Patagonia. Ameghiniana, 21(2–4), 259–265.
   Google Scholar
• Brusatte, S. L., Benson, R. B. J., Carr, T. D., Williamson, T. E., & Sereno, P. C. (2007). The systematic utility of theropod enamel wrinkles. Journal of Vertebrate Paleontology, 27(4), 1052–1056. https://doi.org/10.1671/0272-4634(2007)27[1052:TSUOTE]2.0.CO;2
   Web of Science ®Google Scholar
• Brusatte, S. L., & Carr, T. D. (2016). The phylogeny and evolutionary history of tyrannosauroid dinosaurs. Scientific Reports, 6, 20252. https://doi.org/10.1038/srep20252
   PubMed Web of Science ®Google Scholar
• Brusatte, S. L., & Sereno, P. C. (2007). A new species of Carcharodontosaurus (Dinosauria: Theropoda) from the Cenomanian of Niger and a revision of the genus. Journal of Vertebrate Paleontology, 27(4), 902–916. https://doi.org/10.1671/0272-4634(2007)27[902:ANSOCD]2.0.CO;2
   Web of Science ®Google Scholar
• Buckley, L. G., Larson, D. W., Reichel, M., & Samman, T. (2010). Quantifying tooth variation within a single population of Albertosaurus sarcophagus (Theropoda: Tyrannosauridae) and implications for identifying isolated teeth of tyrannosaurids. Canadian Journal of Earth Sciences, 47(9), 1227–1251.
   Web of Science ®Google Scholar
• Buffetaut, E. (1989). New remains of the enigmatic dinosaur Spinosaurus from the Cretaceous of Morocco and the affinities between Spinosaurus and Baryonyx. Neues Jahrbuch Für Geologie Und Paläontologie, Monatshefte, 2, 79–87.
   Google Scholar
• Buffetaut, E., & Ingavat, R. (1986). Unusual theropod dinosaur teeth from the Upper Jurassic of Phu Wiang, northeastern Thailand. Revue de Paléobiologie, 5(2), 217–220.
   Google Scholar
• 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(5), 415–421. https://doi.org/10.2113/173.5.415
   Google Scholar
• Canale, J. I., Novas, F. E., & Pol, D. (2015). Osteology and phylogenetic relationships of Tyrannotitan chubutensis Novas, de Valais, Vickers-Rich and Rich, 2005 (Theropoda: Carcharodontosauridae) from the Lower Cretaceous of Patagonia, Argentina. Historical Biology, 27(1), 1–32. https://doi.org/10.1080/08912963.2013.861830
   Web of Science ®Google Scholar
• 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. https://doi.org/10.1080/14772019.2011.630927
   Web of Science ®Google Scholar
• Carrano, M. T., & Sampson, S. D. (2008). The phylogeny of Ceratosauria (Dinosauria: Theropoda). Journal of Systematic Palaeontology, 6(2), 183–236. https://doi.org/10.1017/S1477201907002246
   Web of Science ®Google Scholar
• Cau, A., Beyrand, V., Voeten, D. F. A. E., Fernandez, V., Tafforeau, P., Stein, K., Barsbold, R., Tsogtbaatar, K., Currie, P. J., & Godefroit, P. (2017). Synchrotron scanning reveals amphibious ecomorphology in a new clade of bird-like dinosaurs. Nature, 552(7685), 395–399. https://doi.org/10.1038/nature24679
   PubMed Web of Science ®Google Scholar
• Cau, A., Dalla Vecchia, F. M., & Fabbri, M. (2013). A thick-skulled theropod (Dinosauria, Saurischia) from the Upper Cretaceous of Morocco with implications for carcharodontosaurid cranial evolution. Cretaceous Research, 40, 251–260. https://doi.org/10.1016/j.cretres.2012.09.002
   Web of Science ®Google Scholar
• Cau, A., Vecchia, F. M. D., & Fabbri, M. (2012). Evidence of a new carcharodontosaurid from the Upper Cretaceous of Morocco. Acta Palaeontologica Polonica, 57(3), 661–665. https://doi.org/10.4202/app.2011.0043
   Web of Science ®Google Scholar
• Cavin, L., Tong, H., Boudad, L., Meister, C., Piuz, A., Tabouelle, J., Aarab, M., Amiot, R., Buffetaut, E., Dyke, G., Hua, S., & Le Loeuff, J. (2010). Vertebrate assemblages from the early Late Cretaceous of southeastern Morocco: An overview. Journal of African Earth Sciences, 57(5), 391–412.
   Web of Science ®Google Scholar
• Chiarenza, A. A., & Cau, A. (2016). A large abelisaurid (Dinosauria, Theropoda) from Morocco and comments on the Cenomanian theropods from North Africa. PeerJ, 4, e1754.
   PubMed Web of Science ®Google Scholar
• Choubert, G., Clariond, L., & Hindermeyer, J. (1952). Livret-guide de l’excursion C 36. Anti-Atlas central et oriental. Livret-guide de l’excursion C 36. Anti-Atlas central et oriental. Congrès Géologique International, XIXe Session (Alger 1952), Série: Maroc, No. 11.
   Google Scholar
• Cillari, A. (2010). Teeth of Theropoda (Dinosauria, Saurischia): Morphology, function and classification. [Unpublished Ph.D. Dissertation]. Scienze della Terra, Sapienza Universitá di Roma.
   Google Scholar
• Coria, R. A., & Currie, P. J. (2006). A new carcharodontosaurid (Dinosauria, Theropoda) from the Upper Cretaceous of Argentina. Geodiversitas, 28(1), 71–118.
   Web of Science ®Google Scholar
• 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 University Press.
   Google Scholar
• 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), 2037–2081. https://doi.org/10.1139/e93-179
   Web of Science ®Google Scholar
• 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. https://doi.org/10.1671/0272-4634(2005)025[0888:NIOTSO]2.0.CO;2
   Web of Science ®Google Scholar
• D’Amore, D. C. (2009). A functional explanation for denticulation in theropod dinosaur teeth. The Anatomical Record: Advances in Integrative Anatomy and Evolutionary Biology, 292(9), 1297–1314. https://doi.org/10.1002/ar.20977
   Web of Science ®Google Scholar
• de Lapparent, A. F. (1960). Les dinosauriens du “Continental Intercalaire” du Sahara Central. Mémoire de La Société Géologique de France, 88A, l–57.
   Google Scholar
• del Corro, G. (1966). Un nuevo dinosaurio carnívoro del Chubut (Argentina). Communicaciones del Museo Argentino de Ciencias Naturales “Bernardino Rivadavia” e Institutio Nacional de Investigacion de Las Ciencias Naturales: Paleontologia, 1, 1–4.
   Google Scholar
• Depéret, C. (1896). Note sur les dinosauriens sauropodes et théropodes du Crétacé supérieur de Madagascar. Bulletin de La Société Géologique de France, 21, 176–194.
   Google Scholar
• Depéret, C., & Savornin, J. (1925). Sur la découverte d’une faune de vertébrés albiens à Timimoun (Sahara occidental). Comptes Rendus Hebdomadaires Des Séances de l’Académie Des Sciences à Paris, 181, 1108–1111.
   Google Scholar
• Ding, A., Pittman, M., Upchurch, P., O’Connor, J., Field, D. J., & Xu, X. (2020). The biogeography of coelurosaurian theropods and its impact on their evolutionary history. In M. Pittman & X. Xu (Eds.), Pennaraptoran Theropod Dinosaurs Past Progress and New Frontiers. Bulletin of the American Museum of Natural History (Vol. 40, Issue 440, pp. 117–157). American Museum of Natural History.
   Google Scholar
• D’Orazi Porchetti, S., Nicosia, U., Biava, A., & Maganuco, S. (2011). New abelisaurid material from the Upper Cretaceous (Cenomanian) of Morocco. Rivista Italiana Di Paleontologia e Stratigrafia, 117(3), 463–472.
   Web of Science ®Google Scholar
• Dutheil, D. B. (1999). An overview of the freshwater fish fauna from the Kem Kem beds (Late Cretaceous: Cenomanian) of southeastern Morocco. In: G. Arratia & H.-P. Schultze (Eds.), Mesozoic Fishes 2. Systematics and the Fossil Record (pp. 553–563). Verlag Dr. Friedrich Pfeil.
   Google Scholar
• Evans, D. C., Barrett, P. M., Brink, K. S., & Carrano, M. T. (2015). Osteology and bone microstructure of new, small theropod dinosaur material from the early Late Cretaceous of Morocco. Gondwana Research, 27(3), 1034–1041. https://doi.org/10.1016/j.gr.2014.03.016
   Web of Science ®Google Scholar
• Evans, S. E., & Milner, A. R. (1994). Middle Jurassic microvertebrate assemblages from the British Isles. In N. C. Fraser & H. D. Sues (Eds.), In the Shadow of the Dinosaurs: Early Mesozoic Tetrapods (pp. 303–321). Cambridge University Press.
   Google Scholar
• 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. https://doi.org/10.7717/peerj.1323
   PubMed Web of Science ®Google Scholar
• Ezcurra, M. (2017). A new early coelophysoid neotheropod from the Late Triassic of Northwestern Argentina. Ameghiniana, 54(5), 506–538.
   Web of Science ®Google Scholar
• Fanti, F., Cau, A., Martinelli, A., & Contessi, M. (2014). Integrating palaeoecology and morphology in theropod diversity estimation: A case from the Aptian-Albian of Tunisia. Palaeogeography, Palaeoclimatology, Palaeoecology, 410, 39–57. https://doi.org/10.1016/j.palaeo.2014.05.033
   Web of Science ®Google Scholar
• 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(1), 155–166.
   Web of Science ®Google Scholar
• Fitzinger, L. J. F. J. (1843). Systema reptilium. Fasciculus primus: Amblyglossae (Apud Braumüller and Seidel Bibliopolas).
   Google Scholar
• Forey, P. L., López-Arbarello, A., & MacLeod, N. (2011). A new species of Lepidotes (Actinopterygii: Semiontiformes) from the Cenomanian (Upper Cretaceous) of Morocco. Palaeontologia Electronica, 14(1), 1–12.
   Web of Science ®Google Scholar
• Galton, P. M., & Carpenter, K. (2006). Teeth of ornithischian dinosaurs (mostly Ornithopoda) from the Morrison Formation (Upper Jurassic) of Western United States. In: K. Carpenter (Ed.), Horns and Beaks. Ceratopsian and Ornithopod dinosaurs (pp. 17–47). Indiana University Press.
   Google Scholar
• Gates, T. A., Zanno, L. E., & Makovicky, P. J. (2015). Theropod teeth from the upper Maastrichtian Hell Creek Formation “Sue” Quarry: New morphotypes and faunal comparisons. Acta Palaeontologica Polonica, 60(1), 131–139.
   Web of Science ®Google Scholar
• Gauthier, J. (1986). Saurischian monophyly and the origin of birds. In K. Padian (Ed.), The origin of Birds and the Evolution of Flight (pp. 1–55). Memoirs of the California Academy of Sciences Number 8.
   Google Scholar
• 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. https://doi.org/10.1371/journal.pone.0158334
   PubMed Web of Science ®Google Scholar
• Goloboff, P. A., & Catalano, S. A. (2016). TNT version 1.5, including a full implementation of phylogenetic morphometrics. Cladistics, 32(3), 221–238. https://doi.org/10.1111/cla.12160
   PubMed Web of Science ®Google Scholar
• Hammer, Ø., Harper, D. A. T., & Ryan, P. D. (2001). Past: Paleontological Statistics Software Package for education and data analysis. Palaeontologia Electronica, 4(1), 1–9.
   Google Scholar
• Hassler, A., Martin, J. E., Amiot, R., Tacail, T., Godet, F. A., Allain, R., & Balter, V. (2018). Calcium isotopes offer clues on resource partitioning among Cretaceous predatory dinosaurs. Proceedings of the Royal Society B, 285(1876), 20180197. https://doi.org/10.1098/rspb.2018.0197
   PubMed Web of Science ®Google Scholar
• Hastie, M. T., Tibshirani, R., Leisch, F., Hornik, K., Ripley, B. D., & Narasimhan, B. (2020). Mda: Mixture and Flexible Discriminant Analysis. R package version 0.5-2. CRAN R Project.
   Google Scholar
• Heckeberg, N. S., & Rauhut, O. W. M. (2020). Histology of spinosaurid dinosaur teeth from the Albian-Cenomanian of Morocco: Implications for tooth replacement and ecology. Palaeontologia Electronica, 23(3), a48.
   Web of Science ®Google Scholar
• 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. https://doi.org/10.11646/zootaxa.3759.1.1
   PubMed Web of Science ®Google Scholar
• Hendrickx, C., Hartman, S. A., & Mateus, O. (2015a). An overview of non-avian theropod discoveries and classification. PalArch’s Journal of Vertebrate Palaeontology, 12(1), Article 1.
   Google Scholar
• Hendrickx, C., Mateus, O., & Araújo, R. (2015b). A proposed terminology of theropod teeth (Dinosauria, Saurischia). Journal of Vertebrate Paleontology, 35(5), e982797. https://doi.org/10.1080/02724634.2015.982797
   Web of Science ®Google Scholar
• Hendrickx, C., Mateus, O., & Araújo, R. (2015c). The dentition of megalosaurid theropods. Acta Palaeontologica Polonica, 60(3), 627–642. https://doi.org/10.4202/app.00056.2013
   Web of Science ®Google Scholar
• Hendrickx, C., Mateus, O., & Buffetaut, E. (2016). Morphofunctional analysis of the quadrate of Spinosauridae (Dinosauria: Theropoda) and the presence of Spinosaurus and a second spinosaurine taxon in the Cenomanian of North Africa. PLoS ONE, 11(1), e0144695. https://doi.org/10.1371/journal.pone.0144695
   PubMed Web of Science ®Google Scholar
• Hendrickx, C., Mateus, O., Araújo, R., & Choiniere, J. (2019). The distribution of dental features in non-avian theropod dinosaurs: Taxonomic potential, degree of homoplasy, and major evolutionary trends. Palaeontologia Electronica, 22(3), 1–110. https://doi.org/10.26879/820
   Web of Science ®Google Scholar
• Hendrickx, C., Stiegler, J., Currie, P. J., Han, F., Xu, X., Choiniere, J. N., & Wu, X.-C. (2020b). Dental anatomy of the apex predator Sinraptor dongi (Theropoda: Allosauroidea) from the Late Jurassic of China. Canadian Journal of Earth Sciences, 57(9), 1127–1147. https://doi.org/10.1139/cjes-2019-0231
   Web of Science ®Google Scholar
• Hendrickx, C., Tschopp, E., & Ezcurra, M. D. (2020a). Taxonomic identification of isolated theropod teeth: The case of the shed tooth crown associated with Aerosteon (Theropoda: Megaraptora) and the dentition of Abelisauridae. Cretaceous Research, 108, 104312. https://doi.org/10.1016/j.cretres.2019.104312
   Web of Science ®Google Scholar
• Hislop, S. (1861). Remarks on the geology of Nágpur. Journal of the Bombay Branch of the Royal Asiatic Society, 6, 194–206.
   Google Scholar
• Holwerda, F. M. (2020). Sauropod dinosaur fossils from the Kem Kem and extended ‘Continental Intercalaire’ of North Africa: A review. Journal of African Earth Sciences, 163, 103738. https://doi.org/10.1016/j.jafrearsci.2019.103738
   Web of Science ®Google Scholar
• Holwerda, F. M., Díaz, V. D., Blanco, A., Montie, R., & Reumer, J. W. F. (2018). Late Cretaceous sauropod tooth morphotypes may provide supporting evidence for faunal connections between North Africa and Southern Europe. PeerJ, 6, e5925. https://doi.org/10.7717/peerj.5925
   PubMed Web of Science ®Google Scholar
• Hone, D. W. E., & Holtz, T. R. J. (2017). A century of spinosaurs—a review and revision of the Spinosauridae, with comments on their ecology. Acta Geologica Sinica - English Edition, 91(3), 1120–1132. https://doi.org/10.1111/1755-6724.13328
   Web of Science ®Google Scholar
• Hone, D. W. E., & Holtz, T. R. J. (2021). Evaluating the ecology of Spinosaurus: Shoreline generalist or aquatic pursuit specialist? Palaeontologia Electronica, 24(1), a03.
   Web of Science ®Google Scholar
• Hwang, S. H. (2007). Phylogenetic patterns of enamel microstructure in dinosaur teeth [Unpublished Ph.D. Dissertation]. Columbia University, New York, New York.
   Google Scholar
• Ibrahim, N., Sereno, P. C., Sasso, C. D., Maganuco, S., Fabbri, M., Martill, D. M., Zouhri, S., Myhrvold, N., & Iurino, D. A. (2014b). Semiaquatic adaptations in a giant predatory dinosaur. Science, 345(6204), 1613–1616. https://doi.org/10.1126/science.1258750
   PubMed Web of Science ®Google Scholar
• Ibrahim, N., Varricchio, D. J., Sereno, P. C., Wilson, J. A., Dutheil, D. B., Martill, D. M., Baidder, L., & Zouhri, S. (2014a). Dinosaur footprints and other ichnofauna from the Cretaceous Kem Kem Beds of Morocco. PLoS ONE, 9(3), e90751. https://doi.org/10.1371/journal.pone.0090751
   PubMed Web of Science ®Google Scholar
• Ibrahim, N., Maganuco, S., Dal Sasso, C., Fabbri, M., Auditore, M., Bindellini, G., Martill, D. M., Zouhri, S., Mattarelli, D. A., Unwin, D. M., Wiemann, J., Bonadonna, D., Amane, A., Jakubczak, J., Joger, U., Lauder, G. V., & Pierce, S. E. (2020b). Tail-propelled aquatic locomotion in a theropod dinosaur. Nature, 581(7806), Article 7806. https://doi.org/10.1038/s41586-020-2190-3
   PubMed Web of Science ®Google Scholar
• Ibrahim, N., Sereno, P. C., Varricchio, D. J., Martill, D. M., Dutheil, D. B., Unwin, D. M., Baidder, L., Larsson, H. C. E., Zouhri, S., & Kaoukaya, A. (2020a). Geology and paleontology of the Upper Cretaceous Kem Kem Group of eastern Morocco. ZooKeys, 928, 1–216. https://doi.org/10.3897/zookeys.928.47517
   PubMedGoogle Scholar
• Janensch, W. (1925). Die Coelurosaurier und Theropoden der Tendaguruschichten Deutsch-Ostafrikas. Palaeontographica 7 (Supplement 1), 1–100.
   Google Scholar
• Kellner, A. W., & Mader, B. J. (1997). Archosaur teeth from the Cretaceous of Morocco. Journal of Paleontology, 71(3), 525–527.
   Web of Science ®Google Scholar
• Kuhn, M. (2008). Building predictive models in R using the caret package. Journal of Statistical Software, 28(5), 1–26.
   PubMed Web of Science ®Google Scholar
• Kuhn, M., Weston, S., Culp, M., Coulter, N., & Quinlan, R. (2018). C50: C5. 0 Decision Trees and Rule-Based Models (Version 0.1.2).
   Google Scholar
• Lacerda, M. B., Aragão, P. R. L., Vieira, F. S., Sales, M. A. F., & Liparini, A. (2023). On the first Baryonychinae (Theropoda, Spinosauridae) teeth from South America. Zootaxa, 5264(4), 526–544.
   PubMed Web of Science ®Google Scholar
• 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. https://doi.org/10.1016/j.cretres.2018.09.012
   Web of Science ®Google Scholar
• Läng, E., Boudad, L., Maio, L., Samankassou, E., Tabouelle, J., Tong, H., & Cavin, L. (2013). Unbalanced food web in a Late Cretaceous dinosaur assemblage. Palaeogeography, Palaeoclimatology, Palaeoecology, 381–382, 26–32. https://doi.org/10.1016/j.palaeo.2013.04.011
   Web of Science ®Google Scholar
• 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. https://doi.org/10.1371/journal.pone.0054329
   PubMed Web of Science ®Google Scholar
• Lavocat, R. (1948). Découverte de Crétacé à vertébrés dans le soubassement de la Hammada du Guir (Sud marocain). Comptes Rendus de l’Académie Des Sciences, 226(16), 1291–1292.
   Google Scholar
• Lavocat, R. (1954a). Reconnaissance géologique dans les Hammadas des confins algéro-marocains du Sud. Notes et Mémoires Du Service Géologique Du Maroc, 116, 1–147.
   Google Scholar
• Lavocat, R. (1954b). Sur les dinosauriens du Continental Intercalaire des Kem-Kem de la Daoura. 19th International Geological Congress, 15, 65–68.
   Google Scholar
• Leidy, J. (1856). Notice of remains of extinct reptiles and fishes, discovered by Dr. F.V. Hayden in the Bad Lands of the Judith River, Nebraska Territory. Proceedings of the Academy of Natural Sciences of Philadelphia, 8, 72–73.
   Google Scholar
• Liaw, A., & Wiener, M. (2002). Classification and regression by randomForest. R News, 2(3), 18–22.
   Google Scholar
• Longrich, N. R., Pereda-Suberbiola, X., Jalil, N.-E., Khaldoune, F., & Jourani, E. (2017). An abelisaurid from the latest Cretaceous (late Maastrichtian) of Morocco, North Africa. Cretaceous Research, 76, 40–52. https://doi.org/10.1016/j.cretres.2017.03.021
   Web of Science ®Google Scholar
• Lydekker, R. (1890). Note on certain vertebrate remains from the Nagpur district. Records of the Geological Survey of India, 23(1), 20–24.
   Google Scholar
• Maganuco, S., & Sasso, C. D. (2018). The smallest biggest theropod dinosaur: A tiny pedal ungual of a juvenile Spinosaurus from the Cretaceous of Morocco. PeerJ, 6, e4785. https://doi.org/10.7717/peerj.4785
   PubMed Web of Science ®Google Scholar
• Mahler, L. (2005). Record of Abelisauridae (Dinosauria: Theropoda) from the Cenomanian of Morocco. Journal of Vertebrate Paleontology, 25(1), 236–239.
   Web of Science ®Google Scholar
• Malafaia, E., Escaso, F., Mocho, P., Serrano-Martínez, A., Torices, A., Cachão, M., & Ortega, F. (2017). Analysis of diversity, stratigraphic and geographical distribution of isolated theropod teeth from the Upper Jurassic of the Lusitanian Basin, Portugal. Journal of Iberian Geology, 43(2), 257–291. https://doi.org/10.1007/s41513-017-0021-7
   Web of Science ®Google Scholar
• Mannion, P. D., & Barrett, P. M. (2013). Additions to the sauropod dinosaur fauna of the Cenomanian (early Late Cretaceous) Kem Kem beds of Morocco: Palaeobiogeographical implications of the mid-Cretaceous African sauropod fossil record. Cretaceous Research, 45, 49–59. https://doi.org/10.1016/j.cretres.2013.07.007
   Web of Science ®Google Scholar
• Marsh, O. C. (1878). Notice of new dinosaurian reptiles. American Journal of Science, 87, 241–244.
   Google Scholar
• Marsh, O. C. (1881). Principal characters of American Jurassic dinosaurs. Part V. American Journal of Science (Series 3), 21, 417–423.
   Google Scholar
• Marsh, O. C. (1884). The classification and affinities of dinosaurian reptiles. Nature, 31, 68–69.
   Google Scholar
• McFeeters, B. (2020). New mid-cervical vertebral morphotype of Spinosauridae from the Kem Kem Group of Morocco. Vertebrate Anatomy Morphology Palaeontology, 8, 182–193. https://doi.org/10.18435/vamp29370
   Google Scholar
• McFeeters, B., Ryan, M. J., Hinic-Frlog, S., & Schröder-Adams, C. (2013). A reevaluation of Sigilmassasaurus brevicollis (Dinosauria) from the Cretaceous of Morocco. Canadian Journal of Earth Sciences, 50(6), 636–649. https://doi.org/10.1139/cjes-2012-0129
   Web of Science ®Google Scholar
• McGowan, A. J., & Dyke, G. J. (2009). A surfeit of theropods in the Moroccan Late Cretaceous? Comparing diversity estimates from field data and fossil shops. Geology, 37(9), 843–846. https://doi.org/10.1130/G30188A.1
   Web of Science ®Google Scholar
• Meso, J. G., Hendrickx, C., Baiano, M. A., Canale, J. I., Salgado, L., & Martinez, I. D. (2021). Isolated theropod teeth associated with a sauropod skeleton from the Late Cretaceous Allen Formation of Río Negro, Patagonia, Argentina. Acta Palaeontologica Polonica, 66(2), 409–423. https://doi.org/10.4202/app.00847.2020
   Web of Science ®Google Scholar
• Milner, A. C. (2003). Fish-eating theropods: A short review of the systematics, biology and palaeobiogeography of spinosaurs. In: H. Hurtado & F. Torcida Férnandes-Baldor (eds) Actas de las II Jornadas Internacionales sobre Paleontología de Dinosaurios y sus Entornos. Salas de los Infantes, Burgos, España, Septiembre de 2001: 129–138.
   Google Scholar
• Motta, M. J., Aranciago Rolando, A. M., Rozadilla, S., Agnolín, F. E., Chimento, N. R., Egli, F. B., & Novas, F. E. (2016). New theropod fauna from the Upper Cretaceous (Huincul Formation) of northwestern Patagonia, Argentina. New Mexico Museum of Natural History and Science Bulletin, 71, 231–253.
   Google Scholar
• Müller, R. T., Langer, M. C., Bronzati, M., Pacheco, C. P., Cabreira, S. F., & Dias-Da-Silva, S. (2018). Early evolution of sauropodomorphs: Anatomy and phylogenetic relationships of a remarkably well-preserved dinosaur from the Upper Triassic of southern Brazil. Zoological Journal of the Linnean Society, 184(4), 1187–1248. https://doi.org/10.1093/zoolinnean/zly009
   Web of Science ®Google Scholar
• Niedźwiedzki, G., & Gierliński, G. (2002). Isolated theropod teeth from the Cretaceous strata of Khouribga, Morocco. Geological Quarterly, 46(1), 97–100.
   Google Scholar
• Owen, R. (1842). Report on British fossil reptiles. Report of the British Association for the Advancement of Science, 11, 60–294.
   Google Scholar
• Owen, S. R. (1854). On some fossil reptilian and mammalian remains from the Purbeck. Quarterly Journal of the Geological Society, 10(1–2), 420–433. https://doi.org/10.1144/GSL.JGS.1854.010.01-02.48
   Google Scholar
• Paterna, A., & Cau, A. (2022). New giant theropod material from the Kem Kem Compound Assemblage (Morocco) with implications on the diversity of the mid-Cretaceous carcharodontosaurids from North Africa. Historical Biology, 35(11), 2036–2044. https://doi.org/10.1080/08912963.2022.2131406
   Web of Science ®Google Scholar
• Paul, G. S. (2002). Dinosaurs of the Air: The Evolution and Loss of Flight in Dinosaurs and Birds. Johns Hopkins University Press.
   Google Scholar
• Rauhut, O. W. M., Hübner, T., & 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), 1–65.
   Web of Science ®Google Scholar
• Rauhut, O. W. M. (2003). The interrelationships and evolution of basal theropod dinosaurs. Special Papers in Palaeontology, 69, 1–213.
   Google Scholar
• Rauhut, O. W. M., & Carrano, M. T. (2016). The theropod dinosaur Elaphrosaurus bambergi Janensch, 1920, from the Late Jurassic of Tendaguru, Tanzania. Zoological Journal of the Linnean Society, 178(3), 546–610. https://doi.org/10.1111/zoj.12425
   Web of Science ®Google Scholar
• Rauhut, O. W. M., & Foth, C. (2020). The Origin of Birds: Current Consensus, Controversy, and the Occurrence of Feathers. In C. Foth & O. W. M. Rauhut (Eds.), The Evolution of Feathers: From Their Origin to the Present (pp. 27–45). Springer International Publishing. https://doi.org/10.1007/978-3-030-27223-4_3
   Google Scholar
• Rauhut, O. W. M., Foth, C., Tischlinger, H., & Norell, M. A. (2012). Exceptionally preserved juvenile megalosauroid theropod dinosaur with filamentous integument from the Late Jurassic of Germany. Proceedings of the National Academy of Sciences, 109(29), 11746–11751. https://doi.org/10.1073/pnas.1203238109
   PubMed Web of Science ®Google Scholar
• Rauhut, O. W. M., & Kriwet, J. (1994). Teeth of a big theropod dinosaur from Porto das Barcas (Portugal). Berliner Geowissenschaftliche Abhandlungen, E, 13, 179–185.
   Google Scholar
• 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(1), 155–195. https://doi.org/10.1111/j.1096-3642.2009.00591.x
   Web of Science ®Google Scholar
• 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(3), 475–489. https://doi.org/10.1007/BF02987808
   Google Scholar
• Ribeiro, T. B., da Costa Pereira, P. V. L. G., Brusatte, S. L., dos Anjos Candeiro, C. R., & Bergqvist, L. P. (2022). An eye for an eye, a tooth for a tooth: Archosaurian teeth from the Açu Formation (Albian–Cenomanian), Potiguar Basin, Northeast Brazil. Cretaceous Research, 129, 105005. https://doi.org/10.1016/j.cretres.2021.105005
   Web of Science ®Google Scholar
• Richter, U., Mudroch, A., & Buckley, L. G. (2013). Isolated theropod teeth from the Kem Kem Beds (Early Cenomanian) near Taouz, Morocco. Paläontologische Zeitschrift, 87(2), 291–309. https://doi.org/10.1007/s12542-012-0153-1
   Google Scholar
• RStudio Team. (2020). RStudio: Integrated Development for R. RStudio, PBC, Boston, MA URL http://www.rstudio.com/.
   Google Scholar
• Russell, D. A. (1996). Isolated dinosaur bones from the Middle Cretaceous of the Tafilalt, Morocco. Bulletin Du Muséum National d’Histoire Naturelle, 4(18), 349–402.
   Google Scholar
• Russell, D. A., & Paesler, M. A. (2003). Environments of Mid-Cretaceous Saharan dinosaurs. Cretaceous Research, 24(5), 569–588. https://doi.org/10.1016/S0195-6671(03)00072-7
   Web of Science ®Google Scholar
• Sankey, J. T. (2008). Diversity of Latest Cretaceous (Late Maastrichtian) small theropods and birds: Teeth from the Lance and Hell Creek Formations, USA. In J. T. Sankey & S. Baszio (Eds.), Vertebrate Microfossil Assemblages: Their Role in Paleoecology and Paleobiogeography (pp. 117–134). Indiana University Press.
   Google Scholar
• Sereno, P. C., Beck, A. L., Dutheil, D. B., Gado, B., Larsson, H. C. E., Lyon, G. H., Marcot, J. D., Rauhut, O. W. M., Sadleir, R. W., Sidor, C. A., Varricchio, D. D., Wilson, G. P., & Wilson, J. A. (1998). A long-snouted predatory dinosaur from Africa and the evolution of spinosaurids. Science, 282(5392), 1298–1302. https://doi.org/10.1126/science.282.5392.1298
   PubMed Web of Science ®Google Scholar
• Sereno, P. C., Dutheil, D. B., Larochene, 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(5264), 986–991. https://doi.org/10.1126/science.272.5264.986
   PubMed Web of Science ®Google Scholar
• Sereno, P. C., Wilson, J. A., & Conrad, J. L. (2004). New dinosaurs link southern landmasses in the Mid–Cretaceous. Proceedings of the Royal Society of London. Series B: Biological Sciences, 271(1546), 1325–1330. https://doi.org/10.1098/rspb.2004.2692
   PubMed Web of Science ®Google Scholar
• Serrano-Brañas, C. I., Gutiérrez-Blando, C., Bigurra, R. D., & González-León, C. M. (2017). First occurrence of tyrannosaurid theropods from the Corral de Enmedio Formation (Upper Cretaceous) Sonora, México. Cretaceous Research, 75, 81–93. https://doi.org/10.1016/j.cretres.2017.03.015
   Web of Science ®Google Scholar
• 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. https://doi.org/10.4202/app.00101.2014
   Web of Science ®Google Scholar
• Sharma, A., Hendrickx, C., & Singh, S. (2023). First theropod record from the marine Bathonian of Jaisalmer Basin, Tethyan coast of Gondwanan India. Rivista Italiana Di Paleontologia e Stratigrafia, 129(1), Article 1. https://doi.org/10.54103/2039-4942/18306
   Web of Science ®Google Scholar
• Smith, J. B. (2007). Dental morphology and variation in Majungasaurus crenatissimus (Theropoda: Abelisauridae) from the Late Cretaceous of Madagascar. Journal of Vertebrate Paleontology, 27(sup2), 103–126. https://doi.org/10.1671/0272-4634(2007)27[103:DMAVIM]2.0.CO;2
   Web of Science ®Google Scholar
• 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(2), 699–736. https://doi.org/10.1002/ar.a.20206
   Google Scholar
• Smith, R. E., & Martill, D. M. (2023). An unusual dental pathology in a tooth of Spinosaurus (Dinosauria, Theropoda) from the mid-Cretaceous of Morocco. Cretaceous Research, 146, 105499. https://doi.org/10.1016/j.cretres.2023.105499
   Web of Science ®Google Scholar
• Smyth, R. S. H., Ibrahim, N., Kao, A., & Martill, D. M. (2020a). Abelisauroid cervical vertebrae from the Cretaceous Kem Kem beds of Southern Morocco and a review of Kem Kem abelisauroids. Cretaceous Research, 108, 104330. https://doi.org/10.1016/j.cretres.2019.104330
   Web of Science ®Google Scholar
• Smyth, R. S. H., Ibrahim, N., & Martill, D. M. (2020b). Sigilmassasaurus is Spinosaurus: A reappraisal of African spinosaurines. Cretaceous Research, 114, 104520. https://doi.org/10.1016/j.cretres.2020.104520
   Web of Science ®Google Scholar
• Soto, M., Toriño, P., & Perea, D. (2019). A large sized megalosaurid (Theropoda, Tetanurae) from the Late Jurassic of Uruguay and Tanzania. Journal of South American Earth Sciences, 102458. https://doi.org/10.1016/j.jsames.2019.102458
   Google Scholar
• Stromer, E. (1915). Ergebnisse der Forschungsreisen Prof. E. Stromers in den Wüsten Ägyptens. II. Wirbeltier-Reste der Baharîje-Stufe (unterstes Cenoman). 3. Das Original des Theropoden Spinosaurus aegyptiacus nov. gen., nov. spec. Abhandlungen der Königlichen Bayerischen Akademie der Wissenschaften, Mathematisch- Physikalische Klasse, 28, 1–32.
   Google Scholar
• Stromer, E. (1931). Wirbeltier-Reste der Baharije-Stufe (unterstes Cenoman). 10. Ein Skelett-Rest von Carcharodontosaurus nov. Gen. Abhandlungen der Bayerischen Akademie der Wissenschaften, Mathematisch-Naturwissenschaftliche Abteilung, 9, 1–31.
   Google Scholar
• 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. https://doi.org/10.1671/0272-4634(2002)022[0535:ICASDT]2.0.CO;2
   Web of Science ®Google Scholar
• Taquet, P., & Russell, D. A. (1998). New data on spinosaurid dinosaurs from the Early Cretaceous of the Sahara. Comptes Rendus de l’Académie Des Sciences-Series IIA-Earth and Planetary Science, 327(5), 347–353.
   Google Scholar
• Team, R. C. (2020). R: A language and environment for statistical computing. R Foundation for Statistical Computing website.
   Google Scholar
• Torices, A., Reichel, M., & Currie, P. J. (2014). Multivariate analysis of isolated tyrannosaurid teeth from the Danek Bonebed, Horseshoe Canyon Formation, Alberta, Canada. Canadian Journal of Earth Sciences, 51(11), 1045–1051. https://doi.org/10.1139/cjes-2014-0072
   Web of Science ®Google Scholar
• Underwood, C. J., & Ward, D. (2018). Site specific limitations on the use of palaeontological resources. The Geological Curator, 10(10), 617–631.
   Google Scholar
• Walker, A. D. (1964). Triassic reptiles from the Elgin Area: Ornithosuchus and the origin of carnosaurs. Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences, 248(744), 53–134. https://doi.org/10.1098/rstb.1964.0009
   Web of Science ®Google Scholar
• Wang, S., Stiegler, J., Amiot, R., Wang, X., Du, G., Clark, J. M., & Xu, X. (2017). Extreme ontogenetic changes in a ceratosaurian theropod. Current Biology, 27(1), 144–148. https://doi.org/10.1016/j.cub.2016.10.043
   PubMed Web of Science ®Google Scholar
• Wellnhofer, P., & Buffetaut, E. (1999). Pterosaur remains from the Cretaceous of Morocco. Paläontologische Zeitschrift, 73(1–2), 133–142. https://doi.org/10.1007/BF02987987
   Google Scholar
• 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. https://doi.org/10.1371/journal.pone.0093190
   PubMed Web of Science ®Google Scholar
• Wills, S., Underwood, C. J., & Barrett, P. M. (2021). Learning to see the wood for the trees: Machine learning, decision trees, and the classification of isolated theropod teeth. Palaeontology, 64(1), 75–99. https://doi.org/10.1111/pala.12512
   Web of Science ®Google Scholar
• Wills, S., Underwood, C. J., & Barrett, P. M. (2023). Machine learning confirms new records of maniraptoran theropods in Middle Jurassic UK microvertebrate faunas. Papers in Palaeontology. https://doi.org/10.1002/spp2.1487
   Web of Science ®Google Scholar
• Wilson, J. A., & Allain, R. (2015). Osteology of Rebbachisaurus garasbae Lavocat, 1954, a diplodocoid (Dinosauria, Sauropoda) from the early Late Cretaceous–aged Kem Kem beds of southeastern Morocco. Journal of Vertebrate Paleontology, 35(4), e1000701. https://doi.org/10.1080/02724634.2014.1000701
   Web of Science ®Google Scholar
• Young, C. M. E., Hendrickx, C., Challands, T. J., Foffa, D., Ross, D. A., Butler, I. B., & Brusatte, S. L. (2019). New theropod dinosaur teeth from the Middle Jurassic of the Isle of Skye, Scotland. Scottish Journal of Geology, 55(1), 7–19. https://doi.org/10.1144/sjg2018-020
   Google Scholar
• Zinke, J. (1998). Small theropod teeth from the Upper Jurassic coal mine of Guimarota (Portugal). Paläontologische Zeitschrift, 72(1), 179–189.
   Google Scholar
• Zinke, J., & Rauhut, O. W. M. (1994). Small theropods (Dinosauria, Saurischia) from the Upper Jurassic and Lower Cretaceous of the Iberian Peninsula. Berliner Geowissenschaftliche Abhandlugen (E), 13, 163–177.
   Google Scholar
• Zitouni, S., Laurent, C., Dyke, G., & Jalil, N.-E. (2019). An abelisaurid (Dinosauria: Theropoda) ilium from the Upper Cretaceous (Cenomanian) of the Kem Kem beds, Morocco. PLoS ONE, 14(4), e0214055. https://doi.org/10.1371/journal.pone.0214055
   PubMed Web of Science ®Google Scholar