Csiki-Sava, Z., Buffetaut, E., Ősi, A., Pereda-Suberbiola, X. & Brusatte, S. L. Island life in the Cretaceous — faunal composition, biogeography, evolution, and extinction of land-living vertebrates on the Late Cretaceous European archipelago. Zookeys 469, 1–161 (2015).
Weishampel, D. B., Jianu, C.-M., Csiki, Z. & Norman, D. B. Osteology and phylogeny of Zalmoxes (n. g.), an unusual euornithopod dinosaur from the latest Cretaceous of Romania. J. Syst. Palaeontol. 1, 65–123 (2003).
Augustin, F., Ősi, A. & Csiki-Sava, Z. The Rhabdodontidae (Dinosauria, Ornithischia), an enigmatic dinosaur group endemic to the Late Cretaceous European Archipelago. Fossil Rec. 26, 171–189 (2023).
Czepiński, Ł & Madzia, D. Osteology, phylogenetic affinities, and palaeobiogeographic significance of the bizarre ornithischian dinosaur Ajkaceratops kozmai from the Late Cretaceous European archipelago. Zoo. J. Linn. Soc. 202, 1–25 (2024).
Ősi, A., Butler, R. J. & Weishampel, D. B. A Late Cretaceous ceratopsian dinosaur from Europe with Asian affinities. Nature 465, 466–468 (2010).
Ryan, M. J., Chinnery-Allgeier, B. J. & Eberth, D. A. New Perspectives on Horned Dinosaurs (Indiana Univ. Press, 2010).
You, H. & Dodson, P. in The Dinosauria (eds Weishampel, D. B., Dodson, P. and Osmólska, H.) 478–493 (Univ. California Press, 2004).
Norman, D. B., Sues, H.-D., Witmer, L. M. & Coria, R. A. in The Dinosauria (eds Weishampel, D. B., Dodson, P. and Osmólska, H.) 393–412 (Univ. California Press, 2004).
Maidment, S. C. R. & Barrett, P. M. Does morphological convergence imply functional similarity? A test using the evolution of quadrupedalism in ornithischian dinosaurs. Proc. R. Soc. B 279, 3765–3771 (2012).
Nabavizadeh, A. Evolutionary trends in the jaw adductor mechanics of ornithischian dinosaurs. Anat. Rec. 299, 271–294 (2016).
Virág, A. & Ősi, A. Morphometry, microstructure, and wear pattern of neornithischian dinosaur teeth from the Upper Cretaceous Iharkút locality (Hungary). Anat. Rec. 300, 1439–1463 (2017).
Farke, A. A., Maxwell, W. D., Cifelli, R. L. & Wedel, M. J. A ceratopsian dinosaur from the Lower Cretaceous of western North America, and the biogeography of Neoceratopsia. PLoS ONE 9, e112055 (2014).
Longrich, N. R. A ceratopsian dinosaur from the Late Cretaceous of eastern North America, and implications for dinosaur biogeography. Cret. Res. 57, 199–207 (2016).
Morschhauser, E. M. et al. Phylogenetic history of Auroraceratops rugosus (Ceratopsia: Ornithischia) from the Lower Cretaceous of Gansu Province, China. J. Vertebr. Paleontol. 38, 117–147 (2019).
Godefroit, P. & Lambert, O. A re-appraisal of Craspedodon lonzeensis Dollo, 1883 from the Upper Cretaceous of Belgium: the first record of a neoceratopsian dinosaur in Europe? Bull. l’Inst. R. Sci. Naturelles Belgique Sci. Terre 77, 83–93 (2007).
Lindgren, J. et al. The first neoceratopsian dinosaur remains from Europe. Palaeontology 50, 929–937 (2007).
Han, F., Forster, C. A., Xu, X. & Clark, J. M. Postcranial anatomy of Yinlong downsi (Dinosauria: Ceratopsia) from the Upper Jurassic Shishugou Formation of China and the phylogeny of basal ornithischians. J. Syst. Palaeontol. 16, 1159–1187 (2018).
Butler, R. J. & Sullivan, R. M. The phylogenetic position of Stenopelix valdensis from the Lower Cretaceous of Germany and the early fossil record of Pachycephalosauria. Acta Palaeontol. Pol. 54, 21–34 (2009).
Bertozzo, F., Kecheng, N., Gillette, N. V. & Godefroit, P. Anatomical description and digital reconstruction of the skull of Jeholosaurus shangyuanensis (Dinosauria, Ornithopoda) from China. PLoS ONE 20, e0312519 (2025).
Matheron, P. in Mémoires de l’Académie Impériale des Sciences, Belles – Lettres et Arts de Marseille 1–39 (Académie des Sciences, Lettres et Arts de Marseille, Marseille, 1869).
Pincemaille-Quillevere, M. Description d’un squelette partiel de Rhabdodon priscus (Euornithopoda) du Crétacé supérieur de Vitrolles (Bouches du Rhône, France). Oryctos 4, 39–70 (2002).
Ősi, A., Prondvai, E., Butler, R. J. & Weishampel, D. B. Phylogeny, histology and inferred body size evolution in a new rhabdodontid dinosaur from the Late Cretaceous of Hungary. PLoS ONE 7, e44318 (2012).
Magyar, J., Csiki-Sava, Z., Ősi, A., Augustin, F. J. & Botfalvai, G. Rhabdodontid (Dinosauria, Ornithopoda) diversity suggested by the first documented occurrence of associated cranial and postcranial material at Vălioara (uppermost Cretaceous Densuș-Ciula Formation, Hațeg Basin, Romania). Cret. Res. 156, 105810 (2024).
Czepiński, Ł. Ontogeny and variation of a protoceratopsid dinosaur Bagaceratops rozhdestvenskyi from the Late Cretaceous of the Gobi Desert. Hist. Biol. 32, 1394–1421 (2020).
Gilmore, C. W. Osteology of the Jurassic reptile Camptosaurus with a revision of the species of the genus and descriptions of two new species. Proc. US Nat. Mus. 36, 197–302 (1909).
Thomas, D. A. The cranial anatomy of Tenontosaurus tilletti Ostrom, 1970 (Dinosauria, Ornithopoda). Palaeontol. Electron. 18.2.37A, 1–99 (2015).
Porro, L. B., Witmer, L. M. & Barrett, P. M. Digital preparation and osteology of the skull of Lesothosaurus diagnosticus (Ornithischia: Dinosauria). PeerJ 3, e1494 (2015).
Horner, J. R. & Goodwin, M. B. Ontogeny of cranial epi-ossifications in Triceratops. J. Vertebr. Paleontol. 28, 134–144 (2008).
Loewen, M. A. et al. Lokiceratops rangiformis gen. et sp. nov. (Ceratopsidae: Centrosaurinae) from the Campanian Judith River Formation of Montana reveals rapid regional radiations and extreme endemism within centrosaurine dinosaurs. PeerJ 12, e17224 (2024).
Longrich, N. R. & Field, D. J. Torosaurus is not Triceratops: ontogeny in chasmosaurine ceratopsids as a case study in dinosaur taxonomy. PLoS ONE 7, e32623 (2012).
Tykoski, R. S., Fiorillo, A. R. & Chiba, K. New data and diagnosis for the Arctic ceratopsid dinosaur Pachyrhinosaurus perotorum. J. Syst. Palaeontol. 17, 1397–1416 (2019).
Zhao, X., Cheng, Z. & Xu, X. The earliest ceratopsian from the Tuchengzi Formation of Liaoning, China. J. Vertebr. Paleontol. 19, 681–691 (2009).
Sereno, P. C. in New Perspectives on Horned Dinosaurs (Ryan, M. J., Chinnery-Allgeier, B. J. & Eberth, D. A.) 21–58 (Indiana Univ. Press, 2010).
Maryańska, T. & Osmólska, H. Protoceratopsidae (Dinosauria) of Asia. Palaeontol. Pol. 33, 133–181 (1975).
You, H. & Dong, Z. A new protoceratopsid (Dinosauria: Neoceratopsia) from the Late Cretaceous of Inner Mongolia, China. Acta Geol. Sin. 77, 299–303 (2003).
Wolfe, D. G. et al. in New Perspectives on Horned Dinosaurs (Ryan, M. J., Chinnery-Allgeier, B. J. & Eberth, D. A.) 91–98 (Indiana Univ. Press, 2010).
Kirkland, J. I. & DeBlieux, D. D. in New Perspectives on Horned Dinosaurs (Ryan, M. J., Chinnery-Allgeier, B. J. & Eberth, D. A.) 117–140 (Indiana Univ. Press, 2010).
Han, F., Forster, C. A., Clark, J. M. & Xu, X. Cranial anatomy of Yinlong downsi (Ornithischia: Ceratopsia) from the Upper Jurassic Shishugou Formation of Xinjiang, China. J. Vertebr. Paleontol. 36, e1029579 (2016).
Morschhauser, E. M., Li, D., You, H. & Dodson, P. Cranial anatomy of the basal neoceratopsian Auroraceratops rugosus (Ornithischia: Ceratopsia) from the Yujingzi Basin, Gansu Province, China. J. Vertebr. Paleontol. 38, 36–68 (2019).
Hübner, T. & Rauhut, O. W. M. A juvenile skull of Dysalotosaurus lettowvorbecki (Ornithischia: Iguanodontia), and implications for cranial ontogeny, phylogeny, and taxonomy in ornithopod dinosaurs. Zoo. J. Linn. Soc. 160, 366–396 (2010).
Galton, P. M. The ornithischian dinosaur Hypsilophodon from the Wealden of the Isle of Wight. Bull. Brit. Mus. Nat. Hist. Geol. 25, 1–152 (1974).
Brown, B. Leptoceratops, a new genus of Ceratopsia from the Edmonton Cretaceous of Alberta. Bull. Am. Mus. Nat. Hist. 33, 567–580 (1914).
Makovicky, P. J. & Norell, M. A. Yamaceratops dorngobiensis, a new primitive ceratopsian (Dinosauria: Ornithischia) from the Cretaceous of Mongolia. Am. Mus. Novit. 3530, 1–42 (2006).
Tanoue, K., Grandstaff, B. S., You, H. L. & Dodson, P. Jaw mechanics in basal Ceratopsia. Anat. Rec. 292, 1352–1369 (2009).
Butler, R. J., Upchurch, P. & Norman, D. B. The phylogeny of the ornithischian dinosaurs. J. Syst. Palaeontol. 6, 1–40 (2008).
Dodson, P., Forster, C. A. & Sampson, S. A. in The Dinosauria (eds Weishampel, D. B., Dodson, P. & Osmólska. H.) 494–516 (Univ. California Press, 2004).
Dieudonné, P.-E., Cruzado-Caballero, P., Godefroit, P. & Tortosa, T. A new phylogeny of cerapodan dinosaurs. Hist. Biol. 33, 2335–2355 (2021).
Maidment, S. C. R. & Barrett, P. M. Osteological correlates for quadrupedality in ornithischian dinosaurs. Acta Palaeontol. Pol. 59, 53–70 (2014).
Chanthasit, P. The ornithopod dinosaur Rhabdodon from the Late Cretaceous of France: anatomy, systematics and paleobiology. PhD Thesis: Université Claude Bernard (2010).
Csiki, Z., Vremir, M., Brusatte, S. L. & Norell, M. A. An aberrant island-dwelling theropod dinosaur from the Late Cretaceous of Romania. Proc. Natl Acad. Sci. USA 107, 15357–15361 (2010).
Wolfe, D. G. & Kirkland, J. I. Zuniceratops christopheri n. gen & n. sp., a ceratopsian dinosaur from the Moreno Hill Formation (Cretaceous, Turonian) of west-central New Mexico. New Mex. Mus. Nat. Hist. Sci. Bull. 14, 303–317 (1998).
Sues, H. D. & Averianov, A. Turanoceratops tardabilis — the first ceratopsid dinosaur from Asia. Naturwissenschaften 96, 645–652 (2009).
Herbst, E. C., Meade, L. E., Lautenschlager, S., Fioritti, N. & Scheyer, T. M. A toolbox for the retrodeformation and muscle reconstruction of fossil specimens in Blender. Roy. Soc. Open Sci. 9, 220519 (2022).
Goloboff, P. A. & Catalano, S. A. TNT version 1.5, including a full implementation of phylogenetic morphometrics. Cladistics 32, 221–238 (2016).
Maddison, W. P. & Maddison, D. R. Mesquite: a modular system for evolutionary analysis. Version 3.04. Mesquite https//www.mesquiteproject.org (2025).
Ronquist, F. et al. MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Syst. Biol. 61, 539–542 (2012).
Lewis, P. O. A likelihood approach to estimating phylogeny from discrete morphological character data. Syst. Biol. 50, 913–925 (2001).
Nylander, J. A. A., Ronquist, F., Huelsenbeck, J. & Nieves-Aldrey, J. L. Bayesian phylogenetic analysis of combined data. Syst. Biol. 53, 47–67 (2004).
Kass, R. E. & Raftery, A. E. Bayes factor. J. Am. Stat. Assoc. 90, 773–795 (1995).
Maidment, S. C. R., Butler, R. J., Brusatte, S. L., Meade, L. E., Augustin, F. J.,Csiki-Sava, Z. & Ősi, A. Data for “A hidden diversity of ceratopsian dinosaurs in Late Cretaceous Europe”. Zenodo https://doi.org/10.5281/zenodo.17098636 (2025)
Słowiak, J., Tereshchenko, V. S. & Fostowicz-Frelik, Ł Appendicular skeleton of Protoceratops andrewsi (Dinosauria: Ornithischia): comparative morphology, ontogenetic changes, and implications for non-ceratopsid ceratopsian locomotion. PeerJ 7, e7324 (2019).

