Sues, H.-D. Functional morphology of the dome in pachycephalosaurid dinosaurs. N. Jb. Geol. Palaontol. Mh. 8, 459–472 (1978).
Snively, E. & Theodor, J. M. Common functional correlates of head-strike behavior in the pachycephalosaur Stegoceras validum (Ornithischia, Dinosauria) and combative artiodactyls. PLoS ONE 6, e21422 (2011).
Peterson, J. E., Dischler, C. & Longrich, N. R. Distributions of cranial pathologies provide evidence for head-butting in dome-headed dinosaurs (Pachycephalosauridae). PLoS ONE 8, e68620 (2013).
Evans, D. C., Schott, R. K., Larson, D. W., Brown, C. M. & Ryan, M. J. The oldest North American pachycephalosaurid and the hidden diversity of small-bodied ornithischian dinosaurs. Nat. Commun. 4, 1828 (2013).
Dyer, A. D., Powers, M. J. & Currie, P. J. Problematic putative pachycephalosaurids: synchrotron µCT imaging shines new light on the anatomy and taxonomic validity of Gravitholus albertae from the Belly River Group (Campanian) of Alberta, Canada. Vertebr. Anat. Morphol. Palaeontol. 10, 65–110 (2023).
Horner, J. R., Goodwin, M. B. & Evans, D. C. A new pachycephalosaurid from the Hell Creek Formation, Garfield County, Montana, U.S.A. J. Vertebr. Paleontol. 42, e2190369 (2023).
Woodruff, D. C., Schott, R. K. & Evans, D. C. Two new species of small-bodied pachycephalosaurine (Dinosauria, Marginocephalia) from the uppermost Cretaceous of North America suggest hidden diversity in well-sampled formations. Pap. Palaeontol. 9, e1535 (2023).
Knapp, A., Knell, R. J. & Hone, D. W. E. Three-dimensional geometric morphometric analysis of the skull of Protoceratops andrewsi supports a socio-sexual signaling role for the ceratopsian frill. Proc. R. Soc. B 288, 20202938 (2021).
Main, R. P., De Ricqlès, A., Horner, J. R. & Padian, K. The evolution and function of thyreophoran dinosaur scutes: implications for plate function in stegosaurs. Paleobiology 31, 291–314 (2005).
Arbour, V. M., Zanno, L. E. & Evans, D. C. Palaeopathological evidence for intraspecific combat in ankylosaurid dinosaurs. Biol. Lett. 18, 20220404 (2022).
Brown, C. M., Campione, N. E., Mantilla, G. P. W. & Evans, D. C. Size-driven preservational and macroecological biases in the latest Maastrichtian terrestrial vertebrate assemblages of North America. Paleobiology 48, 210–238 (2022).
Hayashi, S., Carpenter, K., Watabe, M. & McWhinney, L. A. Ontogenetic histology of Stegosaurus plates and spikes. Palaeontology 55, 145–161 (2012).
Butler, R. J. & Zhao, Q. The small-bodied ornithischian dinosaurs Micropachycephalosaurus hongtuyanensis and Wannanosaurus yansiensis from the Late Cretaceous of China. Cretaceous Res. 30, 63–77 (2009).
Schott, R. K. et al. Cranial ontogeny in Stegoceras validum (Dinosauria: Pachycephalosauria): a quantitative model of pachycephalosaur dome growth and variation. PLoS ONE 6, e21092 (2011).
Campione, N. E., Evans, D. C., Brown, C. M., Carrano, M. T. & Revell, L. Body mass estimation in non-avian bipeds using a theoretical conversion to quadruped stylopodial proportions. Methods Ecol. Evol. 5, 913–923 (2014).
Ito, M., Matsukawa, M., Saito, T. & Nichols, D. J. Facies architecture and paleohydrology of a synrift succession in the Early Cretaceous Choyr Basin, southeastern Mongolia. Cretaceous Res. 27, 226–240 (2006).
Jerzykiewicz, T. & Russell, D. A. Late Mesozoic stratigraphy, and vertebrates of the Gobi Basin. Cretaceous Res. 12, 345–377 (1991).
Nichols, D. J., Matsukawa, M. & Ito, M. Palynology and age of some Cretaceous nonmarine deposits in Mongolia and China. Cretaceous Res. 27, 241–251 (2006).
Maryańska, T. & Osmólska, H. Pachycephalosauria, a new suborder of ornithischian dinosaurs. Acta Palaeontol. Pol. 30, 45–102 (1974).
Perle, A., Maryańska, T. & Osmólska, H. Goyocephale lattimorei gen. et sp. n., a new flat-headed pachycephalosaur (Ornithischia, Dinosauria) from the Upper Cretaceous of Mongolia. Acta Palaeontol. Pol. 27, 115–132 (1982).
Gilmore, C. W. On Troodon validus, an orthopodous dinosaur from the Belly River Cretaceous of Alberta, Canada. Alberta Univ. Bull. 1, 1–43 (1924).
Woodruff, D. C., Goodwin, M. B., Lyson, T. R. & Evans, D. C. Ontogeny and variation of the pachycephalosaurine dinosaur Sphaerotholus buchholtzae, and its systematics within the genus. Zool. J. Linn. Soc. 193, 563–601 (2021).
Brown, B. & Schlaikjer, E. M. A study of the troodont dinosaurs with the description of a new genus and four new species. Bull. Am. Mus. Nat. Hist. 82, 115–150 (1943).
Sullivan, R. M. A taxonomic review of the Pachycephalosauridae (Dinosauria: Ornithischia). Bull. New Mexico Mus. Nat. Hist. Sci. 35, 347–365 (2006).
Williamson, T. E. & Carr, T. D. A new genus of derived pachycephalosaurian from western North America. J. Vertebr. Paleontol. 22, 779–801 (2002).
Schott, R. K. & Evans, D. C. Squamosal ontogeny and variation in the pachycephalosaurian dinosaur Stegoceras validum Lambe, 1902, from the Dinosaur Park Formation, Alberta. J. Vertebr. Paleontol. 32, 903–913 (2012).
Schott, R. K. & Evans, D. C. Cranial variation and systematics of Foraminacephale brevis gen. nov. and the diversity of pachycephalosaurid dinosaurs (Ornithischia: Cerapoda) in the Belly River Group of Alberta, Canada. Zool. J. Linn. Soc. 179, 865–906 (2016).
Sullivan, R. M. Revision of the dinosaur Stegoceras Lambe (Ornithischia, Pachycephalosauridae). J. Vertebr. Paleontol. 23, 181–207 (2003).
Han, F.-L., 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 (2015).
Evans, D. C., Brown, C. M., You, H. & Campione, N. E. Description and revised diagnosis of Asia’s first recorded pachycephalosaurid, Sinocephale bexelli gen. nov., from the Upper Cretaceous of Inner Mongolia, China. Can. J. Earth Sci. 58, 981–992 (2021).
Goodwin, M. B. & Horner, J. R. Cranial histology of pachycephalosaurs (Ornithischia: Marginocephalia) reveals transitory structures inconsistent with head-butting behavior. Paleobiology 30, 253–267 (2004).
Evans, D. C. et al. Morphology and histology of new cranial specimens of Pachycephalosauridae (Dinosauria: Ornithischia) from the Nemegt Formation, Mongolia. Palaeogeogr. Palaeoclimatol. Palaeoecol. 494, 121–134 (2018).
Hou, L. A new primitive Pachycephalosauria from Anhui, China. Vertebrata PalAsiatica 15, 198–202 (1977).
Owen, R. Monograph on the Fossil Reptilia of the Wealden and Purbeck Formations. Part V. Lacertilia. Palaeontogr. Soc. Monogr. 12, 31–39 (1861).
Thulborn, R. A. A new heterodontosaurid dinosaur (Reptilia: Ornithischia) from the Upper Triassic Red Beds of Lesotho. Zool. J. Linn. Soc. 55, 151–175 (1974).
Sues, H.-D. & Galton, P. M. Anatomy and classification of the North American Pachycephalosauria (Dinosauria: Ornithischia). Palaeontographica 198, 1–40 (1987).
Brown, C. M. & Russell, A. P. Homology and architecture of the caudal basket of Pachycephalosauria (Dinosauria: Ornithischia): the first occurrence of myorhabdoi in Tetrapoda. PLoS ONE 7, e30212 (2012).
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 (2006).
Wings, O. & Sander, P. M. No gastric mill in sauropod dinosaurs: new evidence from analysis of gastrolith mass and function in ostriches. Proc. R. Soc. B: Biol. Sci. 274, 635–640 (2007).
Takasaki, R. & Kobayashi, Y. Effects of diet and gizzard muscularity on grit use in domestic chickens. PeerJ 8, e10277 (2020).
Takasaki, R. Herbivorous Adaptations of Dinosauria: Hadrosaurid Foraging Strategy and Archosaur Gastroliths. PhD thesis, Hokkaido Univ. (2019).
Horner, J. R. & Goodwin, M. B. Extreme cranial ontogeny in the Upper Cretaceous dinosaur Pachycephalosaurus. PLoS ONE 4, e7626 (2009).
Goodwin, M. B., Buchholtz, E. A. & Johnson, R. E. Cranial anatomy and diagnosis of Stygimoloch spinifer (Ornithischia: Pachycephalosauria) with comments on cranial display structures in agonistic behavior. J. Vertebr. Paleontol. 18, 363–375 (1998).
De Margerie, E., Cubo, J. & Castanet, J. Bone typology and growth rate: testing and quantifying ‘Amprino’s rule’ in the mallard (Anas platyrhynchos). C.R. Biol. 325, 221–230 (2002).
Goodwin, M. B. & Evans, D. C. The early expression of squamosal horns and parietal ornamentation confirmed by new end-stage juvenile Pachycephalosaurus fossils from the Upper Cretaceous Hell Creek Formation, Montana. J. Vertebr. Paleontol. 36, 1–8 (2016).
Prondvai, E. et al. Radial porosity profiles: a new bone histological method for comparative developmental analysis of diametric limb bone growth. R. Soc. Open Sci. 9, 211893 (2022).
Griffin, C. T. et al. Assessing ontogenetic maturity in extinct saurian reptiles. Biol. Rev. 96, 470–525 (2021).
Knell, R. J., Naish, D., Tomkins, J. L. & Hone, D. W. Sexual selection in prehistoric animals: detection and implications. Trends Ecol. Evol. 28, 38–47 (2013).
Erickson, G. M., Curry, R. K., Varricchio, D. J., Norell, M. A. & Xu, X. Growth patterns in brooding dinosaurs reveals the timing of sexual maturity in non-avian dinosaurs and genesis of the avian condition. Biol. Lett. 3, 558–561 (2007).
Andrew, H., Lee, A. H. & Werning, S. Sexual maturity in growing dinosaurs does not fit reptilian growth models. Proc. Natl Acad. Sci. USA 105, 582–587 (2008).
Prondvai, E., Godefroit, P., Adriaens, D. & Hu, D. Intraskeletal histovariability, allometric growth patterns, and their functional implications in bird-like dinosaurs. Sci. Rep. 8, 258 (2018).
Cullen, T. M., Simon, D. J., Benner, E. K. & Evans, D. C. Morphology and osteohistology of a large-bodied caenagnathid (Theropoda, Oviraptorosauria) from the Hell Creek Formation (Montana): implications for size-based classifications and growth reconstruction in theropods. Pap. Palaeontol. 7, 751–767 (2021).
Monfroy, Q. T. & Kundrát, M. The osteohistological variability in the evolution of basal avialans. Acta Zool. 103, 1–28 (2022).
Wick, S. L. & Lehman, T. M. A rare ‘flat-headed’ pachycephalosaur (Dinosauria: Pachycephalosauridae) from West Texas, USA, with morphometric and heterochronic considerations. Geobios 86, 89–106 (2024).
Nirody, J. A. et al. Quantifying vascularity in the frontoparietal dome of Stegoceras validum (Dinosauria: Pachycephalosauridae) from high-resolution CT scans. J. Vertebr. Paleontol. 41, e2036991 (2021).
Fonseca, A. O. et al. A comprehensive phylogenetic analysis on early ornithischian evolution. J. Syst. Palaeontol. 22, 2346577 (2024).
Sereno, P. C. in The Age of Dinosaurs in Russia and Mongolia (eds Benton, M. J. et al.) 480–516 (Cambridge Univ. Press, 2000).
Huang, L.-K. & Wang, M.-J. J. Image thresholding by minimizing the measures of fuzziness. Pattern Recognit. 28, 41–51 (1995).
Madzia, et al. The phylogenetic nomenclature of ornithischian dinosaurs. PeerJ 9, e12362 (2021).
Lamm, E. T. in Bone Histology of Fossil Tetrapods (eds Padian, K. & Lamm, E. T.) 55–160 (California Univ. Press, 2013)
Xu, X., Forster, C. A., Clark, J. M. & Mo, J. A basal ceratopsian with transitional features from the Late Jurassic of northwestern China. Proc. Biol. Sci. 273, 2135–2140 (2006).
Osi, A., Prondvai, E., Butler, R. & 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).
Boyd, C. A. The systematic relationships and biogeographic history of ornithischian dinosaurs. PeerJ 3, e1523 (2015).
Dieudonné, P. E., Cruzado-Caballero, P., Godefroit, P. & Tortosa, T. A new phylogeny of cerapodan dinosaurs. Hist. Biol. 33, 2335–2355 (2020).
Ohashi, T. & Barrett, P. M. A new ornithischian dinosaur from the Lower Cretaceous Kuwajima Formation of Japan. J. Vertebr. Paleontol. 29, 748–757 (2009).
Butler, R. J., Upchurch, P. & Norman, D. B. The phylogeny of the ornithischian dinosaurs. J. Syst. Palaeontol. 6, 1–40 (2008).
Longrich, N. R., Sankey, J. & Tanke, D. Texacephale langstoni, a new genus of pachycephalosaurid (Dinosauria: Ornithischia) from the upper Campanian Aguja Formation, southern Texas, USA. Cretaceous Res. 31, 274–284 (2010).
Hill, R. V., Witmer, L. M. & Norell, M. A. A new specimen of Pinacosaurus grangeri (Dinosauria: Ornithischia) from the Late Cretaceous of Mongolia: ontogeny and phylogeny of ankylosaurs. Am. Mus. Novit. 3395, 1–29 (2003).
Han, F., Forster, C. A., Clark, J. M. & Xu, X. A new taxon of basal ceratopsian from China and the early evolution of Ceratopsia. PLoS ONE 10, e0148689 (2015).
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).
Goloboff, P. A. & Catalano, S. A. TNT version 1.5, including a full implementation of phylogenetic morphometrics. Cladistics 32, 221–238 (2016).
Ronquist, F. & Huelsenbeck, J. P. MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics 19, 1572–1574 (2003).
Goloboff, P. A. Extended implied weighting. Cladistics 30, 260–272 (2014).
Diego, P. & Goloboff, P. A. The impact of unstable taxa in coelurosaurian phylogeny and resampling support measures for parsimony analyses. Bull. Am. Mus. Nat. Hist. 440, 97–115 (2020).
Brusatte, S. L., Benton, M. J., Ruta, M. & Lloyd, G. T. Superiority, competition, and opportunism in the evolutionary radiation of dinosaurs. Science 321, 1485–1488 (2008).
Bapst, D. W. paleotree: an R package for paleontological and phylogenetic analyses of evolution. Methods Ecol. Evol. 3, 803–807 (2012).
Bell, M. A., Lloyd, G. T. & Smith, A. strap: an R package for plotting phylogenies against stratigraphy and assessing their stratigraphic congruence. Palaeontology 58, 379–389 (2015).
Zhang, C., Stadler, T., Klopfstein, S., Heath, T. A. & Ronquist, F. Total-evidence dating under the fossilized birth–death process. Syst. Biol. 65, 228–249 (2016).
Kikins, R., Pieper, S. & Vosburgh, K. G. in Intraoperative Imaging and Image-Guided Therapy (ed. Jolesz, F.) 277–289 (Springer, 2014).
Schneider, C. A., Rasband, W. S. & Eliceiri, K. W. NIH Image to ImageJ: 25 years of image analysis. Nat. Methods 9, 671–675 (2012).
Williamson, T. E. & Brusatte, S. L. Pachycephalosaurs (Dinosauria: Ornithischia) from the Upper Cretaceous (upper Campanian) of New Mexico: a reassessment of St. validum novomexicanum. Cretaceous Res. 62, 29–43 (2016).
Julie, J. & Francois, H. missMDA: a package for handling missing values in multivariate data analysis. J. Stat. Softw. 70, 1–31 (2016).
Warton, D. I., Duursma, R. A., Falster, D. S. & Taskinen, S. smatr 3—an R package for estimation and inference about allometric lines. Methods Ecol. Evol. 3, 257–259 (2012).
Elliott, N. G., Haskard, K. & Koslow, J. A. Morphometric analysis of orange roughly (Hoplostethus atlanticus) off the continental slope of southern Australia. J. Fish Biol. 46, 202–220 (1995).
Mannion, P. D. & Upchurch, P. Completeness metrics and the quality of the sauropodomorph fossil record through geological and historical time. Paleobiology 36, 283–302 (2010).
Chinzorig, T. et al. Figshare https://doi.org/10.6084/m9.figshare.28509500 (2025).
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