A new pachycephalosaurid from the Hell Creek Formation, Garfield County, Montana, U.S.A.

LITERATURE CITED

• Alexander, R. M. (1997). “Engineering a dinosaur.” In J. O. Farlow & M. K. Brett-Surman (Eds.), Biology of the Dinosaurs (p. 414–425). Indiana University Press.
   Google Scholar
• Brown, B., & Schlaikjer, E. M. (1943). A study of the troödont dinosaurs with the description of a new genus and four new species. Bulletin of the American Museum of Natural History, 82(5), 121–149.
   Google Scholar
• Brown, C. M., Campione, N. E., Mantilla, G. P. W., & Evans, D. C. (2021). Size-driven preservational and macroecological biases in the latest Maastrichtian terrestrial vertebrate assemblages of North America. Paleobiology, 48(2), 210–238. https://doi.org/10.1017/pab.2021.35
   Web of Science ®Google Scholar
• Brown, C. M., Evans, D. C., Campione, N. E., O’Brien, L. J., & Eberth, D. A. (2013). Evidence for taphonomic size bias in the Dinosaur Park Formation (Campanian, Alberta), a model Mesozoic terrestrial alluvial-paralic system. Palaeogeography, Palaeoclimatology, Palaeontology, 372, 132–141. https://doi.org/10.1016/j.palaeo.2012.06.027
   Web of Science ®Google Scholar
• Brown, C. M., Ryan, M. J., Evans, D. C., Bininda-Emonds, O. R. P., Powell, G. L., Jamniczky, H. A, Bauer, A. M., & Theodor, J. (2015). “A census of Canadian dinosaurs: more than a century of discovery.” In O. R. P. Bininda-Emonds, G. L. Powell, H. A. Jamniczky, A. M. Bauer, & J. Theodor. BIS-Verlag der Carl von Ossietzky (Eds.), (p. 151–209). Universität Oldenburg.
   Google Scholar
• Carpenter, K. (1997). Agonistic behavior in pachycephalosaurs (Ornithischia, Dinosauria); a new look at head-butting behavior. Rocky Mountain Geology, 32(1), 19–25.
   Google Scholar
• Colbert, E. H. (1955). Evolution of the vertebrates. Wiley.
   Google Scholar
• de Camp, L. S. (1956). A gun for dinosaur. Galaxy Science Fiction, 11(5), 6–49.
   Google Scholar
• Dieudonné, P. E., Cruzado-Caballero, P., Godefroit, P., & Tortosa, T. (2020). A new phylogeny of Cerapodan dinosaurs. Historical Biology, 33(10), 2335–2355.
   Web of Science ®Google Scholar
• Dodson, P. (1975). Taxonomic implications of relative growth in lambeosaurine hadrosaurs. Systematic Zoology, 24(1), 37–54
   Google Scholar
• Dyer, A. D., LeBlanc, A. R. H., Doschak, M. R., & Currie, P. J. (2021). Taking a crack at the dome: histopathology of a pachycephalosaurid (Dinosauria: Ornithischia) frontoparietal dome. Biosis: Biological Systems, 2(2), 248–270.
   Google Scholar
• Evans, D. C. (2010). Cranial anatomy and systematics of Hypacrosaurus altispinus, and a comparative analysis of skull growth in lambeosaurine hadrosaurids (Dinosauria: Ornithischia). Zoological Journal of the Linnean Society, 159(2), 398–434.
   Web of Science ®Google Scholar
• Evans, D. C., Schott, R. K., Larson, D. W., Brown, C. M. and Ryan, M. J. (2013). The oldest North American pachycephalosaurid and the hidden diversity of small-bodied ornithischian dinosaurs. Nature communications, 4(1), 1–11.
   Google Scholar
• Evans, D. C., Hayashi, S., Chiba, K., Watabe, M., Ryan, M. J., Lee, Y-N., Currie, P. J., Tsogtbaatar, K., & Barsbold, R. (2018). Morphology and histology of new cranial specimens of Pachycephalosauridae (Dinosauria: Ornithischia) from the Nemegt Formation, Mongolia. Palaeogeography, Palaeoclimatology, Palaeoecology 494, 121–134.
   Web of Science ®Google Scholar
• Farke, A. A. (2008). Frontal sinuses and head-butting in goats: a finite element analysis. Journal of Experimental Biology, 211(19), 3085–3094.
   PubMed Web of Science ®Google Scholar
• Farke, A. A. (2014). Evaluating combat in ornithischian dinosaurs. Journal of Zoology, 292(4), 242–249.
   Web of Science ®Google Scholar
• Galton, P. M. (1970). Pachycephalosaurids: dinosaurian battering rams. Discovery, 6(1), 23–32.
   Google Scholar
• Galton, P. M. (1971). A primitive dome-headed dinosaur (Ornithischia: Pachycephalosauridae) from the lower Cretaceous of England and the function of the dome of pachycephalosaurids. Journal of Paleontology, 45(1), 40–47.
   Google Scholar
• Galton, P. M., & Sues, H.-D. (1983). New data on pachycephalosaurid dinosaurs (Reptilia: Ornithischia) from North America. Canadian Journal of Earth Sciences, 20(3), 462–472.
   Web of Science ®Google Scholar
• Giffin, E. B. (1989). Notes on pachycephalosaurs (Ornithischia). Journal of Paleontology, 63(4), 525–529.
   Web of Science ®Google Scholar
• Gilmore, C. W. (1924). On Troödon validus: An ornithopodous dinosaur from the Belly River Cretaceous of Alberta, Canada. University of Alberta Press.
   Google Scholar
• Goodwin, M. B., & Horner, J. R. (2004). Cranial histology of pachycephalosaurs (Ornithischia: Marginocephalia) reveals transitory structures inconsistent with head-butting behavior. Paleobiology, 30(2), 253–267.
   Web of Science ®Google Scholar
• Goodwin, M. B., & Evans, D. C. (2016). 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. Journal of Vertebrate Paleontology, 36(2), e1078343.
   Web of Science ®Google Scholar
• Horner, J. R., & Marshall, C. (2002). Keratin covered dinosaur skulls [Abstract]. Journal of Vertebrate Paleontology, 22(3), 31A.
   Google Scholar
• Horner, J. R., & Goodwin, M. B. (2009). Extreme cranial ontogeny in the Upper Cretaceous dinosaur Pachycephalosaurus. PLoS One, 4(10), e7626.
   PubMed Web of Science ®Google Scholar
• Horner, J. R., Goodwin, M. B., & Myhrvold, N. (2011). Dinosaur census reveals abundant Tyrannosaurus and rare ontogenetic stages in the Upper Cretaceous Hell Creek Formation (Maastrichtian), Montana, USA. PLoS One, 6(2), e16574.
   PubMed Web of Science ®Google Scholar
• Lee, A. H., & Werning, S. (2008). Sexual maturity in growing dinosaurs does not fit reptilian growth models. Proceedings of the National Academy of Sciences, 105(2), 582–587.
   PubMed Web of Science ®Google Scholar
• Mallon, J. C., Evans, D. C., Tokaryk, T. T., & Currie, M. L. (2015). First pachycephalosaurid (Dinosauria: Ornithischia) from the Frenchman Formation (upper Maastrichtian) of Saskatchewan, Canada. Cretaceous Research, 56, 426–431.
   Web of Science ®Google Scholar
• Maryańska, T., & Osmólska, H. (1974). Pachycephalosauria, a new suborder of ornithischian dinosaurs. Palaeontologia Polonica, 30, 45–102.
   Google Scholar
• Moore, B. R. S., Roloson, M. J., Currie, P. J., Ryan, M. R., Patterson, R. T., & Mallon, J. C. (2022). The appendicular myology of Stegoceras validum (Ornithischia: Pachycephalosauridae) and implications for the head-butting hypothesis. PLoS One, 17(9), e0268144.
   PubMed Web of Science ®Google Scholar
• Nirody, J. A., Goodwin, M. B., Horner, J. R., Huynh, T. L., Colbert, M. W., Smith, D. K., & Evans, D. C. (2022). Quantifying vascularity in the frontoparietal dome of Stegoceras validum (Dinosauria: Pachycephalosauridae) from high resolution CT scans. Journal of Vertebrate Paleontology, 41(5), e2036991.
   Web of Science ®Google Scholar
• Owen, R. (1842). Report on British fossil reptiles. part II. Report for the British Association for the Advancement of Science, Plymouth, 1841, 60–294.
   Google Scholar
• Peterson, J. E., & Vittore, C. P. (2012). Cranial pathologies in a specimen of Pachycephalosaurus. PLoS One, 7(4), e36227.
   PubMed Web of Science ®Google Scholar
• Peterson, J. E., Dischler, C., & Longrich, N. R. (2013). Distributions of cranial pathologies provide evidence for head-butting in dome-headed dinosaurs (Pachycephalosauridae). PLoS One 8(7), e68620.
   PubMed Web of Science ®Google Scholar
• Porter, W. M. R., & Witmer., L. M. (2016). Avian cephalic vasculature anatomy, sites of thermal exchange, and the rete ophthalmicum. The Anatomical Record, 299(11), 1461–1486.
   Google Scholar
• Scannella J. B., & Horner, J. R. (2010). Torosaurus Marsh, 1891, is Triceratops Marsh, 1889 (Ceratopsidae: Chasmosaurinae): synonymy through ontogeny. Journal of Vertebrate Paleontology, 30(4), 1157–1168.
   Web of Science ®Google Scholar
• Schott, R. K., & Evans, D. C. (2012). Squamosal ontogeny and variation in the pachycephalosaurian dinosaur Stegoceras validum Lambe, 1902, from the Dinosaur Park Formation, Alberta. Journal of Vertebrate Paleontology, 32(4), 903–913.
   Web of Science ®Google Scholar
• Schott, R. K., & Evans, D. C. (2017). 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. Zoological Journal of the Linnean Society, 179(4), 865–906.
   Web of Science ®Google Scholar
• Schott, R. K., Evans, D. C., Williamson, T. E., Carr, T. D., & Goodwin, M. B. (2009). The anatomy and systematics of Colepiocephale lambei (Dinosauria: Pachycephalosauridae). Journal of Vertebrate Paleontology, 29(3), 771–786.
   Web of Science ®Google Scholar
• Schott, R. K, Evans, D. C., Goodwin, M. B., Horner, J. R., Brown, C. M., & Longrich, N. R. (2011). Cranial ontogeny in Stegoceras validum (Dinosauria: Pachycephalosauria): A quantitative model of pachycephalosaur dome growth and variation. PLoS One, 6(6), e21092.
   PubMed Web of Science ®Google Scholar
• Seeley, H. G. (1888). I. On the classification of the fossil animals commonly named Dinosauria. Proceedings of the Royal Society of London, 43(258–265), 165–171.
   Google Scholar
• Sereno, P. C. (1997). The origin and evolution of dinosaurs. Annual Review of Earth and Planetary Sciences, 25(1), 435–489.
   Web of Science ®Google Scholar
• Snively, E., & Cox, A. (2008). Structural mechanics of pachycephalosaur crania permitted head-butting behavior. Palaeontologica electronica, 11(1), 3A, 1–17.
   Web of Science ®Google Scholar
• Snively, E., & Theodor, J. M. (2011). Common functional correlates of head-strike behavior in the pachycephalosaur Stegoceras validum (Ornithischia, Dinosauria) and combative artiodactyls. PLoS One 6(6), e21422.
   PubMed Web of Science ®Google Scholar
• Sternberg, C. M. (1945). Pachycephalosauridae proposed for dome-headed dinosaurs, Stegoceras lambei, n. sp., described. Journal of Paleontology, 19, 534–538.
   Google Scholar
• Sues, H.-D. (1978). Functional morphology of the dome in pachycephalosaurid dinosaurs. Neues Jahrbuch für Geologie und Paläontologie, Monatshefte, 8, 459–472.
   Google Scholar
• Swofford, D. L. (1998). PAUP*. Phylogenetic Analysis Using Parsimony (*and Other Methods). Version 4. Sinauer Associates.
   Google Scholar
• Weaver, V., Goodwin, M. B., & Horner, J. R. (2014). Histology of pachycephalosaurid crania reveals evidence of multiple tissue types and variable origin of apical pathologies [Abstract]. Journal of Vertebrate Paleontology, 27A.
   Google Scholar
• Williamson, T. E., & Brusatte, S. L., (2016). Pachycephalosaurs (Dinosauria: Ornithischia) from the Upper Cretaceous (upper Campanian) of New Mexico: a reassessment of Stegoceras novomexicanum. Cretaceous Research, 62, 29–43.
   Web of Science ®Google Scholar
• Williamson, T. E., & Carr, T. D., (2002). A new genus of derived pachycephalosaurian from western North America. Journal of Vertebrate Paleontology, 22(4), 779–801.
   Web of Science ®Google Scholar
• Woodruff, D. C. (2022). Osteology and kinematics of the pachycephalosaurid vertebral column provides further evidence refuting the agonistic head-butting hypothesis [Abstract]. Journal of Vertebrate Paleontology, 370.
   Google Scholar
• Woodruff, D. C., Goodwin, M. B., Lyson, T. R., & Evans, D. C. (2021). Ontogeny and variation of the pachycephalosaurine dinosaur Sphaerotholus buchholtzae, and its systematics within the genus. Zoological Journal of the Linnean Society, 193(2), 563–601.
   Web of Science ®Google Scholar
• Zonneveld, J.-P., AbdelGawad, M. K., & Miller, E. R. (2022). Ectoparasite borings, mesoparasite borings, and scavenging traces in early Miocene turtle and tortoise shell: Moghra Formation, Wadi Moghra, Egypt. Journal of Paleontology, 96(2), 304–322.
   Web of Science ®Google Scholar
• Zonneveld, J.-P., Bartels, W. S., Gunnell, G. F., & McHugh, L. P. (2015). Borings in early Eocene turtle shell from the Wasatch Formation, South Pass, Wyoming. Journal of Paleontology, 89(5), 802–820.
   Web of Science ®Google Scholar