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Phylogenetic relationships, chronostratigraphic, and paleoecological implications of M. intrepidus. a Graphic illustrating temporal range of North American tyrannosauroids including species-level range prior to the discovery of M. intrepidus, extension of current range, and hypothesized range based on isolated teeth12. The current gap in the North American tyrannosauroid record spans from the Tithonian to the Aptian. Faunal composition of Late Cretaceous ecosystems was established between the Albian and Turonian, as recognized by the stratigraphic appearance of major clades (see refs. 7,12 and references therein). b generalized phylogenetic relationships of Tyrannosauroidea, showing the appearance of select traits related to cursoriality in tyrannosaurs that are newly optimized as a result of the discovery of M. intrepidus. Tree topology follows this study using the modified dataset of Carr and colleagues27. Coelurus and Tanycolagreus are grafted as basal tyrannosauroids following Brusatte and colleagues5. c Stratigraphic distribution of Allosauria in North America (incl. Megaraptora but see ref. 70 for alternative hypotheses regarding this clade) documents overlap with M. intrepidus in early Late Cretaceous ecosystems leading to (d) refined calibration on the origin of late diverging tyrannosauroids and clade-level faunal turnover within apex predator roles throughout the Late Jurassic–Late Cretaceous of North America. Colored polygons are stylized call-outs and are not intended to reflect two-dimensional data. Temporal data corresponding to this figure are available in Supplementary Table 5
Drawings of Dryolestida left upper−molar, modified from the original descriptions. A–C, Kimmeridgian; D–F, Tithonian–Berriasian; G–H, early Barremian; I, late Barremian. A. Left M4 or M5 of Comotherium richi Prothero, 1981, from the Upper Jurassic Morrison Formation, Como Bluff locality, Wyoming, USA. B. Right M5, reversed, of Dryolestes leiriensis Martin, 1999, from Guimarota, Portugal.C. Left M6 of Krebsotherium lusitanicum Martin, 1999, from Guimarota, Portugal. D. Right M4 or M 6 (reversed) of Portopinheirodon asymmetricus Martin, 1999, Porto Pinheiro, Lourinha, Portugal. E. Right M5 (reversed) of Laolestes andresi Martin, 1999, from Porto Pinheiro, Lourinha, Portugal. F. Right upper molar of Donodon perscriptoris Sigogneau−Russell, 1991, Anoual, Morocco.G. Left M4 orM5 of Crusafontia amoae sp. nov., holotype, Cuesta Corrales 2, El Castellar Formation, Galve, Teruel, Spain. H. Right M6 orM7 (reversed) of Crusafontia amoae sp. nov., first described as upper molar of Crusafontia cuencana by Krebs (1993), P−2 H4 Pelejón 2, Galve, Teruel, Spain. I. Left M2 or M3, of Crusafontia cuencana Henkel and Krebs, 1969, Uña, La Huérguina Formation, Cuenca, Spain, (from Krebs 1993). Scale bars 1 mm.
Left ilium of Stokesosaurus clevelandi, Madsen 1974 (UMNH VP 7473), Morrison Formation, Utah, USA, Late Jurassic (early Tithonian).