Large neotheropods from the Upper Triassic of North America and the early evolution of large theropod body sizes

Large body sizes among nonavian theropod dinosaurs is a major feature in the evolution of this clade, with theropods reaching greater sizes than any other terrestrial carnivores. However, the early evolution of large body sizes among theropods is obscured by an incomplete fossil record, with the largest Triassic theropods represented by only a few individuals of uncertain ontogenetic stage. Here I describe two neotheropod specimens from the Upper Triassic Bull Canyon Formation of New Mexico and place them in a broader comparative context of early theropod anatomy. These specimens possess morphologies indicative of ontogenetic immaturity (e.g., absence of femoral bone scars, lack of co-ossification between the astragalus and calcaneum), and phylogenetic analyses recover these specimens as early-diverging neotheropods in a polytomy with other early neotheropods at the base of the clade. Ancestral state reconstruction for body size suggests that the ancestral theropod condition was small (∼240 mm femur length), but the ancestral neotheropod was larger (∼300–340 mm femur length), with coelophysoids experiencing secondary body size reduction, although this is highly dependent on the phylogenetic position of a few key taxa. Theropods evolved large body sizes before the Triassic–Jurassic extinction, as hypothesized in most other ancestral state reconstructions of theropod body sizes, but remained rare relative to smaller theropods until the Jurassic.