The Phylogeny of Basal Coelurosaurian Theropods (Archosauria: Dinosauria) and Patterns of Morphological Evolution during the Dinosaur-Bird Transition

Theropod dinosaurs are an iconic and familiar group of extinct species that include predators such as Tyrannosaurus and Velociraptor, as well as an array of other Mesozoic taxa. Carnivorous theropods are the evolutionary ancestors of birds, and the evolutionary transition between theropods and birds is a textbook example of a major evolutionary transformation in the history of life. Despite a flurry of research on early birds and their dinosaurian relatives, however, several questions still remain. First, the anatomy of some major theropod groups has yet to be described in detail. Second, there is little consensus on the phylogenetic relationships of the basal members of a theropod subgroup called Coelurosauria: the clade of birds and their closest relatives (defined as all taxa closer to birds than to Allosaurus). Third, there has been little synthetic work on large-scale macroevolutionary patterns during theropod evolution. This dissertation includes three chapters that touches on these three major issues. Chapter 1 is a detailed description of the Late Cretaceous tyrannosaurid theropod Alioramus altai, based on its holotype specimen from the Tsaagan Khuushu locality in the Maastrichtian Nemegt Formation of Mongolia. This monographic description provides further evidence that Alioramus is an unusual long-snouted, gracile, and slender-limbed taxon with an unpredecented degree of cranial ornamentation among tyrannosaurids and an extremely pneumatized skeleton. Anatomical comparisons indicate that the long skull of Alioramus is an autapomorphic feature that is proportionally longer (relative to femur length) than in any other known tyrannosaurid specimen, including juveniles, and that Alioramus is morphologically distinctive relative to similarly-sized individuals of the contemporary and sympatric Tarbosaurus. The coexistence of the long-snouted Alioramus and robust and deep-snouted Tarbosaurus, which are found together at the Tsaagan Khuushu locality, demonstrate that multiple large tyrannosaurids were able to live in sympatry, likely because of niche partitioning due to differences in craniofacial morphology and functional behavior. Chapter 2 presents a comprehensive new phylogenetic analysis of coelurosaurian theropods, which is an updated version (and thus the latest iteration) of the long-standing Theropod Working Group (TWiG) analysis. The new analysis incoroporates a wealth of new taxa and character data into the TWiG matrix for the first time, most of which is relevant to basal (non-maniraptoran) coelurosaurs such as tyrannosauroids and ornithomimosaurs, which had previously been the subject of only cursory character and taxon sampling in TWiG studies. The full dataset was analyzed under parsimony, and the resulting phylogeny includes several well supported relationships and agrees with previous analyses in many aspects. As a result, it is argued that a consensus view of basal coelurosaurian relationships has emerged, including: 1) the monophyly of major subclades such as Tyrannosauroidea, Compsognathidae, and Ornithomimosauria; 2) the position of the singleton genera Bicentenaria, Zuolong, and Tugulusaurus near the base of Coelurosauria; 3) the placement of Tyrannosauroidea as the most basal major coelurosaurian subclade; 4) the inclusion of Guanlong, Dilong, and Proceratosaurus within Tyrannosauroidea; 5) the existence of a derived maniraptoran clade that includes alvarezsauroids, therizinosauroids, oviraptorosaurs, and paravians to the exclusion of ornithomimosaurs and tyrannosauroids. Remaining areas of uncertainty include the phylogenetic position of Compsognathidae and the singleton genus Ornitholestes, and relationships at the base of the Ornithomimosauria + Maniraptora clade and Maniraptora itself. The phylogeny indicates that much of the early history of Coelurosauria has yet to be sampled in the fossil record, that coelurosaurs originated at small body size, and that the evolution of the iconic Tyrannosaurus-like bauplan occurred only towards the end of the Cretaceous. Chapter 3 presents a geometric morphometric analysis that is used to study broad patterns in theropod skull shape variation and compare the distribution of taxa in cranial morphospace (form) to both phylogeny and quantitative metrics of biting behaviour (function). The analysis finds that theropod skulls primarily differ in relative anteroposterior length and snout depth and to a lesser extent in orbit size and depth of the cheek region, and that oviraptorosaurs deviate most strongly from the "typical" and ancestral theropod morphologies. Noncarnivorous taxa generally fall out in distinct regions of morphospace and exhibit greater overall disparity than carnivorous taxa, whereas large-bodied carnivores independently converge on the same region of morphospace. The distribution of taxa in morphospace is strongly correlated with phylogeny but only weakly correlated with functional biting behaviour. These results imply that phylogeny, not biting function, was the major determinant of theropod skull shape.