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Numerous paleontologists have noted wear facets on tyrannosaurid lateral teeth over the past century. While several workers have proposed explanations for these features, there remains to this day no consensus concerning their etiology. Here we report on an examination of wear surfaces on these teeth from the Upper Cretaceous (mid−Campanian) Judith River Group of southern Alberta, Canada. This study reveals two distinct types of wear features on the labial and lingual sides of tyrannosaurid lateral teeth: irregular “spalled” surfaces and wear facets. The irregular spalled surfaces typically extend to the apex of the tooth, which evidently reflects flaking of enamel resulting from forces produced during contact between tooth and food. These surfaces are often rounded, presumably from antemortem wear following spalling. Wear striations on these surfaces are oriented heterogeneously. The wear facets, in contrast, occur on only one side of the tooth and are typically elliptical in outline and evince parallel wear striations. Similar patterns of parallel wear striations in extant mammals reflect tooth−tooth contact. We therefore propose that wear facets in tyrannosaurids were formed by repeated tooth−tooth contact between the lingual side of maxillary teeth and labial side of dentary teeth. It remains unclear whether this contact was serendipitous or adaptive, though it appears to be unusual for reptiles, as we have found no evidence for wear facets in extant varanids and crocodilians.
Tyrannosaurid theropods display several unusual adaptations of the skulls and teeth. Their nasals are fused and vaulted, suggesting that these elements braced the cranium against high feeding forces. Exceptionally high strengths of maxillary teeth in Tyrannosaurus rex indicate that it could exert relatively greater feeding forces than other tyrannosaurids. Areas and second moments of area of the nasals, calculated from CT cross−sections, show higher nasal strengths for large tyrannosaurids than for Allosaurus fragilis. Cross−sectional geometry of theropod crania reveals high second moments of area in tyrannosaurids, with resulting high strengths in bending and torsion, when compared with the crania of similarly sized theropods. In tyrannosaurids trends of strength increase are positively allomeric and have similar allometric exponents, indicating correlated progression towards unusually high strengths of the feeding apparatus. Fused, arched nasals and broad crania of tyrannosaurids are consistent with deep bites that impacted bone and powerful lateral movements of the head for dismembering prey.
Recent discoveries of more than ten new species of tyrannosauroid theropods are helping to understand the origin and evolu− tion of colossal body size and other characteristic features of Tyrannosaurus rex and its terminal Cretaceous relatives. Partic− ularly important has been the discovery and reinterpretation of Late Jurassic tyrannosauroids from Europe and North Amer− ica, which are intermediate in size and phylogenetic position between small basal tyrannosauroids and the largest Late Cre− taceous species. The fragmentary nature of these Jurassic specimens, however, has frustrated attempts to understand their systematics and phylogeny. A new specimen from the Late Jurassic of England was recently named as a new species (Stokesosaurus langhami) of the genus Stokesosaurus, which is known from several fragmentary fossils from North Amer− ica. We review the systematics and phylogeny of these European and North American specimens and show that there are no unequivocal synapomorphies uniting them. Furthermore, a revised phylogenetic analysis does not recover them as sister taxa. This necessitates a taxonomic revision of this material, and we name a new genus (Juratyrant) for theBritish specimen.
Premaxillary tooth count tends to be stable amongst toothed dinosaurs, and most theropods have four teeth in each premaxilla. Only one case of bilaterally asymmetric variation is known in theropod premaxillary dentition, and there is no record of ontogenetic or individual variation in premaxillary tooth count. Based on these observations, a tyrannosaurid left premaxilla with three teeth (TMP 2007.20.124) is an interesting deviation and represents an unusual individual of Daspletosaurus sp. with a developmental abnormality. The lower number of teeth is coupled with relatively larger alveoli, each of which is capable of hosting a larger than normal tooth. This indicates that tooth size and dental count vary inversely, and instances of reduction in tooth count may arise from selection for increased tooth size. On the other hand, the conservative number of premaxillary teeth in most theropods implies strong developmental constraints and a functional trade−off between the dimensions of the premaxillary alveolar margin and the size of the teeth. In light of recent advances in the study of tooth morphogenesis, tooth count is a function of two parameters: dimensions of an odontogenic field for a tooth series, and dimensions of tooth positions. A probable developmental cause for the low tooth count of TMP 2007.20.124 is that the dimensions of the alveoli expanded by approximately a third during tooth morphogenesis. Numerical traits such as tooth count are difficult to treat in a phylogenetic analysis. When formulating a phylogenetic character, a potential alternative to simply counting is to rely on the morphological signature for developmental parameters that control the number of the element in question.
The costovertebral articulation is integral to constrain the thoracic kinematics and to infer the breathing mechanism in the respect with costal aspiration. However, the structure of the costovertebral articulation in non−avian theropods has not been studied in great detail before. This study highlights the Tyrannosauridae, which is represented by numerous complete specimens. Costovertebral articulations of ten tyrannosaurid specimens, including two nearly in−situ articulated fossils, were investigated and compared with those in extant Archosauria. For extant archosaurs, dissections were conducted to rationalize the soft tissue anatomy in tyrannosaurids. This study shows that the rib articulates ventrally or posteroventrally with the distal end of the corresponding vertebral transverse process in the tyrannosaurid ribcage. A ligament (ligamentum costotransversarium) can be reconstructed to connect the rib tuberculum to the transverse process in each articulation. The scar for lig. costotransversarium is recognizable in many theropod skeletons, and this rugosity can be used to identify the rotational axis for the rib. This result provides a cornerstone for exploring the evolution of the ribcage and breathing mechanisms across the theropod lineage leading to birds.
Beautifully preserved, nearly complete theropod skeletons from Alberta (Canada) allow re−evaluation of the taxonomic status of North American tyrannosaurids. It is concluded that the most parsimonious interpretation of relationships leads to the separation of the two species of Albertosaurus(sensu Russell 1970) into Gorgosaurus libratusfrom the Campanian Dinosaur Park Formation and Albertosaurus sarcophagus from the upper Campanian/lower Maastrichtian Horseshoe Canyon Formation. Albertosaurus and Gorgosaurus are closely related, but can be distinguished from each other by more characters than are known to justify generic distinction within another tyrannosaurid clade that includes Daspletosaurus, Tarbosaurus and Tyrannosaurus. Daspletosaurus is known from multiple species that cover extensive geographic, ecological and temporal ranges, and it is sensible to maintain its generic distinction from Tyrannosaurus. All tyrannosaurid species have consistent ontogenetic trends. However, one needs to be cautious in assessing ontogenetic stage because many characters are size−dependent rather than age−dependent. There are relatively few osteological differences that can distinguish tyrannosaurid species at any age. For example, Nanotyrannus lancensis is probably a distinct species from Tyrannosaurus rex because there is no evidence of ontogenetic reduction of tooth counts in any other tyrannosaurid species. Some characters that are good for separating mature tyrannosaurids, such as differences in the sizes and shapes of maxillary fenestrae, are not useful for identifying the species of juveniles.
The skull of a newly prepared Tarbosaurus bataar is described bone by bone and compared with a disarticulated skull of Tyrannosaurus rex. Both Tarbosaurus bataar and Tyrannosaurus rex skulls are deep in lateral view. In dorsal view, the skull of T. rex is extremely broad posteriorly but narrows towards the snout; in Ta. bataarthe skull is narrower (especially in its ventral part: the premaxilla, maxilla, jugal, and the quadrate complex), and the expansion of the posterior half of the skull is less abrupt. The slender snout of Ta. bataaris reminiscent of more primitive North American tyrannosaurids. The most obvious difference between T. rex and Ta. bataar is the doming of the nasal in Ta. bataar which is high between the lacrimals and is less attached to the other bones of the skull, than in most tyrannosaurids. This is because of a shift in the handling of the crushing bite in Ta. bataar. We propose a paleogeographically based division of the Tyrannosaurinae into the Asiatic forms (Tarbosaurus and possibly Alioramus) and North American forms (Daspletosaurus and Tyrannosaurus). The division is supported by differences in anatomy of the two groups: in Asiatic forms the nasal is excluded from the major series of bones participating in deflecting the impact in the upper jaw and the dentary−angular interlocking makes a more rigid lower jaw.
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Tyrannosauridae can be subdivided into two distinct subfamilies—the Albertosaurinae and the Tyrannosaurinae. Previously recognized subdivisions Aublysodontinae and Shanshanosaurinae are rejected because they are based on insufficient material and juvenile specimens. Our results are based upon a phylogenetic analysis using PAUP program (Swofford 1999) of 77 skull characters and seven genera (Albertosaurus, Alioramus, Daspletosaurus, Gorgosaurus, Nanotyrannus, Tarbosaurus, and Tyrannosaurus); with Allosaurus as outgroup. Of the 77 characters used, more than half were parsimony informative. Asingle most parsimonious tree was obtained with the Tree Length being 88. The analysis of cranial characters and comparison of postcranial features reveal that Tarbosaurus bataar is not the sister taxon of Tyrannosaurus rex (contra Holtz 2001). Their similarities are partially due to the fact that both are extremely large animals. Thus, Tarbosaurus should be considered a genus distinct from Tyrannosaurus.
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