Tyrannosaurids are a well-supported clade of very large predatory dinosaurs of Late Cretaceous Asiamerica. Traditional dinosaurian systematics place these animals within the infraorder Carnosauria with the other large theropods (allosaurids, megalosaurids). A new cladistic analysis indicates that the tyrannosaurs were in fact derived members of the Coelurosauria, a group of otherwise small theropods. Despite certain gross cranial similarities with the large predators of the Jurassic and Early Cretaceous, the Late Cretaceous tyrannosaurids are shown to be the sister group to ornithomimids and troodontids, which share a derived condition of the metatarsus. This clade is found to be nested within Maniraptora, which is a more inclusive taxon than previously recognized. The atrophied carpal structure found in tyrannosaurids and ornithomimids is derived from a maniraptoran condition with a large semilunate carpal, rather than from the plesiomorphic theropod morphology.The taxa “Carnosauria” and “Deinonychosauria” (Dromaeosauridae plus Troodontidae) are shown to be polyphyletic, and the Late Jurassic African form Elaphrosaurus is found to be the sister taxon to Abelisauridae rather than a primitive ornithomimosaur. Purported allosaurid-tyrannosaurid synapomorphies are seen to be largely size-related, present in the larger members of both clades, but absent in smaller members of the Tyrannosauridae. The remaining giant tetanurine theropods (Megalosaurus and Torvosaurus) were found to be progressively distant outgroups to an allosaurid-coelurosaur clade. The inclusion of the Tyrannosauridae within Maniraptora suggests a major adaptive radiation of coelurosaurs within Cretaceous Asiamerica comparable to contemporaneous radiations in various herbivorous dinosaurian clades.
The Late Cretaceous dinosaur Tyrannosaurus rex was recently split into three species based on the premise that variation in the T. rex hypodigm is exceptional, indicating cryptic species and “robust” and “gracile” morphs. The morphs are based on proportional ratios throughout the skeleton. The species are claimed to be stratigraphically separate, with an early robust species followed by robust and gracile descendants. There are problems with the hypothesis: the taxon diagnoses are based on two features that overlap between the species; several skulls cannot be identified based on the diagnoses; proportional comparisons between Tyrannosaurus and other theropods are based on incomparable samples; the tooth data are problematic; the stratigraphic framework divides the Hell Creek Formation into thirds, without the stratigraphic position of each specimen, or independent age control showing the subdivisions are coeval over the entire geographic area; previous work found variation in T. rex, but it cannot be parsed into discrete categories. We tested for “gracile” and “robust” morphs by analyzing the femoral and tooth ratios that were published in the multiple species study using agglomerative hierarchical clustering. The results found that each set of ratios are explained by one cluster, showing that dimorphism is not supported. We tested for exceptional variation of the femoral ratio of Tyrannosaurus; we calculated the mean intraspecific robusticity for 112 species of living birds and 4 nonavian theropods. The results showed that the absolute variation in Tyrannosaurus is unexceptional and it does not indicate cryptic diversity. We conclude that “T. regina” and “T. imperator” are subjective junior synonyms of T. rex.
The spinosaurids represent an enigmatic and highly unusual form of large tetanuran theropods that were first identified in 1915. A recent flurry of discoveries and taxonomic revisions of this important and interesting clade had added greatly to our knowledge, however, spinosaur body fossils are generally rare and most species are known from only limited skeletal remains. Their unusual anatomical adaptations to the skull, limbs and axial column all differ from other large theropods and point to an unusual ecological niche and a lifestyle intimately linked to water.
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