Mark A. Norell

Two feathered dinosaurs from northeastern China

Current controversy over the origin and early evolution of birds centres on whether or not they are derived from coelurosaurian theropod dinosaurs. Here we describe two theropods from the Upper Jurassic/Lower Cretaceous Chaomidianzi Formation of Liaoning province, China. Although both theropods have feathers, it is likely that neither was able to fly. Phylogenetic analysis indicates that they are both more primitive than the earliest known avialan (bird), Archaeopteryx. These new fossils represent stages in the evolution of birds from feathered, ground-living, bipedal dinosaurs.

A basal dromaeosaurid and size evolution preceding avian flight

Fossil evidence for changes in dinosaurs near the lineage leading to birds and the origin of flight has been sparse. A dinosaur from Mongolia represents the basal divergence within Dromaeosauridae. The taxons small body size and phylogenetic position imply that extreme miniaturization was ancestral for Paraves (the clade including Avialae, Troodontidae, and Dromaeosauridae), phylogenetically earlier than where flight evolution is strongly inferred. In contrast to the sustained small body sizes among avialans throughout the Cretaceous Period, the two dinosaurian lineages most closely related to birds, dromaeosaurids and troodontids, underwent four independent events of gigantism, and in some lineages size increased by nearly three orders of magnitude. Thus, change in theropod body size leading to flights origin was not unidirectional.

Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs

How evolutionary changes in body size are brought about by variance in developmental timing and/or growth rates (also known as heterochrony) is a topic of considerable interest in evolutionary biology. In particular, extreme size change leading to gigantism occurred within the dinosaurs on multiple occasions. Whether this change was brought about by accelerated growth, delayed maturity or a combination of both processes is unknown. A better understanding of relationships between non-avian dinosaur groups and the newfound capacity to reconstruct their growth curves make it possible to address these questions quantitatively. Here we study growth patterns within the Tyrannosauridae, the best known group of large carnivorous dinosaurs, and determine the developmental means by which Tyrannosaurus rex, weighing 5,000 kg and more, grew to be one of the most enormous terrestrial carnivorous animals ever. T. rex had a maximal growth rate of 2.1 kg d-1, reached skeletal maturity in two decades and lived for up to 28 years. T. rexs great stature was primarily attained by accelerating growth rates beyond that of its closest relatives.

Basal tyrannosauroids from China and evidence for protofeathers in tyrannosauroids

Tyrannosauroids are one of the last and the most successful large-bodied predatory dinosaur groups, but their early history remains poorly understood. Here we report a new basal tyrannosauroid from the Early Cretaceous Yixian Formation of western Liaoning, China, which is small and gracile and has relatively long arms with three-fingered hands. The new taxon is the earliest known unquestionable tyrannosauroid found so far. It shows a mosaic of characters, including a derived cranial structure resembling that of derived tyrannosauroids and a primitive postcranial skeleton similar to basal coelurosaurians. One of the specimens also preserves a filamentous integumentary covering similar to that of other coelurosaurian theropods from western Liaoning. This provides the first direct fossil evidence that tyrannosauroids had protofeathers.

The Fossil Record and Evolution: Comparing Cladistic and Paleontologic Evidence for Vertebrate History

The fossil record offers the only direct evidence of extinct life and thus has figured prominently in considerations of evolutionary patterns. But the incomplete nature of the fossil record has also been emphasized in arguments that fossils play only a secondary role in the recovery of phylogenetic histories based on extant taxa. Although these criticisms recently have been countered, there is no general understanding of the correspondence between the fossil record and phylogeny. An empirical survey of recently published studies suggests no basis for assuming that the stratigraphic occurrence of fossils always provides a precise reflection of phylogeny. Nevertheless, our survey of a sample of taxa shows a tendency for positive correlation between age and clade rank and, hence, a degree of correspondence between phylogenetic pattern and the paleontologic record.

A new troodontid dinosaur from China with avian-like sleeping posture

Discovering evidence of behaviour in fossilized vertebrates is rare. Even rarer is evidence of behaviour in non-avialan dinosaurs that directly relates to stereotypical behaviour seen in extant birds (avians) and not previously predicted in non-avialan dinosaurs. Here we report the discovery of a new troodontid taxon from the Early Cretaceous Yixian Formation of western Liaoning, China. Numerous other three-dimensionally preserved vertebrate fossils have been recovered recently at this locality, including some specimens preserving behavioural information. The new troodontid preserves several features that have been implicated in avialan origins. Notably, the specimen is preserved in the stereotypical sleeping or resting posture found in extant Aves. Evidence of this behaviour outside of the crown group Aves further demonstrates that many bird features occurred early in dinosaurian evolution.

Combined support for wholesale taxic atavism in gavialine crocodylians.

Morphological and molecular data sets favor robustly supported, contradictory interpretations of crocodylian phylogeny. A longstanding perception in the field of systematics is that such significantly conflicting data sets should be analyzed separately. Here we utilize a combined approach, simultaneous analyses of all relevant character data, to summarize common support and to reconcile discrepancies among data sets. By conjoining rather than separating incongruent classes of data, secondary phylogenetic signals emerge from both molecular and morphological character sets and provide solid evidence for a unified hypothesis of crocodylian phylogeny. Simultaneous analyses of four gene sequences and paleontological data suggest that putative adaptive convergences in the jaws of gavialines (gavials) and tomistomines (false gavials) offer character support for a grouping of these taxa, making Gavialinae an atavistic taxon. Simple new methods for measuring the influence of extinct taxa on topological support indicate that in this vertebrate order fossils generally stabilize relationships and accentuate hidden phylogenetic signals. Remaining inconsistencies in minimum length trees, including concentrated hierarchical patterns of homoplasy and extensive gaps in the fossil record, indicate where future work in crocodylian systematics should be directed.

A basal troodontid from the Early Cretaceous of China.

Troodontid dinosaurs form one of the most avian-like dinosaur groups. Their phylogenetic position is hotly debated, and they have been allied with almost all principal coelurosaurian lineages. Here we report a basal troodontid dinosaur, Sinovenator changii gen. et sp. nov., from the lower Yixian Formation of China. This taxon has several features that are not found in more derived troodontids, but that occur in dromaeosaurids and avialans. The discovery of Sinovenator and the examination of character distributions along the maniraptoran lineage indicate that principal structural modifications toward avians were acquired in the early stages of maniraptoran evolution.

A Theropod Dinosaur Embryo and the Affinities of the Flaming Cliffs Dinosaur Eggs

An embryonic skeleton of a nonavian theropod dinosaur was found preserved in an egg from Upper Cretaceous rocks in the Gobi Desert of Mongolia. Cranial features identify the embryo as a member of Oviraptoridae. Two embryo-sized skulls of dromaeosaurids, similar to that of Velociraptor, were also recovered in the nest. The eggshell microstructure is similar to that of ratite birds and is of a type common in the Djadokhta Formation at the Flaming Cliffs (Bayn Dzak). Discovery of a nest of such eggs at the Flaming Cliffs in 1923, beneath the Oviraptor philoceratops holotype, suggests that this dinosaur may have been a brooding adult.

A basal tyrannosauroid dinosaur from the Late Jurassic of China.

The tyrannosauroid fossil record is mainly restricted to Cretaceous sediments of Laurasia, although some very fragmentary Jurassic specimens have been referred to this group. Here we report a new basal tyrannosauroid, Guanlong wucaii gen. et sp. nov., from the lower Upper Jurassic of the Junggar Basin, northwestern China. G. wucaii is the oldest known tyrannosauroid and shows several unexpectedly primitive pelvic features. Nevertheless, the limbs of G. wucaii share several features with derived coelurosaurs, and it possesses features shared by other coelurosaurian clades. This unusual combination of character states provides an insight into the poorly known early radiation of the Coelurosauria. Notably, the presumed predatory Guanlong has a large, fragile and highly pneumatic cranial crest that is among the most elaborate known in any non-avian dinosaur and could be comparable to some classical exaggerated ornamental traits among vertebrates.