Gregory M. Erickson

Definitive fossil evidence for the extant avian radiation in the Cretaceous

Long-standing controversy surrounds the question of whether living bird lineages emerged after non-avian dinosaur extinction at the Cretaceous/Tertiary (K/T) boundary or whether these lineages coexisted with other dinosaurs and passed through this mass extinction event. Inferences from biogeography and molecular sequence data (but see ref. 10) project major avian lineages deep into the Cretaceous period, implying their ‘mass survival’ at the K/T boundary. By contrast, it has been argued that the fossil record refutes this hypothesis, placing a ‘big bang’ of avian radiation only after the end of the Cretaceous. However, other fossil data—fragmentary bones referred to extant bird lineages—have been considered inconclusive. These data have never been subjected to phylogenetic analysis. Here we identify a rare, partial skeleton from the Maastrichtian of Antarctica as the first Cretaceous fossil definitively placed within the extant bird radiation. Several phylogenetic analyses supported by independent histological data indicate that a new species, Vegavis iaai, is a part of Anseriformes (waterfowl) and is most closely related to Anatidae, which includes true ducks. A minimum of five divergences within Aves before the K/T boundary are inferred from the placement of Vegavis; at least duck, chicken and ratite bird relatives were coextant with non-avian 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.

The ontogeny of bite‐force performance in American alligator (Alligator mississippiensis)

American alligators Alligator mississippiensis undergo major transformations in morphology and ecology during development. These include several thousand-fold changes in body mass, modified snout and dental proportions, and shifts in diet from small, delicate foodstuffs to the inclusion of increasingly larger, more robust prey. How these changes in anatomical form contribute to actual physical performance and niche use is largely unknown. In the present study, bite-force measurements for 41 specimens of A. mississipiensis, were made throughout ontogeny (hatchling–older adults) using a series of precision force transducers. How this performance indicator scaled with respect to cranial and whole-body measurements was determined. Bite-force production throughout development was contrasted with ontogenetic changes in trophic ecology. The influences of this performance measure on these changes were then analysed. The results showed a 800-fold range (12–9452 N) of bite forces with values positively correlating with increases in body size. Scaling of biting forces through ontogeny showed positive allometry with respect to body mass, head length, jaw length, snout–vent length and total length. These patterns may be attributable to allometric growth of individual skeletal elements (and associated musculature), and/or progressive fusion and ossification of skull and jawbones during development. The overall pattern of force increase throughout ontogeny did not vary in association with major shifts in diet. Notably, the bite-force values for adult A. mississippiensis are the highest measured for any living animal and represent the first measures for a large crocodilian. Additionally, these data provide the first documentation of how bite force changes during ontogeny in a reptile. By bridging the rich morphological and ecological databases for these animals, this study opens the door to a comprehensive understanding of feeding in A. mississippiensis. Furthermore, it provides groundwork for standardized comparative studies of feeding among crocodilian, reptilian, or other gnathostome vertebrates.

A Jurassic ceratosaur from China helps clarify avian digital homologies

Theropods have traditionally been assumed to have lost manual digits from the lateral side inward, which differs from the bilateral reduction pattern seen in other tetrapod groups. This unusual reduction pattern is clearly present in basal theropods, and has also been inferred in non-avian tetanurans based on identification of their three digits as the medial ones of the hand (I-II-III). This contradicts the many developmental studies indicating II-III-IV identities for the three manual digits of the only extant tetanurans, the birds. Here we report a new basal ceratosaur from the Oxfordian stage of the Jurassic period of China (156–161 million years ago), representing the first known Asian ceratosaur and the only known beaked, herbivorous Jurassic theropod. Most significantly, this taxon possesses a strongly reduced manual digit I, documenting a complex pattern of digital reduction within the Theropoda. Comparisons among theropod hands show that the three manual digits of basal tetanurans are similar in many metacarpal features to digits II-III-IV, but in phalangeal features to digits I-II-III, of more basal theropods. Given II-III-IV identities in avians, the simplest interpretation is that these identities were shared by all tetanurans. The transition to tetanurans involved complex changes in the hand including a shift in digit identities, with ceratosaurs displaying an intermediate condition.

A king-sized theropod coprolite

Fossil faeces (coprolites) provide unique trophic perspectives on ancient ecosystems. Yet, although thousands of coprolites have been discovered, specimens that can be unequivocally attributed to carnivorous dinosaurs are almost unknown. A few fossil faeces have been ascribed to herbivorous dinosaurs, but it is more difficult to identify coprolites produced by theropods because other carnivorous taxa coexisted with dinosaurs and most faeces are taxonomically ambiguous. Thus sizeable (up to 20 cm long and 10 cm wide) phosphatic coprolites from Belgium and India, that have been attributed to dinosaurs might have been produced by contemporaneous crocodylians or fish. But there is no ambiguity about the theropod origin of the Cretaceous coprolite we report here. This specimen is more than twice as large as any previously reported carnivore coprolite, and its great size and temporal and geographic context indicate that it was produced by a tyrannosaur, most likely Tyrannosaurus rex. The specimen contains a high proportion (30–50%) of bone fragments, and is rare tangible evidence of theropod diet and digestive processes.

Evolution of the biomechanical material properties of the femur

The biomechanical performance of long bones is dictated by four key factors: element size, element shape, loading conditions, and material properties. Our understanding of the latter of these has been mostly limited to eutherian mammals and birds, which show similarity. Whether their possession of comparable material properties reflects common ancestry or independent evolution is uncertain. In the present analysis, we tested the bending strength, modulus, and failure strains of the femur and its pterygiophore homolog in actinpterygian fish. Sixty‐nine specimens representing basal character states in seven major vertebrate crown clades were tested. These data were then coupled with avian and mammalian data from the literature and analyzed in an evolutionary context using phylogenetic character analysis. Mean values of 188 MPa for yield strength, 22.4 GPa for Youngs modulus, and 8,437 μ∈ for yield strain were obtained for the long bones. Analysis of variance (ANOVA) revealed comparable values between clades that span a 30,000‐fold range of body mass. We conclude that material properties of the first long bones 475 million years ago were conserved throughout evolution. Major locomotory challenges to femora during vertebrate evolution were almost solely accomplished by modifications of element size and shape. Anat Rec 268:115–124, 2002.

Was dinosaurian physiology inherited by birds? Reconciling slow growth in archaeopteryx.

Background Archaeopteryx is the oldest and most primitive known bird (Avialae). It is believed that the growth and energetic physiology of basalmost birds such as Archaeopteryx were inherited in their entirety from non-avialan dinosaurs. This hypothesis predicts that the long bones in these birds formed using rapidly growing, well-vascularized woven tissue typical of non-avialan dinosaurs. Methodology/Principal Findings We report that Archaeopteryx long bones are composed of nearly avascular parallel-fibered bone. This is among the slowest growing osseous tissues and is common in ectothermic reptiles. These findings dispute the hypothesis that non-avialan dinosaur growth and physiology were inherited in totality by the first birds. Examining these findings in a phylogenetic context required intensive sampling of outgroup dinosaurs and basalmost birds. Our results demonstrate the presence of a scale-dependent maniraptoran histological continuum that Archaeopteryx and other basalmost birds follow. Growth analysis for Archaeopteryx suggests that these animals showed exponential growth rates like non-avialan dinosaurs, three times slower than living precocial birds, but still within the lowermost range for all endothermic vertebrates. Conclusions/Significance The unexpected histology of Archaeopteryx and other basalmost birds is actually consistent with retention of the phylogenetically earlier paravian dinosaur condition when size is considered. The first birds were simply feathered dinosaurs with respect to growth and energetic physiology. The evolution of the novel pattern in modern forms occurred later in the groups history.

Growth patterns in brooding dinosaurs reveals the timing of sexual maturity in non-avian dinosaurs and genesis of the avian condition

The timing of sexual maturation in non-avian dinosaurs is not known. In extant squamates and crocodilians it occurs in conjunction with the initial slowing of growth rates as adult size is approached. In birds (living dinosaurs) on the other hand, reproductive activity begins well after somatic maturity. Here we used growth line counts and spacing in all of the known brooding non-avian dinosaurs to determine the stages of development when they perished. It was revealed that sexual maturation occurred well before full adult size was reached—the primitive reptilian condition. In this sense, the life history and physiology of non-avian dinosaurs was not like that of modern birds. Palaeobiological ramifications of these findings include the potential to deduce reproductive lifespan, fecundity and reproductive population sizes in non-avian dinosaurs, as well as aid in the identification of secondary sexual characteristics.

Tyrannosaur Paleobiology: New Research on Ancient Exemplar Organisms

Tyrannosaurs Revisited Tyrannosaurs represent some of the most successful and largest carnivores in Earths history. An expanding fossil record has allowed studies of their evolution and behavior that now allow broader comparisons with other groups, not just dinosaurs. Brusatte et al. (p. 1481) review the biology and evolutionary history of tyrannosaurs and update their phylogenetic relations to include several new fossils. The analysis suggests that tyrannosaurs remained relatively small (less than about 5 meters long) until the Late Cretaceous (about 80 million years ago). Tyrannosaurs, the group of dinosaurian carnivores that includes Tyrannosaurus rex and its closest relatives, are icons of prehistory. They are also the most intensively studied extinct dinosaurs, and thanks to large sample sizes and an influx of new discoveries, have become ancient exemplar organisms used to study many themes in vertebrate paleontology. A phylogeny that includes recently described species shows that tyrannosaurs originated by the Middle Jurassic but remained mostly small and ecologically marginal until the latest Cretaceous. Anatomical, biomechanical, and histological studies of T. rex and other derived tyrannosaurs show that large tyrannosaurs could not run rapidly, were capable of crushing bite forces, had accelerated growth rates and keen senses, and underwent pronounced changes during ontogeny. The biology and evolutionary history of tyrannosaurs provide a foundation for comparison with other dinosaurs and living organisms.