François Therrien

High-precision U-Pb zircon geochronology of the Late Triassic Chinle Formation, Petrified Forest National Park (Arizona, USA): Temporal constraints on the early evolution of dinosaurs

The Triassic successions of the Colorado Plateau preserve an important record of vertebrate evolution and climate change, but correlations to a global Triassic framework are hampered by a lack of geochronological control. Tuffaceous sandstones and siltstones were collected from the Upper Triassic Chinle Formation exposed in the Petrified Forest National Park, Arizona, USA, within a refined stratigraphic context of 31 detailed measured sections. U-Pb analyses by the isotope dilution–thermal ionization mass spectrometry (ID-TIMS) method constrain maximum depositional ages for nine tuffaceous beds and provide new insights into the depositional history of the Chinle fluvial system. The base of the Blue Mesa Member of the Chinle Formation is placed at ca. 225 Ma, and the top of the Petrified Forest Member is placed at 208 Ma or younger, bracketing an ∼280-m-thick section that spans nearly the entire Norian Stage of the Late Triassic. Estimated sediment accumulation rates throughout the section reflect extensive hiatuses and/or sediment removal by channel erosion. The new geochronology for the Chinle Formation underscores the potential pitfalls of correlation of fluvial units based solely on lithostratigraphic criteria. A mid-Norian age (ca. 219–213 Ma) for the distinctive Sonsela conglomeratic sandstone bed constrains the Adamanian-Revueltian land vertebrate faunachron boundary. Our new data permit a significant time overlap between the lower Chinle sequence and the dinosauromorph-rich Ischigualasto Formation of northwestern Argentina. Near-contemporaneity of the trans-American deposits and their faunal similarities imply that early dinosaur evolution occurred rapidly across the Americas.

Evolution of olfaction in non-avian theropod dinosaurs and birds

Little is known about the olfactory capabilities of extinct basal (non-neornithine) birds or the evolutionary changes in olfaction that occurred from non-avian theropods through modern birds. Although modern birds are known to have diverse olfactory capabilities, olfaction is generally considered to have declined during avian evolution as visual and vestibular sensory enhancements occurred in association with flight. To test the hypothesis that olfaction diminished through avian evolution, we assessed relative olfactory bulb size, here used as a neuroanatomical proxy for olfactory capabilities, in 157 species of non-avian theropods, fossil birds and living birds. We show that relative olfactory bulb size increased during non-avian maniraptoriform evolution, remained stable across the non-avian theropod/bird transition, and increased during basal bird and early neornithine evolution. From early neornithines through a major part of neornithine evolution, the relative size of the olfactory bulbs remained stable before decreasing in derived neoavian clades. Our results show that, rather than decreasing, the importance of olfaction actually increased during early bird evolution, representing a previously unrecognized sensory enhancement. The relatively larger olfactory bulbs of earliest neornithines, compared with those of basal birds, may have endowed neornithines with improved olfaction for more effective foraging or navigation skills, which in turn may have been a factor allowing them to survive the end-Cretaceous mass extinction.

MY THEROPOD IS BIGGER THAN YOURS … OR NOT: ESTIMATING BODY SIZE FROM SKULL LENGTH IN THEROPODS

Abstract To develop a widely applicable method to estimate body size in theropods, the scaling relationship between skull length, body length, and body mass was investigated using 13 strictly carnivorous, non-avialan theropod taxa ranging in size from the 1-m Sinosauropteryx prima to the 12-m Tyrannosaurus rex. Body length was obtained from the literature for complete to nearly-complete specimens and body mass was obtained from three-dimensional mathematical slicing of those same specimens to ensure accurate body length-body mass associations. Least-square regressions reveal a tight correlation between skull length and body length (SK-BL) and skull length and body mass (SK-BM). The SK-BL regression is negatively allometric, which indicates that skulls become longer relative to body length with increasing body size. In contrast, the SK-BM regression is positively allometric, indicating that body mass increases faster than skull length with increasing body size. These conclusions confirm that the common practice of scaling isometrically smaller relatives of a given taxon to obtain body length and body mass estimates is not valid. Although predictive equations derived from the regressions fail to predict accurately body size in abelisaurids and juvenile theropods due to their different head/body proportions, they produce accurate body size estimates for theropods of known body size, thus validating their applicability. Body size estimates for Carcharodontosaurus and Giganotosaurus, approaching 13 m and 14 tons, suggest that they may have surpassed Tyrannosaurus in size. A revised body size estimate for a large Spinosaurus specimen suggests a much shorter and heavier animal than recently suggested.

Feathered Non-Avian Dinosaurs from North America Provide Insight into Wing Origins

Fancy Feathers In the past few decades, an increasing number of dinosaurs have been shown to have possessed feathers. While it seems likely that feathers themselves may have evolved for thermoregulation, the original function of wings has been less clear and remained a matter of debate. Based on examination of three Theropod specimens from the genus Ornothomimus, Zelenitsky et al. (p. 510) conclude that the feathered wing may have evolved not for locomotion or prey capture (the animals were herbivores), but rather as a courtship display. All of the individuals examined had a covering consisting of short filamentous feathers, but the adult specimens, which would have reached sexual maturity, also had long shafted feathers on their forelimbs. Analysis of feathered theropod fossils from North America suggests that feathers may have first evolved for display. Previously described feathered dinosaurs reveal a fascinating record of feather evolution, although substantial phylogenetic gaps remain. Here we report the occurrence of feathers in ornithomimosaurs, a clade of non-maniraptoran theropods for which fossilized feathers were previously unknown. The Ornithomimus specimens, recovered from Upper Cretaceous deposits of Alberta, Canada, provide new insights into dinosaur plumage and the origin of the avian wing. Individuals from different growth stages reveal the presence of a filamentous feather covering throughout life and winglike structures on the forelimbs of adults. The appearance of winglike structures in older animals indicates that they may have evolved in association with reproductive behaviors. These specimens show that primordial wings originated earlier than previously thought, among non-maniraptoran theropods.

Olfactory acuity in theropods: palaeobiological and evolutionary implications.

This research presents the first quantitative evaluation of the olfactory acuity in extinct theropod dinosaurs. Olfactory ratios (i.e. the ratio of the greatest diameter of the olfactory bulb to the greatest diameter of the cerebral hemisphere) are analysed in order to infer the olfactory acuity and behavioural traits in theropods, as well as to identify phylogenetic trends in olfaction within Theropoda. A phylogenetically corrected regression of olfactory ratio to body mass reveals that, relative to predicted values, the olfactory bulbs of (i) tyrannosaurids and dromaeosaurids are significantly larger, (ii) ornithomimosaurs and oviraptorids are significantly smaller, and (iii) ceratosaurians, allosauroids, basal tyrannosauroids, troodontids and basal birds are within the 95% CI. Relative to other theropods, olfactory acuity was high in tyrannosaurids and dromaeosaurids and therefore olfaction would have played an important role in their ecology, possibly for activities in low-light conditions, locating food, or for navigation within large home ranges. Olfactory acuity was the lowest in ornithomimosaurs and oviraptorids, suggesting a reduced reliance on olfaction and perhaps an omnivorous diet in these theropods. Phylogenetic trends in olfaction among theropods reveal that olfactory acuity did not decrease in the ancestry of birds, as troodontids, dromaeosaurids and primitive birds possessed typical or high olfactory acuity. Thus, the sense of smell must have remained important in primitive birds and its presumed decrease associated with the increased importance of sight did not occur until later among more derived birds.

A NEW TURTLE FROM THE ARUNDEL CLAY FACIES (POTOMAC FORMATION, EARLY CRETACEOUS) OF MARYLAND, U.S.A.

Abstract A new paracryptodiran turtle, Arundelemys dardeni, gen. et sp. nov., is described on the basis of an isolated, nearly complete skull from the Early Cretaceous Arundel Clay facies of Maryland, USA. The basicranial region exhibits the paracryptodiran condition of a single foramen for the canalis caroticus internus located midway along the basisphenoid. As revealed by CT scans, the basicranial region of Arundelemys is unusual in that the right and left canales carotici interni merge just before reaching the sella turcica and the canalis caroticus lateralis is very small or absent. A phylogenetic analysis places Arundelemys dardeni as the basal-most member of the Paracryptodira. Within the Paracryptodira, Arundelemys dardeni is most similar to Compsemys victa in general proportions.

Homology of the “Ethmoid Complex” of Tyrannosaurids and its Implications for the Reconstruction of the Olfactory Apparatus of Non-Avian Theropods

Abstract The “ethmoid complex” is an enigmatic element of the anterior portion of the braincase first described in Tyrannosaurus rex in 1912, which has since been recognized in many non-avian theropods. Because the “ethmoid complex” is a landmark for the reconstruction of the olfactory apparatus of non-avian theropods, we clarify the homology of this structure among archosaurs. The “ethmoid complex” consists of a trough-shaped element that is attached to an anteriorly-located median septum capped by a dorsal plate. Based on anatomical comparisons with the olfactory region of extant birds and crocodylians, the components of the “ethmoid complex” are shown to have cartilaginous or osteological homologues among extant archosaurs: the trough is homologous to the anterior portion of the planum supraseptale of crocodylians and embryonic birds, whereas the median septum and overlying dorsal plate are homologous to the avian mesethmoid and to the nasal septum and tectum nasi of crocodylians. Based on the location and ossification of olfactory region structures in non-avian theropods, the most appropriate terms for elements of the “ethmoid complex” are the sphenethmoid for the trough and the mesethmoid for the median septum and dorsal plate. The olfactory bulbs of nonavian theropods were housed within the sphenethmoid, which restricted the maximum size of the olfactory bulbs to a size smaller than the cerebral hemispheres.

Eggshell Porosity Provides Insight on Evolution of Nesting in Dinosaurs.

Knowledge about the types of nests built by dinosaurs can provide insight into the evolution of nesting and reproductive behaviors among archosaurs. However, the low preservation potential of their nesting materials and nesting structures means that most information can only be gleaned indirectly through comparison with extant archosaurs. Two general nest types are recognized among living archosaurs: 1) covered nests, in which eggs are incubated while fully covered by nesting material (as in crocodylians and megapodes), and 2) open nests, in which eggs are exposed in the nest and brooded (as in most birds). Previously, dinosaur nest types had been inferred by estimating the water vapor conductance (i.e., diffusive capacity) of their eggs, based on the premise that high conductance corresponds to covered nests and low conductance to open nests. However, a lack of statistical rigor and inconsistencies in this method render its application problematic and its validity questionable. As an alternative we propose a statistically rigorous approach to infer nest type based on large datasets of eggshell porosity and egg mass compiled for over 120 extant archosaur species and 29 archosaur extinct taxa/ootaxa. The presence of a strong correlation between eggshell porosity and nest type among extant archosaurs indicates that eggshell porosity can be used as a proxy for nest type, and thus discriminant analyses can help predict nest type in extinct taxa. Our results suggest that: 1) covered nests are likely the primitive condition for dinosaurs (and probably archosaurs), and 2) open nests first evolved among non-avian theropods more derived than Lourinhanosaurus and were likely widespread in non-avian maniraptorans, well before the appearance of birds. Although taphonomic evidence suggests that basal open nesters (i.e., oviraptorosaurs and troodontids) were potentially the first dinosaurs to brood their clutches, they still partially buried their eggs in sediment. Open nests with fully exposed eggs only became widespread among Euornithes. A potential co-evolution of open nests and brooding behavior among maniraptorans may have freed theropods from the ground-based restrictions inherent to covered nests and allowed the exploitation of alternate nesting locations. These changes in nesting styles and behaviors thus may have played a role in the evolutionary success of maniraptorans (including birds).

First fossil gravid turtle provides insight into the evolution of reproductive traits in turtles

Here we report on the first discovery of shelled eggs inside the body cavity of a fossil turtle and on an isolated egg clutch, both referable to the Cretaceous turtle Adocus. These discoveries provide a unique opportunity to gain insight into the reproductive traits of an extinct turtle and to understand the evolution of such traits among living turtles. The gravid adult and egg clutch indicate that Adocus laid large clutches of rigid-shelled spherical eggs and established their nests near rivers, traits that are shared by its closest living relatives, the soft-shelled turtles. Adocus eggshell, however, was probably more rigid than that of living turtles, based on its great thickness and structure, features that may represent unique adaptations to intense predation or to arid nest environments. In light of the reproductive traits observed in Adocus, the distribution of reproductive traits among turtles reveals that large clutches of rigid-shelled eggs are primitive for hidden-necked turtles (cryptodirans) and that spherical eggs may have evolved independently within this group.

ASSIGNMENT OF YAMACERATOPS DORNGOBIENSIS AND ASSOCIATED REDBEDS AT SHINE US KHUDAG (EASTERN GOBI, DORNGOBI PROVINCE, MONGOLIA) TO THE REDESCRIBED JAVKHLANT FORMATION (UPPER CRETACEOUS)

DAVID A. EBERTH, YOSHITSUGU KOBAYASHI, YUONG-NAM LEE, OCTÁVIO MATEUS, FRANÇOIS THERRIEN, DARLA K. ZELENITSKY, and MARK A. NORELL; Royal Tyrrell Museum, Box 7500, Drumheller, Alberta, Canada, david.eberth@gov.ab.ca; Hokkaido University, Hokkaido, Japan; Korean Institute of Geoscience & Mineral Resources, Daejeon, Korea; Museu da Lourinhã & Universidade Nova de Lisboa, Lourinhã, Portugal; Department of Geoscience, University of Calgary, Calgary, Alberta, Canada; Division of Paleontology, American Museum of Natural History, New York City, USA