The skull evolution of oviraptorosaurian dinosaurs: the role of niche partitioning in diversification

Oviraptorosaurs are bird‐like theropod dinosaurs that thrived in the final pre‐extinction ecosystems during the latest Cretaceous, and the beaked, toothless skulls of derived species are regarded as some of the most peculiar among dinosaurs. Their aberrant morphologies are hypothesized to have been caused by rapid evolution triggered by an ecological/biological driver, but little is known about how their skull shapes and functional abilities diversified. Here, we use quantitative techniques to study oviraptorosaur skull form and mandibular function. We demonstrate that the snout is particularly variable, that mandibular form and upper/lower beak form are significantly correlated with phylogeny, and that there is a strong and significant correlation between mandibular function and mandible/lower beak shape, suggesting a form–function association. The form–function relationship and phylogenetic signals, along with a moderate allometric signal in lower beak form, indicate that similar mechanisms governed beak shape in oviraptorosaurs and extant birds. The two derived oviraptorosaur clades, oviraptorids and caenagnathids, are significantly separated in morphospace and functional space, indicating that they partitioned niches. Oviraptorids coexisting in the same ecosystem are also widely spread in morphological and functional space, suggesting that they finely partitioned feeding niches, whereas caenagnathids exhibit extreme disparity in beak size. The diversity of skull form and function was likely key to the diversification and evolutionary success of oviraptorosaurs in the latest Cretaceous.     

Latirhinus uitstlani,a ‘broad-nosed’ saurolophine hadrosaurid (Dinosauria,Ornithopoda) from the late Campanian (Cretaceous) of northern Mexico

We describe a new genus and species of hadrosaurid dinosaur, Latirhinus uitstlani, from upper Campanian strata of the Cerro del Pueblo Formation in northern Mexico. The new taxon differs from other hadrosaurids, except Gryposaurus, in possessing a deep and arcuate dorsal process of the nasal; it differs from Gryposaurus, as well as from all other saurolophines, in having a broader narial foramen, and an anterodorsally curved deltoid crest of the scapula (convergent with Lambeosaurinae within the context of Hadrosauridae). L. uitstlani appears to be allied to ‘kritosaurs’, particularly to species of Gryposaurus, and it represents one of the southernmost saurolophines of North America. Likewise, L. uitstlani adds to the diversity of saurolophines in the southern Western Interior Basin and its occurrence in northern Mexico is consistent with the apparent predominance of ‘kritosaurs’ in southern Laramidia during late Campanian times.     

A remarkable short-snouted horned dinosaur from the Late Cretaceous (late Campanian) of southern Laramidia

The fossil record of centrosaurine ceratopsids is largely restricted to the northern region of western North America (Alberta, Montana and Alaska). Exceptions consist of single taxa from Utah (Diabloceratops) and China (Sinoceratops), plus otherwise fragmentary remains from the southern Western Interior of North America. Here, we describe a remarkable new taxon, Nasutoceratops titusi n. gen. et sp., from the late Campanian Kaiparowits Formation of Utah, represented by multiple specimens, including a nearly complete skull and partial postcranial skeleton. Autapomorphies include an enlarged narial region, pneumatic nasal ornamentation, abbreviated snout and elongate, rostrolaterally directed supraorbital horncores. The subrectangular parietosquamosal frill is relatively unadorned and broadest in the mid-region. A phylogenetic analysis indicates that Nasutoceratops is the sister taxon to Avaceratops, and that a previously unknown subclade of centrosaurines branched off early in the group''s history and persisted for several million years during the late Campanian. As the first well-represented southern centrosaurine comparable in age to the bulk of northern forms, Nasutoceratops provides strong support for the provincialism hypothesis, which posits that Laramidia—the western landmass formed by inundation of the central region of North America by the Western Interior Seaway—hosted at least two coeval dinosaur communities for over a million years of late Campanian time.     

Thyreophoran vertebrae from the Callovian (Middle Jurassic) of the Vaches Noires cliffs (Normandy,France), with remarks on the dinosaur assemblage from the Vaches Noires

Two associated incomplete thyreophoran dorsal vertebrae from the Callovian Marnes de Dives Formation of the Vaches Noires cliffs, on the Normandy coast, are referred to an indeterminate stegosaur that appears to be different from Lexovisaurus, previously reported from the Callovian of western France. These vertebrae are the first evidence of thyreophorans from the Vaches Noires and complement the dinosaur assemblage from this locality, which hitherto consisted of several theropod taxa and an indeterminate sauropod. The dinosaur record from the Vaches Noires is heavily dominated by theropods and this imbalance is difficult to explain. A possible explanation may be that the dinosaur sample from the Vaches Noires is too small to be statistically significant and representative of the original faunal assemblage from which it is derived.     

A juvenile skull of Dysalotosaurus lettowvorbecki (Ornithischia: Iguanodontia), and implications for cranial ontogeny,phylogeny, and taxonomy in ornithopod dinosaurs

A juvenile skull of Dysalotosaurus lettowvorbecki, a basal iguanodontian from the Middle Dinosaur Member (Kimmeridgian) of the Tendaguru Beds (Tanzania), is described and reconstructed in detail. Further preparation and computed tomography scans have uncovered additional and formerly unknown skull elements, especially of the lower jaw. The reconstruction of the skull reveals ontogenetic differences to earlier reconstructions, which were based on specimens of relatively older individuals. The most notable ontogenetic changes in Dysalotosaurus are a relative lengthening of the muzzle, a decrease in the relative size of the orbit, and a straightening of the posterior skull roof. Additionally, ontogenetic variations were found in many single elements of the skull, all of which reflect the three main tendencies described above. Furthermore, there might have been an ontogenetic change in the diet, from omnivorous juveniles to fully herbivorous adults. The results of this study will help to evaluate ontogenetic stages in other ornithopods, and will shed light on a crucial stage of ornithopod evolution that culminated in the highly specialized and diverse hadrosaurs of the Cretaceous. Many of the evolutionary changes seen in this lineage can be attributed to peramorphism, and probably reflect a perfection of the adaptation towards an obligatory herbivorous diet. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160 , 366–396.     

Analysis of forelimb function in basal ceratopsians

Here, I present the first study of forelimb function in basal ceratopsians [Correction added after publication 28 July 2007: in the preceding sentence 'Here, I present the first study of forelimb function in basal ceratopsians Dinosauria Orthischia ' was corrected to 'Here, I present the first study of forelimb function in basal ceratopsians']. I examined forelimb bones and casts of Psittacosaurus neimongoliensis , Psittacosaurus mongoliensis , Leptoceratops gracilis and Protoceratops andrewsi . For Ps. neimongoliensis and L. gracilis , I used manual manipulations of bones and casts to determine the range of motion at available forelimb joints. I then used range of motion and morphology to test the predictions of several hypotheses of forelimb function. Forelimb morphology and range of motion indicate that Psittacosaurus was an obligate biped and that Leptoceratops and Protoceratops were capable of quadrupedal locomotion. Forelimb mobility was too limited in Psittacosaurus for the hands to reach the mouth. Leptoceratops and Protoceratops are members of an evolutionary radiation in which an extension of the glenoid enabled the forelimbs to sprawl laterally for transverse pivoting, perhaps for display, but quadrupedal locomotion was accomplished with the elbows tucked in. In Protoceratops , the radius crosses over the ulna, causing the palms to face caudally. In Leptoceratops , the radius does not cross over the ulna; the palms face largely medially and the fingers have been reoriented so that flexion produces a caudal, propulsive force, even without caudally facing palms.     

FORELIMB FUNCTION IN ORNITHOLESTES HERMANNI OSBORN (DINOSAURIA, THEROPODA)

Abstract:  Ornitholestes hermanni is a Late Jurassic theropod dinosaur from North America. This kinematic study of Ornitholestes uses manual manipulations of forelimb casts to determine range of motion. The manual phalanges of the O. hermanni holotype, previously unidentified, are here identified as phalanges I-1, I-2 (ungual), II-2 and II-3 (ungual). At all represented manual joints, hyperextensibility is small or absent, whereas flexion is strong, as in most other theropods. The elbow can be strongly flexed beyond a right angle. When data on range of forelimb motion in Ornitholestes are added to such data from other theropods, high elbow flexion is present in maniraptoriform coelurosaurs but not in basal theropods. Forelimb functions requiring strong elbow flexion (such as holding objects to the chest, or tucking the forearms in for their protection or to reduce wind resistance or heat loss) were therefore available to maniraptoriform coelurosaurs but not to basal theropods.     

Phylogenetic eigenvectors and nonstationarity in the evolution of theropod dinosaur skulls

Despite the long‐standing interest in nonstationarity of both phenotypic evolution and diversification rates, only recently have methods been developed to study this property. Here, we propose a methodological expansion of the phylogenetic signal‐representation (PSR) curve based on phylogenetic eigenvectors to test for nonstationarity. The PSR curve is built by plotting the coefficients of determination R² from phylogenetic eigenvector regression (PVR) models increasing the number of phylogenetic eigenvectors against the accumulated eigenvalues. The PSR curve is linear under a stationary model of trait evolution (i.e. the Brownian motion model). Here we describe the distribution of shifts in the models R² and used a randomization procedure to compare observed and simulated shifts along the PSR curve, which allowed detecting nonstationarity in trait evolution. As an applied example, we show that the main evolutionary pattern of variation in the theropod dinosaur skull was nonstationary, with a significant shift in evolutionary rates in derived oviraptorosaurs, an aberrant group of mostly toothless, crested, birdlike theropods. This result is also supported by a recently proposed Bayesian‐based method (AUTEUR). A significant deviation between Ceratosaurus and Limusaurus terminal branches was also detected. We purport that our new approach is a valuable tool for evolutionary biologists, owing to its simplicity, flexibility and comprehensiveness.     

Dispersal and diversity in the earliest North American sauropodomorph dinosaurs, with a description of a new taxon

Sauropodomorph dinosaurs originated in the Southern Hemisphere in the Middle or Late Triassic and are commonly portrayed as spreading rapidly to all corners of Pangaea as part of a uniform Late Triassic to Early Jurassic cosmopolitan dinosaur fauna. Under this model, dispersal allegedly inhibited dinosaurian diversification, while vicariance and local extinction enhanced it. However, apomorphy-based analyses of the known fossil record indicate that sauropodomorphs were absent in North America until the Early Jurassic, reframing the temporal context of their arrival. We describe a new taxon from the Kayenta Formation of Arizona that comprises the third diagnosable sauropodomorph from the Early Jurassic of North America. We analysed its relationships to test whether sauropodomorphs reached North America in a single sweepstakes event or in separate dispersals. Our finding of separate arrivals by all three taxa suggests dispersal as a chief factor in dinosaurian diversification during at least the early Mesozoic. It questions whether a 'cosmopolitan' dinosaur fauna ever existed, and corroborates that vicariance, extinction and dispersal did not operate uniformly in time or under uniform conditions during the Mesozoic. Their relative importance is best measured in narrow time slices and circumscribed geographical regions.     

Footprints pull origin and diversification of dinosaur stem lineage deep into Early Triassic

The ascent of dinosaurs in the Triassic is an exemplary evolutionary radiation, but the earliest phase of dinosaur history remains poorly understood. Body fossils of close dinosaur relatives are rare, but indicate that the dinosaur stem lineage (Dinosauromorpha) originated by the latest Anisian (ca 242-244 Ma). Here, we report footprints from the Early-Middle Triassic of Poland, stratigraphically well constrained and identified using a conservative synapomorphy-based approach, which shifts the origin of the dinosaur stem lineage back to the Early Olenekian (ca 249-251 Ma), approximately 5-9 Myr earlier than indicated by body fossils, earlier than demonstrated by previous footprint records, and just a few million years after the Permian/Triassic mass extinction (252.3 Ma). Dinosauromorph tracks are rare in all Polish assemblages, suggesting that these animals were minor faunal components. The oldest tracks are quadrupedal, a morphology uncommon among the earliest dinosauromorph body fossils, but bipedality and moderately large body size had arisen by the Early Anisian (ca 246 Ma). Integrating trace fossils and body fossils demonstrates that the rise of dinosaurs was a drawn-out affair, perhaps initiated during recovery from the Permo-Triassic extinction.