First Known Feeding Trace of the Eocene Bottom-Dwelling Fish Notogoneus osculus and Its Paleontological Significance

BACKGROUND: The Green River Formation (early Eocene, about 42-53 Ma) at and near Fossil Butte National Monument in Wyoming, USA, is world famous for its exquisitely preserved freshwater teleost fish in the former Fossil Lake. Nonetheless, trace fossils attributed to fish interacting with the lake bottom are apparently rare, and have not been associated directly with any fish species. Here we interpret the first known feeding and swimming trace fossil of the teleost Notogoneus osculus Cope (Teleostei: Gonorynchidae), which is also represented as a body fossil in the same stratum. METHODOLOGY/PRINCIPAL FINDINGS: A standard description of the trace fossil, identified as Undichna cf. U. simplicatas, was augmented by high-resolution digital images and spatial and mathematical analyses, which allowed for detailed interpretations of the anatomy, swimming mode, feeding behavior, and body size of the tracemaker. Our analysis indicates that the tracemaker was about 45 cm long; used its caudal, anal, and pelvic fins (the posterior half of its body) to make the swimming traces; and used a ventrally oriented mouth to make overlapping feeding marks. We hypothesize that the tracemaker was an adult Notogoneus osculus. CONCLUSIONS/SIGNIFICANCE: Our results are the first to link a specific teleost tracemaker with a trace fossil from the Green River Formation, while also interpreting the size and relative age of the tracemaker. The normal feeding and swimming behaviors indicated by the trace fossil indicate temporarily oxygenated benthic conditions in the deepest part of Fossil Lake, counter to most paleoecological interpretations of this deposit. Lastly, our spatial and mathematical analyses significantly update and advance previous approaches to the study of teleost trace fossils

Evolutionary Divergence in Brain Size between Migratory and Resident Birds

Despite important recent progress in our understanding of brain evolution, controversy remains regarding the evolutionary forces that have driven its enormous diversification in size. Here, we report that in passerine birds, migratory species tend to have brains that are substantially smaller (relative to body size) than those of resident species, confirming and generalizing previous studies. Phylogenetic reconstructions based on Bayesian Markov chain methods suggest an evolutionary scenario in which some large brained tropical passerines that invaded more seasonal regions evolved migratory behavior and migration itself selected for smaller brain size. Selection for smaller brains in migratory birds may arise from the energetic and developmental costs associated with a highly mobile life cycle, a possibility that is supported by a path analysis. Nevertheless, an important fraction (over 68%) of the correlation between brain mass and migratory distance comes from a direct effect of migration on brain size, perhaps reflecting costs associated with cognitive functions that have become less necessary in migratory species. Overall, our results highlight the importance of retrospective analyses in identifying selective pressures that have shaped brain evolution, and indicate that when it comes to the brain, larger is not always better

3-D Modelling of Megaloolithid Clutches: Insights about Nest Construction and Dinosaur Behaviour

BACKGROUND: Megaloolithid eggs have long been associated with sauropod dinosaurs. Despite their extensive and worldwide fossil record, interpretations of egg size and shape, clutch morphology, and incubation strategy vary. The Pinyes locality in the Upper Cretaceous Tremp Formation in the southern Pyrenees, Catalonia provides new information for addressing these issues. Nine horizons containing Megaloolithus siruguei clutches are exposed near the village of Coll de Nargó. Tectonic deformation in the study area strongly influenced egg size and shape, which could potentially lead to misinterpretation of reproductive biology if 2D and 3D maps are not corrected for bed dip that results from tectonism. METHODOLOGY/FINDINGS: Detailed taphonomic study and three-dimensional modelling of fossil eggs show that intact M. siruguei clutches contained 20-28 eggs, which is substantially larger than commonly reported from Europe and India. Linear and grouped eggs occur in three superimposed levels and form an asymmetric, elongate, bowl-shaped profile in lateral view. Computed tomography data support previous interpretations that the eggs hatched within the substrate. Megaloolithid clutch sizes reported from other European and Indian localities are typically less than 15 eggs; however, these clutches often include linear or grouped eggs that resemble those of the larger Pinyes clutches and may reflect preservation of incomplete clutches. CONCLUSIONS/SIGNIFICANCE: We propose that 25 eggs represent a typical megaloolithid clutch size and smaller egg clusters that display linear or grouped egg arrangements reported at Pinyes and other localities may represent eroded remnants of larger clutches. The similarity of megaloolithid clutch morphology from localities worldwide strongly suggests common reproductive behaviour. The distinct clutch geometry at Pinyes and other localities likely resulted from the asymmetrical, inclined, and laterally compressed titanosaur pes unguals of the female, using the hind foot for scratch-digging during nest excavation

Cartilaginous Epiphyses in Extant Archosaurs and Their Implications for Reconstructing Limb Function in Dinosaurs

Extinct archosaurs, including many non-avian dinosaurs, exhibit relatively simply shaped condylar regions in their appendicular bones, suggesting potentially large amounts of unpreserved epiphyseal (articular) cartilage. This “lost anatomy” is often underappreciated such that the ends of bones are typically considered to be the joint surfaces, potentially having a major impact on functional interpretation. Extant alligators and birds were used to establish an objective basis for inferences about cartilaginous articular structures in such extinct archosaur clades as non-avian dinosaurs. Limb elements of alligators, ostriches, and other birds were dissected, disarticulated, and defleshed. Lengths and condylar shapes of elements with intact epiphyses were measured. Limbs were subsequently completely skeletonized and the measurements repeated. Removal of cartilaginous condylar regions resulted in statistically significant changes in element length and condylar breadth. Moreover, there was marked loss of those cartilaginous structures responsible for joint architecture and congruence. Compared to alligators, birds showed less dramatic, but still significant changes. Condylar morphologies of dinosaur limb bones suggest that most non-coelurosaurian clades possessed large cartilaginous epiphyses that relied on the maintenance of vascular channels that are otherwise eliminated early in ontogeny in smaller-bodied tetrapods. A sensitivity analysis using cartilage correction factors (CCFs) obtained from extant taxa indicates that whereas the presence of cartilaginous epiphyses only moderately increases estimates of dinosaur height and speed, it has important implications for our ability to infer joint morphology, posture, and the complicated functional movements in the limbs of many extinct archosaurs. Evidence suggests that the sizes of sauropod epiphyseal cartilages surpassed those of alligators, which account for at least 10% of hindlimb length. These data suggest that large cartilaginous epiphyses were widely distributed among non-avian archosaurs and must be considered when making inferences about locomotor functional morphology in fossil taxa

Didactyl Tracks of Paravian Theropods (Maniraptora) from the ?Middle Jurassic of Africa

BACKGROUND: A new dinosaur tracksite from ?Middle Jurassic sediments of the Irhazer Group on the plains of Agadez (Rep. Niger, northwest Africa) revealed extraordinarily well preserved didactyl tracks of a digitigrade bipedal trackmaker. The distinct morphology of the pes imprints indicates a theropod trackmaker from a paravian maniraptoran closely related to birds. METHODOLOGY/PRINCIPAL FINDINGS: The early age and the morphological traits of the tracks allow for description of the new ichnotaxon Paravipus didactyloides. A total of 120 tracks are assigned to 5 individual trackways. The 'medium-sized' tracks with an average footprint length of 27.5 cm and footprint width of 23.1 cm are deeply imprinted into the track bearing sandstone. CONCLUSIONS/SIGNIFICANCE: A comparison with other didactyl tracks gives new insights into the foot morphology of advanced maniraptoran theropods and contributes to knowledge of their evolutionary history. The new ichnotaxon takes an important position in the ichnological fossil record of Gondwana and the mid-Jurassic biota worldwide, because it is among the earliest known records of paravian maniraptorans and of didactyl theropod tracks from Africa

Cranial Growth and Variation in Edmontosaurs (Dinosauria: Hadrosauridae): Implications for Latest Cretaceous Megaherbivore Diversity in North America

The well-sampled Late Cretaceous fossil record of North America remains the only high-resolution dataset for evaluating patterns of dinosaur diversity leading up to the terminal Cretaceous extinction event. Hadrosaurine hadrosaurids (Dinosauria: Ornithopoda) closely related to Edmontosaurus are among the most common megaherbivores in latest Campanian and Maastrichtian deposits of western North America. However, interpretations of edmontosaur species richness and biostratigraphy have been in constant flux for almost three decades, although the clade is generally thought to have undergone a radiation in the late Maastrichtian. We address the issue of edmontosaur diversity for the first time using rigorous morphometric analyses of virtually all known complete edmontosaur skulls. Results suggest only two valid species, Edmontosaurus regalis from the late Campanian, and E. annectens from the late Maastrichtian, with previously named taxa, including the controversial Anatotitan copei, erected on hypothesized transitional morphologies associated with ontogenetic size increase and allometric growth. A revision of North American hadrosaurid taxa suggests a decrease in both hadrosaurid diversity and disparity from the early to late Maastrichtian, a pattern likely also present in ceratopsid dinosaurs. A decline in the disparity of dominant megaherbivores in the latest Maastrichtian interval supports the hypothesis that dinosaur diversity decreased immediately preceding the end Cretaceous extinction event

Dinosaur Speed Demon: The Caudal Musculature of Carnotaurus sastrei and Implications for the Evolution of South American Abelisaurids

In the South American abelisaurids Carnotaurus sastrei, Aucasaurus garridoi, and, to a lesser extent Skorpiovenator bustingorryi, the anterior caudal ribs project at a high dorsolateral inclination and have interlocking lateral tips. This unique morphology facilitated the expansion of the caudal hypaxial musculature at the expense of the epaxial musculature. Distinct ridges on the ventrolateral surfaces of the caudal ribs of Aucasaurus garridoi are interpreted as attachment scars from the intra caudofemoralis/ilio-ischiocaudalis septa, and confirm that the M. caudofemoralis of advanced South American abelisaurids originated from a portion of the caudal ribs. Digital muscle models indicate that, relative to its overall body size, Carnotaurus sastrei had a substantially larger M. caudofemoralis than any other theropod yet studied. In most non-avian theropods, as in many extant sauropsids, the M. caudofemoralis served as the primary femoral retractor muscle during the locomotive power stroke. This large investment in the M. caudofemoralis suggests that Carnotaurus sastrei had the potential for great cursorial abilities, particularly short-burst sprinting. However, the tightly interlocking morphology of the anterior caudal vertebrae implies a reduced ability to make tight turns. Examination of these vertebral traits in evolutionary context reveals a progressive sequence of increasing caudofemoral mass and tail rigidity among the Abelisauridae of South America

A New Troodontid Theropod Dinosaur from the Lower Cretaceous of Utah

BACKGROUND: The theropod dinosaur family Troodontidae is known from the Upper Jurassic, Lower Cretaceous, and Upper Cretaceous of Asia and from the Upper Jurassic and Upper Cretaceous of North America. Before now no undisputed troodontids from North America have been reported from the Early Cretaceous. METHODOLOGY/PRINCIPAL FINDINGS: Herein we describe a theropod maxilla from the Lower Cretaceous Cedar Mountain Formation of Utah and perform a phylogenetic analysis to determine its phylogenetic position. The specimen is distinctive enough to assign to a new genus and species, Geminiraptor suarezarum. Phylogenetic analysis places G. suarezarum within Troodontidae in an unresolved polytomy with Mei, Byronosaurus, Sinornithoides, Sinusonasus, and Troodon+(Saurornithoides+Zanabazar). Geminiraptor suarezarum uniquely exhibits extreme pneumatic inflation of the maxilla internal to the antorbital fossa such that the anterior maxilla has a triangular cross-section. Unlike troodontids more closely related to Troodon, G. suarezarum exhibits bony septa between the dental alveoli and a promaxillary foramen that is visible in lateral view. CONCLUSIONS/SIGNIFICANCE: This is the first report of a North American troodontid from the Lower Cretaceous. It therefore contributes to a fuller understanding of troodontid biogeography through time. It also adds to the known dinosaurian fauna of the Cedar Mountain Formation

Intra-trackway morphological variations due to substrate consistency: the El Frontal dinosaur tracksite (Lower Cretaceous, Spain).

An ichnological and sedimentological study of the El Frontal dinosaur tracksite (Early Cretaceous, Cameros basin, Soria, Spain) highlights the pronounced intra-trackway variation found in track morphologies of four theropod trackways. Photogrammetric 3D digital models revealed various and distinct intra-trackway morphotypes, which reflect changes in footprint parameters such as the pace length, the track length, depth, and height of displacement rims. Sedimentological analyses suggest that the original substrate was non-homogenous due to lateral changes in adjoining microfacies. Multidata analyses indicate that morphological differences in these deep and shallow tracks represent a part of a continuum of track morphologies and geometries produced by a gradient of substrate consistencies across the site. This implies that the large range of track morphologies at this site resulted from similar trackmakers crossing variable facies. The trackways at the El Frontal site present an exemplary case of how track morphology, and consequently potential ichnotaxa, can vary, even when produced by a single trackmaker

Growth Dynamics of Australia's Polar Dinosaurs

Analysis of bone microstructure in ornithopod and theropod dinosaurs from Victoria, Australia, documents ontogenetic changes, providing insight into the dinosaurs' successful habitation of Cretaceous Antarctic environments. Woven-fibered bone tissue in the smallest specimens indicates rapid growth rates during early ontogeny. Later ontogeny is marked by parallel-fibered tissue, suggesting reduced growth rates approaching skeletal maturity. Bone microstructure similarities between the ornithopods and theropods, including the presence of LAGs in each group, suggest there is no osteohistologic evidence supporting the hypothesis that polar theropods hibernated seasonally. Results instead suggest high-latitude dinosaurs had growth trajectories similar to their lower-latitude relatives and thus, rapid early ontogenetic growth and the cyclical suspensions of growth inherent in the theropod and ornithopod lineages enabled them to successfully exploit polar regions