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Here I assess the use of osteoderms in systematics with comparative material from fossil and extant tetrapod taxa. Putative differences among three groups (ankylosaurid, nodosaurid, and polacanthid) were evaluated. Archosaur osteoderms have cortices surrounding a cancellous core. Ankylosaurs are united by a superficial cortex distinguishable from the core, lack of Sharpeys fibers, and mineralized structural fibers. Nododsaurids lack a deep cortex and have dense superficial cortical fibres. Ankylosaurid osteoderms are thinner than those of other ankylosaurs. Polacanthids (and some nodosaurids and ankylosaurids) have a cancellous core. Cortical thickness overlaps among groups, so a thick cortex is not diagnostic for polacanthids. Modified elements diverge histologically from the primitive condition for specific functions. Haversian bone in the core is not indicative of any group. Some shapes and superficial textures are diagnostic for specific taxa. Parsimony analyses show support for the Ankylosauridae and Nodosauridae, but not a monophyletic polacanthid clade. / Systematics and Evolution

Understanding the changes undergone during the life of an organism is often crucial to properly interpreting the evolutionary history of a group. For extinct organisms, this process can only be directly studied through growth series of fossils representing individuals at different stages of maturity. The growth patterns of the earliest dinosaurs (230–190 million years ago), in particular the morphological changes undergone during the life history of an individual (i.e., ontogeny) is poorly understood. To tackle this problem, I studied the changes undergone during growth of two early theropod dinosaurs, Coelophysis bauri and Megapnosaurus rhodesiensis. To reconstruct the growth of these dinosaurs I used ontogenetic sequence analysis (OSA). I found that, unlike living birds, early dinosaurs possessed an extremely high amount of intraspecific variation in growth. This variation had been previously interpreted as sexual difference; however, I found no evidence of this. Because this variation is widespread among early dinosaurs and their relatives, I hypothesize that this is the ancestral condition of dinosaurian growth, and that this was lost along the evolution to birds. These ontogenetic events are conserved through evolution, and I used this to assess the maturity of large Triassic theropods: I suggest that all known large-bodied Triassic theropods were still growing rapidly at death, and that the maximum body size of Triassic theropods was higher than previously supposed. Theropods were large before the end Triassic mass extinction, unlike what has been previously hypothesized. / Master of Science

Thesis (M.S.)--University of Texas at El Paso, 2007. / Title from title screen. Vita. CD-ROM. Includes bibliographical references. Also available online.

Longitudinal profile, channel cross-section geometry, and depositional patterns of the Green River in its course through the eastern Uinta Mountains are each strongly influenced by river -level geology and tributary sediment delivery processes. We surveyed channel cross sections at 1-km intervals, mapped surficial geology, and measured size and characteristics of bed material in order to evaluate the geomorphic organization of the 70- km study reach. Canyon reaches that are of high gradient and narrow channel geometry are associated with the most resistant lithologies exposed at river level and the most frequent occurrences of tributary debris fans. Meandering reaches that are characterized by low gradient and wide channel geometry are associated. with river-level lithology that is of moderate to low resistance and very low debris fan frequency. The channel is in contact with bedrock or talus along only 42 percent of the bank length in canyon reaches and there is an alluvial fill of at least 12 m that separates the channel bed from bedrock at three borehole sites. The influence of lithology primarily operates through the presence of resistant boulders in debris fans that are delivered by debris flows from steep tributaries. The depositional settings created by debris fans consist of (1) channel-margin deposits in the backwater above the debris fan, (2) eddy bars in the zone of recirculating flow below the constriction, and (3) expansion gravel bars in the expansion below the zone of recirculating flow. These fan-eddy complexes are the storage location of about 70 percent. by area, of all fine- and coarse-grained alluvium contained within the canyons above the low-water stage. Immediately adjacent meandering reaches contain an order of magnitude more alluvium by area but have no debris fan-created depositional settings. This study also describes the flood-plain and terrace stratigraphy of the Green River in the eastern Uinta Mountains and changes due to the operations of Flaming Gorge Dam, upstream from the study area. These landforms are vertically aggrading deposits that are longitudinally correlative throughout the 65-km study reach. The suite of surfaces identified includes a terrace that is inundated by rare pre- or post-dam floods, an intermediate bench that is inundated by rare post-dam floods, and a post-dam flood plain that is inundated by the post-dam mean annual flood. Analysis of historical photographs in the study reach shows that both the intermediate bench and post-dam flood plain are landforms that were not present in any of the 6 years for which photographs were examined between 1871 and 1954. Photographic replications also show that gravel bars consisting of bare gravel in 1922 and earlier photographs are now covered by fine-grained alluvium and vegetation . Decreased gravel-bar mobility is indicated by estimates of critical and average boundary shear stress. Comprehensive surficial geologic mapping of the study area indicates that the bankfull channel has decreased in width by an average of about 20 percent.

Longitudinal profile , channel cross-section geometry, and depositional patterns of the Green River in its course through the eastern Uinta Mountains are each strongly influenced by river-level geology and tributary sediment delivery processes. We surveyed channel cross sections at 1-km intervals, mapped surficial geology , and measured size and characteristics of bed material in order to evaluate the geomorphic organization of the 70- km study reach . Canyon reaches that are of high gradient and narrow channel geometry are associated with the most resistant lithologies exposed at river level and the most frequent occurrences of tributary debris fans. Meandering reaches that are characterized by low gradient and wide channel geometry are associated with river-level lithology that is of moderate to low resistance and very low debris fan frequency. The channel is in contact with bedrock or talus along only 42 percent of the bank length in canyon reaches and there is an alluvial fill of at least 12 m that separates the channe l bed from bedrock at three borehole sites. The influence of lithology primarily operates through the presence of resistant boulders in debris fans that are delivered by debris flows from steep tributaries. The depositional settings created by debris fans consist of (1) channel-margin deposits in the backwater above the debris fan, (2) eddy bars in the zone of recirculating flow below the constriction, and (3) expansion gravel bars in the expansion below the zone of recirculating flow. These fan-eddy complexes are the storage location of about 70 percent, by area, of all fine- and coarse-grained alluvium contained within the canyons above the low-water stage. Immediately adjacent meandering reaches contain an order of magnitude more alluvium by area but have no debris fan-created depositional settings. This study also describes the flood-plain and terrace stratigraphy of the Green River in the eastern Uinta Mountains and changes due to the operations of Aarning Gorge Dam, upstream from the study area. These landforms are vertically aggrading deposits that are longiuidinally correlative throughout the 65-km study reach. The suite of surfaces identified includes a terrace that is inundated by rare pre- or post-dam floods, an intermediate bench that is inundated by rare post-dam floods, and a post-dam flood plain that is inundated by the post-dam mean annual flood. Analysis of historical photographs in the study reach shows that both the intermediate bench and post-dam flood plain are landforms that were not present in any of the 6 years for which photographs were examined between 1871 and 1954. Photographic replications also show that gravel bars consisting of bare gravel in 1922 and earlier photographs are now covered by fine-grained alluvium and vegetation. Decreased gravel-bar mobility is indicated by estimates of critical and average boundary shear stress. Comprehensive surficial geologic mapping of the study area indicates that the bankfull channel has decreased in width by an average of about 20 percent.

Patterns of organismal development—including embryonic morphogenesis, postnatal ontogenetic trends, and growth—are fundamental to understanding the evolution of phenotypes. Consideration of these processes is necessary to construct a holistic and rigorous account of morphological evolution. Therefore, the integration of both the fossil record (actualistic observations of changes in morphology through Earth history) and observations of extant clades (comparative patterns of pre- and postnatal development) is required to study the evolution of form across deep time. The pelvic complex plays a central role in tetrapod locomotion with the exception of taxa lacking hindlimbs (e.g., whales, snakes, caecilians). This complex, composed of the pelvic girdle (ilia, ischia, and pubes) and the sacrum (vertebrae articulated with the ilia via sacral ribs) is in some aspects highly conserved through amniote evolution: all terrestrial amniotes with functional hindlimbs retain the same bones of the pelvic girdle in roughly the same configuration with each other and a sacrum consisting of at least two vertebrae, the ancestral amniote condition. Despite the retention of this basic plan, the pelvic complex of many groups is characterized by extreme deviations from the ancestral condition. Therefore, the evolution of the pelvis and sacrum across amniotes—especially among archosaurs, which possess highly derived variations of the pelvic complex in response to the evolution of novel baupläne—provides an excellent case study to explore the influence of developmental patterns on the evolution of morphology. The first chapter of my dissertation reviews the methods that have been used to study reptilian growth and maturity in the fossil record. I clarify often ambiguous terminology and highlight clade-specific best practices for assessing growth in extinct reptiles. The second chapter uses the dicynodont sacrum as a case study to explore the evolution of novel morphologies along the normally constrained synapsid lineage that are convergent on several archosaurian innovations, integrating comparative anatomy, comparative phylogenetic methods, and developmental biology. The third chapter utilizes a new technique (modified CLARITY protocol) for imaging the soft tissues of developing embryos to explore the morphogenesis of the avian pelvic complex, finding that the derived avian condition arrives late in development, with plesiomorphic—more typically "dinosaurian"—character states being present at the outset of musculoskeletal development. / Doctor of Philosophy / Every animal's anatomy is determined by the developmental processes they undergo during their life history. Because of this, understanding these developmental processes is a key part of determining how animals have evolved into the many disparate forms we see today. Conversely, it is only through the fossil record that the actual history of anatomical evolution can be observed, so a holistic account of the pattern and process underlying evolution must integrate both developmental biology and paleontology. The pelvic complex—the hips and the vertebrae articulated with the hips—plays a fundamental role in how all nearly land vertebrates move (with the exception of legless groups). In some ways, the pelvic complex is very similar across all groups of mammals, reptiles, and their extinct close relatives: all retain the same three bones of the pelvis in roughly the same configuration with each other, and all possess a sacrum consisting of at least two vertebrae, the ancestral condition. Despite the retention of this basic plan, the pelvic complex of many groups is characterized by extreme deviations from the ancestral condition, with the three pelvic bones having highly disparate shapes and proportions. Therefore, the evolution of the pelvis and sacrum across mammals, reptiles, and their extinct relatives—especially among archosaurs (the reptilian group containing crocodylians and birds), which possess highly derived variations of the pelvic complex in response to the evolution of novel body plans—provides an excellent case study to explore the influence of developmental patterns on the evolution of animal form. The first chapter of my dissertation reviews the methods that have been used to study reptilian growth and maturity in the fossil record. I clarify often ambiguous terminology and highlight group-specific best practices for assessing growth in extinct reptiles. The second chapter uses the evolution of the sacrum among extinct mammal relatives as a case study to explore the evolution of novel anatomy along the normally conservative mammalian lineage, integrating comparative anatomy, evolutionary statistics, and developmental biology. The third chapter utilizes a new technique for imaging the soft tissues of developing embryos to explore the origins of the highly unusual pelvic complex in birds, finding that the distinctive bird anatomy arrives late in development, with ancestral—more typically "dinosaurian"—features being present at early-stage embryos of birds.

Fossil tracks represent the only direct record of behaviour and locomotion of extinct animals. A computer model using finite element analysis (FEA) has been developed to simulate vertebrate track formation in cohesive substrates. This model has been designed for, and successfully run on, high performance computing (HPC) resources. A number of individual studies were carried out using the computer model to simulate both abstract indenters and virtual dinosaur autopodia. In addition to the simulation studies, two fossil tracks were described, including the first report of bird tracks at the Mammoth Site of Hot Springs, South Dakota (USA) and a re-description of a 'dinosaur tail drag' as the trace of a crocodilian. Using the computer model, it has been shown that in a wet, soft mud the indentation of a non-webbed virtual tridactyl foot created a resultant track with features analogous to 'webbing' between digits. This 'webbing' was a function of sediment deformation and subsequent failure in 3D, specific to rheology. Apparent webbing impressions were clearly developed only within a limited range of sediment conditions and pedal geometry. Indenter (pedal) geometry and morphology affect track depth independently of substrate and loading parameters. More complex morphologies interact with the cohesive substrate creating a lower effective load than that applied. In non-cohesive substrates such as sand, this effect is reversed, and it is the more compact morphologies that indent to a lesser degree. Virtual sauropod tracks were modelled, based on published soft tissue reconstructions of autopodia anatomy, and published mass/centre of mass estimates. It was shown that foot morphology and differential loading between fore- and hind- limbs leads to a range of substrates in which only the manus or pes are able to generate tracks. This offers a new mechanism for the formation of manus-only sauropod trackways, previously interpreted as having been made by swimming dinosaurs. A series of tracks were simulated using input data (loads, pedal morphologies) from four different dinosaurs (Brachiosaurus, Tyrannosaurus, Struthiomimus, and Edmontosaurus). The cohesive substrates used displayed a 'Goldilocks' effect, allowing the formation for tracks only for a very limited range of loads for any given foot. In addition, there was a strong bias toward larger animals, both in homogeneous and theoretically heterogeneous substrates. These findings imply that interpretations from track assemblages must consider that only a small proportion of the total fauna present may be recorded as a track assemblage due to substrate properties. The use of FEA to simulate dinosaur track formation has been shown to be successful, and offers a number of advantages over physical modelling including; consistency between experiments, specific control over input variables, rapid undertaking of repeatable experiments, and the ability to view subsurface deformation non-destructively. It is hoped that this work will lead to an increased interest in modelling tracks, and offer a quantitative method for studying fossil tracks.

The Cedar Mountain Formation contains the most diverse record of Early Cretaceous dinosaurs in the western hemisphere. However, analyses of its faunas have been hindered because 1) most taxa are based on incomplete/fragmentary materials or incomplete descriptions, 2) most sites and some horizons preserve few taxa, and 3) the stratigraphy and geochronology are poorly understood. To help resolve these stratigraphic and correlation problems U-Pb LA-ICP-MS detrital zircon ages were obtained at significant sites and horizons. These dates indicate all sites at or near the base of the formation are no older than 122 to 124 Ma, thus all basal stratigraphic packages are time equivalent. Detrital zircons coarsely bracket the temporal span of the Ruby Ranch Member between about 115 Ma to 111 Ma while the base of the Mussentuchit Member is dated between 108 to 104 Ma and the top of the member is Cenomanian in age. Multivariate analyses utilizing Simpson and Raup-Crick similarity index and pair-group moving algorithms reveal that formationfs faunas fall into two groups. These groups are compared statistically with European, Asian, and Morrison faunas. Results indicate (1) that there is no close relationship between the Yellow Cat fauna and the Morrison Formation fauna and (2) corroborate long-standing hypotheses that the Yellow Cat fauna has European ties and the Mussentuchit fauna has Asian ties. Detrital zircon LA-ICP-MS U-Pb ages were used in this study to approximate the time of deposition of strata because volcanic ashes are rarely preserved in the formation. The ability to select the youngest crystals in a sample prior to applying analytical methods could substantially reduce the number of crystals and cost required to obtain these dates. To this end, the hypothesis that the most pristine, unabraded crystals should be younger than abraded crystals was tested by imaging detrital zircons via SEM, ranking the crystals by the degree of abrasion, and determining their ages. Results of this study partly corroborate the hypothesis in that there is a correlation between the degree of abrasion and ages – obviously abraded crystals are most likely the oldest while pristine to slightly abraded crystals are usually the youngest in a given sample.

Nature is full of mysterious creatures which fascinate and spark imagination. In my final project, What Crawls Beneath, I take a closer look at what drives my interest in creatures that simultaneously attract and repel. Drawing on inspiration from parasites and dinosaurs, I was able to create a piece that represents the danger and beauty that nature so masterfully brings together. The importance of process is discussed as I consider the traditional methods of working with clay and how they affect the outcome of the artwork.