VTechWorks Repository :: Browsing by Author "Nesbitt, Sterling J."

Browsing by Author "Nesbitt, Sterling J."

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Are 'exceptionally' preserved skeletal fossils necessarily exceptional chemically and cytologically?
Korneisel, Dana Elaine (Virginia Tech, 2019-09-19)
At the macroscopic scale, vertebrate fossils are considered exceptional when non-biomineralized (soft) tissues are preserved. Histologically, high quality is defined by trueness to original shape of a bone, preservation of fine details (e.g. canaliculi), and presence or absence of matrix material in void spaces. Some fossils are hypothesized to preserve cells and durable organelles. Traditionally, cytological details and biomolecular remains have been sought in exceptional fossils. Durable cytological features such as melanosomes do appear to follow feather preservation, but traditionally exceptional fossils are not necessarily exceptional on a microscopic scale. Here, we analyze a feathered dinosaur specimen from the Jehol Lagerstätte to assess claims of blood cell preservation and the state of potential biomolecular preservation. Beipiaosaurus inexpectus is a fairly complete specimen with preserved feathers. Though crushed, fine details in thin section are prevalent. Using Raman spectroscopy, Energy Dispersive X-ray Spectrometry, and Time-of-Flight Secondary Ion Mass Spectroscopy we found no evidence of exceptional molecular preservation. Instead, we found evidence that the vasculature, once hypothesized to contain preserved red blood cells, is filled with clay minerals, with the purported cells chemically indistinguishable from materials of other shapes infilling the vessels. Despite yielding exceptional fossils, the preservational environment of the Jehol biota does not necessarily preserve exceptional details cytologically or biomolecularly. Consequently, we conclude that a systematic approach to biomolecular and cytological preservation studies should rely on traits other than classic exceptional preservation.
The axial skeleton of Poposaurus langstoni (Pseudosuchia: Poposauroidea) and its implications for accessory intervertebral articulation evolution in pseudosuchian archosaurs
Stefanic, Candice M.; Nesbitt, Sterling J. (PeerJ, 2018-02-14)
Dinosaurs and their close relatives grew to sizes larger than any other terrestrial animal in the history of life on Earth, and many enormous dinosaurs (e.g., Diplodocus, Spinosaurus, Tyrannosaurus) have accessory intervertebral articulations that have been suggested to support these large body sizes. Some pseudosuchian archosaurs have been reported to have these articulations as well, but few have been characterized in these taxa because of a lower abundance of complete, threedimensional pseudosuchian vertebral material in relation to dinosaurs. We describe the axial column of the large (similar to 4-5 m) poposauroid pseudosuchian Poposaurus langstoni from the Upper Triassic of Texas (TMM Locality 31025 of the Otis Chalk localities; Dockum Group, Howard County, TX, USA). P. langstoni was originally named from pelvic girdle elements and vertebrae; here we describe newly discovered and prepared presacral vertebrae and a presacral rib from the original excavation of the holotype in the 1940s. The well-preserved vertebrae have well-defined vertebral laminae and clear hyposphene-hypantrum intervertebral articulations, character states mentioned in pseudosuchians but rarely described. The new material demonstrates variation present in the hyposphene-hypantrum articulation through the vertebral column. We compared these morphologies to other pseudosuchians with and without the hyposphene-hypantrum articulation. Based on these careful comparisons, we provide an explicit definition for the hyposphene-hypantrum articulation applicable across Archosauria. Within Pseudosuchia, we find the hyposphene-hypantrum appeared independently in the clade at least twice, but we also see the loss of these structures in clades that had them ancestrally. Furthermore, we found the presence of large body sizes (femoral lengths >similar to 300 mm) and the presence of the hyposphene-hypantrum is correlated in most non-crocodylomorph pseudosuchian archosaurs with a few exceptions. This result suggests that the presence of the hyposphene-hypantrum is controlled by the increases and decreases in body size and not strictly inheritance.
Contributions to Exceptional Fossil Preservation
Muscente, Anthony Drew (Virginia Tech, 2016-04-21)
Exceptionally preserved fossils—or fossils preserved with remains of originally non-biomineralized (i.e. soft) tissues—constitute a key resource for investigating the history of the biosphere. In comparison to fossils of biomineralized skeletal elements, which represent the majority of the fossil record but only a fraction of the total diversity that existed in the past, exceptionally preserved fossils are comparatively rare because soft tissues are rapidly destroyed in typical depositional environments. Assemblages of such fossils, nonetheless, have received special attention among scientists in multiple fields of Earth and life sciences because they represent relatively 'complete' windows to past life. Through such windows, researchers are able to reconstruct original biological features (e.g. soft tissue anatomies) of extinct organisms and to describe the structures and compositions of ancient soft-bodied paleocommunities. To accomplish these goals, however, researchers must incorporate background information regarding the pre- and post-burial histories of exceptionally preserved fossils. In this context, my dissertation focuses on the environmental settings, diagenetic conditions, geomicrobiological activities, and weathering processes, which influence the conservation of original biological features within exceptionally preserved fossils and control their occurrences in time and space. An improved understanding of these critical factors involved in exceptional fossil preservation will ultimately our advance our knowledge regarding the history of the biosphere and the Earth system as a whole. Each chapter of original research in this dissertation includes an innovative and distinct approach for studying exceptional fossil preservation. The second chapter describes environmental and geologic overprints in the exceptional fossil record, as revealed by a comprehensive statistical meta-analysis of a global dataset of exceptionally preserved fossil assemblages. Moving from global to specimen-based perspectives, the second and third chapters focus on minerals (products of geomicrobioloigcal, diagenetic, and weathering processes) and carbonaceous materials replicating exceptionally preserved fossils. The third chapter examines the causes of preservational variations observed among organophosphatic tubular shelly Sphenothallus fossils in the lower Cambrian of South China using an experimental approach. (Although Sphenothallus is not an exceptionally preserved fossil sensu stricto, its conservation of original organic matrix tissues in South China provides key insights into the preservation of carbonaceous material within fossils.) Lastly, the fourth chapter presents data acquired using various in situ nanoscale analytical techniques to test the hypothesis that microstructures within exceptionally preserved microfossils of the Ediacaran Doushantuo Formation of South China are some of the oldest putative cylindrical siliceous demosponge spicules in the fossil record. Collectively, these chapters describe environmental, authigenic, diagenetic, and weathering processes that affect exceptional fossil preservation, and highlight innovative methods and approaches for testing major paleobiologic and geobiologic hypotheses regarding exceptionally preserved fossils.
A diverse diapsid tooth assemblage from the Early Triassic (Driefontein locality, South Africa) records the recovery of diapsids following the end-Permian mass extinction
Hoffman, Devin K.; Hancox, John P.; Nesbitt, Sterling J. (Public Library of Science, 2023-05)
Mass extinctions change the trajectory of evolution and restructure ecosystems. The largest mass extinction, the end-Permian, is a particularly interesting case due to the hypothesized delay in the recovery of global ecosystems, where total trophic level recovery is not thought to have occurred until 5-9 million years after the extinction event. Diapsids, especially archosauromorphs, play an important role in this recovery, filling niches left vacant by therapsids and anapsids. However, the nature of lineage and ecological diversification of diapsids is obscured by the limited number of continuous, well-dated stratigraphic sections at the Permian-Triassic boundary and continuing through the first half of the Triassic. The Karoo Basin of South Africa is one such record, and particularly the late Early Triassic (Olenekian) Driefontein locality fills this gap in the diapsid fossil record. We collected a total of 102 teeth of which 81 are identified as diapsids and the remaining 21 as identified as temnospondyls. From the sample, seven distinct tooth morphotypes of diapsids are recognized, six of which are new to the locality. We used a combination of linear measurements, 3D geomorphometrics, and nMDS ordination to compare these morphotypes and made inferences about their possible diets. Although the morphotypes are readily differentiated in nMDS, the overall morphological disparity is low, and we infer five morphotypes are faunivorous with the other two potentially omnivorous or piscivorous based on their morphological similarities with dentitions from extant diapsids, demonstrating an unsampled taxonomic and ecological diversity of diapsids in the Early Triassic based on teeth. Although ecological specialization at Driefontein may be low, it records a diversity of diapsid taxa, specifically of archosauromorph lineages.
The earliest equatorial record of frogs from the Late Triassic of Arizona
Stocker, Michelle R.; Nesbitt, Sterling J.; Kligman, Ben T.; Paluh, Daniel J.; Marsh, Adam D.; Blackburn, David C.; Parker, William G. (Royal Society Publishing, 2019-02-01)
Crown-group frogs (Anura) originated over 200 Ma according to molecular phylogenetic analyses, though only a few fossils from high latitudes chronicle the first approximately 60 Myr of frog evolution and distribution. We report fossils that represent both the first Late Triassic and the earliest equatorial record of Salientia, the group that includes stem and crown-frogs. These small fossils consist of complete and partial ilia with anteriorly directed, elongate and distally hollow iliac blades. These features of these ilia, including the lack of a prominent dorsal protuberance and a shaft that is much longer than the acetabular region, suggest a closer affinity to crown-group Anura than to Early Triassic stem anurans Triadobatrachus from Madagascar and Czatkobatrachus from Poland, both high-latitude records. The new fossils demonstrate that crown anurans may have been present in the Late Triassic equatorial region of Pangea. Furthermore, the presence of Early Jurassic anurans in the same stratigraphic sequence (Prosalirus bitis from the Kayenta Formation) suggests that anurans survived the climatic aridification of this region in the early Mesozoic. These fossils highlight the importance of the targeted collection of microfossils and provide further evidence for the presence of crown-group representatives of terrestrial vertebrates prior to the end-Triassic extinction.
Early crocodylomorph increases top tier predator diversity during rise of dinosaurs
Zanno, Lindsay E.; Drymala, Susan; Nesbitt, Sterling J.; Schneider, Vincent P. (Springer Nature, 2015-03-19)
Triassic predatory guild evolution reflects a period of ecological flux spurred by the catastrophic end-Permian mass extinction and terminating with the global ecological dominance of dinosaurs in the early Jurassic. In responding to this dynamic ecospace, terrestrial predator diversity attained new levels, prompting unique trophic webs with a seeming overabundance of carnivorous taxa and the evolution of entirely new predatory clades. Key among these was Crocodylomorpha, the largest living reptiles and only one of two archosaurian lineages that survive to the present day. In contrast to their existing role as top, semi-aquatic predators, the earliest crocodylomorphs were generally small-bodied, terrestrial faunivores, occupying subsidiary (meso) predator roles. Here we describe Carnufex carolinensis a new, unexpectedly large-bodied taxon with a slender and ornamented skull from the Carnian Pekin Formation (similar to 231 Ma), representing one of the oldest and earliest diverging crocodylomorphs described to date. Carnufex bridges a problematic gap in the early evolution of pseudosuchians by spanning key transitions in bauplan evolution and body mass near the origin of Crocodylomorpha. With a skull length of >50 cm, the new taxon documents a rare instance of crocodylomorphs ascending to top-tier predator guilds in the equatorial regions of Pangea prior to the dominance of dinosaurs.
The evolution and role of the hyposphene-hypantrum articulation in Archosauria: phylogeny, size and/or mechanics?
Stefanic, Candice M.; Nesbitt, Sterling J. (Royal Society, 2019-08-22)
Living members of Archosauria, the reptile clade containing Crocodylia and Aves, have a wide range of skeletal morphologies, ecologies and body size. The range of body size greatly increases when extinct archosaurs are included, because extinct Archosauria includes the largest members of any terrestrial vertebrate group (e.g. 70-tonne titanosaurs, 20-tonne theropods). Archosaurs evolved various skeletal adaptations for large body size, but these adaptations varied among clades and did not always appear consistently with body size or ecology. Modification of intervertebral articulations, specifically the presence of a hyposphene-hypantrum articulation between trunk vertebrae, occurs in a variety of extinct archosaurs (e.g. non-avian dinosaurs, pseudosuchians). We surveyed the phylogenetic distribution of the hyposphene-hypantrum to test its relationship with body size. We found convergent evolution among large-bodied clades, except when the clade evolved an alternative mechanism for vertebral bracing. For example, some extinct lineages that lack the hyposphene-hypantrum articulation (e.g. ornithischians) have ossified tendons that braced their vertebral column. Ossified tendons are present even in small taxa and in small-bodied juveniles, but largebodied taxa with ossified tendons reached those body sizes without evolving the hyposphene-hypantrum articulation. The hyposphene-hypantrum was permanently lost in extinct crownward members of both major archosaur lineages (i.e. Crocodylia and Aves) as they underwent phyletic size decrease, changes in vertebral morphology and shifts in ecology.
The evolution of convergence, growth, and diet under an adaptive landscape framework
Wynd, Brenen Michael (Virginia Tech, 2022-03-23)
Macroevolutionary patterns of adaptation are a product of natural selection acting on genetic and developmental variation within populations, the basis of microevolution. In microevolution, an adaptive landscape is used to visualize the relationship between phenotype and fitness, through a series of peaks and valleys. The adaptive landscape, as a concept suggests that there is some phenotypic optimum, or a combination of phenotypes, that result in a maximum fitness. This peak is not stable but is a reflection of interactions between the environment and the flora and fauna within. To expand the adaptive landscape to macroevolutionary scales is to assume that there is some optimum that a species or population is adapted to, and that numerous species can be compared to one another on the same landscape. The world of phylogenetic comparative methods uses the theory of the adaptive landscape in investigating the trajectory of trait change but is often limited to extant organisms. The fossil record often represents a major gap in the use of adaptive landscape theory, due in part to the incomplete nature of specimens or difficulties in untangling evolutionary relationships. Within this gap, the Triassic Period (252.2 – 201.5 MA) is sparsely represented, due to the often highly incomplete nature of Triassic fossils and our constantly evolving understanding of their phylogenetic relationships. However, the Triassic Period is bookended by mass extinctions, and is thus a useful case study to explore the utility of adaptive landscape theory for organisms in a time of rapid environmental change. My dissertation explores convergence and growth through an adaptive landscape framework, to reconstruct how species were evolving, or populations adapting, to a changing environment. The first chapter of my dissertation explores the evolution of a long snout in reptiles, with exploration of convergent evolution for both extant and extinct reptiles across the tree of life. The second chapter of my dissertation explores a statistical method to incorporate variation due to fossilization in estimating and quantifying growth curves. This second chapter was necessary to explore the third chapter of my dissertation, the ontogeny of a large-bodied mammal relative, Exaeretodon argentinus. With my third chapter, I quantify growth curves, compare them across other proto mammals closely related to Exaeretodon, and explore how diet may have changed over the lifetime of a single individual. These chapters focus on the adaptive landscape over different scales (population vs clade), and serve as a basis for future work in estimating dietary evolution.
The evolution of intraspecific variation, growth, and body size in early theropod dinosaurs
Griffin, Christopher T. (Virginia Tech, 2016-06-24)
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.
Femoral specializations to locomotor habits in early archosauriforms
Pintore, Romain; Houssaye, Alexandra; Nesbitt, Sterling J.; Hutchinson, John R. (Wiley, 2022-05)
The evolutionary history of archosaurs and their closest relatives is characterized by a wide diversity of locomotor modes, which has even been suggested as a pivotal aspect underlying the evolutionary success of dinosaurs vs. pseudosuchians across the Triassic-Jurassic transition. This locomotor diversity (e.g., more sprawling/erect; crouched/upright; quadrupedal/bipedal) led to several morphofunctional specializations of archosauriform limb bones that have been studied qualitatively as well as quantitatively through various linear morphometric studies. However, differences in locomotor habits have never been studied across the Triassic-Jurassic transition using 3D geometric morphometrics, which can relate how morphological features vary according to biological factors such as locomotor habit and body mass. Herein, we investigate morphological variation across a dataset of 72 femora from 36 different species of archosauriforms. First, we identify femoral head rotation, distal slope of the fourth trochanter, femoral curvature, and the angle between the lateral condyle and crista tibiofibularis as the main features varying between bipedal and quadrupedal taxa, all of these traits having a stronger locomotor signal than the lesser trochanter's proximal extent. We show a significant association between locomotor mode and phylogeny, but with the locomotor signal being stronger than the phylogenetic signal. This enables us to predict locomotor modes of some of the more ambiguous early archosauriforms without relying on the relationships between hindlimb and forelimb linear bone dimensions as in prior studies. Second, we highlight that the most important morphological variation is linked to the increase of body size, which impacts the width of the epiphyses and the roundness and proximodistal position of the fourth trochanter. Furthermore, we show that bipedal and quadrupedal archosauriforms have different allometric trajectories along the morphological variation in relation to body size. Finally, we demonstrate a covariation between locomotor mode and body size, with variations in femoral bowing (anteroposterior curvature) being more distinct among robust femora than gracile ones. We also identify a decoupling in fourth trochanter variation between locomotor mode (symmetrical to semi-pendant) and body size (sharp to rounded). Our results indicate a similar level of morphological disparity linked to a clear convergence in femoral robusticity between the two clades of archosauriforms (Pseudosuchia and Avemetatarsalia), emphasizing the importance of accounting for body size when studying their evolutionary history, as well as when studying the functional morphology of appendicular features. Determining how early archosauriform skeletal features were impacted by locomotor habits and body size also enables us to discuss the potential homoplasy of some phylogenetic characters used previously in cladistic analyses as well as when bipedalism evolved in the avemetatarsalian lineage. This study illuminates how the evolution of femoral morphology in early archosauriforms was functionally constrained by locomotor habit and body size, which should aid ongoing discussions about the early evolution of dinosaurs and the nature of their evolutionary "success" over pseudosuchians.
The first ornithosuchid from Brazil and its macroevolutionary and phylogenetic implications for Late Triassic faunas in Gondwana
Muller, Rodrigo T.; Belen von Baczko, M.; Desojo, Julia B.; Nesbitt, Sterling J. (2020-01-31)
Ornithosuchidae is one of the most enigmatic clades of Triassic pseudosuchians. The group is composed by three carnivorous species that were excavated from Upper Triassic beds of Scotland and Argentina. We describe the first ornithosuchid from the Upper Triassic sediments of Brazil and explore its phylogenetic affinities and implications for the evolution of the group. Dynamosuchus collisensis gen. et sp. nov. was found as the sister taxon of the Argentinean form Venaticosuchus rusconii. These relationships reject a potential endemic radiation of ornithosuchids from the Ischigualasto-Villa Union Basin and would better support multiple diversification events. Our findings with ornithosuchids is consistent with the pattern reported for proterochampsid and erpetosuchid archosaurifonns from Ischigualasto-Villa Union and the Parana basins. In addition, the presence of an ornithosuchid in the Late Triassic of Brazil suggests that ornithosuchids were more widespread than previously thought in the southern hemisphere. The new ornithosuchid further demonstrates a faunistic link between the Argentinean and Brazilian basins during the Carnian. Finally, the discovery of the new species provides the first clue of a putative necrophagous vertebrate from the oldest dinosaur-bearing beds and expands our knowledge regarding the trophic structure of the Late Triassic of Brazil.
Including Distorted Specimens in Allometric Studies: Linear Mixed Models Account for Deformation
Wynd, Brenen M.; Uyeda, Josef C.; Nesbitt, Sterling J. (Oxford Academic, 2021-05-18)
Allometry—patterns of relative change in body parts—is a staple for examining how clades exhibit scaling patterns representative of evolutionary constraint on phenotype, or quantifying patterns of ontogenetic growth within a species. Reconstructing allometries from ontogenetic series is one of the few methods available to reconstruct growth in fossil specimens. However, many fossil specimens are deformed (twisted, flattened, and displaced bones) during fossilization, changing their original morphology in unpredictable and sometimes undecipherable ways. To mitigate against post burial changes, paleontologists typically remove clearly distorted measurements from analyses. However, this can potentially remove evidence of individual variation and limits the number of samples amenable to study, which can negatively impact allometric reconstructions. Ordinary least squares (OLS) regression and major axis regression are common methods for estimating allometry, but they assume constant levels of residual variation across specimens, which is unlikely to be true when including both distorted and undistorted specimens. Alternatively, a generalized linear mixed model (GLMM) can attribute additional variation in a model (e.g., fixed or random effects). We performed a simulation study based on an empirical analysis of the extinct cynodont, Exaeretodon argentinus, to test the efficacy of a GLMM on allometric data. We found that GLMMs estimate the allometry using a full dataset better than simply using only non-distorted data. We apply our approach on two empirical datasets, cranial measurements of actual specimens of E. argentinus (n = 16) and femoral measurements of the dinosaur Tawa hallae (n = 26). Taken together, our study suggests that a GLMM is better able to reconstruct patterns of allometry over an OLS in datasets comprised of extinct forms and should be standard protocol for anyone using distorted specimens.
Inferring Extinct Reptilian Response To Global Warming: Insights From Modern Stable Isotope Ratios
Riegler, Mitchell Scott (Virginia Tech, 2018-06-04)
Lizard ecology through time is largely unknown. Understanding ecology is important because of today’s drastic climate change, but this is not a unique event. Early Cenozoic hyperthermals were comparable to the perturbations currently experienced by living species. Understanding ecology through time must acknowledge the dynamic relationship between an organism and its environment on multiple scales. Ecological inferences can be based on form equaling function, correlating certain features (e.g. leaf-shaped dentition) with certain behaviors (e.g. herbivorous diet). Though this applies to specific taxa, there are confounding examples. Ecology can also be inferred through indirect means, but these are disconnected from the taxon of interest. Stable isotope geochemistry, however, provides an independent test. I analyzed stable isotope ratios (δ¹⁸O, δ¹³C) from enamel, providing new data on the connection between morphology, diet, and environment. I find a trophic separation in δ¹³C, and indications of aridity through δ¹⁸O. I applied this framework to extinct lizards from an Early Eocene (Wa4) assemblage, a key time between two major global warming events (Paleocene-Eocene Thermal Maximum and Early Eocene Climatic Optimum). I identify xenosaurid and glyptosaurine squamates and alethinophidian snakes. The xenosaurid is one of the youngest representatives of Restes rugosus, and I provide the first testable hypothesis of its ecology. These δ¹⁸O values corroborate hypotheses of a wet, tropical environment, and the δ¹³C values indicate an insectivorous or carnivorous diet for both taxa. My study provides an independent test of ecology of both extant and fossil lizards, with implications for differing survivorship throughout the early Cenozoic.
The influence of developmental patterns on vertebrate evolution, with the evolution of the sacrum and pelvis as a case study
Griffin, Christopher Thomas (Virginia Tech, 2020-06-03)
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.
Integrating gross morphology and bone histology to assess skeletal maturity in early dinosauromorphs: new insights from Dromomeron (Archosauria: Dinosauromorpha)
Griffin, Christopher T.; Bano, Lauren S.; Turner, Alan H.; Smith, Nathan D.; Irmis, Randall B.; Nesbitt, Sterling J. (PeerJ, 2019-02-11)
Understanding growth patterns is central to properly interpreting paleobiological signals in tetrapods, but assessing skeletal maturity in some extinct Glades may be difficult when growth patterns are poorly constrained by a lack of ontogenetic series. To overcome this difficulty in assessing the maturity of extinct archosaurian reptiles-crocodylians, birds and their extinct relatives-many studies employ bone histology to observe indicators of the developmental stage reached by a given individual. However, the relationship between gross morphological and histological indicators of maturity has not been examined in most archosaurian groups. In this study, we examined the gross morphology of a hypothesized growth series of Dromomeron romeri femora (96.6-144.4 mm long), the first series of a non-dinosauriform dinosauromorph available for such a study. We also histologically sampled several individuals in this growth series. Previous studies reported that D. romeri lacks well-developed rugose muscle scars that appear during ontogeny in closely related dinosauromorph taxa, so integrating gross morphology and histological signal is needed to determine reliable maturity indicators for early bird-line archosaurs. We found that, although there are small, linear scars indicating muscle attachment sites across the femur, the only rugose muscle scar that appears during ontogeny is the attachment of the M. caudofemoralis longus, and only in the largest-sampled individual. This individual is also the only femur with histological indicators that asymptotic size had been reached, although smaller individuals possess some signal of decreasing growth rates (e.g., decreasing vascular density). The overall femoral bone histology of D. romeri is similar to that of other early bird-line archosaurs (e.g., woven-bone tissue, moderately to well-vascularized, longitudinal vascular canals). All these data indicate that the lack of well-developed femoral scars is autapomorphic for this species, not simply an indication of skeletal immaturity. We found no evidence of the high intraspecific variation present in early dinosaurs and other dinosauriforms, but a limited sample size of other early bird-line archosaur growth series make this tentative. The evolutionary history and phylogenetic signal of gross morphological features must be considered when assessing maturity in extinct archosaurs and their close relatives, and in some groups corroboration with bone histology or with better-known morphological characters is necessary.
Intervertebral Articulation and the Evolution of Large Body Size in Archosauria
Stefanic, Candice Marie (Virginia Tech, 2017-06-26)
Dinosaurs are the largest animals to ever walk on the continents and some reached body sizes of up to 70 tons. Observation of their closest living relatives, birds and crocodylians, could never allow for prediction of gigantic dinosaurian forms. Therefore, the fossil record is crucial to understanding the evolutionary changes of these animals through time, including body size trends. The reptile group Archosauria encompasses living and extinct birds and crocodylians as well as non-avian dinosaurs and crocodile relatives called pseudosuchians. My research focuses on studying fossils of extinct archosaurs to determine how the morphology of their skeletons allowed for growth to large body sizes. I am specifically interested in how the vertebral column fits together and how the structures that articulate vertebrae change throughout the phylogeny (i.e. family tree) of Archosauria. Although major body size trends are well known for archosaurs, less research has focused on skeletal features that are associated with the evolution of large body size in that group. I hypothesize that the vertebral column will have these features. To answer the question of how vertebrae morphology is related to body size, I first described eight vertebrae from a large pseudosuchian archosaur Poposaurus langstoni. This animal possesses an accessory articulation between the vertebrae in its trunk region (i.e. between the neck and hips) called the hyposphene-hypantrum articulation. I then surveyed vertebrae from across Archosauria and found a close fit of presence of the articulation with large sizes and that it evolves independently in several archosaur groups.
Is the presence of biomolecules evidence for molecular preservation in the fossil record?
Colleary, Caitlin (Virginia Tech, 2019-05-06)
The molecular components of life (i.e., biomolecules such as DNA, proteins, lipids) have the potential to preserve in animals that have been extinct for millions of years, offering a scale of analysis previously inaccessible from the fossil record. As new technology (e.g., high resolution mass spectrometry) has been incorporated into fossil analyses, researchers have begun to detect biomolecules in terrestrial vertebrates dating back to the Triassic Period (~230 Ma). However, these biomolecules have not been demonstrated to be the biological remains of these ancient animals and may instead be exogenous organic contaminants. Here, I developed a series of analytical techniques to detect and interpret the preservation of the degraded remains of the most common protein in bone, collagen, in terrestrial vertebrates from two time slices that represent the two ends of the preservation spectrum: a "shallow time" study of fossils <150,000 years old from different burial environments (i.e., permafrost, fluvial and hot springs) and a deep time study of dinosaurs (~212 - 66 Ma) from the same burial environment (i.e., fluvial), representing the current limit of the reported protein preservation in the fossil record. Unlike previous studies that have focused on organic extractions to detect biomolecules, I studied intact fossil bones and the rocks they were found in, to understand more about the effect of burial conditions on preservation and potential alternative sources of organic compounds. I found endogenous amino acids (the degradation products of proteins) and lipids in the mammoth bones, although they were already heavily degraded in fluvial environments, even on such short timescales. I also found that there were amino acids and lipids preserved in the dinosaur bones, however tests on the age of the amino acids and the types of lipids present, demonstrate that they are not original to the animals in this study. Therefore, fluvial environments, one of the most common depositional environments preserved in the geologic record, are not conducive to the preservation of proteins on long timescales and researchers should be cautious when using these biomolecules to make interpretations about the biology of ancient animals.
A long-necked tanystropheid from the Middle Triassic Moenkopi Formation (Anisian) provides insights into the ecology and biogeography of tanystropheids
Formoso, Kiersten; Nesbitt, Sterling J.; Pritchard, Adam C.; Stocker, Michelle R.; Parker, William G. (Society of Vertebrate Paleontology, 2019-11)
Archosauromorphs are a diverse and successful group of reptiles that radiated into a series of groups around the time of the end-Permian extinction. One of these groups of archosauromorphs, tanystropheids, consists of diverse forms, and some of the largest members of the group possessed extremely elongated cervical vertebrae (greater than five times longer than tall), resulting in a hyperelongate neck. These derived tanystropheids have been found in Tethyan marine deposits of Pangaea. Four partial cervical vertebrae from a hyperelongate-necked tanystropheid from the Middle Triassic Moenkopi Formation of Arizona and New Mexico are described in this paper. These cervical vertebrae are assigned to Tanystropheidae, specifically the clade that includes the hyperelongate-necked Tanystropheus based on character states, which include an elongate centrum (length to height ratio of 6.2), the presence of epipophyses, and an elongate axial centrum. The Moenkopi tanystropheid elements were found in lower latitude fluvial sequences without any marine influence, corresponding to western Pangaea, whereas Tanystropheus-like tanystropheids are typically associated with marginal marine environments in middle to high latitudes of eastern Pangaea. These fossils suggest that hyperelongate-necked, Tanystropheus-like tanystropheids were perhaps behaviorally bound to general semi-aquatic environments, both marine and freshwater, due to their unique morphology. These fossils also greatly extend the biogeographic range of large tanystropheids and increase the anatomical diversity of tanystropheids known from North America demonstrating that the clade persisted in a wide variety of environments throughout the Triassic Period.
Molecular preservation in mammoth bone and variation based on burial environment
Colleary, Caitlin; Lamadrid, Hector M.; O'Reilly, Shane S.; Dolocan, Andrei; Nesbitt, Sterling J. (2021-01-29)
Biomolecules preserved in fossils are expanding our understanding of the biology and evolution of ancient animals. Molecular taphonomy seeks to understand how these biomolecules are preserved and how they can be interpreted. So far, few studies on molecular preservation have considered burial context to understand its impact on preservation or the potentially complementary information from multiple biomolecular classes. Here, we use mass spectrometry and other analytical techniques to detect the remains of proteins and lipids within intact fossil mammoth bones of different ages and varied depositional setting. By combining these approaches, we demonstrate that endogenous amino acids, amides and lipids can preserve well in fossil bone. Additionally, these techniques enable us to examine variation in preservation based on location within the bone, finding dense cortical bone better preserves biomolecules, both by slowing the rate of degradation and limiting the extent of exogenous contamination. Our dataset demonstrates that biomolecule loss begins early, is impacted by burial environment and temperature, and that both exogenous and endogenous molecular signals can be both present and informative in a single fossil.
Muted Blue
Ngo, Huy Quoc (Virginia Tech, 2020-07-01)
Muted Blue (Whale Level) is a Virtual Reality educational experience that explores the possibility of creating an alternative learning method in an informal learning setting such as a museum. This project seeks to merge art, technology and education to create an immersive, interactive learning experience that is geared toward Gen Z and Millennial but can be enjoyed by every age group. Unreal 4 game engine along with other 3D packages made it possible to develop a Virtual Reality experience that can be used as a complimentary piece that can possibly exist along side a museum exhibit.

Browsing by Author "Nesbitt, Sterling J."

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