Proefschrift--Groningen. / Summary in English. "Stellingen": [2] leaves inserted. Bibliography: p. 147-153.

I have collected a series of linear measurements of the postcranial skeletons of 43 extant crocodylian and lepidosaur taxa to determine if those engaging in similar locomotor behavior display similar morphology despite phylogenetic differences. Stepwise discriminant function analyses reveal reptile locomotor mode can be accurately predicted (over 80% correct) based on morphology. Semi-aquatic taxa are distinguished by a longer ischium relative to pubis length, a longer scapula relative to humerus length, and a broader acetabulum than terrestrial and arboreal taxa. Arboreal taxa display a more elongate, gracile humerus and a smaller acetabulum. This morphometric data can potentially be used to predict the locomotor behavior of a wide range of extinct reptile taxa. Within this study, Hyposaurus rogersii, Necrosuchus ionensis, Alligator sp. of the Gray Fossil Site, Crocodylus affinis, and Allognathosuchus mooki were examined and all were inferred to be semi-aquatic by the discriminant function analysis.

The latest-Permian mass extinction was the greatest biotic crisis of the Phanerozoic. The extinction decimated both marine and terrestrial communities, and changed the evolutionary trajectory of multicellular life on the planet. The unique nature of the extinction’s aftermath has prompted attention from paleontologists seeking to understand the timing and pattern of the Triassic recovery. With this dissertation I have sought to shed additional light on the terrestrial side of the extinction by examining different patterns by which its survivors responded to the extinction. Temnospondyl amphibians were one of the few tetrapod clades that were able to take advantage of the extinction to expand their diversity. In Chapter 1 I examine the relationship between taxonomic and ecological diversity of temnospondyls across the Permian-Triassic (P-Tr) boundary in the Karoo Basin of South Africa. Ecomorphological diversity, as implied by differences in cranial shape, was incorporated into the study by the use of a landmark-based geometric morphometric analysis. Both taxonomic diversity and cranial disparity were low during the Permian and increased across the Permian-Triassic boundary. Taxonomic diversity was stable through the Triassic, but disparity showed subsequent increases during the Olenekian and Anisian. Temnospondyls were restricted in size immediately following the extinction, but size range fully rebounded by the Olenekian. Tests of phylogenetic signal demonstrate that cranial shape was heavily influenced by phylogenetic relatedness, and the observed increases in disparity may be partly the result of decreases in the net relatedness of coeval Karoo stereospondylomorph temnospondyls in younger faunas. The increase in community-level taxonomic diversity for temnospondyls in the Karoo following the latest-Permian mass extinction was likely facilitated by an influx of distantly related and ecologically distinct species from other parts of Pangea. In Chapter 2, I discuss the merits of different potential methods for quantifying rates of dispersal within clades. I then apply some of these methods to two very different scenarios, first the dispersal of crocodylians across oceanic barriers during the Late Cretaceous and Cenozoic, and second the dispersal of different groups of tetrapods across Pangea during the Permo-Triassic interval. For crocodylians, because they were dispersing across substantial geographic obstacles, I opted for a direct measurement approach utilizing the optimization of discrete dispersals onto phylogenies. I examined the history of crocodylian biogeography using both parsimony and maximum likelihood on three distinct topologies with several different methods for estimating branch lengths. Across all analyses, members of the clade Alligatoroidea consistently dispersed across oceanic barriers less frequently than did non-alligatoroids. This is consistent with the hypothesis that the greater degree of salt tolerance observed in extant crocodyloids and gavialoids played a role in shaping crocodylian biogeography. The phylogenetic and temporal distribution of high dispersal rates points to an acquisition of greater salt tolerance early in the history of Crocodyloidea and Gavialoidea, potentially near the base of Longirostres if the combined evidence topology is correct. Patterns observed for changes in dispersal rate within individual clades can be largely attributed to changes in global climate and continental configuration over their history. The greater geographic ambiguity represented by the Permo-Triassic continental configuration makes a direct measurement approach inappropriate. For this study I instead opted for a proxy measurement approach, using the phylogenetic clustering of taxa within a community, measured using the Net Relatedness Index. I examined temporal changes in the phylogenetic clustering of five major tetrapod clades that span the Permian-Triassic boundary (Stereospondylomorpha, Parareptilia, Neodiapsida, Anomodontia, and Eutheriodontia) in order to examine patterns of extinction and origination through time, as well as rates of geographic dispersal. Some clades (Stereospondylomorpha, Parareptilia, and Neodiapsida) show evidence of phylogenetically selective extinction across the boundary, but this is not a universal pattern. Only one clade, Stereospondylomorpha, shows an unambiguous increase in dispersal rate following the mass extinction event. Other clades either show no change in dispersal rate, or have results that are mixed, depending on the parameters used in the analysis. These results show that stereospondylomorph temnospondyls were dispersing between geographical regions at increased rates during the Early Triassic, and this may explain much of their apparent increase in diversity following the latest-Permian mass extinction. In Chapter 3, I perform a comparison between the timing of the Triassic recovery with that following the Cretaceous-Paleogene (K-Pg) mass extinction. Three terrestrial fossil-bearing successions were examined, the Lower Triassic Beaufort Group in South Africa and Cis-Ural succession in Russia, and the Paleocene faunas of the American northern Great Plains. A comparison of generic diversity of tetrapods through time for the post-extinction intervals reveals a temporal disparity between the length of terrestrial recovery after the latest-Permian and K-Pg extinctions. Both Permo-Triassic successions show a period of low taxonomic richness (4-5Myr) after the extinction event, followed by an eventual rise in richness. The North American K-Pg succession shows a different pattern, with an immediate rise in richness culminating in a plateau shortly after the extinction (1-3 Myr). This disparity in recovery times may result from prolonged deleterious environmental conditions following the P-Tr events, although several important differences exist between these sequential fossil assemblages that may be affecting the apparent speed of recovery.

The early Pliocene Gray Fossil Site (GFS) is a biodiverse site with a unique faunal assemblage that represents one of few sites of its age in eastern North America. A community structure analysis of the mammals at GFS was done to characterize species and better understand the paleoenvironment. Data and was gathered from twenty modern communities and five late Neogene sites to compare with GFS. Species from these 26 sites were categorized by body size, locomotor mode, cheek tooth crown height, and diet to characterize niches occupied. Descriptive statistics contrasted proportions of species within categories across communities. Discriminant function analyses (DFA) determined characteristics that best differentiate communities and classified fossil site habitat types. DFA results indicate that the GFS paleoenvironment was a habitat most similar to modern temperate forest regions in the eastern U.S. and China. Characteristics of GFS are dissimilar from other late Neogene sites examined in this study.

Based on a thorough examination of museum and field Climactichnites specimens, two species of this trace, which is restricted to North America, are recognized, each representing a unique behavioral variant. C. wilsoni represents surface-produced trackways, whereas C. youngi is re-erected for subsurface burrows. Burrowing behavior is supported by the presence of C. youngi within beds, the orientation of burrows inclined to bedding, and the presence of distinct burrow fills. Burrows are distinguished from surface traces by characteristics including the absence of lateral ridges and the presence of mm-sized striations superimposed on the trace. Burrowing behavior was previously unknown and represents a new behavior for the animal. A new ichnospecies, Musculopodus sedentarius, is erected for sedentary impressions of the animal. In the future Musculopodus may be expanded to include the resting traces of other soft-bodied animals known from fossils. Analysis of Climactichnites indicates that the trackmaker was elongate, bilaterally symmetric, dorsoventrally flattened, and soft-footed. These characteristics are consistent with the trackmaker being a primitive mollusk or mollusk-like animal. Unlike other Neoproterozoic and Cambrain mollusks, such as Matthevia, Wiwaxia, and Odontogriphus, the Climactichnites trackmaker could reach considerable size. At up to 29 cm wide and possibly 67 cm long, it was one of the largest animals of its time. During locomotion, the animal generated muscular waves along the sole of its foot, which was extended and clamped into the substrate. Contraction of pedal muscles then pulled the body forward. This method of locomotion is similar to that employed by some gastropods, including Bullia and Polinices, which make Climactichnites-like trackways in exposed intertidal settings today. However, these modern trackways are not preserved because they are eroded by wind, waves, tides and subsequent bioturbation, as experiments confirm. Abundant microbial sedimentary structures associated with Cambrian occurrences suggests that microbial binding may have mediated the preservation of Climactichnites. Two lines of evidence suggest that the Climactichnites trackmaker may have been one of the first animals to venture onto land: the co-occurrence of subaerially-produced sedimentary structures, such as adhesion structures and raindrop impressions, and trackways that span shoreline depth gradients and exhibit variable preservation quality along their length.

Cricetidae, which includes a wide range of mice and rats, is the second-most diverse and abundant family of mammals. Though presently distributed across the Americas and Eurasia and also known from many fossil sites in western North America, few records have been studied from the east prior to the Pleistocene. This study has identified seven species of cricetids from the Gray Fossil Site (GFS), an early Pliocene site located in the northeast corner of Tennessee well-known for preserving fossils of many large mammals. While a few studies had previously noted the occurrence of cricetids at GFS, this study provides a detailed description of the cricetid assemblage and helps expand our understanding of small mammals from this site. Specimens were examined under stereomicroscope and DinoLite digital microscope camera, allowing qualitative assessment of occlusal morphology and quantitative comparison to modern cricetids and published fossil records from across North America. The most common taxa at the site included the deer mouse Peromyscus, which is currently the most abundant and widely distributed mammal in North America, deer mouse-like Postcopemys, and the woodrat Neotoma, which is larger rat with high-crowned molars that still lives in the region today. Other less common ones, like Sigmodon, Repomys, and Neotomodon, are also present. While most of the GFS specimens can be categorized into known genera, distinctive morphological features suggest the presence of new species. This assemblage of cricetids differs from those found at contemporaneous sites, emphasizing the spatial and temporal uniqueness of GFS. The Pliocene cricetid taxa at GFS exhibit diverse body sizes and dietary preferences, providing insights into the paleoecology of the region. As a biodiversity hotspot in the Appalachian region today, GFS was likely an important habitat for cricetid evolution during the Pliocene.

Over the past decade, the study of organismal or systematic paleobiology has been progressing into a new age of digital paleobiology, in which advanced instrumentation is utilized for primary data collection and analyses. Having been progressing throughout this field of study, advanced instruments–commonly electron- and ion- microbeam equipment–have been employed for numerous fossils over the entire range of geologic time, from microfossils to macrofossils and from the Archean (beginning at 3800 Ma) to the Cenozoic (ending at the recent). These techniques, predominantly used for geochemical, morphological, and ultra-/micro-structural analyses, have unlocked an incredible amount of detail contributing to our understanding of fossil organisms, their modes of life, and their biological affinities. But further, as these techniques continue to grow and become popularized in various fields of paleobiological study, they are certain to significantly progress our comprehension and knowledge of the evolution of life through time. While the chapters presented in this dissertation may not have a unifying theme in terms of a distinct fossil organism or specific time in Earth's history, furthering the use of electron- and ion- microbeam instrumentation and expanding the paleo-genres to which digital paleobiological approaches may be applied encompasses the fundamental intention of my research. Two of the chapters reported here focus on the geochemical, ultrastructural, and microstructural investigation of organic-walled microfossils, or acritarchs, from the Paleoproterozoic (2500–1600 Ma) and Mesoproterozoic (1600–1000 Ma), using a range of advanced instrumentation including field emission scanning electron microscopy, transmission electron microscopy, laser Raman spectroscopy, electron microprobe, secondary ion mass spectroscopy, and focused ion beam electron microscopy. Moving into the Neoproterozoic (1000–542 Ma), the third primary research chapter utilizes field emission scanning electron microscopy for high-resolution, high magnification imaging and quantitative evaluation of an entire fossil assemblage–from acritarchs and algal fossils to the earliest metazoan embryos. This study was conducted in an effort to examine and describe the phosphatization taphonomic window of the Doushantuo Formation of South China, which is a prime example of exceptional preservation. Finally, the fourth primary research chapter reported here uses field emission scanning electron microscopy and environmental scanning electron microscopy in a field of paleobiology in which advanced instrumentation has been highly underutilized – predatory-prey interactions. This research examines microstructural characteristics of predatory drill holes in both modern and fossil organisms in an attempt to mitigate the identification of predation traces in the fossil record. / Ph. D.

Bison spp. (bison) fossils are scarce on the Colorado Plateau, especially within the greater Grand Canyon region. Because of the poor fossil record for bison on the plateau and in Grand Canyon National Park, various resource managers have surreptitiously designated bison a nonnative and human-introduced species. The lack of evidence for bison seems to be the result of collection bias rather than a true lack of bison remains. Today, Grand Canyon National Park has a neighboring herd of 350 bison that have meandered unwantedly onto National Park lands from neighboring Forest Service and State of Arizona lands. This study spatiotemporally illustrates bison are recently native to the greater Grand Canyon area based on previously misidentified specimens in archaeological collections. Data here may require resource managers to reconsider whether or not bison should be reconsidered a native species to the Grand Canyon National Park and elsewhere on the Colorado Plateau.

Sciurids are one of the most diverse mammalian groups today, but the fossil record does not reflect that diversity. The purpose of this project was to identify sciurids from two late Miocene sites—the Gray Fossil Site (GFS) of Tennessee, and Tyner Farm of Florida—and use those identifications to infer the ecology of the two. From the GFS four sciurids were identified: Eutamias or Neotamias, Glaucomys, and two tree squirrels (Sciurini). Two sciurids were identified from Tyner Farm: a chipmunk attributed to Tamiina, and a ground squirrel attributed to Marmotina. Paleoenvironmental inferences based on the sciurids present at both sites are consistent with previous work. The GFS Eutamias or Neotamias is the first record of a western chipmunk population in eastern North America; the GFS tree squirrels are the earliest record of a tree squirrel in eastern North America. The GFS Glaucomys is the earliest record of the genus.

During the last two decades, a broad spectrum of short- and long-term studies on different taxonomic groups has enriched our understanding about how dynamics of taxonomic and ecological diversification have changed through geologic time. There are two major issues that have impacted these studies: the quality and quantity of data used are often insufficient in various ways and the methods used may produce results that are more equivocal than supposed. To investigate these issues more fully, this dissertation focuses on studies on two major aspects: 1) short-term studies examining the nature of successful and unsuccessful predatory attacks on Plio-Pleistocene bivalves; and 2) a Phanerozoic-scale project examining trends in bivalve richness and ecological differentiation. The short-term studies, focusing on shell-breaking predation on bivalves, have shown that the existing methodologies which only study either successful or unsuccessful component of predation in isolation are fraught with potential issues in developing effective interpretations. When these two components (i.e., successful and unsuccessful) are studied in tandem as was done here, however, traces of predation can be used to better constrain potential paleoecological interpretations related to predation intensity, predator's attack strategies, and predator-prey dynamics. The long-term project includes two Phanerozoic studies on bivalves' taxonomic and ecological richness. The taxonomic study has shown how the elements included in various datasets used can affect the Phanerozoic richness trajectory of bivalves. The revised and newly compiled dataset developed here reveals that bivalves showed three major episodes of diversification - a Ordovician radiation of orders and families, a Mesozoic diversification of families, and a dramatic Cenozoic rise in the total number of genera - all of which were synchronous with ecological diversification in terms of appearances of new life forms capable of colonizing new ecospace (i.e., cubes). However, these synchronous changes in taxonomic-ecologic richness were influenced by many biotic (e.g., predation, competition, and adaptive innovations) and abiotic (e.g., nutrient availability, sea level, and temperature) components, for which I propose a multilevel mixed model such that all these components can be studied in tandem.