The Cretaceous/Tertiary (K/Pg) mass extinction has long been viewed as a pivotal event in mammalian evolutionary history, in which the extinction of non-avian dinosaurs allowed mammals to rapidly expand from small-bodied, generalized insectivores to a wide array of body sizes and ecological specializations. Many studies have used global- or continental-scale taxonomic databases to analyze this event on coarse temporal scales, but few studies have documented morphological diversity of mammalian paleocommunities on fine spatiotemporal scales in order to examine ecomorphological selectivity and ecospace filling across this critical transition. Focusing on well-sampled and temporally well-constrained mammalian faunas across the K/Pg boundary in northeastern Montana, I quantified dental-shape disparity and morphospace occupancy via landmark- and semilandmark-based geometric morphometrics and mean body size, body-size disparity, and body-size structure via body-mass estimates.

My results reveal several key findings: (1) latest Cretaceous mammals, particularly metatherians and multituberculates, had a greater ecomorphological diversity than is generally appreciated, occupying regions of the morphospace that are interpreted as strict carnivory, plant-dominated omnivory, and herbivory; (2) the decline in dental-shape disparity and body-size disparity across the K/Pg boundary shows a pattern of constructive extinction selectivity against larger-bodied dietary specialists, particularly strict carnivores and taxa with plant-based diets, that suggests the kill mechanism was related to depressed primary productivity rather than a globally instantaneous event; (3) the ecomorphological recovery in the earliest Paleocene was fueled by immigrants, namely three multituberculate families (taeniolabidids, microcosmodontids, eucosmodontids) and to a lesser extent archaic ungulates; and (4) despite immediate increases in the taxonomic richness of eutherians, their much-celebrated post-K/Pg ecomorphological expansion had a slower start than is generally perceived and most likely only began 400,000 to 1 million years after the extinction event.

Using tangential thin sections, we examined variation in porosity and water vapor conductance across two eggs of Troodon formosus, a small (∼50 kg) theropod dinosaur from the North American Upper Cretaceous, testing two hypotheses of egg incubation: (1) full burial within sediments or vegetation and (2) partial burial with exposed upper egg portions. We divided and sampled the eggs in five zones, 1 through 5 from blunt top to more pointed bottom. A geometric model composed of a hemisphere, cone, and paraboloid was used to estimate total and zonal volumes and surface areas. The 138 × 67 mm idealized Troodon egg has a volume, surface area, and mass of 296.4 cm3, 239.23 cm2, and 314.2 g, respectively. Zonal surface areas and volumes highlight the strongly asymmetric and elongate form of the Troodon egg. Geometric modeling provides better estimates of volume and surface area where egg shape diverges markedly from that of a typical bird egg. Porosity varies significantly across both Troodon eggs, with zones 2 and 3 having the largest pores and a majority (70–78%) of total conductance, whereas zone 5 has very low conductance. Total water vapor conductance in the two eggs are 31.85 and 40.62 mg H2O day− Torr−, values 76% and 97% of those predicted for an avian egg of similar size. Low total conductance compares favorably to values in extant birds and non-avian reptiles that incubate in open nests, arguing against full burial incubation. Together with nesting site evidence, low conductance values favor partial burial and incubation by a Troodon adult. Asymmetric egg shape concentrates volume, surface area, and conductance near or at the point of subaerial exposure. Among non-avian dinosaurs, the eggs of Troodon and troodontids are most similar to those of modern birds in having an asymmetric shape, low porosity, no ornamentation, and three structural eggshell layers.

The presumed affinities of the Terminal Neoproterozoic Ediacara biota have been much debated. However, even in the absence of concrete evidence for phylogenetic affinity, numerical paleoecological approaches can be effectively used to make inferences about organismal biology, the nature of biotic interactions, and life history. Here, we examine the population structure of three Ediacaran rangeomorph taxa (Fractofusus, Beothukis, and Pectinifrons), and one non-rangeomorph taxon (Thectardis) across five fossil surfaces around the Avalon Peninsula, Newfoundland, through analysis of size-frequency distributions using Bayesian Information Criterion (BIC). Best-supported models resolve communities of all studied Ediacaran taxa at Mistaken Point as single cohorts with wide variance. This result is best explained in terms of a “continuous reproduction” model, whereby Ediacaran organisms reproduce aseasonally, so that multiple size modes are absent from preserved communities. Modern benthic invertebrates (both as a whole and within specific taxonomic groups) in deeper-water settings reproduce both seasonally and aseasonally; distinguishing between biological (i.e., continuous reproductive strategies) and environmental (lack of a seasonal trigger) causes for this pattern is therefore difficult. However, we hypothesize that the observed population structure could reflect the lack of a trigger for reproduction in deepwater settings (i.e., seasonal flux of organic matter), until the explosive appearance of mesozooplankton near the base of the Cambrian.

Reconciliation of paleontological and molecular phylogenetic evidence holds great promise for a better understanding of the temporal succession of cladogenesis and character evolution, especially for taxa with a fragmentary fossil record and uncertain classification. In zoology, studies of this kind have largely been restricted to Bilateria. Hexactinellids (glass sponges) readily lend themselves to test such an approach for early-branching (non-bilaterian) animals: they have a long and rich fossil record, but for certain taxa paleontological evidence is still scarce or ambiguous. Furthermore, there is a lack of consensus for taxonomic interpretations, and discrepancies exist between neontological and paleontological classification systems. Using conservative fossil calibration constraints and the largest molecular phylogenetic data set assembled for this group, we infer divergence times of crown-group Hexactinellida in a Bayesian relaxed molecular clock framework. With some notable exceptions, our results are largely congruent with interpretations of the hexactinellid fossil record, but also indicate long periods of undocumented evolution for several groups. This study illustrates the potential of an integrated molecular/paleobiological approach to reconstructing the evolution of challenging groups of organisms.

Nectocaridids are soft-bodied early to middle Cambrian organisms known from Burgess Shale-type deposits in Canada, China, and Australia. Originally described as unrelated species, they have recently been interpreted as a clade; their flexible tentacles, camera-type eyes, lateral fins, internal gills, axial cavity, and funnel point to a relationship with the cephalopods. However, aspects of this reinterpretation, including the relevance of the group to cephalopod evolution, have been called into question.

Here, I examine new and existing nectocaridid material, including a large new form that may represent a sexual dimorph of Nectocaris pteryx. Differences between existing taxa largely represent taphonomic variation between sites and specimens—which provides further constraint on the organisms' anatomy. I revise the morphology of the tentacles and fins, and describe mouthparts and phosphatized gills for the first time. A mathematical analysis supports the presence of the earliest known camera-type eyes, and fluid mechanical considerations suggest that the funnel is optimized for efficient jet propulsion in a low Reynolds number flow regime.

Nectocaridids closely resemble coleoid cephalopods, but a position deeper within Cephalopoda raises fewer stratigraphic challenges. Whether its coleoid-like construction reflects common ancestry or profound convergence, the Nectocaris body plan adds substantially to Cambrian disparity, demonstrating the rapid colonization of nektobenthic niches after the Cambrian explosion.

We infer the body-size scaling slope of metabolic rate in a trilobite by applying a cell-size model that has been proposed to explain metabolic scaling in living organisms. This application is especially tractable in fossil arthropods with well-preserved compound eyes because the number and size of eye facets appear to be useful proxies for the relative number and size of cells in the body. As a case study, we examined the ontogenetic scaling of facet size and number in a ∼390-Myr-old local assemblage of the trilobite Eldredgeops rana, which has well-preserved compound eyes and a wide body-size range. Growth in total eye lens area resulted from increases in both facet area and number in relatively small (presumably young) specimens, but only from increases in facet area in large (presumably more mature) specimens. These results suggest that early growth in E. rana involved both cell multiplication and enlargement, whereas later growth involved only cell enlargement. If the cell-size model is correct, then metabolic rate scaled allometrically in E. rana, and the scaling slope of log metabolic rate versus log body mass decreased from ∼0.85 to 0.63 as these animals grew. This inferred age-specific change in metabolic scaling is consistent with similar changes frequently observed in living animals. Additional preliminary analyses of literature data on other trilobites also suggest that the metabolic scaling slope was <1 in benthic species, but ∼1 in pelagic species, as has also been observed in living invertebrates. The eye-facet size (EFS) method featured here opens up new possibilities for examining the bioenergetic allometry of extinct arthropods.

Synapsids dominated the terrestrial realm between the late Pennsylvanian and the Triassic. Their early evolution includes some of the first amniotes to evolve large size, herbivory, and macro-predators. However, little research has focused on the changes in diversity occurring during this early phase in their evolutionary history, with more effort concentrating on later events such the Permo-Triassic extinction. Here we assess synapsid diversity, at both the species and genus levels, between the Carboniferous (Moscovian) and the Middle Permian (Capitanian). A raw, taxic diversity (richness) estimate is generated, and we use two separate methods to correct for sampling biases in this curve. To remove the effect of anthropogenic sampling bias, we apply a recently published modification of the residual diversity method, and then generate a supertree, using matrix representation with parsimony to infer ghost lineages and obtain a phylogenetic diversity estimate. The general diversity pattern reflects the initial diversification of synapsids in the late Pennsylvanian and early Cisuralian, which was followed by an extinction event during the Sakmarian. Diversity recovered during the Artinskian and Kungurian, coinciding with the radiation of Caseidae, although other families begin to decline. A second extinction event occurred across the Kungurian/Roadian boundary, in which Edaphosauridae and Ophiacodontidae died out although Caseidae and Therapsida diversified. The sampling-corrected curves reveal further extinction during the Roadian, although therapsids were again unaffected. Pelycosaurian-grade synapsids survived during the Wordian and Capitanian, but were a minor part of an otherwise therapsid-dominated fauna. Evidence of significant anthropogenic sampling bias calls into question previous diversity studies that have not employed sampling correction.

The turnover-pulse hypothesis (TPH) makes explicit predictions concerning the potential responses of species to climate change, which is considered to be a major cause of faunal turnover (extinction, speciation, and migration). Previous studies have tested the TPH primarily by examining temporal correlations between turnover pulses and climatic events. It is rarely possible to dissect such correlations and observe turnover as it is occurring or to predict how different lineages will respond to climate change. Thus, whether climate change drives faunal turnover in the manner predicted by the TPH remains unclear. In this study, we test the underlying mechanics of the TPH using well-dated Quaternary ungulate records from southern Africa's Cape Floristic Region (CFR). Changes in sea level, vegetation, and topographic barriers across glacial-interglacial transitions in southern Africa caused shifts in habitat size and configuration, allowing us to generate specific predictions concerning the responses of ungulates characterized by different feeding habits and habitat preferences. Examples from the CFR show how climatically forced vegetation change and allopatry can drive turnover resulting from extinction and migration. Evidence for speciation is lacking, suggesting either that climate change does not cause speciation in these circumstances or that the evolutionary outcome of turnover is contingent on the nature and rate of climate change. Migrations and extinctions are observed in the CFR fossil record over geologically short time intervals, on the order of Milankovitch-scale climate oscillations. We propose that such climate oscillations could drive a steady and moderate level of faunal turnover over 104-year time scales, which would not be resolved in paleontological records spanning 105 years and longer. A turnover pulse, which is a marked increase in turnover relative to previous and subsequent time periods, requires additional, temporally constrained climatic forcing or other processes that could accelerate evolutionary change, perhaps mediated through biotic interactions.

Understanding morphological integration is one of the central goals of evolutionary developmental biology. Despite its applicability to questions of paleontological interest, there are few studies on integration in fossil vertebrates. In this study, we examine limb integration in the Lower Jurassic ichthyosaur Stenopterygius quadriscissus, with the aim of examining the effect of ontogeny and anagenetic changes over short geological time spans on metrics of limb integration. Both ontogenetic and stratigraphic effects had a significant influence on measured values of integration, the identity of strongly integrated elements, and some common ratio values such as the relative integration of the forelimb to the hind limb, or within-limb to between-limb integration. Ontogenetic effects were relatively greater, although this could be linked to sample size. Although adults showed the lowest levels of overall integration, they possessed high levels of integration between serially homologous elements, something that was unexpected due to strong divergence in limb size and perhaps functional differences in derived ichthyosaurs. Ontogenetic differences in the relative integration of the forelimb to the hind limb are probably related to early locomotor demands on the forelimb. We conclude that if samples are pooled, the resulting pattern of integration may not reflect any one subsample but will be a composite created through the superposition of several variables. Pooling data in paleontological studies of integration has a non-trivial effect on the results obtained.

The Paleocene-Eocene Thermal Maximum (PETM; ca. 55.8 Ma) is thought to coincide with a profound but entirely transient change among nannoplankton communities throughout the ocean. Here we explore the ecology of nannoplankton during the PETM by using multivariate analyses of a global data set that is based upon the distribution of taxa in time and space. We use these results, coupled with stable isotope data and geochemical modeling, to reinterpret the ecology of key genera. The results of the multivariate analyses suggest that the community was perturbed significantly in coastal and high-latitudes sites compared to the open ocean, and the relative influence of temperature and nutrient availability on the assemblage varies regionally. The open ocean became more stratified and less productive during the PETM and the oligotrophic assemblage responded primarily to changes in nutrient availability. Alternatively, assemblages at the equator and in the Southern Ocean responded to temperature more than to nutrient reduction. In addition, the assemblage change at the PETM was not merely transient—there is evidence of adaptation and a long-term change in the nannoplankton community that persists after the PETM and results in the disappearance of a high-latitude assemblage. The long-term effect on communities caused by transient warming during the PETM has implications for modern-day climate change, suggesting similar permanent changes to nannoplankton community structure as the oceans warm.

The deep-sea Cenozoic planktonic microfossil record has the unique characteristics of continuously well-preserved populations of most species, with virtually unlimited sample size, and therefore constitutes, in principle, a major resource for macroevolutionary research. Antarctic Neogene radiolarians in particular, are diverse, abundant and consistently well-preserved and evolved rapidly. This fauna is, in theory, a near-perfect testing ground for paleodiversity reconstructions. In this study we determined the diversity history of these faunas from a new quantitative, taxonomically complete data set from Neogene and Quaternary sections at several Antarctic sites. The pattern retrieved by our whole-fauna data set shows a significant, largely extinctionless ecological change in faunal composition and decrease in the evenness of species' abundances during the late Miocene, followed 3 Myr later, at around 5 Ma, by a significant drop in diversity. We tentatively associate this ecological event with a synchronous, regional change in the composition of the primary producers, but as yet cannot identify any environmental changes associated with the later extinction. Further, our whole-fauna diversity history was compared to diversity computed from much less complete, biostratigraphically oriented studies of species' occurrences, compiled in the Neptune database and reconstructed by using subsampling methodologies. Comparison of our whole-fauna and subsampling-reconstructed diversity patterns shows that the first-order trends are the same in both, suggesting that, to some degree, such literature compilations can be used to explore diversity history of plankton. However, our results also highlight substantial errors and distortions in the reconstructed diversity which make it poorly suited to more-detailed studies (e.g., for comparison of diversity history with paleoenvironmental history). We conclude that detailed studies of plankton diversity, and particularly those attempting to understand the relation between diversity and paleoceanographic change, should be based on taxonomically comprehensive, quantitative data.

Significant warming of Earth's climate in the near term seems increasingly likely. If significant enough, this climatic regime could, in the long term, come to resemble previous greenhouse intervals in earth history. Consequently, analysis of the fossil record during periods of extreme warmth may provide important lessons for species biology, including biogeography, in a much warmer world. To explore this issue, we analyzed the biogeographic response of 63 molluscan species to the long-term global warmth in the Late Cretaceous Western Interior Seaway (WIS) of North America, using Geographic Information Systems (GIS) to quantitatively measure changes in range size and distribution throughout this interval. We specifically considered the role that geographic range size played in mediating extinction resistance and invasion potential of these WIS species. We found no relationship between geographic range size and survivorship. However, endemic species with small range sizes were more likely to become invasive. Finally, mollusks did not experience a poleward shift in range out of the tropics during this warm regime. To the extent that these patterns are representative, and the WIS and taxa considered constitute a reasonable ancient analogue to a warmer future world, these results suggest that some biogeographic “rules” may not prevail under greenhouse conditions of long-term, equable warmth. They also suggest that other factors beyond geographic range size, including distinctive niche characteristics, may play quite important roles in species survival and invasion potential. This potentially complicates predictions regarding the future responses of extant species to long-term warming.

Whether or not climate plays a causal role in mammal body-size evolution is one of the longest-standing debates in ecology. Bergmann's Rule, the longest-standing modeladdressing this topic, posits that geographic body-mass patterns are driven by temperature, whereas subsequent research has suggested that other ecological variables, particularly precipitation and seasonality, may be the major drivers of body-size evolution. While paleoecological data provide a unique and crucial perspective on this debate, paleontological tests of Bergmann's rule and its corollaries have been scarce. We present a study of body-size evolution in three ecologically distinct families of mammal (equids, canids, and sciurids) during the Oligo-Miocene of the northwest United States, an ideal natural laboratory for such studies because of its rich fossil and paleoclimatic records. Body-size trends are different in all three groups, and in no case is a significant relationship observed between body size and any climatic variable, counter to what has been observed in modern ecosystems. We suggest that for most of the Cenozoic, at least in the Northwest, body mass has not been driven by any one climatic factor but instead has been the product of complex interactions between organisms and their environments, though the nature of these interactions varies from taxon to taxon. The relationship that exists between climate and body size in many groups of modern mammals, therefore, is the exception to the rule and may be the product of an exceptionally cool and volatile global climate. As anthropogenic global warming continues and ushers in climatic conditions more comparable to earlier intervals of the Cenozoic than to the modern day, models of corresponding biotic variables such as body size may lose predictive power if they do not incorporate paleoecological data.

The profound evolutionary success of mammals has been linked to behavioral and life-history traits, many of which have been tied to brain size. However, studies of the evolution of this key trait have yet to explore the full potential of the fossil record, being limited by the difficulty of obtaining endocranial data from fossils. Using measurements of endocranial volume, length, height, and width of the braincase in 503 adult specimens from 199 extant species, representing 99 of 133 extant mammalian families, we expand upon a simple method of using multiple regression to develop a formula for estimating brain size from external skull measurements. We also examined non-mammalian synapsids to assess the phylogenetic limits of our model's application. Model-predicted volume correlates strongly with measured volume (R2 = 0.993) and prediction error is between 16% and 19%. Error decreases if models developed for well-sampled subclades such as primates or rodents are used, demonstrating that some differential evolution of the relationship between brain size and skull size has occurred. However, reanalysis using phylogenetically independent contrasts demonstrates weak phylogenetic dependency, indicating that our model is appropriate for estimating the endocranial volume of species of unknown phylogenetic affinity. Thus, the model represents a generally applicable, fast and cost-efficient way to dramatically expand the taxonomic and temporal scope of mammalian brain size data sets. Even endocranial volumes of taxa with highly derived crania, such as cetaceans and monotremes, can be estimated confidently. However, the model works best for generalized placental crania. Fundamental differences in cranial architecture suggest that the model cannot provide accurate estimates of endocranial volume in non-mammalian synapsids more basal than Morganucodon (ca. 200 Ma). Therefore, use of the model for taxa phylogenetically distant from the mammalian crown group is not warranted, but it might be used to establish relative brain sizes between closely related subgroups.