This paper aims at assessing the influence of the bone ornamentation and, specifically, the associated loss of bone mass on the mechanical response of the crocodylomorph osteoderms. To this end, we have performed three-dimensional (3D) modeling and a finite element analysis on a sample that includes both extant dry bones and well-preserved fossils tracing back to the Late Triassic. We simulated an external attack under various angles on the apical surface of each osteoderm and further repeated the simulation on an equivalent set of smoothed 3D-modeled osteoderms. The comparative results indicated that the presence of an apical sculpture has no significant influence on the von Mises stress distribution in the osteoderm volume, although it produces a slight increase in its numerical score. Moreover, performing parametric analyses, we showed that the Young's modulus of the osteoderm, which may vary depending on the bone porosity, the collagen fiber orientation, or the calcification density, has no impact on the von Mises stress distribution inside the osteoderm volume. As the crocodylomorph bone ornamentation is continuously remodeled by pit resorption and secondary bone deposition, we assume that the apical sculpture may be the outcome of a trade-off between the bone mechanical resistance and the involvement in physiological functions. These physiological functions are indeed based on the setup of a bone superficial vessel network and/or the recurrent release of mineral elements into the plasma: heat transfers during basking and respiratory acidosis buffering during prolonged apnea in neosuchians and teleosaurids; compensatory homeostasis in response to general calcium deficiencies. On a general morphological basis, the osteoderm geometric variability within our sample leads us to assess that the global osteoderm geometry (whether square or rectangular) does not influence the von Mises stress, whereas the presence of a dorsal keel would somewhat reduce the stress along the vertical axis.

Naturally time-averaged accumulations of skeletal remains—death assemblages—provide reliable, albeit temporally coarse, information on the species composition and structure of communities in diverse settings, and their mismatch with local living communities usually signals recent human-driven ecological change. Here, we present the first test of live–dead mismatch as an indicator of human stress using ostracodes. On three islands along a gradient of human population density in the Bahamas, we compared the similarity of living and death assemblages in 10 lakes with relatively low levels of human stress to live–dead similarity in 11 physically comparable lakes subject to industrial, agricultural, or other human activities currently or in the past. We find that live–dead agreement in pristine lakes is consistently excellent, boding well for using death assemblages in modern-day and paleolimnological biodiversity assessments. In most comparison of physically similar paired lakes, sample-level live–dead mismatch in both taxonomic composition and species’ rank abundance is on average significantly greater in the stressed lakes; live–dead agreement is not lower in all samples from stressed lakes, but is more variable. When samples are pooled for lake-level and island-level comparisons, stressed lakes still yield lower live–dead agreement, but the significance of the difference with pristine lakes decreases—species that occur dead-only (or alive-only) in one sample are likely to occur alive (or dead) in other samples. Interisland differences in live–dead agreement are congruent with, but not significantly correlated with, differences in human population density. This situation arises from heterogeneity in the timing and magnitudes of stresses and in the extent of poststress recovery. Live–dead mismatch in ostracode assemblages thus may be a reliable indicator of human impact at the sample level with the potential to be a widely applicable tool for identifying impacted habitats and, perhaps, monitoring the progress of their recovery.

Mean adult body mass of mammal taxa is a fundamental ecological variable. Variability in the distributions of body masses of a mammal fauna suggest variability in habitat structure. Mammal remains from the Marmes archaeological site in southeastern Washington State date between 13,200 and 10,400 b.p., during the Pleistocene–Holocene transition (PHT). Known environmental history prompts the expectations that the Marmes PHT mammal remains should represent greater species richness and a larger array of body-mass sizes than modern faunas in the Marmes locale and in open shrub-steppe habitats, and lower species richness and a smaller array of body-mass sizes than modern faunas in closed forest habitats; species richness and the array of body-mass sizes should be similar to that for a mixed habitat of cool shrub-steppe with scattered conifers. The Marmes PHT cenogram meets these expectations. Body-mass clumps displayed by the Marmes PHT mammal fauna fall between those of closed forests and open shrub-steppe habitats in terms of clump richness and breadth, and in terms of gap width. Marmes PHT body-mass clumps are very similar to those for the mixed habitat. Cenograms and body-mass clumps confirm conclusions drawn 40 years ago that the Marmes PHT habitat was much like that of today but cooler and with more plant biomass and greater structural diversity than today.

Describing patterns of connectivity among organs is essential for identifying anatomical homologies among taxa. It is also critical for revealing morphogenetic processes and the associated constraints that control the morphological diversification of clades. This is particularly relevant for studies of organisms with skeletons made of discrete elements such as arthropods, vertebrates, and echinoderms. Nonetheless, relatively few studies devoted to morphological disparity have considered connectivity patterns as a level of morphological organization or developed comparative frameworks with proper tools. Here, we analyze connectivity patterns among apical plates in Atelostomata, the most diversified clade among irregular echinoids. The clade comprises approximately 1600 fossil and Recent species (e.g., 25% of post-Paleozoic species of echinoids) and shows high levels of morphological disparity. Plate connectivity patterns were analyzed using tools and statistics of graph theory. To describe and explore the diversity of connectivity patterns among plates, we symbolized each pattern as a graph in which plates are coded as nodes that are connected pairwise by edges. We then generated a comparative framework as a morphospace of connections, in which the disparity of plate patterns observed in nature was mapped and analyzed. Main results show that apical plate patterns are both highly disparate between and within atelostomate groups and limited in number; overall, they also constitute small, compact, and simple structures compared to possible random patterns. Main traits of the evolution of apical plate patterns reveal the existence of strong morphogenetic constraints that are phylogenetically determined. In contrast, evolutionary radiations within atelostomates were accompanied by a clear increase in disparity, suggesting a release of some constraints at the origin of clades.

Oceanic anoxic events (OAEs) are contemporaneous with 11 of the 18 largest Phanerozoic extinction events, but the magnitude and selectivity of their paleoecological impact remains disputed. OAEs are associated with abrupt, rapid warming and increased CO2 flux to the atmosphere; thus, insights from this study may clarify the impact of current anthropogenic climate change on the biosphere. We investigated the influence of the Late Cretaceous Bonarelli event (OAE2; Cenomanian/Turonian stage boundary; ~94 Ma) on generic- and species-level molluscan diversity, extinction rates, and ecological turnover. Cenomanian/Turonian results were compared with changes across all Cretaceous stage boundaries, some of which are coincident with less severe OAEs. We found increased generic turnover, but not species-level turnover, associated with several Cretaceous OAEs. The absence of a species-level pattern may reflect species occurrence data that are too temporally coarse to robustly detect patterns. Five hypotheses of ecological selectivity relating anoxia to survivorship were tested across stage boundaries with respect to faunality, mobility, and diet using generalized linear models. Interestingly, benthic taxa were consistently selected against throughout the Cretaceous regardless of the presence or absence of OAEs. These results suggest that: (1) the Cenomanian/Turonian boundary (OAE2) was associated with a decline in molluscan diversity and increase in extinction rate that were significantly more severe than Cretaceous background levels; and (2) no differential ecological selectivity was associated with OAE-related diversity declines among the variables tested here.

Energy availability influences natural selection on the ontogenetic histories of organisms. However, it remains unclear whether physiological controls on size remain constant throughout ontogeny or instead shift as organisms grow larger. Benthic foraminifera provide an opportunity to quantify and interpret the physicochemical controls on both initial (proloculus) and adult volumes across broad environmental gradients using first principles of cell physiology. Here, we measured proloculus and adult test dimensions of 129 modern rotaliid species from published images of holotype specimens, using holotype size to represent the maximum size of all species’ occurrences across the North American continental margin. We merged size data with mean annual temperature, dissolved oxygen concentration, particulate organic carbon flux, and seawater calcite saturation for 718 unique localities to quantify the relationship between physicochemical variables and among-species adult/proloculus size ratios. We find that correlation of community mean adult/proloculus size ratios with environmental parameters reflects covariation of adult test volume with environmental conditions. Among-species proloculus sizes do not covary identifiably with environmental conditions, consistent with the expectation that environmental constraints on organism size impose stronger selective pressures on adult forms due to lower surface area-to-volume ratios at larger sizes. Among-species adult/proloculus size ratios of foraminifera occurring in resource-limited environments are constrained by the limiting resource in addition to temperature. Identified limiting resources are food in oligotrophic waters and oxygen in oxygen minimum zones. Because among-species variations in adult/proloculus size ratios from the North American continental margin are primarily driven by the local environment’s influence on adult sizes, the evolution of foraminiferal sizes over the Phanerozoic may have been strongly influenced by changing oceanographic conditions. Furthermore, lack of correspondence between among-species proloculus sizes and environmental conditions suggests that offspring sizes in foraminifera are rarely limited by physiological constraints and are more susceptible to selection related to other aspects of fitness.

To a first approximation, the logarithmic spiral is a close fit to the biconvex lateral outline of three terebratellid genera from Australasia. Investigation of spiral parameters confirms the value of the biconvex morphospace of McGhee (1980), but not the method for estimating the spiral itself. Deviations from the estimated spiral are episodic and consistent with Rudwick's (1959) observations of allometry in brachiopod growth. Morphospace location is postulated to be indicative of a species' substrate and lifestyle. The logarithmic spiral offers much opportunity for gaining better understanding of external shape and its cause in articulate brachiopods.

Faunal similarity among regions is often used as a means of identifying regions of endemism in fossil faunas. At least two large-scale taphonomic effects can affect apparent faunal similarity: stratigraphic and facies mismatching. In stratigraphic mismatching, an unconformity represents removal of most or all of a complete assemblage zone in one region, and the constituent taxa are mistakenly interpreted as having never inhabited that region. In facies mismatching, environmental differences between two regions (possibly unrecognized) cause the inference of a barrier that never existed. The two types of mismatching can work in concert if a facies was originally represented in a single stratigraphic interval that has been completely removed from one region. Analysis of faunal similarity via multivariate analysis of individual localities, coupled with comparison of the regions as single samples, may indicate mismatching if the results differ significantly.

We view these two problems as part of a suite of taphonomic effects that are not evident in paleobiological analyses of smaller geographic scope. First, there is ambiguity in the notion of “barrier,” even when a candidate is obvious. Second, barriers in paleobiogeography are often hidden and must be inferred from their effects rather than observed. Third, stratigraphic and facies mismatching produce effects on regional faunas similar to those produced by barriers. Anyone using barriers to explain faunal disruptions should address these three points.

Upper Eocene faunas of central Florida seem taxonomically distinct from those of the remainder of the Gulf Coastal Plain. This has historically been attributed to a known paleogeographic feature, the Suwannee Strait, which acts as a barrier. The amount of dissimilarity of the echinoid faunas is greater than the amount predicted as a result of sampling problems. Comparison of the results of multivariate and whole-region analyses suggests that mismatching of the two faunas, rather than a true barrier, causes the distinction. Principally facies, but also strata are mismatched. Early Late Eocene faunas inhabited terrigenous sands to the north of the strait and carbonate sands to the south and show the highest distinctiveness. Middle Late Eocene faunas inhabited primarily carbonate sands to the south and both carbonate sands and muds to the north. Overall similarity is higher for both local and regional analyses, and the faunas of northern sands are more similar to those of the southern region than to the northern mud faunas. Overall similarity across the strait is highest in the late Late Eocene strata when both regions had carbonate mud and sand facies. The faunas exhibit greater similarity within facies than they do within regions. Upper Late Eocene strata are poorly preserved north of the strait because of post-Eocene erosion.

In considering the history of biodiversity paleontologists have focused on exploratory investigations of empirical data derived from the fossil record. Starting with the pioneering work of Philips (1860), and continuing at an increasing pace through today, this inductive approach has dominated diversity research. In contrast, deductive theoretical considerations that focus on the expected history of biodiversity, and develop independently of empirical knowledge, have remained under-explored. Appreciating the need for a nomothetic paleobiology (Gould 1980), we here reconsider the history of biodiversity, using deductive models constrained by a few, self-evident parameters. This analysis centers on the marine fossil record, the primary target of most previous empirical studies on the geological history of global biodiversity (e.g., Valentine 1969; Raup 1972, 1976; Sepkoski et al. 1981; Alroy et al. 2008).

Accretionary outlines of many brachiopods are closely matched by one or more episodes of logarithmic spiral growth where each episode has a constant growth gradient. An episodic model supports the observations of abrupt change, but not that of gradual change in accretionary growth. A reliable guide to the number of growth episodes is given by the pattern of residuals after fitting a single spiral to an outline. An episodic model exposes subtle variations in growth that are not easily described by visual examination. A three-episode model clarifies a shape difference between two species of the endemic New Zealand genus Neothyris. Direct observation, along with linear measurements and their ratios, was unable to pinpoint that the well-documented dorsoventral compression in one species arises from a growth episode in the ventral valve. Logarithmic spirals about a single coiling axis are not suited for large, disjunct changes in outline for which the moving-frame methods or aperture-deposition models are applicable.

The deep-sea microfossil record is characterized by an extraordinarily high density and abundance of fossil specimens, and by a very high degree of spatial and temporal continuity of sedimentation. This record provides a unique opportunity to study evolution at the species level for entire clades of organisms. Compilations of deep-sea microfossil species occurrences are, however, affected by reworking of material, age model errors, and taxonomic uncertainties, all of which combine to displace a small fraction of the recorded occurrence data both forward and backwards in time, extending total stratigraphic ranges for taxa. These data outliers introduce substantial errors into both biostratigraphic and evolutionary analyses of species occurrences over time. We propose a simple method—Pacman—to identify and remove outliers from such data, and to identify problematic samples or sections from which the outlier data have derived. The method consists of, for a large group of species, compiling species occurrences by time and marking as outliers calibrated fractions of the youngest and oldest occurrence data for each species. A subset of biostratigraphic marker species whose ranges have been previously documented is used to calibrate the fraction of occurrences to mark as outliers. These outlier occurrences are compiled for samples, and profiles of outlier frequency are made from the sections used to compile the data; the profiles can then identify samples and sections with problematic data caused, for example, by taxonomic errors, incorrect age models, or reworking of sediment. These samples/sections can then be targeted for re-study.

Mass extinctions can play a role in shaping macroevolutionary trends through time, but the contribution of recoveries to this process has yet to be examined in detail. This study focuses on the effects of three extinction events, the end-Cretaceous (K/T), mid-Eocene (mid-E), and end-Eocene (E/O), on long-term patterns of body size in veneroid bivalves. Systematic data were collected for 719 species and 140 subgenera of veneroids from the Late Cretaceous through Oligocene of North America and Europe. Centroid size measures were calculated for 101 subgenera and global stratigraphic ranges were used to assess extinction selectivity and preferential recovery. Veneroids underwent a substantial extinction at the K/T boundary, although diversity recovered to pre-extinction levels by the early Eocene. The mid-E and E/O events were considerably smaller and their recovery intervals much shorter. None of these events were characterized by significant extinction selectivity according to body size at the subgenus level; however, all three recoveries were strongly size biased. The K/T recovery was biased toward smaller veneroids, whereas both the mid-E and E/O recoveries were biased toward larger ones. The decrease in veneroid size across the K/T recovery actually reinforced a Late Cretaceous trend toward smaller sizes, whereas the increase in size resulting from the Eocene recoveries was relatively short-lived. Early Cenozoic changes in predation, temperature, and/or productivity may explain these shifts.

The data on the longevity of genera of articulate brachiopods for the Upper Ordovician-Devonian interval (Boucot 1975) are averaged for the taxa of each order. The generic longevity-frequency distributions form three significantly different clusters. The orthids have the highest mean generic longevity value and display a semi-log survivorship profile that is more convex than the other orders. The spiriferid and strophomenid orders have mean generic longevity values higher than and slopes in the survivorship curves less steep than the pentamerids, rhynchonellids and terebratulids. The appreciable differences in the inferred generic extinction rates for the clusters are attributed to the heterogeneity of the ecologies of the orders. The pentamerids had a proportionately higher number of genera concentrated in narrow-niched reefs wherein higher extinction rates depressed the mean generic longevity of this order relative to the orthids, strophomenids and spiriferids. Enhanced longevity of many offshore (non-reef) genera possibly explains the greater mean generic longevity of the orthids, spiriferids and strophomenids relative to the rhynchonellids and terebratulids which included a proportionately higher number of genera concentrated in nearshore habitats. Alternatively, the depressed mean generic longevity values for the rhynchonellids and terebratulids may reflect systematic oversplitting and temporal bias, i.e., origination and initial diversification of order.

The quantity of biomass in an ecosystem is constrained by energy availability. It is less clear, however, how energy availability constrains taxonomic and functional diversity. Competing models suggest biodiversity is either resource-limited or far from any bound. We test the hypothesis that functional diversity in marine bivalve communities is constrained by energy availability, measured as particulate organic carbon (POC) flux, in the modern oceans. We find that POC flux predicts the relative prevalence of ecological modes in both the Atlantic and Pacific Oceans. Moreover, the associations of ecological modes with POC fluxes are similar between the Atlantic and Pacific despite being based on independent sets of species, indicating a direct causal relationship. We then use the relationship between POC flux and the prevalence of functional groups in the modern to test the hypothesis that the trend of increasing functional diversity in bivalves across the past 500 Myr has occurred in response to increased POC flux. We find no evidence that the earliest-appearing modes of life are preferentially associated with low-POC environments or that the mean POC flux experienced by marine bivalves has increased across geological time. To reconcile the close association between ecological mode and POC flux in the modern oceans with the lack of evidence for increasing POC fluxes across time, we propose that POC flux has not increased substantially over time but, rather, the increase in bivalve functional diversity enabled bivalves to become more abundant, to occupy a broader range of environments, and to capture a greater fraction of the total POC flux. The results here suggest at the geographic scale of oceans and through geologic time bivalve diversity was not bounded by food availability.

Over the last 50 years, access to new data and analytical tools has expanded the study of analytical paleobiology, contributing to innovative analyses of biodiversity dynamics over Earth's history. Despite—or even spurred by—this growing availability of resources, analytical paleobiology faces deep-rooted obstacles that stem from the need for more equitable access to data and best practices to guide analyses of the fossil record. Recent progress has been accelerated by a collective push toward more collaborative, interdisciplinary, and open science, especially by early-career researchers. Here, we survey four challenges facing analytical paleobiology from an early-career perspective: (1) accounting for biases when interpreting the fossil record; (2) integrating fossil and modern biodiversity data; (3) building data science skills; and (4) increasing data accessibility and equity. We discuss recent efforts to address each challenge, highlight persisting barriers, and identify tools that have advanced analytical work. Given the inherent linkages between these challenges, we encourage discourse across disciplines to find common solutions. We also affirm the need for systemic changes that reevaluate how we conduct and share paleobiological research.

Fossil bivalve shells are well-suited for landmark/semilandmark morphometric analysis because they preserve both traces of the internal anatomy and the whole shell outline. Utilizing landmarks and semilandmarks, we have characterized internal and external shape variation in a monophyletic clade of Cenozoic New Zealand and Australian crassatellid bivalves, to test the contiguity in morphospace of species-level taxa and to quantitatively examine the “Concept of Independent Entities” of Yonge (1953). Thirteen species from two genera (Spissatella Finlay 1926 and Eucrassatella Iredale 1924) are investigated. Spissatella n. sp. C is confirmed as forming a contiguous group separate to S. trailli and S. clifdenensis. Shell outline and internal anatomy are found to covary in shape, refuting the “Concept of Independent Entities” in the study group.

In a recent contribution, Rightmire (1981) claims that “No significant trends can be observed” in several characters measured in skulls of Homo erectus. While I sympathize with the frustrating problem of small sample size in hominid finds, one must ask the question: could you ever see a trend, given the quality of your information? If sample size is low, if trends are of relatively small magnitude, and if sample variance is high, one can always safely show that nothing (i.e., stasis) has happened. This form of inference falls into a classic statistical trap: attempting to prove statistical homogeneity. You cannot do it without a concomitant estimate of least significant difference, that is, estimating the magnitude of variation within which you cannot distinguish among estimates of a parameter.

This paper investigates trends in the evolution of body size and shape in the Plesiosauria, a diverse clade of Mesozoic marine reptiles. Using measures from well-preserved plesiosaur specimens, we document and interpret evolutionary patterns in relative head size, body size, and locomotor variables. Size increase is a significant trend in the clade as a whole, and in constituent clades. The trend in relative head size is of variance increase; observed head sizes are both smaller and larger than ancestral values. In the locomotor system, changes in propodial and girdle proportions appear concomitant with body size increase and are interpreted as allometric responses to the physical constraints of large body size. Other trends in the locomotor system are significantly correlated with both body size and relative head size. These locomotor trends evolved convergently in several clades of plesiosaurs, and may have had an ecomorphological basis, although data are lacking to constrain speculation on this point. The evolution of the locomotor system in plesiosaurs sheds new light on the response of aquatic tetrapods to the physical constraints of foraging at large body size.

The warm, equable, and ice-free early Eocene Epoch permits investigation of ecosystem function and macro-ecological patterns during a very different climate regime than exists today. It also provides insight into what the future may entail, as anthropogenic CO2 release drives Earth toward a comparable hothouse condition. Studying plant–insect herbivore food webs during hothouse intervals is warranted, because these account for the majority of nonmicrobial terrestrial biodiversity. Here, we report new plant and insect herbivore damage census data from two floodplain sites in the Wind River Basin of central Wyoming, one in the Aycross Formation (50–48.25 Ma) at the basin edge (WRE) and the second in the Wind River Formation in the interior of the basin (WRI). The WRI site is in stratigraphic proximity to a volcanic ash that is newly dated to 52.416 ± 0.016/0.028/0.063 (2σ). We compare the Wind River Basin assemblages to published data from a 52.65 Ma floodplain flora in the neighboring Bighorn (BH) Basin and find that only 5.6% of plant taxa occur at all three sites and approximately 10% occur in both basins. The dissimilar floras support distinct suites of insect herbivores, as recorded by leaf damage. The relatively low-diversity BH flora has the highest diversity of insect damage, contrary to hypotheses that insect herbivore diversity tracks floral diversity. The distinctiveness of the WRE flora is likely due to its younger age and cooler reconstructed paleotemperature, but these factors are nearly identical for the WRI and BH floras. Site-specific microenvironmental factors that cannot be measured easily in deep time may account for these differences. Alternatively, the Owl Creek Mountains between the two basins may have provided a formidable barrier to the thermophilic organisms that inhabited the basin interiors, supporting Janzen's hypothesis that mountain passes appear higher in tropical environments.

Presented here is a coupled model of the nonmarine fossil record, based on a geometric model of deposition, a random-branching model of evolution, and an ecological model based on an elevation gradient. This model provides testable predictions about the stratigraphy and fossil occurrences in coastal nonmarine settings under three scenarios of sea-level change. A slow relative rise in sea level causes a declining ratio of channel to floodplain deposits, plus changes in community composition that reflect an upward increase in elevation relative to sea level. A rapid relative rise in sea level drives increasing aggradation rates, decreases the ratio of channel to floodplain deposits, and triggers a shift from higher-elevation (more inland) to lower-elevation (more coastal) communities. A fall in sea level produces an unconformity, manifested by valleys separated by interfluves. The resumption of deposition following the sea-level fall causes an abrupt shift in community composition across the unconformity, reflecting the duration of the hiatus and the increased elevation relative to sea level. This produces a cluster of first and last occurrences at the unconformity, and it is the only sequence-stratigraphic source of such clusters in a nonmarine system, in contrast to the multiple mechanisms for generating these clusters in marine systems. A central prediction of these models is that the nonmarine fossil record preserves systematic changes in community composition that reflect elevation (or equivalently, distance from shore). Diagnosing these gradients in ancient systems is a promising avenue of future research.