strap (Stratigraphic Tree Analysis for Palaeontology) is a new package for the freely available statistical programming language R designed to perform three main tasks: (1) to time-scale phylogenies of fossil taxa; (2) to plot those time-scaled trees against stratigraphy; and (3) to assess congruence between phylogenies and stratigraphy. Time-scaling is performed with the DatePhylo function, with three approaches offered. Plotting trees against a choice of five different geological time scales is possible using the geoscalePhylo function. Finally, the function StratPhyloCongruence calculates stratigraphic congruence measures for one or more input phylogenies, with no taxon limit. All three major congruence measures are offered: Stratigraphic Consistency Index (SCI), Manhattan Stratigraphic Measure (MSM*) and the gap excess ratio (GER; including GERt and GER*), as well as the pseudocongruence measure, the Relative Completeness Index (RCI). Each measure has an accompanying significance test that works by comparing the input trees against a userdefined number of randomly generated topologies with the same taxon set and age ranges. Additional options for generating these random topologies allow the user to fix the outgroup or retain the input tree shape to make fairer comparisons. A tutorial that assumes no prior knowledge of R showcases all three functions using two different example data sets.

The geographic distribution of life on Earth supports a general pattern of increase in biodiversity with increasing temperature. However, some previous analyses of the 540-million-year Phanerozoic fossil record found a contrary relationship, with paleodiversity declining when the planet warms. These contradictory findings are hard to reconcile theoretically. We analyze marine invertebrate biodiversity patterns for the Phanerozoic Eon while controlling for sampling effort. This control appears to reverse the temporal association between temperature and biodiversity, such that taxonomic richness increases, not decreases, with temperature. Increasing temperatures also predict extinction and origination rates, alongside other abiotic and biotic predictor variables. These results undermine previous reports of a negative biodiversity-temperature relationship through time, which we attribute to paleontological sampling biases. Our findings suggest a convergence of global scale macroevolutionary and macroecological patterns for the biodiversity-temperature relationship.climate change | Court Jester | mass extinction | Red Queen | rock record B eyond small geographical scales, biodiversity consistently decreases with latitude (1-3), reflecting a strong, probably causal, association between warmer climates and standing richness in both the terrestrial, water availability permitting (4), and marine realms (5, 6). Our understanding of the association between biodiversity and warm climates in space contrasts strongly with our models of how climate explains global diversity through time (7). One analysis of compendia of fossil taxa suggests that biodiversity declines with increasing global temperatures (8), but the focus on temperature as the driver, without reference to other variables, has drawn criticism (7). Analyses at smaller scales (geographic, temporal, or taxonomic) are equivocal (9). How can it be that warm temperatures should apparently have negative effects on biodiversity through time while also having positive effects across space (10) (i.e., contrasting macroevolutionary and macroecological patterns)? Recently, however, our understanding of past changes in biodiversity has been transformed by the application of techniques to control for sampling bias in paleodiversity data (11). Here we apply more-robust measures of fossil diversity, origination, and extinction through time to reevaluate the role of temperature in the context of other potential environmental drivers.Much effort to understand macroevolutionary changes through the Phanerozoic has focused on marine invertebrates, first through Sepkoski's genus-level compendium (12) and latterly via the Paleobiology Database Project (PaleoDB) (11). Putative factors proposed to drive temporal fluctuation in biodiversity include biotic drivers, such as competition between taxa (13, 14) and predation intensity (15). Alternatively, abiotic variables such as sea level change (16,17), nutrient inputs and shelf redox conditions (17, 18), plate tectonic events (19), volcan...

Metriorhynchids were a peculiar group of fully marine Mesozoic crocodylomorphs, some of which reached large body size and were probably apex predators. The estimation of their total body length in the past has proven problematic. Rigorous size estimation was provided using five complete metriorhynchid specimens, by means of regression equations derived from basicranial and femoral length against total body length. The use of the Alligator femoral regression equation as a proxy to estimate metriorhynchid total body length led to a slight underestimation, whereas cranial regression equations of extant genera resulted in an overestimation of body length. Therefore, the scaling of crania and femora to total body length of metriorhynchids is noticeably different from that of extant crocodylians, indicating that extant crocodylians are not ideal proxies for size reconstruction of extinct taxa that deviate from their semi-aquatic morphotype. The lack of a correlation between maximum, minimum, or the range of generic body lengths with species richness demonstrates that species diversification is driven by factors other than just variation in body size. Maximum likelihood modelling also found no evidence for directionality in body size evolution. However, niche partitioning in Metriorhynchidae is mediated not only by craniodental differentiation, as shown by previous studies, but also by body size variation.

There is a well-established discrepancy between paleontological and molecular data regarding the timing of the origin and diversification of placental mammals. Molecular estimates place interordinal diversification dates in the Cretaceous, while no unambiguous crown placental fossils have been found prior to the end-Cretaceous mass extinction. Here, the completeness of the eutherian fossil record through geological time is evaluated to assess the suggestion that a poor fossil record is largely responsible for the difference in estimates of placental origins. The completeness of fossil specimens was measured using the character completeness metric, which quantifies the completeness of fossil taxa as the percentage of phylogenetic characters available to be scored for any given taxon. Our data set comprised 33 published cladistic matrices representing 445 genera, of which 333 were coded at the species level.There was no significant difference in eutherian completeness across the Cretaceous/Paleogene (K/Pg) boundary. This suggests that the lack of placental mammal fossils in the Cretaceous is not due to a poor fossil record but more likely represents a genuine absence of placental mammals in the Cretaceous. This result supports the “explosive model” of early placental evolution, whereby placental mammals originated around the time of the K/Pg boundary and diversified soon after.No correlation was found between the completeness pattern observed in this study and those of previous completeness studies on birds and sauropodomorph dinosaurs, suggesting that different factors affect the preservation of these groups. No correlations were found with various isotope proxy measures, but Akaike information criterion analysis found that eutherian character completeness metric scores were best explained by models involving the marine-carbonate strontium-isotope ratios (87Sr/86Sr), suggesting that tectonic activity might play a role in controlling the completeness of the eutherian fossil record.

Metriorhynchid crocodylomorphs were the only group of archosaurs to fully adapt to a pelagic lifestyle. During the Jurassic and Early Cretaceous, this group diversified into a variety of ecological and morphological types, from large super-predators with a broad short snout and serrated teeth to specialized piscivores/teuthophages with an elongate tubular snout and uncarinated teeth. Here, we use an integrated repertoire of geometric morphometric (form), biomechanical finite-element analysis (FEA; function) and phylogenetic data to examine the nature of craniofacial evolution in this clade. FEA stress values significantly correlate with morphometric values representing skull length and breadth, indicating that form and function are associated. Maximum-likelihood methods, which assess which of several models of evolution best explain the distribution of form and function data on a phylogenetic tree, show that the two major metriorhynchid subclades underwent different evolutionary modes. In geosaurines, both form and function are best explained as evolving under 'random' Brownian motion, whereas in metriorhynchines, the form metrics are best explained as evolving under stasis and the function metric as undergoing a directional change (towards most efficient low-stress piscivory). This suggests that the two subclades were under different selection pressures, and that metriorhynchines with similar skull shape were driven to become functionally divergent.

During and after the Cambrian explosion, very large marine invertebrate species have evolved in several groups. Gigantism in Carboniferous land invertebrates has been explained by a peak in atmospheric oxygen concentrations, but Palaeozoic marine invertebrate gigantism has not been studied empirically and explained comprehensively. By quantifying the spatiotemporal distribution of the largest representatives of some of the major marine invertebrate clades (orthoconic cephalopods, ammonoids, trilobites, marine eurypterids), we assessed possible links between environmental parameters (atmospheric or oceanic oxygen concentrations, ocean water temperature or sea level) and maximum body size, but we could not find a straightforward relationship between both. Nevertheless, marine invertebrate gigantism within these groups was temporally concentrated within intervals of high taxonomic diversity (Ordovician, Devonian) and spatially correlated with latitudes of high occurrence frequency. Regardless of whether temporal and spatial variation in sampled diversity and occurrence frequency reflect true biological patterns or sampling controls, we find no evidence that the occurrences of giants in these groups were controlled by optimal conditions other than those that controlled the group as a whole; if these conditions shift latitudinally, occurrences of giants will shift as well. It is tempting to attribute these shifts to contemporary changes in temperature, oxygen concentrations in the atmosphere and the oceans as well as global palaeogeography over time, but further collection‐based studies are necessary on finer stratigraphic and phylogenetic resolution to corroborate such hypotheses and rule out sampling or collection biases.

Aim The aim was to investigate those factors that influenced the differentiation of high‐latitude and polar marine faunas on both ecological and evolutionary time‐scales. Can a focus on a greenhouse world provide some important clues? Location World‐wide, but with particular emphasis on the evolution of Antarctic marine faunas. Time period Early Cenozoic era and present day. Major taxa studied Mollusca, especially Neogastropoda. Methods The Early Cenozoic global radiation of one of the largest extant marine clades, Neogastropoda, was examined, and detailed comparisons were made between two tropical localities and Antarctica. High‐ to low‐latitude faunal differentiation was assessed using Sørensen's dissimilarity index, and component species in each of the three faunas were assigned to 29 families and family groups. Relative diversity distributions were fitted to these three faunas and two modern ones to assess the contrast in evenness between high‐ and low‐latitude assemblages. Results By the Middle Eocene, a distinct high‐latitude neogastropod fauna had evolved in Antarctica. In addition, the distribution of species within families in this fauna is statistically significantly less even than that in the tropics. Indeed, there is no detectable difference in the scale of this separation from that seen today. Exactly as in the modern fauna, Middle Eocene Antarctic neogastropods are dominated by a small number of trophic generalist groups. Main conclusions As the hyperdiverse Neogastropoda clade radiated globally through the Early Cenozoic, it differentiated into distinct high‐ and low‐latitude components. The fact that it did so in a greenhouse world strongly suggests that something else besides temperature was involved in this process. The predominance of generalist feeding types in the Antarctic fossil faunas is linked to the phenomenon of a seasonally pulsed food supply, exactly as it is today. Seasonality in primary productivity may act as a fundamental control on the evolution of large‐scale biodiversity patterns.

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