Data that accurately capture the spatial structure of biodiversity are required for many paleobiological questions, from assessments of changing provinciality and the role of geographic ranges in extinction and originations, to estimates of global taxonomic or morphological diversity through time. Studies of temporal changes in diversity and global biogeographic patterns have attempted to overcome fossil sampling biases through sampling standardization protocols, but such approaches must ultimately be limited by available literature and museum collections. One approach to evaluating such limits is to compare results from the fossil record with models of past diversity patterns informed by modern relationships between diversity and climatic factors. Here we use present-day patterns for marine bivalves, combined with data on the geologic ages and distributions of extant taxa, to develop a model for Pliocene diversity patterns, which is then compared with diversity patterns retrieved from the literature as compiled by the Paleobiology Database (PaleoDB). The published Pliocene bivalve data (PaleoDB) lack the first-order spatial structure required to generate the modern biogeography within the time available (<3 Myr). Instead, the published data (raw and standardized) show global diversity maxima in the Tropical West Atlantic, followed closely by a peak in the cool-temperate East Atlantic. Either today's tropical West Pacific diversity peak, double that of any other tropical region, is a purely Pleistocene phenomenon—highly unlikely given the geologic ages of extant genera and the topology of molecular phylogenies—or the paleontological literature is such a distorted sample of tropical Pliocene diversity that current sampling standardization methods cannot compensate for existing biases. A rigorous understanding of large-scale spatial and temporal diversity patterns will require new approaches that can compensate for such strong bias, presumably by drawing more fully on our understanding of the factors that underlie the deployment of diversity today.

Previous genetic studies of extant planktonic foraminifera have provided evidence that the traditional, strictly morphological definition of species in these organisms underestimates their biodiversity. Here, we report the first case where this pattern is reversed. The modern (sub)tropical species plexus Globigerinoides sacculifer is characterized by large morphological variability, which has led to the proliferation of taxonomic names attributed to morphological end-members within the plexus. In order to clarify the taxonomic status of its morphotypes and to investigate the genetic connectivity among its currently partly disjunct (sub)tropical populations, we carried out a global survey of two ribosomal RNA regions (SSU and ITS-1) in all recent morphotypes of the plexus collected throughout (sub)tropical surface waters of the global ocean. Unexpectedly, we find an extremely reduced genetic variation within the plexus and no correlation between genetic and morphological divergence, suggesting taxonomical overinterpretation. The genetic homogeneity within the morphospecies is unexpected, considering its partly disjunct range in the (sub)tropical Atlantic and Indo-Pacific and its old age (early Miocene). A sequence variant in the rapidly evolving ITS-1 region indicates the existence of an exclusively Atlantic haplotype, which suggests an episode of relatively recent (last glacial) isolation, followed by subsequent resumption of unidirectional gene flow from the Indo-Pacific into the Atlantic. This is the first example in planktonic foraminifera where the morphological variability in a morphospecies exceeds its rDNA genetic variability. Such evidence for inconsistent scaling of morphological and genetic diversity in planktonic foraminifera could complicate the interpretation of evolutionary patterns in their fossil record.

The regeneration abilities of crinoids not only are important to understanding crinoid ecology, but also can serve as the basis for assessing the pressure exerted on crinoids by predators both in the Recent and in the geologic past. This is especially true of regenerating arms, because arm loss, and subsequent regeneration, is thought to result from interactions with predators, primarily fish. However, the commonly used regeneration-based proxy for predation pressure (proportion of individuals with regenerating arms) does not provide a measure of the rate at which those events occurred. Here we present a method for reconstructing the arm-loss rate per individual, a more direct proxy of predation pressure. This metric accounts for differences in arm length, arm number, and branching pattern, features highly variable among taxa, among environments, and through geologic time. Normalizing for those characters permits the transformation of observed proportions of regenerating arms to rates that can be compared across samples of morphologically distinct crinoids. Applying this method to a Recent crinoid (Cenometra bella) reveals that this shallow-water comatulid loses arms at a rate of about once every ten days. The same approach reveals that Mississippian shallow-water crinoids (Rhodocrinites kirbyi) experienced arm loss much less frequently, approximately once every 36 days, suggesting that predation pressure on crinoids today is greater than it was in the Mississippian.

Over the past quarter-century there has been considerable innovation in methods for assessing the tempo and mode of evolution in paleobiological data sets. The current literature of these methods centers on three competing hypotheses—stasis, random walk, and directional trend—corresponding to an increasing scaling of variance with time interval (unchanging, for stasis; linear, for random walk; quadratic, for trend). For applications to a single trait there are powerful methods for discriminating among these hypotheses; but for multivariate data sets, especially the very high-dimensional multivariate data arising in image-feature-based and morphometric studies, current statistical approaches appear to be of less help. This paper proves that in the limiting case of high-dimensional morphospaces, the principal component or principal coordinate ordination of every sufficiently lengthy isotropic random walk tends to the same geometrical shape, which is not that of an ellipsoid and for which the principal components or coordinates are not independent even though they are uncorrelated. Specifically, the “scatter” of PC1 against PC2 is just a parabolic curve. The quantitative characteristics of this specific shape are not described appropriately by the corresponding “covariance structure” or Gaussian model, and the discrepancy may be pertinent to much of the existing literature of methods for differentiating among those three models of evolutionary multivariate time series. From a close examination of this common geometry of the ideal random walk model as seen in its principal components, I suggest a test for stasis, along with a mixed model illustrated by a reanalysis of some data of Gunz et al., and a related test for directional trend. These comments are intended to apply to all high-dimensional morphospaces, not just those arising in geometric morphometrics. Applications of principal components in this context distort high-dimensional data in ways that have a tendency to mislead; but these distortions can be intercepted so that studies of tempo and mode can nevertheless proceed.

Mollusks in general and ammonoids in particular are known to display a sometimes profound morphological intraspecific variability of their shell. Although this phenomenon is of greatest importance, it has rarely been investigated and quantified. It is especially crucial for taxonomy and incidentally for biodiversity analyses to account for it, because otherwise, the number of described species might exceed that of actual species within any group. Early ammonoids (Early Devonian, Paleozoic) typically suffer from this bias. For instance, most specimens from the same layer and the same region (e.g., the Erbenoceras beds of the Moroccan eastern Anti-Atlas studied here) differ morphologically from each other. Depending on the importance given to certain morphological characters, therefore, one could create a new species for almost every specimen. In this study, we measured nearly 100 such specimens from a restricted stratigraphic interval and quantified their intraspecific variability. There is a variable but strong overlap of the quantified shell characters at most ontogenetic stages, and only two species are here separated rather than the four previously recognized in Morocco. When ontogenetic trajectories of the Moroccan specimens are compared with coeval faunas from other regions (assigned to other species), a strong overlap between the morphospaces occupied by these taxa becomes apparent. The justification of some of these latter species is thus questionable even if their mean values in some conch parameters differ considerably from the mean values of the Moroccan species. Hence, the number of currently valid species of these loosely coiled early ammonoids is probably much too high. Extreme caution must therefore be taken when examining the diversity of groups in which the intraspecific variability is poorly known.

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.

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.

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.

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.

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.