Evolutionary stasis has often been explained by stabilizing selection, intrinsic constraints, or, more recently, by spatially patterned population dynamics. To distinguish which of these mechanisms explains a given case of stasis in the fossil record, stasis must first be rigorously documented in a high-resolution stratigraphic time series of fossil specimens. Furthermore, past studies of evolutionary mode in fossil mammalian lineages have often been limited to univariate traits (e.g., molar crown area). It is reasonable to assume that tooth shape, a multivariate trait, reflects important additional aspects of tooth form and function. Here we present the results of a geometric morphometric analysis of the lower dentition of the Paleocene-Eocene condylarth species Ectocion osbornianus collected from the Bighorn and Clarks Fork Basins of northwestern Wyoming. Tooth margin shape, cusp configuration, and shearing crest shape were digitized for the last lower premolar, p4, and for two lower molars, m1 and m3. Multivariate statistical tests of evolutionary mode were used to analyze the change in shape variance over time in addition to the magnitude and direction of shape change. Test results characterize the shape time series as consisting of counteracting changes with less change than expected under a random walk (i.e., stasis). The temporal structure of shape variance implies that the sampled E. osbornianus most likely represent a single population, which is not concordant with the population dynamic mechanism of stasis. Stabilizing selection and/or intrinsic constraints remain as the mechanisms that could explain stasis in the lower dental shape of E. osbornianus despite the variable environmental conditions of the Paleocene–Eocene.

Leaf mass per area (MA) is a central ecological trait that is intercorrelated with leaf life span, photosynthetic rate, nutrient concentration, and palatability to herbivores. These coordinated variables form a globally convergent leaf economics spectrum, which represents a general continuum running from rapid resource acquisition to maximized resource retention. Leaf economics are little studied in ancient ecosystems because they cannot be directly measured from leaf fossils. Here we use a large extant data set (65 sites; 667 species-site pairs) to develop a new, easily measured scaling relationship between petiole width and leaf mass, normalized for leaf area; this enables MA estimation for fossil leaves from petiole width and leaf area, two variables that are commonly measurable in leaf compression floras. The calibration data are restricted to woody angiosperms exclusive of monocots, but a preliminary data set (25 species) suggests that broad-leaved gymnosperms exhibit a similar scaling. Application to two well-studied, classic Eocene floras demonstrates that MA can be quantified in fossil assemblages. First, our results are consistent with predictions from paleobotanical and paleoclimatic studies of these floras. We found exclusively low-MA species from Republic (Washington, U.S.A., 49 Ma), a humid, warm-temperate flora with a strong deciduous component among the angiosperms, and a wide MA range in a seasonally dry, warm-temperate flora from the Green River Formation at Bonanza (Utah, U.S.A., 47 Ma), presumed to comprise a mix of short and long leaf life spans. Second, reconstructed MA in the fossil species is negatively correlated with levels of insect herbivory, whether measured as the proportion of leaves with insect damage, the proportion of leaf area removed by herbivores, or the diversity of insect-damage morphotypes. These correlations are consistent with herbivory observations in extant floras and they reflect fundamental trade-offs in plant-herbivore associations. Our results indicate that several key aspects of plant and plant-animal ecology can now be quantified in the fossil record and demonstrate that herbivory has helped shape the evolution of leaf structure for millions of years.

Understanding what drives global diversity requires knowledge of the processes that control diversity and turnover at a variety of geographic and temporal scales. This is of particular importance in the study of mass extinctions, which have disproportionate effects on the global ecosystem and have been shown to vary geographically in extinction magnitude and rate of recovery.

Here, we analyze regional diversity and turnover patterns for the paleocontinents of Laurentia, Baltica, and Avalonia spanning the Late Ordovician mass extinction and Early Silurian recovery. Using a database of genus occurrences for inarticulate and articulate brachiopods, bivalves, anthozoans, and trilobites, we show that sampling-standardized diversity trends differ for the three regions. Diversity rebounded to pre-extinction levels within 5 Myr in the paleocontinent of Laurentia, compared with 15 Myr or longer for Baltica and Avalonia. This increased rate of recovery in Laurentia was due to both lower Late Ordovician extinction rates and higher Early Silurian origination rates relative to the other continents. Using brachiopod data, we dissected the Rhuddanian recovery into genus origination and invasion. This analysis revealed that standing diversity in the Rhuddanian consisted of a higher proportion of invading taxa in Laurentia than in either Baltica or Avalonia. Removing invading genera from diversity counts caused Rhuddanian diversity to fall in Laurentia. However, Laurentian diversity still rebounded to pre-extinction levels within 10 Myr of the extinction event, indicating that genus origination rates were also higher in Laurentia than in either Baltica or Avalonia. Though brachiopod diversity in Laurentia was lower than in the higher-latitude continents prior to the extinction, increased immigration and genus origination rates made it the most diverse continent following the extinction. Higher rates of origination in Laurentia may be explained by its large size, paleogeographic location, and vast epicontinental seas. It is possible that the tropical position of Laurentia buffered it somewhat from the intense climatic fluctuations associated with the extinction event, reducing extinction intensities and allowing for a more rapid rebound in this region. Hypotheses explaining the increased levels of invasion into Laurentia remain largely untested and require further scrutiny. Nevertheless, the Late Ordovician mass extinction joins the Late Permian and end-Cretaceous as global extinction events displaying an underlying spatial complexity.

If last appearances of marine animal genera are taken as reasonable proxies for true extinctions, then there is appreciable global extinction in every stage of the Phanerozoic. If, instead, backsmearing of extinctions by incomplete sampling is explicitly taken into consideration, a different view of extinction emerges, in which the pattern of extinction is much more volatile and in which quiescent time spans—with little or no global extinction for several million years—are punctuated by major extinction events that are even more extreme than is generally thought. Independent support for this alternative view comes from analysis of genus occurrence data in the Paleobiology Database, which agrees with previous estimates of sampling probability and implies that offsets between extinction and last appearance of one or more stages are quite probable.

Carbon and nitrogen stable isotopic compositions of fossil materials from Lake Messel (47 Ma) in Germany are used to investigate Eocene ecosystem dynamics. Autolithified soft tissues of terrestrial and aquatic vertebrate organisms, as well as plant compression fossils, contain organic material (20–50 wt% C, 1–6 wt% N), which appears to retain precursor compositions. Stable isotopic compositions (δ13C and δ15N) of Messel fossils are similar to those reported for components in modern lacustrine ecosystems. These data show trophically sensible enrichments relative to food sources, reflect multiple feeding strategies for each organism (e.g., omnivory, planctivory, piscivory), and differentiate between benthic and pelagic organic carbon sources. These chemical data broadly confirm existing Messel food web models based on coprolite and gut content analyses. δ13C values for the lacustrine shale range from −30.3 to −26.3‰, pointing to mixed terrestrial and aquatic origins for primary producers in the food web. δ13C values for primary consumers such as insects overlap with those for primary producers but are comparatively enriched in 15N. Secondary and higher consumers (fish, crocodiles, and frogs) are associated with even more positive δ15N values and show a more constrained range of δ13C values. Omnivory appears widespread in both low and high trophic level consumers. Hence, the stable isotopic compositions of Messel fossils are complex and overlap, and must be combined with paleontological investigations in order to be conclusive. This study represents the first comprehensive isotopic reconstruction, featuring tens of components, of an ecosystem of Eocene age. A thorough understanding of trophic structure in Eocene Lake Messel contributes to the global databank of ecological history.

Paleobiological diversity is often expressed as α (within-sample), β (among-sample), and γ (total) diversities. However, when studying the effects of extinction on diversity patterns, only variations in α and γ diversities are typically addressed. A null model that examines changes in β diversity as a function of percent extinction is presented here.

The model examines diversity in the context of a hierarchical sampling strategy that allows for the additive partitioning of γ diversity into mean α and β diversities at varying scales. Here, the sampling hierarchy has four levels: samples, beds, facies, and region; thus, there are four levels of α diversity (α1, α2, α3, α4) and three levels of β diversity (β1, β2, and β3). Taxa are randomly assigned to samples within the hierarchy according to probability of occurrence, and initial mean α and β values are calculated. A regional extinction is imposed, and the hierarchy is resampled from the remaining extant taxa. Post-extinction mean α and β values are then calculated.

Both non-selective and selective extinctions with respect to taxon abundance yield decreases in α, β, and γ diversities. Non-selective extinction with respect to taxon abundance shows little effect on diversity partitioning except at the highest extinction magnitudes (above 75% extinction), where the contribution of α1 to total γ increases at the expense of β3, with β1 and β2 varying little with increasing extinction magnitude. The pre-extinction contribution of α1 to total diversity increases with increased probabilities of taxon occurrence and the number of shared taxa between facies. Both β1 and β2 contribute equally to total diversity at low occurrence probabilities, but β2 is negligible at high probabilities, because individual samples preserve all the taxonomic variation present within a facies. Selective extinction with respect to rare taxa indicates a constant increase in α1 and constant decrease in β3 with increasing extinction magnitudes, whereas selective extinction with respect to abundant taxa yields the opposite pattern of an initial decrease in α1 and increase in β3. Both β1 and β2 remain constant with increasing extinction for both cases of selectivity. By comparing diversity partitioning before and after an extinction event, it may be possible to determine whether the extinction was selective with respect to taxon abundances, and if so, whether that selectivity was against rare or abundant taxa.

Field data were collected across a Late Ordovician regional extinction in the Nashville Dome of Tennessee, with sampling hierarchy similar to that of the model. These data agree with the abundant-selective model, showing declines in α, β, and γ diversities, and a decrease in α1 and increase in β3, which suggests this extinction may have targeted abundant taxa.

We assessed selective extinction patterns in bivalves during a late Neogene mass extinction event observed along the temperate Pacific coast of South America. The analysis of 99 late Neogene and Quaternary fossil sites (recorded from 7°S to 55°S), yielding ∼2800 occurrences and 118 species, revealed an abrupt decline in Lyellian percentages during the late Neogene–Pleistocene, suggesting the existence of a mass extinction that decimated ∼66% of the original assemblage. Using the late Neogene data set (n = 59 species, 1346 occurrences), we tested whether the extinction was nonrandom according to taxonomic structure, life habit, geographic range, and body size. Our results showed that the number of higher taxa that went extinct was not different than expected by random. At first sight, extinction was selective only according to life habit and geographic range. Nevertheless, when phylogenetic effects were accounted for, body size also showed significant selectivity. In general, epifaunal, small-sized (after phylogenetic correction), and short-ranged species tended to have increased probability of extinction. This is verified by strong interactions between the variables herein analyzed, suggesting the existence of nonlinear effects on extinction chances. In the heavily decimated epifaunal forms, survival was not enhanced by widespread ranges or larger body sizes. Conversely, the widespread and large-sized infaunal forms tended to have lower probability of extinction. Overall, the ultimate extinction of late Neogene bivalve species along the Pacific coast of South America seems to have been determined by a complex interplay of ecological and historical (phylogenetic) effects.

The process of evolution hinders our ability to make large-scale ecological comparisons—such as those encompassing marine biotas spanning the Phanerozoic—because the compared entities are taxonomically and morphologically dissimilar. One solution is to focus instead on life habits, which are repeatedly discovered by taxa because of convergence. Such an approach is applied to a comparison of the ecological diversity of Paleozoic (Cambrian–Devonian) and modern marine biotas from deep-subtidal, soft-substrate habitats. Ecological diversity (richness and disparity) is operationalized by using a standardized ecospace framework that can be applied equally to extant and extinct organisms and is logically independent of taxonomy. Because individual states in the framework are chosen a priori and not customized for particular taxa, the framework fulfills the requirements of a universal theoretical ecospace. Unique ecological life habits can be recognized as each discrete, n-dimensional combination of character states in the framework. Although the basic unit of analysis remains the organism, the framework can be applied to other entities—species, clades, or multispecies assemblages—for the study of comparative paleoecology and ecology. Because the framework is quantifiable, it is amenable to analytical techniques used for morphological disparity. Using these methods, I demonstrate that the composite Paleozoic biota is approximately as rich in life habits as the sampled modern biota, but that the life habits in the modern biota are significantly more disparate than those in the Paleozoic; these results are robust to taphonomic standardization. Despite broadly similar distributions of life habits revealed by multivariate ordination, the modern biota is composed of life habits that are significantly enriched, among others, in mobility, infaunality, carnivory, and exploitation of other organisms (or structures) for occupation of microhabitats.

Most evolutionary innovations—power-enhancing phenotypes previously absent in a lineage—have arisen multiple times within major clades. This repetition permits a comparative approach to ask how, where, when, in which clades, and under which circumstances adaptive innovations are acquired and secondarily lost. I use new and literature-based data on the phylogeny, functional morphology, and fossil record of gastropods to explore the acquisition and loss of the siphonal canal and its variations in gastropods. The siphonal indentation, canal, notch, or tube at the front end of the shell is associated in living gastropods with organs that detect chemical signals directionally and at a distance in an anteriorly restricted inhalant stream of water.

Conservative estimates indicate that the siphonate condition arose 23 times and was secondarily lost 17 times. Four siphonate clades have undergone prodigious diversification. All siphonate gastropods have a shell whose axis of coiling lies at a low angle above the plane of the aperture (retroaxial condition). In early gastropods, the siphonal canal was short and more or less confined to the apertural plane. Later (mainly Cretaceous and Cenozoic) variations include a dorsally deflected canal, a long canal, and a closed canal. The closed siphonal canal, in which the edges join to form a tube, arose 15 times, all in the adult stages of caenogastropods with determinate growth.

Gastropods in which the siphonate condition arose were mobile, bottom-dwelling, microphagous animals. Active predaceous habits became associated with the siphonate condition in the Mesozoic and Cenozoic Purpurinidae-Latrogastropoda clade. Loss of the siphonate condition is associated with nonmarine habits, miniaturization, and especially with a sedentary or slow-moving mode of life.

The siphonate condition arose seven times each during the early to middle Paleozoic, the late Paleozoic, and the early to middle Mesozoic, and only once each during the Late Cretaceous and Cenozoic. Well-adapted incumbents prevented most post-Jurassic clades from evolving a siphonal indentation and its associated organs. Dorsally deflected, long, and closed canals are known only from Cretaceous and Cenozoic marine gastropods, and represent improvements in sensation and passive armor.

In a discussion of contrasting ecologies of clades that gained and lost the siphonate condition, I argue that macroevolutionary trends in the comings and goings of innovations and clades must incorporate ecological and functional data. Acquisitions of energy-intensive, complex innovations that yield greater power have a greater effect on ecosystems, communities, and their resident clades than do reversals, which generally reflect energy savings.

Tectonic deformation is an important part of the taphonomic histories of many fossils. Although the effects of deformation, and methods to remove those effects, have been a subject of inquiry for over a century, systematic testing under known parameters has never been used to determine how the effects of deformation and the performance of retrodeformation techniques might vary. Comparative studies of morphology depend on the accurate estimation of variance-covariance structure, so an understanding of the effects of retrodeformation on covariance structure is important in assessing the utility of these methods. Here we address these issues by using geometric morphometric simulations. Nondeformed data sets were generated from specimens of the extant turtle Emys marmorata, which were known by definition to be nondeformed, and which possess a known ontogenetic signal. Deformation was simulated by applying a combination of uniform shear and uniform compression/dilation to the data. Data were retrodeformed by reflection and averaging of bilaterally symmetric landmarks, use of a principal components analysis to identify a deformation component of shape variation, and removal of the affine component of shape variation among specimens. Deformation increased the amount of variance in the data, as well as altering the variance structure. However, low to moderate levels of deformation did not prevent the confident recovery of the known ontogenetic signal in some cases. The tested retrodeformation techniques did not work well. They either removed too little or too much variance from the data, and provided little improvement in variance structure. Retrodeformation often did not improve our ability to extract the ontogenetic signal from the data, and in some cases introduced an arti-factual relationship between size and shape. All of the scrutinized methods showed some properties, such as reducing variance or producing visually appealing images of specimens, that could make them appear to be working in cases where the correct biological signal is not known. This emphasizes the need for simulation testing in the development and evaluation of retrodeformation techniques.

Tyrannosaurus rex and other tyrannosaurid theropods exerted high bite forces, and large muscle attachments suggest that the tyrannosaurid neck was a concomitantly powerful component of the feeding apparatus. We examine accelerative and work-generating capacity (WGC) of neck muscles in adult Tyrannosaurus rex, using a 3-D vector-based method that incorporates aspects of muscle force generation, reconstruction of muscle morphology and moment arms, and rotational inertias of the head and neck. Under conservative assumptions, radial accelerations of the head by large superficial muscles (M. transversospinalis capitis, M. complexus, and M. longissimus capitis superficialis) enabled rapid gaze shifts and imparted high tangential velocities to food sufficient for inertial feeding. High WGC by these and deeper muscles under eccentric contraction indicate high efficacy for tearing flesh, especially with the head and neck in an extended posture. Sensitivity analyses suggest that assigned density of the antorbital region has substantial effects on calculated rotational inertia, and hence on the accuracy of results. However, even with high latitude for estimation errors, the results indicate that adult T. rex could strike rapidly at prey and engage in complexly modulated inertial feeding, as seen in extant archosaurs.

Biotic invasions are a common feature of the fossil record, yet remarkably little is known about them, given their enormous potential to reveal the processes that regulate local and regional diversity over long time scales. We used additive diversity partitioning to examine how diversity structure changed as a result of a marine biotic invasion in tropical, shallow and deep subtidal environments spanning approximately 4 Myr in the Late Ordovician. The biotic invasion increased richness in the regional ecosystem by nearly 40%. Within-habitat turnover diversity accounts for most of the increase in richness, with between-habitat turnover diversity contributing a lesser amount. Increases in these components of diversity were accommodated by increased packing of species along a depth gradient and increased habitat heterogeneity. Diversity metrics that incorporate taxon abundance (Shannon information, Simpson's D) show similar patterns and reveal that many invading taxa were locally abundant and widespread in their occurrence. Extinction of incumbent taxa did not foster the invasion; rather the invasion appears to be linked to a regional or global warming event. Taken together, these observations indicate that these Late Ordovician marine communities were open to invasion and not saturated with species. Moreover, the increase in species diversity caused by the invasion was not ephemeral; instead it lasted for at least 1 Myr. Similar studies of other biotic invasions in the fossil record are necessary to determine (1) the factors, such as extinction of incumbents or resource limitation, that may facilitate or inhibit invasion in ancient ecosystems; (2) how local and regional ecosystems respond to invasion; and (3) the extent to which biotic invasions play a substantial role in ecosystem change through geologic time.

Echinoderms have long been characterized by the presence of ambulacra that exhibit pentaradiate symmetry and define five primary body axes. In reality, truly pentaradial ambulacral symmetry is a condition derived only once in the evolutionary history of echinoderms and is restricted to eleutherozoans, the clade that contains most living echinoderm species. In contrast, early echinoderms have a bilaterally symmetrical 2-1-2 arrangement, with three ambulacra radiating from the mouth. Branching of the two side ambulacra during ontogeny produces the five adult rays. During the Cambrian Explosion and Ordovician Radiation, some 30 clades of echinoderms evolved, many of which have aberrant ambulacral systems with one to four rays. Unfortunately, no underlying model has emerged that explains ambulacral homologies among disparate forms. Here we show that most Paleozoic echinoderms are characterized by uniquely identifiable ambulacra that develop in three distinct postlarval stages. Nearly all “aberrant” echinoderm morphologies can be explained by the paedomorphic ambulacra reduction (PAR) model through the loss of some combination of these growth stages during ontogeny. Superficially similar patterns of ambulacral reduction in distantly related clades have resulted from the parallel loss of homologous ambulacra during ontogeny. Pseudo-fivefold symmetry seen in Blastoidea and the true fivefold symmetry seen in Eleutherozoa result from great reduction and total loss, respectively, of the 2–1–2 symmetry early in ontogeny. These ambulacral variations suggest that both developmental and ecological constraints affect the evolution of novel echinoderm body plans.

Four independent lines of evidence, (1) the quality of specimen preservation, (2) taxonomic collection curves, (3) molecular divergence estimates, and (4) ghost lineage analysis of a genus-level cladogram, point to echinoids having a much poorer fossil record in the Triassic than in the Lower Jurassic. Furthermore, preservational differences between Triassic and Lower Jurassic echinoids have remained a consistent feature over 160 years of discovery. Differences exist in how effectively paleontologists have collected the fauna from available outcrops in the Triassic and Lower Jurassic. Collection curves suggest that rocks have been more efficiently searched for their fossils in Europe than elsewhere in the world, and that Lower Jurassic faunas are better sampled from available outcrop than Triassic faunas. The discovery of Triassic taxa has quickened in pace over the past 4 decades (though largely driven by a single Lagerstätte—the St. Cassian beds) while discoveries of new taxa from the Lower Jurassic have slowed. Molecular analysis of extant families and ghost lineage analysis of Triassic and Lower Jurassic genera both point to poorer sampling of Triassic faunas. This difference in the quality of the fossil record may be partially explained by differences in rock outcrop area, as marine sedimentary rocks are much less common in the Triassic than in the Lower Jurassic. However, improving biomechanical design of the echinoid test over this critical time interval was probably as important, and better explains observed preservational trends. Changes in the quality of the echinoid fossil record were thus driven as much by intrinsic biological factors as by sampling patterns.

Many authors have proposed scenarios for mass extinctions that consist of multiple pulses or stages, but little work has been done on accounting for the Signor-Lipps effect in such extinction scenarios. Here we introduce a method for computing confidence intervals for the time or stratigraphic distance separating two extinction pulses in a pulsed extinction event, taking into account the incompleteness of the fossil record. We base our method on a flexible likelihood ratio test framework that is able to test whether the fossil record is consistent with any extinction scenario, whether simultaneous, pulsed, or otherwise. As an illustration, we apply our method to a data set on marine invertebrates from the Permo-Triassic boundary of Meishan, China. Using this data set, we show that the fossil record of ostracodes and that of brachiopods are each consistent with simultaneous extinction, and that these two extinction pulses are separated by 720,000 to 1.2 million years with 95% confidence. With appropriate data, our method could also be applied in other situations, such as tests of origination patterns, coordinated stasis, and recovery after a mass extinction.