Rates of speciation and extinction are often linked to many ecological factors, traits (emergent and nonemergent) such as environmental tolerance, body size, feeding type, and geographic range. Marine gastropods in particular have been used to examine the role of larval dispersal in speciation. However, relatively few studies have been conducted placing larval modes in species-level phylogenetic context. Those that have, have not incorporated fossil data, while landmark macroevolutionary studies on fossil clades have not considered both phylogenetic context and net speciation (speciation–extinction) rates. This study utilizes Eocene volutid Volutospina species from the U.S. Gulf Coastal Plain and the Hampshire Basin, U.K., to explore the relationships among larval mode, geographic range, and duration. Based on the phylogeny of these Volutospina, we calculated speciation and extinction rates in order to compare the macroevolutionary effects of larval mode. Species with planktotrophic larvae had a median duration of 9.7 Myr, which compared significantly to 4.7 Myr for those with non-planktotrophic larvae. Larval mode did not significantly factor into geographic-range size, but U.S. and U.K. species do differ, indicating a locality-specific component to maximum geographic-range size. Non-planktotrophs (NPTs)were absent among the Volutospina species during the Paleocene–early Eocene. The relative proportions of NPTs increased in the early middle Eocene, and the late Eocene was characterized by disappearance of planktotrophs (PTs). The pattern of observed lineage diversity shows an increasing preponderance of NPTs; however, this is clearly driven by a dramatic extinction of PTs, rather than higher NPT speciation rates during the late Eocene. This study adds nuance to paleontology's understanding of the macroevolutionary consequences of larval mode.

Sharks have a long and rich fossil record that consists predominantly of isolated teeth due to the poorly mineralized cartilaginous skeleton. Tiger sharks (Galeocerdo), which represent apex predators in modern oceans, have a known fossil record extending back into the early Eocene (ca. 56 Ma) and comprise 22 recognized extinct and one extant species to date. However, many of the fossil species remain dubious, resulting in a still unresolved evolutionary history of the tiger shark genus. Here, we present a revision of the fossil record of Galeocerdo by examining the morphological diversity and disparity of teeth in deep time. We use landmark-based geometric morphometrics to quantify tooth shapes and qualitative morphological characters for species discrimination. Employing this combined approach on fossil and extant tiger shark teeth, our results only support six species to represent valid taxa. Furthermore, the disparity analysis revealed that diversity and disparity are not implicitly correlated and that Galeocerdo retained a relatively high dental disparity since the Miocene despite its decrease from four to one species. With this study, we demonstrate that the combined approach of quantitative geometric morphometric techniques and qualitative morphological comparisons on isolated shark teeth provides a useful tool to distinguish between species with highly similar tooth morphologies.

Interpreting the impact of climate change on vertebrates in the fossil record can be complicated by the effects of potential biotic drivers on morphological patterns observed in taxa. One promising area where this impact can be assessed is a high-resolution terrestrial record from the Bighorn Basin, Wyoming, that corresponds to the Paleocene–Eocene thermal maximum (PETM), a geologically rapid (~170 kyr) interval of sustained temperature and aridity shifts about 56 Ma. The PETM has been extensively studied, but different lines of research have not yet been brought together to compare the timing of shifts in abiotic drivers that include temperature and aridity proxies and those of biotic drivers, measured through changes in floral and faunal assemblages, to the timing of morphological change within mammalian species lineages. We used a suite of morphometric tools to document morphological changes in molar crown morphology of three lineages of stem erinaceid eulipotyphlans. We then compared the timing of morphological change to that of both abiotic and other biotic records through the PETM. In all three species lineages, we failed to recover any significant changes in tooth crown shape or size within the PETM. These results contrast with those documented previously for lineages of medium-sized mammals, which show significant dwarfing within the PETM. Our results suggest that biotic drivers such as shifts in community composition may have also played an important role in shaping species-level patterns during this dynamic interval in Earth history.

Lanternfishes currently represent one of the dominant groups of mesopelagic fishes in terms of abundance, biomass, and diversity. Their otolith record dominates pelagic sediments below 200 m in dredges, especially during the entire Neogene. Here we provide an analysis of their diversity and rise to dominance primarily based on their otolith record. The earliest unambiguous fossil myctophids are known based on otoliths from the late Paleocene and early Eocene. During their early evolutionary history, myctophids were likely not adapted to a high oceanic lifestyle but occurred over shelf and upper-slope regions, where they were locally abundant during the middle Eocene. A distinct upscaling in otolith size is observed in the early Oligocene, which also marks their earliest occurrence in bathyal sediments. We interpret this transition to be related to the change from a halothermal deep-ocean circulation to a thermohaline regime and the associated cooling of the deep ocean and rearrangement of nutrient and silica supply. The early Oligocene myctophid size acme shows a remarkable congruence with diatom abundance, the main food resource for the zooplankton and thus for myctophids and whales. The warmer late Oligocene to early middle Miocene period was characterized by an increase in disparity of myctophids but with a reduction in their otolith sizes. A second and persisting secular pulse in myctophid diversity (particularly within the genus Diaphus) and increase in size begins with the “biogenic bloom” in the late Miocene, paralleled with diatom abundance and mysticete gigantism.

Cervids living in high latitudes have evolved to thrive in ecosystems that experience dramatic seasonal changes. Understanding these seasonal adaptations is important for reconstructing cervid life histories, ecosystem dynamics, and responses in the distant and not-so-distant past to changing seasonality caused by climate change. Cervid antlers provide a rare opportunity for insight into faunal seasonal ecology, as they are grown and shed each year. Stable isotopes of carbon and nitrogen measured directly from antlers have the potential to provide seasonal dietary data for individuals. If the isotopic signals in bone and antler are controlled by the same metabolic processes, then the stable carbon and nitrogen isotope compositions of collagen (δ13CColl and δ15NColl) from incrementally grown antler tissue provide time-constrained dietary signals from the spring and summer growth season. Bone, by comparison, provides an average signal over several years. The amino acid (glutamate and phenylalanine) δ15N in antlers from modern captive caribou showed similar trophic discrimination factors to earlier results for other collagenous tissues (bone, tooth dentin, and cementum). Hence, growth rate was not the primary control on the stable isotope composition of antler collagen. We applied this knowledge to assess seasonal shifts in Quaternary fossils of three Cervidae species: elk (Cervus elaphus), moose (Alces alces), and caribou (Rangifer tarandus). Paired antler–bone δ13CColl and δ15NColl from the same individual were used to identify differences between summer and annual diet and ecology. Intra-antler isotopic variability from serially sampled antlers was used to examine seasonal dietary shifts and specialization.

A short stratigraphic interval near Bulin in western Hunan (China) yields multiple specimens of the ~514-Myr-old oryctocarine trilobite Oryctocarella duyunensis. Size data obtained from these specimens indicate that, from meraspid degree 1 onward, degrees represent successive instars. Meraspid growth persisted until a terminal stage was reached, providing the first example of determinate growth in trilobites and, notably, in an early Cambrian species. The sample contains three varieties of such terminal stages, recognized as holaspids, with 9, 10, or 11 thoracic segments, respectively. During the meraspid phase, growth rates were not constant in this species. The pattern of growth seen in the Bulin assemblage differs modestly from that reported in the same species from two other localities, attesting to microevolutionary variation in developmental patterns among these collections.

An increasing number of unexpectedly diverse benthic communities are being reported from microbially precipitated carbonate facies in shallow-marine platform settings after the end-Permian mass extinction. Ostracoda, which was one of the most diverse and abundant metazoan groups during this interval, recorded its greatest diversity and abundance associated with these facies. Previous studies, however, focused mainly on taxonomic diversity and, therefore, left room for discussion of paleoecological significance. Here, we apply a morphometric method (semilandmarks) to investigate morphological variance through time to better understand the ecological consequences of the end-Permian mass extinction and to examine the hypothesis that microbial mats played a key role in ostracod survival. Our results show that taxonomic diversity and morphological disparity were decoupled during the end-Permian extinction and that morphological disparity declined rapidly at the onset of the end-Permian extinction, even though the high diversity of ostracods initially survived in some places. The decoupled changes in taxonomic diversity and morphological disparity suggest that the latter is a more robust proxy for understanding the ecological impact of the extinction event, and the low morphological disparity of ostracod faunas is a consequence of sustained environmental stress or a delayed post-Permian radiation. Furthermore, the similar morphological disparity of ostracods between microbialite and non-microbialite facies indicates that microbial mats most likely represent a taphonomic window rather than a biological refuge during the end-Permian extinction interval.

Analyses of ancient food webs reveal important paleoecological processes and responses to a range of perturbations throughout Earth's history, such as climate change. These responses can inform our forecasts of future biotic responses to similar perturbations. However, previous analyses of ancient food webs rarely accounted for key differences between modern and ancient community data, particularly selective loss of soft-bodied taxa during fossilization. To consider how fossilization impacts inferences of ancient community structure, we (1) analyzed node-level attributes to identify correlations between ecological roles and fossilization potential and (2) applied selective information loss procedures to food web data for extant systems. We found that selective loss of soft-bodied organisms has predictable effects on the trophic structure of “artificially fossilized” food webs because these organisms occupy unique, consistent food web positions. Fossilized food webs misleadingly appear less stable (i.e., more prone to trophic cascades), with less predation and an overrepresentation of generalist consumers. We also found that ecological differences between soft- and hard-bodied taxa—indicated by distinct positions in modern food webs—are recorded in an early Eocene web, but not in Cambrian webs. This suggests that ecological differences between the groups have existed for ≥48 Myr. Our results indicate that accounting for soft-bodied taxa is vital for accurate depictions of ancient food webs. However, the consistency of information loss trends across the analyzed food webs means it is possible to predict how the selective loss of soft-bodied taxa affects food web metrics, which can permit better modeling of ancient communities.

Extant and extinct terebratulide brachiopod species have been defined primarily on the basis of morphology. What is the fidelity of morphological species to biological species? And how can we test this fidelity with fossils? Taxonomically and phylogenetically, the most informative internal feature in the brachiopod suborder Terebratellidina is the geometrically complex long-looped brachidium, which is highly fragile and only rarely preserved in the fossil record. Given this, it is essential to test other sources of morphological data, such as valve outline shape, when trying to recognize and identify species. We analyzed valve outlines and brachidia in the genus Laqueus to explore the utility of shell shape in discriminating extant and fossil species. Using geometric morphometric methods, we quantified valve outline variability using elliptical Fourier methods and tested whether long-looped brachidial morphology correlates with shell outline shape. We then built classification models based on machine learning algorithms using outlines as shape variables to predict fossil species’ identities. Our results demonstrate that valve outline shape is significantly correlated with long-looped brachidial shape and that even relatively simple outlines are sufficiently morphologically distinct to enable extant Laqueus species to be identified, validating current taxonomic assignments. These are encouraging results for the study and delimitation of fossil terebratulide species, and their recognition as biological species. In addition, machine learning algorithms can be successfully applied to help solve species recognition and delimitation problems in paleontology, especially when morphology can be characterized quantitatively and analyzed statistically.

A key question in paleoecology and macroevolution is whether assemblages of species (paleocommunities) are persistent entities that endure over millions of years. While community turnover in the face of abiotic change is the presumed norm, paleocommunities have been shown to persist for long time periods and regardless of environmental disruption. It remains an open question, however, as to what processes allow for this. We investigate these questions by analyzing the Carboniferous brachiopod paleocommunities from the Midcontinent of North America. These diverse communities were subjected to repeated and geologically rapid changes in sea level. Using a suite of statistical techniques, we characterize the nature and scope of changes in these paleocommunities over time. We find that, at the paleocommunity scale, there is no evidence for obdurate ecological stasis, with fluctuations in both taxonomic composition and the associated abundance of taxa. However, at a higher ecological scale, stability is manifest, as diversity patterns remain stable across time, with a consistent number of species that can exist in any given paleocommunity. This suggests ecological rules such as taxon packing are in effect, resulting in a form of ecological stability even in the face of constant disequilibrium, and parallels ecological patterns of disruption and recovery previously observed for invertebrate communities from modern marine systems. Based on these results, we advocate for consideration of different hierarchical entities and scales when interpreting the ecological dynamics of fossil assemblages, as focusing exclusively on changes in taxon identity/abundance or diversity levels can lead to very different results.

Geographic range size and abundance are important determinants of extinction risk in fossil and extant taxa. However, the relationship between these variables and extinction risk has not been tested extensively during evolutionarily “quiescent” times of low extinction and speciation in the fossil record. Here we examine the influence of geographic range size and abundance on extinction risk during the late Paleozoic (Mississippian–Permian), a time of “sluggish” evolution when global rates of origination and extinction were roughly half those of other Paleozoic intervals. Analyses used spatiotemporal occurrences for 164 brachiopod species from the North American midcontinent. We found abundance to be a better predictor of extinction risk than measures of geographic range size. Moreover, species exhibited reductions in abundance before their extinction but did not display contractions in geographic range size. The weak relationship between geographic range size and extinction in this time and place may reflect the relative preponderance of larger-ranged taxa combined with the physiographic conditions of the region that allowed for easy habitat tracking that dampened both extinction and speciation. These conditions led to a prolonged period (19–25 Myr) during which standard macroevolutionary rules did not apply.

Heterochrony describes acceleration, displacement, and/or retardation of descendants’ development events compared with ancestral states and has often been cited as an important process to bring about morphological novelty. It was coined one-and-a-half centuries ago and has been discussed by both paleobiologists and biologists frequently ever since. Many types of fossil organisms preserve aspects of their development histories in their bones or shells that have been used for heterochrony analyses, with body size being used as a developmental age indicator, despite questions being raised regarding this practice. For organisms whose hard structures consist of multiple chambers, or that contain growth lines, age information suggested by these structures independently can facilitate ontogenetic modeling. In this way, relations among size, shape, and age can be established to document patterns of morphological development.

Morphological analysis of pseudoschwagerine fusulinids, a fossil foraminifera group that developed a morphologically novel spherical shell, along with their presumptive triticitid ancestors illustrates this approach to heterochrony analysis. Ontogenetic trajectory comparisons of four major pseudoschwagerine genera, as well as those of triticitids, document relations between their shapes, sizes, and developmental ages. A complex of heterochronic patterns, including peramorphic predisplacement, hypermorphosis, and acceleration, characterize pseudoschwagerine development and appear to be responsible for the novel appearance of large, inflated fusiform and spherical tests in these late Paleozoic benthic foraminifera. The morphometric approach employed in this investigation could be applied widely in the quantitative morphological studies of development histories in a variety of other fossil groups.

Brachiopods dominated the seafloor as a primary member of the Paleozoic fauna. Despite the devastating effects of the end-Permian extinction, the group recovered during the early Mesozoic only to gradually decline from the Jurassic to today. This decline likely had multiple causes, including increased predation and bioturbation-driven substrate disruption, but the role of changing substrate is not well understood. Given the importance of substrate for extant brachiopod habitat, we documented Mesozoic–Cenozoic lithologic preferences and morphological changes to assess how decreasing firm-substrate habitat may have contributed to the brachiopod decline. Compared with bivalves, Mesozoic brachiopods occurred more frequently and were disproportionately abundant in carbonate lithologies. Although patterns in glauconitic or ferruginous sediments are equivocal, brachiopods became more abundant in coarser-grained carbonates and less abundant in fine-grained siliciclastics. During the Jurassic, brachiopod species rarely had abraded beaks but tended to be more convex with a high beak, potentially consistent with a non-analogue lifestyle resting on the seafloor. However, those highly convex morphotypes largely disappeared by the Cenozoic, when more terebratulides had abraded beaks, suggesting closer attachment to hard substrates. Rhynchonellides disproportionately declined to become a minor component of Cenozoic faunas, perhaps because of less pronounced morphological shifts. Trends in lithologic preferences and morphology are consistent with bioturbation-driven substrate disruption, with brachiopods initially using firmer carbonate sediments as refugia before adapting to live primarily attached to hard surfaces. This progressive habitat restriction likely played a role in the final brachiopod decline, as bioturbating ecosystem engineers transformed benthic habitats in the Mesozoic and Cenozoic.

The intersection of paleontology and biomechanics can be reciprocally illuminating, helping to improve paleobiological knowledge of extinct species and furthering our understanding of the generality of biomechanical principles derived from study of extant species. However, working with data gleaned primarily from the fossil record has its challenges. Building on decades of prior research, we outline and critically discuss a complete workflow for biomechanical analysis of extinct species, using locomotor biomechanics in the Triassic theropod dinosaur Coelophysis as a case study. We progress from the digital capture of fossil bone morphology to creating rigged skeletal models, to reconstructing musculature and soft tissue volumes, to the development of computational musculoskeletal models, and finally to the execution of biomechanical simulations. Using a three-dimensional musculoskeletal model comprising 33 muscles, a static inverse simulation of the mid-stance of running shows that Coelophysis probably used more upright (extended) hindlimb postures and was likely capable of withstanding a vertical ground reaction force of magnitude more than 2.5 times body weight. We identify muscle force-generating capacity as a key source of uncertainty in the simulations, highlighting the need for more refined methods of estimating intrinsic muscle parameters such as fiber length. Our approach emphasizes the explicit application of quantitative techniques and physics-based principles, which helps maximize results robustness and reproducibility. Although we focus on one specific taxon and question, many of the techniques and philosophies explored here have much generality to them, so they can be applied in biomechanical investigation of other extinct organisms.

Troodon formosus, a theropod from the Late Cretaceous, is one of the few species of dinosaurs with multiple nest sites uncovered. It has been consistently demonstrated that eggs within these nests would have been partially buried in life—an exceedingly rare state in modern vertebrates. There has been debate over Troodon's capacity to engage in thermoregulatory contact incubation, especially regarding an adult's ability to efficiently supply partially buried eggs with energy. An actualistic investigation was undertaken to determine the thermodynamic efficiency of contact incubating partially buried eggs. An efficient system would keep eggs at temperatures closer to the surrogate parent than the ambient, without prohibitively high energy input. For the experiment, a surrogate dinosaur was created and used in both indoor controlled ambient temperature trials and in an outdoor variant. Even with ambient temperatures that were likely cooler than Cretaceous averages, the results showed that contact incubating partially buried eggs did seem to confer an energetic advantage; egg temperatures remained closer to the surrogate than ambient in both indoor and outdoor tests. Still, critics of contact incubating partially buried eggs are correct in that there is a depth at which adult energy would fail to make much of an impact—perhaps more relevant to buried eggs, as partially buried eggs would be in contact with an adult and likely above the thermal input threshold. Additionally, results from this experiment provide evidence for a possible evolutionary path from guarding behavior to thermoregulatory contact incubation.

Throughout their 250 Myr history, archosaurian reptiles have exhibited a wide array of body sizes, shapes, and locomotor habits, especially in regard to terrestriality. These features make Archosauria a useful clade with which to study the interplay between body size, shape, and locomotor behavior, and how this interplay may have influenced locomotor evolution. Here, digital volumetric models of 80 taxa are used to explore how mass properties and body proportions relate to each other and locomotor posture in archosaurs. One-way, nonparametric, multivariate analysis of variance, based on the results of principal components analysis, shows that bipedal and quadrupedal archosaurs are largely distinguished from each other on the basis of just four anatomical parameters (p < 0.001): mass, center of mass position, and relative forelimb and hindlimb lengths. This facilitates the development of a quantitative predictive framework that can help assess gross locomotor posture in understudied or controversial taxa, such as the crocodile-line Batrachotomus (predicted quadruped) and Postosuchus (predicted biped). Compared with quadrupedal archosaurs, bipedal species tend to have relatively longer hindlimbs and a more caudally positioned whole-body center of mass, and collectively exhibit greater variance in forelimb lengths. These patterns are interpreted to reflect differing biomechanical constraints acting on the archosaurian Bauplan in bipedal versus quadrupedal groups, which may have shaped the evolutionary histories of their respective members.

The Red Queen's hypothesis portrays evolution as a never-ending competition for expansive energy, where one species’ gain is another species’ loss. The Red Queen is neutral with respect to body size, implying that neither small nor large species have a universal competitive advantage. Here we ask whether, and if so how, the Red Queen's hypothesis really can accommodate differences in body size. The maximum population growth in ecology clearly depends on body size—the smaller the species, the shorter the generation length, and the faster it can expand given sufficient opportunity. On the other hand, large species are more efficient in energy use due to metabolic scaling and can maintain more biomass with the same energy. The advantage of shorter generation makes a wide range of body sizes competitive, yet large species do not take over. We analytically show that individuals consume energy and reproduce in physiological time, but need to compete for energy in real time. The Red Queen, through adaptive evolution of populations, balances the pressures of real and physiological time. Modeling competition for energy as a proportional prize contest from economics, we further show that Red Queen's zero-sum game can generate unimodal hat-like patterns of species rise and decline that can be neutral in relation to body size.

The Cretaceous–Paleogene (K/Pg) extinction appears to have been geographically heterogeneous for some organismal groups. Southern Hemisphere K/Pg palynological records have shown lower extinction and faster recovery than in the Northern Hemisphere, but no comparable, well-constrained Southern Hemisphere macrofloras spanning this interval had been available. Here, macrofloral turnover patterns are addressed for the first time in the Southern Hemisphere, using more than 3500 dicot leaves from the latest Cretaceous (Maastrichtian) and the earliest Paleocene (Danian) of Argentine Patagonia. A maximum ca. 90% macrofloral extinction and ca. 45% drop in rarefied species richness is estimated across the K/Pg, consistent with substantial species-level extinction and previously observed extirpation of host-specialized leaf mines. However, prior palynological and taxonomic studies indicate low turnover of higher taxa and persistence of general floral composition in the same sections. High species extinction, decreased species richness, and homogeneous Danian macrofloras across time and facies resemble patterns often observed in North America, but there are several notable differences. When compared with boundary-spanning macrofloras at similar absolute paleolatitudes (ca. 50°S or 50°N) from the Williston Basin (WB) in the Dakotas, both Maastrichtian and Danian Patagonian species richnesses are higher, extending a history of elevated South American diversity into the Maastrichtian. Despite high species turnover, our analyses also reveal continuity and expansion of leaf morphospace, including an increase in lobed and toothed species unlike the Danian WB. Thus, both Patagonian and WB K/Pg macrofloras support a significant extinction event, but they may also reflect geographically heterogeneous diversity, extinction, and recovery patterns warranting future study.

Hadrosaurid dinosaurs, the dominant large-bodied terrestrial herbivores in most Laurasian Late Cretaceous ecosystems, have an exceptional fossil record consisting of many species known from partial ontogenetic series, making them an ideal clade with which to conduct life-history studies. Previous research considered the Dinosaur Park Formation (DPF) of Alberta as an attritional, or time-averaged, sample and interpreted size–frequency distribution of long bones collected from the DPF with three size classes to suggest that hadrosaurids from the DPF attained near-asymptotic body size in under 3 years. This conflicted with previously published osteohistological estimates of 6+ years for penecontemporaneous hadrosaurids from the Two Medicine Formation (TMF) of Montana, suggesting either extreme variation in hadrosaurid growth rates or that size–frequency distributions and/or osteohistology and growth modeling inaccurately estimate ontogenetic age.

We tested the validity of the previously proposed size–age relationship of hadrosaurids from the DPF by significantly increasing sample size and combining data from size–frequency distributions and osteohistology across multiple long-bone elements. The newly constructed size–frequency distributions typically reveal four relatively distinct size–frequency peaks that, when integrated with the osteohistological data, aligned with growth marks. The yearling size class was heavily underrepresented in the size–frequency distribution. If not due to preservation, this suggests that either juvenile (<2 years of age) hadrosaurids from the DPF had increased survivorship following an initially high nestling mortality rate or that yearlings were segregated from adults. A growth-curve analysis revealed asymptotic body size was attained in approximately 7 years, which is consistent with hadrosaurids from the TMF. The data suggest size–frequency distributions of attritional samples underestimate age and overestimate growth rates, but when paired with osteohistology can provide unique life-history insights.

Simulations are playing an increasingly important role in paleobiology. When designing a simulation study, many decisions have to be made and common challenges will be encountered along the way. Here, we outline seven rules for executing a good simulation study. We cover topics including the choice of study question, the empirical data used as a basis for the study, statistical and methodological concerns, how to validate the study, and how to ensure it can be reproduced and extended by others. We hope that these rules and the accompanying examples will guide paleobiologists when using simulation tools to address fundamental questions about the evolution of life.