The seemingly aberrant coiling of heteromorphic ammonoids suggests that they underwent more significant changes in hydrostatic properties throughout ontogeny than their planispiral counterparts. Such changes may have been responses to different selective pressures at different life stages. The hydrostatic properties of three species of Didymoceras (D. stevensoni, D. nebrascense, and D. cheyennense) were investigated by creating virtual 3D models at several stages during growth. These models were used to compute the conditions for neutral buoyancy, hydrostatic stability, orientation during life, and thrust angles (efficiency of directional movement). These properties suggest that Didymoceras and similar heteromorphs lived low-energy lifestyles with the ability to hover above the seafloor. The resultant static orientations yielded a downward-facing aperture in the hatchling and a horizontally facing aperture throughout most of the juvenile stage, before terminating in an upward direction at maturity. Relatively high hydrostatic stabilities would not have permitted the orientation of Didymoceras to be considerably modified with active locomotion. During the helical phase, Didymoceras would have been poorly suited for horizontal movement, yet equipped to pirouette about the vertical axis. Two stages throughout growth, however, would have enhanced lateral mobility: a juvenile stage just after the formation of the first bend in the shell and the terminal stage after completion of the U-shaped hook. These two more mobile phases in ontogeny may have improved juvenile dispersal potential and mate acquisition during adulthood, respectively. In general, life orientation and hydrostatic stability change more wildly for these aberrantly coiled ammonoids than their planispiral counterparts.

The final 10 Myr of the Paleozoic saw two of the biggest biological crises in Earth history: the middlePermian extinction (often termed the Guadalupian–Lopingian extinction [GLE]) that was followed 7–8 Myr later by Earth's most catastrophic loss of diversity, the Permian–Triassic mass extinction (PTME). These crises are not only manifest as sharp decreases in biodiversity and—particularly for the PTME—total ecosystem collapse, but they also drove major changes in biological morphological characteristics such as the Lilliput effect. The evolution of test size among different clades of foraminifera during these two extinction events has been less studied. We analyzed a global database of foraminiferal test size (volume) including 20,226 specimens in 464 genera, 98 families, and 9 suborders from 632 publications. Our analyses reveal significant reductions in foraminiferal mean test size across the Guadalupian/Lopingian boundary (GLB) and the Permian/Triassic boundary (PTB), from 8.89 to 7.60 log10 μm3 (lg μm3) and from 7.25 to 5.82 lg μm3, respectively. The decline in test size across the GLB is a function of preferential extinction of genera exhibiting gigantism such as fusulinoidean fusulinids. Other clades show little change in size across the GLB. In contrast, all Lopingian suborders in our analysis (Fusulinina, Lagenina, Miliolina, and Textulariina) experienced a significant decrease in test size across the PTB, mainly due to size-biased extinction and within-lineage change. The PTME was clearly a major catastrophe that affected many groups simultaneously, and the GLE was more selective, perhaps hinting at a subtler, less extreme driver than the later PTME.

Living crinoids are exclusively passive suspension feeders and benthic as adults. However, in the past they adapted to a broad range of ecological niches. For instance, the stratigraphically important middle Paleozoic scyphocrinoids are hypothesized to have been planktonic, employing their inferred gas-filled globular, chambered structure at the distal end of the stem, the so-called lobolith, as a buoyancy device with the crinoid calyx suspended below it. Here, we evaluate this hypothesis using evidence from skeletal micromorphology and theoretical biomechanical modeling. Lobolith walls are typically composed of ossicles, which are exclusively composed of constructional labyrinthic stereom. In plates from the distal side of the lobolith, this stereom extends into microperforate stereom layer, forming wavy ridges and spines. No microscale adaptations for preventing gas leaks and/or ingress of water (such as internal and external imperforate stereom layers) are known. Furthermore, theoretical calculations suggest that the scyphocrinoid tow-net mode of feeding would have resulted in small relative velocities between the towed filter and the ambient water, thus making it an ineffective passive filter feeder. We suggest that the lobolith of these crinoids acted as a modified holdfast rather than as a floating buoy. Its globular shape and distally positioned microspines served as adaptations for living in unconsolidated sediments, analogous to iceberg- and snowshoe-like strategies used by some mollusks and brachiopods. Like modern isocrinids, scyphocrinoids could have maintained an upright feeding posture by extending the distal portion of the stalk along the bottom. In this recumbent posture, the distal part of the stalk with the lobolith might have functioned as a drag anchor. As a consequence of the ~3-m-long stem, even with this posture, the benthic scyphocrinoids could have risen to the highest epifaunal tier in the Paleozoic.

Taphonomic processes are informative about the magnitude and timing of paleoecological changes but remain poorly understood with respect to freshwater invertebrates in spring-fed rivers and streams. We compared taphonomic alteration among freshwater gastropods in live, dead (surficial shell accumulations), and fossil (late Pleistocene–early Holocene in situ sediments) assemblages from two Florida spring-fed systems, the Wakulla and Silver/Ocklawaha Rivers. We assessed taphonomy of two gastropod species: the native Elimia floridensis (n = 2504) and introduced Melanoides tuberculata (n = 168). We quantified seven taphonomic attributes (aperture condition, color, fragmentation, abrasion, juvenile spire condition, dissolution, and exterior luster) and combined those attributes into a total taphonomic score (TT). Fossil E. floridensis specimens exhibited the greatest degradation (highest TT scores), whereas live specimens of both species were least degraded. Specimens of E. floridensis from death assemblages were less altered than fossil specimens of the same species. Within death assemblages, specimens of M. tuberculata were significantly less altered than specimens of E. floridensis, but highly degraded specimens dominated in both species. Radiocarbon dates on fossils clustered between 9792 and 7087 cal BP, whereas death assemblage ages ranged from 10,692 to 1173 cal BP. Possible explanations for the observed taphonomic patterns include: (1) rapid taphonomic shell alteration, (2) prolonged near-surface exposure to moderate alteration rates, and/or (3) introduction of reworked fossil shells into surficial assemblages. Combined radiocarbon dates and taphonomic analyses suggest that all these processes may have played a role in death assemblage formation. In these fluvial settings, shell accumulations develop as a complex mixture of specimens derived from multiple sources and characterized by multimillennial time-averaging. These findings suggest that, when available, fossil assemblages may be more appropriate than death assemblages for assessing preindustrial faunal associations and recent anthropogenic changes in freshwater ecosystems.

We document a positive and strong correlation between speciation and extinction rates in the Paleozoic zooplankton graptoloid clade, between 481 and 419 Ma. This correlation has a magnitude of ~0.35–0.45 and manifests at a temporal resolution of <50 kyr and, for part of our data set, <25 kyr. It cannot be explained as an artifact of the method used to measure rates, sampling bias, bias resulting from construction of the time series, autocorrelation, underestimation of species durations, or undetected phyletic evolution. Correlations are approximately equal during the Ordovician and Silurian, despite the very different speciation and extinction regimes prevailing during these two periods, and correlation is strongest in the shortest-lived cohorts of species.

We infer that this correlation reflects approximately synchronous coupling of speciation and extinction in the graptoloids on timescales of a few tens of thousands of years. Almost half of graptoloid species in our data set have durations of <0.5 Myr, and previous studies have demonstrated that, during times of background extinction, short-lived species were selectively targeted by extinction. These observations may be consistent with the model of ephemeral speciation, whereby new species are inferred to form constantly and at high rate, but most of them disappear rapidly through extinction or reabsorption into the ancestral lineage. Diversity dependence with a lag of ~1 Myr, also documented elsewhere, may reflect a subsequent and relatively slow, competitive dynamic that governed those species that dispersed beyond their originating water mass and escaped the ephemeral species filter.

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.

Site-selectivity analysis of drilling predation traces may provide useful behavioral information concerning a predator interacting with its prey. However, traditional approaches exclude some spatial information (i.e., oversimplified trace position) and are dependent on the scale of analysis (e.g., arbitrary grid system used to divide the prey skeleton into sectors). Here we introduce the spatial point pattern analysis of traces (SPPAT), an approach for visualizing and quantifying the distribution of traces on shelled invertebrate prey, which includes improved collection of spatial information inherent to drillhole location (morphometric-based estimation), improved visualization of spatial trends (kernel density and hotspot mapping), and distance-based statistics for hypothesis testing (K-, L-, and pair correlation functions). We illustrate the SPPAT approach through case studies of fossil samples, modern beach-collected samples, and laboratory feeding trials of naticid gastropod predation on bivalve prey. Overall results show that kernel density and hotspot maps enable visualization of subtle variations in regions of the shell with higher density of predation traces, which can be combined with the maximum clustering distance metric to generate hypotheses on predatory behavior and anti-predatory responses of prey across time and geographic space. Distance-based statistics also capture the major features in the distribution of traces across the prey skeleton, including aggregated and segregated clusters, likely associated with different combinations of two modes of drilling predation, edge and wall drilling. The SPPAT approach is transferable to other paleoecologic and taphonomic data such as encrustation and bioerosion, allowing for standardized investigation of a wide range of biotic interactions.

Cosmopolitanism occurred recurrently during the geologic past, especially after mass extinctions, but the underlying mechanisms remain poorly known. Three theoretical models, not mutually exclusive, can lead to cosmopolitanism: (1) selective extinction in endemic taxa, (2) endemic taxa becoming cosmopolitan after the extinction and (3) an increase in the number of newly originated cosmopolitan taxa after extinction. We analyzed an updated occurrence dataset including 831 middle Permian to Middle Triassic ammonoid genera and used two network methods to distinguish major episodes of ammonoid cosmopolitanism during this time interval. Then, we tested the three proposed models in these case studies. Our results confirm that at least two remarkable cosmopolitanism events occurred after the Permian–Triassic and late Smithian (Early Triassic) extinctions, respectively. Partitioned analyses of survivors and newcomers revealed that the immediate cosmopolitanism event (Griesbachian) after the Permian–Triassic event can be attributed to endemic genera becoming cosmopolitan (model 2) and an increase in the number of newly originated cosmopolitan genera after the extinction (model 3). Late Smithian cosmopolitanism is caused by selective extinction in endemic taxa (model 1) and an increase in the number of newly originated cosmopolitan genera (model 3). We found that the survivors of the Permian–Triassic mass extinction did not show a wider geographic range, suggesting that this mass extinction is nonselective among the biogeographic ranges, while late Smithian survivors exhibit a wide geographic range, indicating selective survivorship among cosmopolitan genera. These successive cosmopolitanism events during severe extinctions are associated with marked environmental upheavals such as rapid climate changes and oceanic anoxic events, suggesting that environmental fluctuations play a significant role in cosmopolitanism.

A recent study by Saulsbury and colleagues assessed factors influencing growth coefficients in bivalves. Like many macroevolutionary studies that cover a wide range of taxa and environments, this study fails to account for important heterogeneity among taxa and among environments. Rankings of factors depend on the range of values sampled, patterns of allocation of energy to various competing functions, and whether taxa in any given clade are uniform in their growth coefficients. Heterogeneity is universal and must be taken into account in large-scale analyses.

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 Strophomenata, which includes two large orders, the Strophomenida and Productida, is the largest group of Paleozoic brachiopods. Nearly all uncemented strophomenatans possessed an unusual concave brachial valve. Most of these have been considered to have lived epifaunally, but had they rested on the seafloor, not only would they have faced intense predation, but their physical instability would have been fatal. I conclude that nearly all strophomenatans, like similar concavo-convex pectinid bivalves, lived infaunally by ejecting water to create a pit into which they descended, to be protected by sediment covering the concave valve. Strophomenatans have been discovered with this sediment preserved in place. If exhumed and turned upside down, a strophomenatan could have righted itself by squirting water. Many productides had anchoring spines, and some had hinge areas with stabilizing flanges. Small spines on the brachial valves of some productides served to trap disguising sediment. Evolutionary loss of hinge teeth within both the Strophomenida and Productida reduced the friction of valve clapping. Partly because of their slender shape, strophomenides were typically more vulnerable to exhumation than productides. Strophomenides also ejected water less effectively than productides and would have been less adept at righting themselves. The virtual disappearance of the strophomenides during the Devonian can be attributed to their vulnerability to intensifying benthic bulldozing and predation. The success of the productides during the late Paleozoic can be attributed to their relatively deep sequestration in the sediment and ability to right themselves and reburrow effectively when exhumed and overturned.

Reconstructing patterns of macroevolution has become a central endeavor in paleobiology, because it offers insight into evolutionary models shaping the history of life. As the most diverse and abundant animals since the Cambrian period, arthropods provide copious data to elucidate the emergence of body plans in metazoan lineages. However, information provided by fossils on the tempo and mode of this phenomenon has lacked a recent synthesis. Here, I investigate macroevolutionary patterns of morphological evolution in Euarthropoda using a combined extinct and extant dataset optimized for multivariate analyses. Overall ordination patterns between the main morphogroups are consistent with another, independently coded, extant-only dataset providing molecular and morphological rates of evolution. Based on a “deep split” phylogenetic framework, total-group Mandibulata and Arachnomorpha emerge as directional morphoanatomical lineages, with basal fossil morphogroups showing heterogeneously spread-out occupations of the morphospace. In addition to a more homogeneous morphological variation, new morphogroups arose by successive reductions of translation distances; this pattern was interrupted only by terrestrialization events and the origin of pancrustaceans. A displaced optimum type of model is proposed to explain the fast assembly of canalized body plans during the Cambrian, with basal fossil morphogroups fitting intermediate fitness peaks in a moving adaptive landscape. Given time constraints imposed by the paleontological evidence, and owing to the interplay between canalization and modularity, as well as a decoupling between molecular and morphological rates, the rise of euarthropods would support the view that the swiftness of the Cambrian explosion was mostly associated with the buildup of genetic regulatory networks.