The Mazon Creek Lagerstätte (Moscovian Stage, late Carboniferous Period; Illinois, USA) captures a diverse view of ecosystems in delta-influenced coastal settings through exceptional preservation of soft tissues in siderite concretions. The generally accepted paradigm of the Mazon Creek biota has been that of an inferred paleoenvironmental divide between what have been termed the Braidwood and Essex assemblages, wherein the former represents a freshwater ecosystem with terrestrial input and the latter a marine-influenced prodelta setting with abundant cnidarians, bivalves, worm phyla, and diverse arthropods. Here, we revisit the paleoecology of the Mazon Creek biota by analyzing data from nearly 300,000 concretions from more than 270 locations with complementary multivariate ordinations. Our results show the Braidwood assemblage as a legitimate shoreward community and provide evidence for further subdivision of the Essex assemblage into two distinct subassemblages, termed here the Will-Essex and Kankakee-Essex. The Will-Essex represents a benthos dominated by clams and trace fossils along the transition between nearshore and offshore deposits. The Kankakee-Essex is dominated by cnidarians, presenting an ecosystem approaching the geographic margin of this taphonomic window. These new insights also allow a refined taphonomic model, wherein recalcitrant tissues of Braidwood organisms were subject to rapid burial rates, while organisms of the Essex assemblage typically had more labile tissues and were subject to slower burial rates. Consequently, we hypothesize that the Braidwood fossils should record more complete preservation than the Essex, which was exposed for longer periods of aerobic decomposition. This is supported by a higher proportion of non-fossiliferous concretions in the Essex than in the Braidwood.

Over the interval 2008–2023 a large number of studies have been published testing various aspects of punctuated equilibria, including the prevalence of stasis, and also the extent to which most evolutionary change is concentrated at cladogenesis. In the vast majority of studies, punctuated equilibria continued to be strongly validated, as widespread evidence for stasis accumulated, with only some rare incidences of gradual change found. Support for the importance of cladogenetic change has increased, and new analytical approaches to study punctuated equilibria have been developed. Over this time period, there has also been an increase in the number of studies that have concentrated on extant taxa to test for punctuated equilibria, and these have also corroborated its widespread presence. In this respect, punctuated equilibria has served as an important bridge between neontological and paleontological approaches to evolutionary biology. From 2008 to 2023, there has also been some drift in how stasis is defined, such that, in certain studies, the definition diverged from the original 1972 definition in important respects. Notably, it is the few studies that have most changed the definition of what stasis constitutes that have most challenged the validity of punctuated equilibria, indicating it is morphing interpretations and definitions rather than the discovery of data compatible with phyletic gradualism that are most responsible for divergent results.

Ostracoderms, Paleozoic jawless stem-gnathostomes, are characterized by distinctive bony shields covering the front of their bodies. These headshields exhibit significant variations in morphology across species, boasting frontal, lateral, and dorsal processes. Ostracoderms represent pivotal intermediaries between modern jawless and jawed vertebrates, so understanding their biology and ecology is crucial for unraveling the selective pressures that shaped the early evolution and diversification of jawed vertebrates, which now dominate vertebrate diversity. This study employs virtual paleontology techniques and phylogenetic comparative methods to explore the hydrodynamic and ecological implications of these processes, focusing on pteraspidomorphs, the most diverse ostracoderm group. The analysis reveals widespread convergence in the arrangement and development of headshield processes. Lateral processes enhance hydrodynamic efficiency and generate lift, while combined lateral and dorsal processes provide stability in rolling, yawing, and pitching. Frontal processes reduce drag in many cases. These findings illuminate the enigmatic roles of ostracoderm headshields, showing how the dimensions and arrangement of their processes are biomechanically linked to a range of functions and ecological roles. Collectively, this highlights the intricate evolutionary pathways of lifestyles and ecologies within stem-gnathostomes, challenging the idea of a unidirectional trend toward more active lifestyles in vertebrate evolution and suggesting diverse ecological roles for ostracoderms.

Every organism interacts with a host of other organisms of the same and different species throughout its life. These biotic interactions have varying influences on the reproduction and dispersal of the organism, and hence also the population and species lineage to which the organism belongs. By extension, biotic interactions must contribute to the macroevolutionary patterns that we observe in the fossil record, but exactly how, when, and why are research questions we have been asking before the start of the journal Paleobiology. In this contribution for Paleobiology’s 50th anniversary, we present a brief overview of how paleobiologists have studied biotic interactions and their macroevolutionary consequences, recognizing paleontology’s unique position to contribute data and insights to the topic of interspecies interactions. We then explore, in a semi-free-form manner, what promising avenues might be open to those of us who use the fossil record to understand biotic interactions. In general, we emphasize the need for increased effort surrounding the understanding of ecological details, integration of different types of information, and model-based approaches.

Despite its many extensions and implications, we argue that punctuated equilibrium itself has two core, empirical claims: (1) stasis dominates within fossil species; and (2) morphological change is concentrated in pulses that occur associated with speciation. Here we assess the state of the evidence for these two claims, 50 years after punctuated equilibrium’s foundational paper. Spurred by controversy, paleontologists have amassed a large number of case studies in which morphology in species-level lineages is tracked over time. Modern, likelihood-based methods have been used to fit to these data models of stasis, random walks, and directional trends, as well as more complex dynamics. Compilations reveal that the directional trends predicted by gradualist expectations are infrequent. Although stasis is commonly observed, it is favored in less than half of cases, and meandering random walks or more complex models generally account for the majority of cases. The second claim of punctuated equilibrium has received much less empirical scrutiny than the first. Although speciational pulses are plausible in theory, only a few paleontological studies integrate ancestor–descendant time series into a phylogenetic framework as is needed to estimate cladogenetic change and compare it with anagenesis. These studies, as well as more indirect analyses of extant clades, suggest that speciational change can occur, but we cannot yet assess with confidence its frequency or importance compared with anagenetic changes.

The Great Ordovician Biodiversification Event (GOBE) records a global increase in marine biodiversity that reached maximum diversification rates during the Middle Ordovician. The degree to which the causes of the GOBE are regional or global is a question that must be addressed through analysis of regional data. In this study, stratigraphically constrained field-based data from the Middle Ordovician Simpson Group of Oklahoma were collected to identify temporal trends in body volume and determine whether body volume trends are more closely associated regional or global environmental and diversity changes. Anteroposterior–transverse (AT) volume estimations were produced for rhynchonelliform brachiopods at a bedding-plane level of resolution. Time-series analysis was used to establish temporal trends in brachiopod volume. Volume data were then analyzed alongside paired δ18O, Δ13C, 87Sr/86Sr, taxonomic diversity, and lithologic data using a boosted regression model to identify their relative influence on shell volume through time. Results of these analyses indicate that (1) a rapid pulse of brachiopod volume increase occurred coincident with the main diversification pulse in Simpson Group strata and (2) volume increase was not coupled with an increase in brachiopod volume variance. Volume increase was primarily associated with global-scale factors such as age, δ18O (temperature), 87Sr/86Sr (tectonics), and taxonomic diversity trends; whereas local-scale factors of Δ13C (carbon cycle) and lithologic trends were more weakly associated with local volume trends. Notably, all factors had a nonzero influence over brachiopod volume, indicating that local diversification was influenced by multifaceted interactions among abiotic and biotic controls. These results support the argument that Ordovician diversification included a substantial biotic shift during the Middle Ordovician and support the hypothesis that global factors were dominant, influencing diversification patterns during the main phase of the GOBE.

Ichthyosauria, Plesiosauria, and Metriorhynchidae were apex predators in Mesozoic oceanic trophic networks. Previous stable oxygen isotope studies suggested that several taxa belonging to these groups were endothermic and that some of them were homeothermic organisms. However, these conclusions remain contentious owing to the associated uncertainties regarding the δ18O value and oxygen isotope fractionation relative to environmental seawater. Here, we present new bioapatite phosphate δ18O values (δ18Op) of Ichthyosauria, Plesiosauria, and Metriorhynchidae (Middle Jurassic to Early Cretaceous) recovered from mid- to high paleolatitudes to better constrain their thermophysiology and investigate the presence of regional heterothermies. The intraskeletal δ18Op variability failed to reveal distinct heterothermic patterns within any of the specimens, indicating either intrabody temperature homogeneity or an overriding diagenetic overprint of the original biological δ18Op bone record. Body temperature estimates have been reassessed from new and published δ18Op values of well-preserved isolated teeth, recently revised Mesozoic latitudinal δ18O oceanic gradients, and 18O-enrichment factors of fully aquatic air-breathing vertebrates. Our results confirm that Ichthyosauria were homeothermic endotherms (31°C to 41°C), while Plesiosauria were likely poikilothermic endotherms (27°C to 34°C). The new body temperature estimates of the Metriorhynchidae (25°C to 32°C) closely follow ambient temperatures and point to poikilothermic strategy with no or little endothermic ability. These results improve our understanding of Mesozoic marine reptile thermoregulation and indicate that due to their limited body temperature variations, the δ18Op values from Ichthyosauria fossil remains could be used as valuable archives of Mesozoic oceans δ18Osw values that may help improve paleoenvironmental and paleoclimatic reconstructions.

Mass extinctions are natural experiments on the short- and long-term consequences of pushing biotas past breaking points, often with lasting effects on the structure and function of biodiversity. General properties of mass extinctions—exceptionally severe, taxonomically broad, global losses of taxa—are starting to come into focus through comparisons among dimensions of biodiversity, including morphological, functional, and phylogenetic diversity. Notably, functional diversity tends to persist despite severe losses of taxonomic diversity, whereas taxic and morphological losses may or may not be coupled. One of the biggest challenges in synthesizing and extracting general consequences of these events has been that they are often driven by multiple, interacting pressures, and the taxa and their traits vary among events, making it difficult to link single stressors to specific traits. Ongoing improvements in the taxonomic and stratigraphic resolution of these events for multiple clades will sharpen tests for selectivity and help to isolate hitchhiking effects, whereby organismal traits are carried by differential survival or extinction of taxa owing to other organismal or higher-level attributes, such as geographic-range size. Direct comparative analyses across multiple extinction events will also clarify the impacts of particular drivers on taxa, functional traits, and morphologies. It is not just the extinction filter that deserves attention, as the longer-term impact of extinctions derives in part from their ensuing rebounds. More work is needed to uncover the biotic and abiotic circumstances that spur some clades into re-diversification while relegating others to marginal shares of biodiversity. Combined insights from mass extinction filters and their rebounds bring a macroevolutionary view to approaching the biodiversity crisis in the Anthropocene, helping to pinpoint the clades, functional groups, and morphologies most vulnerable to extinction and failed rebounds.

Ammonoid cephalopods are excellent model systems for evolutionary biomechanics due to their volatile evolutionary dynamics and remarkable fossil record. During the Mesozoic marine revolution, natural selection increasingly favored ammonoid shells with specific ranges of ornamentation patterns (projections that influence surface roughness). While this evolutionary pattern has been attributed to enemy-driven evolution (i.e., escalation), many morphologies lack clear defensive roles. Using a combination of 3D modeling, physical experiments, and computer simulations, we investigate these patterns from a hydromechanical perspective. We model theoretical morphologies along a continuum of increasing ornamentation coarseness. Neutrally buoyant, 3D-printed models, weighted to match the mass distribution of their virtual counterparts, demonstrate that coarser patterns progressively attenuate rocking motions. Flow visualization experiments reveal these coarser patterns produce higher energy dissipation rates in the disturbed fluid. Computational fluid dynamics simulations were performed to characterize the hydrodynamic costs of ornamentation patterns over the majority of biologically relevant swimming speeds and shell sizes for planispiral ammonoids. Only the coarsest categories incur substantial increases in hydrodynamic drag. However, ornamentation patterns with intermediate coarseness effectively avoid this physical trade-off, experiencing dynamic stabilization without considerably reducing swimming efficiency. These trade-off-defying morphologies were progressively favored during the Mesozoic, becoming more abundant than others by the end of this era. Ultimately, these experiments highlight important hydromechanical selective pressures involved in ammonoid evolutionary trends and some fundamental constraints on aquatic locomotion more broadly.

Ancient changes in the biosphere, from organismic traits to wholesale ecosystem changes, can be aligned with climate forcing across the Phanerozoic. Clear examples of abrupt climate warming causing biodiversity crises are primarily found between the Permian and Paleogene periods. During these times, catastrophic events occurred, resembling the extreme climate scenarios projected for the near future. The paleobiologic literature around these events generally supports the hypothesis that abrupt climate change was a dominant trigger of extinction and/or ecological crisis. When climate change and climate history are considered, virtually all post-Paleozoic global biotic events can be confidently attributed to climatic change, with abrupt warming (hyperthermal events) leaving the most consistent fingerprint. The combined stress of deoxygenation and warming are sufficient to explain marine extinction patterns across most hyperthermal events. Although ocean acidification may have contributed, the direct role of pH on the extinction toll of organisms is not consistently demonstrated. Future research can enhance the correspondence between the magnitudes of climatic changes and their biological impacts, even though observed rates of change cannot currently be compared across different timescales. Mimicking multi-scale approaches in modern ecology, paleontological approaches to climate impact research will benefit from specifically targeting scaling relationships.

Punctuated equilibria (PE) was presented 50 years ago as an alternative to the widespread assumption that most evolution proceeds by gradual phyletic change within lineages. Unfortunately, PE has been widely misunderstood, misrepresented, and unfairly dismissed since this first publication. We argue that much of this misunderstanding centers around a misinterpretation of the meaning of “mode” in evolution, and its significance, properly understood, for how we understand macroevolutionary processes. PE proposed that most morphospecies do not show significant anagenetic trends through their stratigraphic ranges, and that most new morphospecies that are recognized arise via cladogenesis. To the degree that this is true, most exploration of disparity space must be associated with cladogenesis.

We surveyed a sample of the recent paleontological literature to assess the frequency with which new morphospecies appear in the fossil record via anagenesis versus cladogenesis using a persistence of ancestor criterion and found the overwhelmingly dominant mode of species origin to be cladogenesis. This is a valuable but underutilized approach to this problem, which could be exploited with more studies of species-level phylogenies of fossil taxa. Combined with the conclusions of other studies that stasis or nondirectional change is common, this finding of the dominance of cladogenesis affirms that PE is very much alive and of substantial significance for understanding macroevolutionary patterns.

Toothed whales (odontocetes) make use of high-frequency sounds to echolocate, differing significantly from their sister group baleen whales (mysticetes), which make use of low-frequency sound for long-distance communication. This divergence in auditory ability has led to considerable speculation as to how hearing functioned in the ancestral archaeocetes, and when the specializations of modern species arose. Numerous studies have attempted to infer auditory capabilities from morphological correlates valid in modern species. Here, we build upon these previous methods with a focus on cochlear structures that have well-understood links to function. We combine this with information on the sound conduction apparatus to chart the evolutionary trajectory of cetacean hearing. Our results suggest an initial move toward low-frequency specialization in early Eocene cetaceans, which coincides with the appearance of new sound conduction pathways. This paved the way for the later movement toward higher-frequency hearing in protocetids; however, the ultra-high- and low-frequency hearing specializations of both modern cetacean clades evolved after their divergence. We use these data to test the hypotheses that evolutionary brain size increases in cetaceans were related to the origin of high-frequency echolocation. We show that no shift in relative brain size coincides with any changes toward high-frequency perception. However, this does not rule out a role for other changes in hearing ability such as some simple forms of echolocation, similar to that suggested for hippopotamuses or bowhead whales, which may have been present in even the earliest cetaceans.

Humans have dramatically transformed ecosystems over the previous millennia and are potentially causing a mass extinction event comparable to the others that shaped the history of life. However, only a fraction of these impacts has been directly recorded, limiting conservation actions. Conservation paleobiology leverages geohistorical records to offer a long-term perspective on biodiversity change in the face of anthropogenic stressors. Nevertheless, the field's on-the-ground contributions to conservation outcomes are still developing. Here, we present an overview of directions in which paleobiological research could progress to aid conservation in the coming decades using elasmobranchs (sharks, rays, and skates)—a highly threatened group with a rich fossil record—as a model. These research directions are guided by areas of overlap between an expert-led list of current elasmobranch conservation priorities and available fossil and historical records. Four research topics emerged for which paleobiological research could address open questions in elasmobranch science and conservation: (1) baselines, (2) ecological roles, (3) threats, and (4) conservation priorities. Increasingly rich datasets and novel analytical frameworks present exciting opportunities to apply the elasmobranch fossil record to conservation practice. A similar approach could be extended to other clades. Given the synthetic nature of these research topics, we encourage collaboration across timescales and with conservation practitioners to safeguard the future of our planet's rapidly disappearing species.

Analysis of length-frequency data using the ELEFAN (Electronic Length-Frequency Analysis) approach and software is widely used to quantify the growth, mortality, longevity, and related parameters of Recent marine animals. Here we analyze a sample (n = 211) of the Ediacaran metazoan Parvancorina minchami Glaessner, 1958, from the Vendian siliciclastic marine deposits of the southeastern White Sea region, Russia. The results fit a von Bertalanffy equation with the parameters L∞ = 2 cm, K = yr−1 (with t0 not estimated) and an instantaneous rate of mortality (M) of 1.44 yr−1, implying M/K ≈ 2, as commonly occurs in Recent small invertebrates. These parameter values also imply a longevity for P. minchami of about 4 yr. The concepts and approach used here, previously applied to an Ordovician trilobite and a Cambrian radiodont, suggest that inferences on growth, mortality, longevity, and related parameters can be obtained from suitable size-frequency samples of long-extinct metazoans, opening new vistas on their growth dynamics and functional roles in ancient ecosystems.

Most species exhibit morphological stasis following speciation, and this is a key feature of the concept of punctuated equilibria. Stasis results in species often having long durations on geological timescales. Durational data are fundamental to many types of paleobiological analyses and are ideally based on occurrence data represented by specimens in museum collections. Often, however, durational data are presented without supporting information about voucher specimens that document stratigraphic ranges, including first and last appearances. We use the iconic Devonian trilobite Eldredgeops rana to demonstrate that durational data can be challenging to determine at multiple taxonomic levels. Further, we show that different datasets—including Sepkoski’s published databases, the Paleobiology Database, and iDigBio—give discordant results concerning first and last occurrences. We argue that paleontologists should adopt two general best practices to help address these problems. First, systematists should clearly identify voucher specimens that represent stratigraphic occurrences of species. Second, we recommend that high-quality photographs of occurrence vouchers be placed in open access websites and be assigned public domain licensing before being paywalled by journals. Such voucher images also have a role to play in training artificial intelligence (AI) systems that will be applied to future paleobiological questions.

The fossil record offers important opportunities to reconstruct plant community response to past disturbance events. Yet reconstructions are hindered by limited empirical evidence of successional variation in functional traits measurable on fossil leaves, including leaf morphology and δ13C. In addition, the role the leaf economic spectrum (LES) plays across succession within temperate deciduous forests is unresolved. Finally, it is unclear to what degree disturbance confounds the leaf morphology–climate relationships utilized in paleoclimate proxies.

We utilize a chronosequence spanning forest stands varied by time since logging (4, 21, 44, and 94 years old) and one old-growth stand in North Carolina. Leaf traits of woody non-monocot angiosperm (WNMA) leaves, including all trees and prominent understory plants, were measured to document patterns relating to the LES (e.g., leaf mass per area [LMA]), patterns of leaf morphology and δ13C, and their confounding influence on climatic estimates using the digital leaf physiognomy proxy.

LMA increased across succession among trees, driven by variation in both leaf thickness and leaf density, supporting the role of the LES. The petiole metric (PM), which is biomechanically linked to LMA, increased across succession among trees as hypothesized, as did the proportion of entire-margined leaves and, among tree dominants, leaf margin complexity. Measures of diversity (morphological and species richness, δ13C, and LMA variance) for all WNMAs were often highest in the old-growth stand, reflecting structural and niche complexity, yet peaked in mid-succession among trees, reflecting a mixing of ecological strategies. Other leaf traits had complicated or subtle trends across succession that were difficult to reconcile and tie to function. Changes in leaf morphology across succession did not strongly confound the accuracy of paleoclimate reconstructions. Successional patterns of this study importantly highlight the utility of PM, leaf margin, and leaf morphological richness in interpreting successional dynamics from fossil leaf assemblages sourced from temperate deciduous forests.

Fossil data are subject to inherent biological, geologic, and anthropogenic filters that can distort our interpretations of ancient life and environments. The inevitable presence of incomplete fossils thus requires a holistic assessment of how to navigate the downstream effects of bias on our ability to accurately reconstruct aspects of biology in deep time. In particular, we must assess how biases affect our capacity to infer evolutionary relationships, which are essential to analyses of diversification, paleobiogeography, and biostratigraphy in Earth history. In this study, we use an established completeness metric to quantify the effects of taphonomic filters on the amount of phylogenetic information available in the fossil record of 795 extinct squamate (e.g., lizards, snakes, amphisbaenians, and mosasaurs) species spanning 242 Myr of geologic time. This study found no meaningful relationship between spatiotemporal sampling intensity and fossil record completeness. Instead, major differences in squamate fossil record completeness stem from a combination of anatomy/body size and affinities of different squamate groups to specific lithologies and depositional environments. These results reveal that naturally occurring processes create structural megabiases that filter anatomical and phylogenetic data in the squamate fossil record, while anthropogenic processes play a secondary role.

The Pleistocene/Holocene transition furnishes a classic example of apparent evolutionary stasis during a period of major environmental change, where observed environmental clines might predict evolutionary change. We have previously attempted to assess whether or not body size and shape were static in the extensive Quaternary vertebrate fauna of Rancho La Brea (RLB). However, the validity of time-series studies depends on dating, and there are indications that previous approaches based on pit mean radiocarbon ages may be misleading. Here we have compiled and recalibrated all available RLB radiocarbon ages, reanalyzed our morphometric data using a novel method for bootstrap resampling of calibrated C age distributions to estimate a time series of populations, and fit the time series with a range of simple evolutionary time-series models.

Although the shortness of our time series tends to favor nondirectional models and is insufficient to allow reliable discrimination between punctuated and gradual change, the results can still be clearly interpreted. The population means for most anatomical elements in most species at RLB do genuinely appear to be static through the Pleistocene/Holocene transition, as previously published. Some species exhibit previously undetected changes in population mean size and shape, including Smilodon, Gymnogyps, and Equus. However, the timing of change is variable among the non-static species and generally does not correspond to changes in temperature, and thus resists a Bergmann’s rule interpretation. Considering the species by ecological category may reveal more about the effects of climate regime shifts.

The Neotropics host the highest diversity of plants on the Earth today and have done since at least the late Paleogene (~58 Ma). Several mechanisms have been proposed to explain this elevated diversity, but the empirical patterns of Neotropical plant diversification that would test key aspects of those mechanisms are still unclear. We use an extensive palynological database from northern South America to characterize patterns of extinction, origination, and diversity and their possible drivers since the Paleogene. The foreland Llanos basin of Colombia preserves the evolutionary history of Neotropical vegetation as well as the geological evolution of northern South America, offering a unique opportunity to study the relationship between the geological and fossil records. The palynological record of the Llanos basin has been intensely studied mainly for oil exploration, and we use this information to infer the evolutionary history of Neotropical vegetation in Colombia during the Cenozoic. There is no straightforward relationship between global temperature and Neotropical plant diversity. Nevertheless, environmental change had an important influence on the dynamics of diversification, especially during volatile climate intervals such as the Paleocene–Eocene and the Pleistocene. Pulses of regional extinction were driven by large-scale temperature excursions, including both warming and cooling phases. Time-lagged origination pulses results in rapid floral replacement on a timescale of 1 Myr. Origination and extinction are essentially balanced on long timescales, leading to a near-zero long-term net diversification rate. Regional geological events, like the uplift of the Andean Cordillera, and changes in paleogeography also played an important role in Neotropical plant diversification.

The first compilations of Proterozoic eukaryote diversity, published in the 1980s showed a dramatic peak in the Tonian Period (1000–720 Ma), interpreted as the initial radiation of eukaryotes in the marine realm. Over the decades, new discoveries filled in the older part of the record and the peak diminished, but the idea of a Tonian radiation of eukaryotes has remained strong, and is now widely accepted as fact. We present a new diversity compilation based on 181 species and 713 species occurrences from 145 formations ranging in age from 1890 Ma to 720 Ma and find a significant increase in diversity in the Tonian. However, we also find that the number of eukaryotic species through time is highly correlated with the number of formations in our dataset (i.e. eukaryote-bearing formations) through time. This correlation is robust to interpretations of eukaryote affinity, bin size, and bin boundaries. We also find that within-assemblage diversity—a measure thought to circumvent sampling bias—is related to the number of eukaryote-bearing formations through time. Biomarkers show a similar pattern to body fossils, where the rise of eukaryotic biosignatures correlates with increased sampling. We find no evidence that the proportion of eukaryote-bearing versus all fossiliferous formations changed through the Proterozoic, as might be expected if the correlation reflected an increase in eukaryote diversity driving an increase in the number of eukaryote-bearing formations. Although the correlation could reflect a common cause such as changes in sea level driving both diversification and an increase in sedimentary rock volume, we favor the explanation that the pattern of early eukaryote diversity is driven by variations in paleontological sampling.