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 pollen analysis is an “open-world” problem in paleontology for which there is a long-standing need for automated identification and classification. In the open world, categorical classes are imbalanced, test classes are not known a priori, and test data are captured across different domains. Pollen samples capture large numbers of specimens that include both common and abundant types and rare and sometimes novel taxa. Pollen is diverse morphologically and features can be altered during fossilization. Additionally, there is little standardization in the imaging of pollen samples. Therefore, generalized workflows for automated pollen analysis require techniques that are robust to these differences and can work with microscope images. We focus on a critical first step, the initial detection of pollen specimens on a palynological slide and review how existing methods can be employed to build robust and generalizable analysis pipelines. First, we demonstrate how a mixture-of-experts approach—the fusion of a general pollen detector with an expert model trained on minority classes—can be used to address taxonomic biases in detections, particularly the missed detections of rarer pollen types. Second, we demonstrate the efficiency of domain fine-tuning in addressing domain gaps—differences in image magnification and resolution across microscopes and of taxa across different sample sources. Third, we demonstrate the importance of continual learning workflows, which integrate expert feedback, in training detection models from incomplete data. Finally, we demonstrate how cutting-edge segmentation models can be used to refine and clean detections for downstream deep learning classification models.

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.

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.

Polymorphism, the occurrence of different morphs of a trait within the population of a single species, plays a crucial role in species diversification, genetic variation, and adaptation. Detecting polymorphism in a single character helps us to understand population dynamics, particularly in species that inhabit diverse environments. However, detecting polymorphisms in fossil taxa is challenging due to the fragmentary and incomplete records. Dimorphism, defined as the occurrence of different morphs of a trait within the population of a single species, represents the simplest and most common form of polymorphism. This study focuses on dimorphism instead of polymorphism, which allows for a more streamlined analysis. We use computational simulation experiments to estimate the minimum sample size required to detect bimodal distribution in univariate morphological variables. We describe the morphological diversity of a measured variable (e.g., body mass or skeletal length) as a probability density distribution with specific parameter sets. Subsequently, we simulate the diversity of the measured variable with varying sample sizes and conduct resampling procedures to ensure the robustness. Four key parameters that characterize the probability distribution are identified as having significant influence on the minimum sample size for dimorphism recognition. According to the simulation experiments, a model is built to estimate the minimum sample size for dimorphism recognition based on these parameters. A dataset from extant avian and reptilian species is used to test the model. Furthermore, we calculate a reference for the minimal sample size required for assessing phenotypic dimorphism in fossil avian taxa by applying parameters derived from extant avian species.

The Paleocene–Eocene thermal maximum (PETM) was the largest early Cenozoic hyperthermal event, one of a series of carbon cycle and climate perturbations marked by massive releases of carbon into the atmosphere and spikes in global temperature. Previous studies have documented major changes in the composition of terrestrial plant and animal communities during the PETM, as well as changes in arthropod herbivory. Here, we examine possible changes in pollination mode during the PETM in the Bighorn Basin, Wyoming, USA, as inferred from three lines of evidence: (1) the prevalence of fossil pollen preserved as clumps, (2) the pollination mode of nearest living relatives (NLR), and (3) angiosperm pollen morphological diversity. These suggest animal pollination became more common and wind pollination less common during the PETM. The decrease in wind pollination during the PETM reflects the basin-scale extirpation of wind-pollinated lineages and their replacement by dominantly animal-pollinated lineages concomitant with rapid warming and drying. The hotter and seasonally drier climates not only facilitated the northward range shift of plant taxa, but likely their insect and/or vertebrate pollinators as well. The dramatic floral changes during the PETM in the Bighorn Basin may also have changed available resources for insect and/or vertebrate pollinators.

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.

Multituberculate extinction is often cited as a classic case of competitive exclusion, coinciding with the first rodent arrivals in the late Paleocene. Analyzing 124 North American multituberculate last occurrence records during the Eocene from 56 to 34 million years ago, this study aimed to differentiate Eocene multituberculate and coeval rodent floral associations through geographic spatial analysis to understand niche overlap between the two groups. If competitive exclusion with rodents was a factor in multituberculate extinction, both multituberculates and rodents would be predicted to share similar forest habitat preferences and have competed for similar ecological niches regarding their forest associations. Using spatial analysis, this study found that Eocene rodents and multituberculates did not overlap in their forest associations. The findings indicate that multituberculates were unique in inhabiting a specific type of ancient forest habitat, favoring forests composed of Metasequoia, Glyptostrobus, and Alnus, and thus thrived in wetter northern temperate forest communities during the Eocene. Metasequoia and Glyptostrobus declined significantly in North America during the later Cenozoic, coinciding with multituberculate decline and extinction as the global climate shifted toward colder and drier climates around the Eocene/Oligocene boundary. In contrast, the success of rodents is attributed to their much broader forest affinity. These preferences align with the widespread distribution of rodents today, contributing to their modern success. The absence of any similar reconstructed forest habitat preferences between rodents and multituberculates suggests that changing forest structure, rather than competitive exclusion, drove multituberculate extinction.

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.

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.

Organismal metabolic rate is linked to environmental temperature and oxygen consumption, and as such, may be a useful predictor of extinction risk. This is especially true during major climate-driven extinctions, given the tightly linked stressors of warming and hypoxia. However, metabolic attributes can be quantified in different ways, highlighting differing aspects of organisms’ ecology. Here, we estimate resting whole-body and mass-specific metabolic rates in post-Carboniferous bivalve taxa using body size, seawater paleotemperature, and a taxon-specific adjustment factor to assess how metabolic rate correlates with survival both during and outside intervals of rapid climate warming, or “hyperthermals.” Accounting for the effects of geographic range size, we find a pattern of preferential extinction of bivalves with lower total calorific needs, consistent with increasing body size and the postulated ramping up of ecosystem energetics over the Meso-Cenozoic. Contrary to expectations, extinction selectivity based on total calorific needs, which emphasizes body size, does not differ between hyperthermals and other time intervals. However, a higher metabolic rate per gram of tissue—which is more strongly determined by environmental temperature than by body size—consistently increases the probability of extinction during hyperthermals relative to baseline conditions, particularly within the paleotropics. This serves to highlight the potential significance of environmental temperature on metabolic performance, particularly in organisms that are already living close to their thermal limits. In tandem with previously documented patterns of extinction selectivity based on relative activity levels, including motility and feeding style, these results enhance our understanding of the role of metabolic rate through time and during climate-driven extinctions. When standardized by mass, metabolic rate may represent a useful metric through which to predict the effects of anthropogenic climate change on modern marine faunas.

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.

A primary tenet of punctuated equilibria (PE) is that stasis, that is, little to no net morphological change, characterizes the histories of species. In the past ~50 years since PE was proposed, stasis has been recognized in the evolutionary histories of many species, but consensus has not been reached concerning its causes.

One unresolved issue is whether viable ecological mechanisms for stasis exist. We argue that a promising potential ecological explanation for stasis is coevolutionary alternation, which addresses how antagonists (e.g., predators or parasites and their groups of victims) coevolve over eco-evolutionary time across broad spatial scales. Coevolutionary alternation predicts different patterns of predator preferences and prey defenses within different populations and alternation of high and low levels of prey defenses as predator preferences evolve. The geographic structure of populations experiencing different environmental pressures and coevolutionary dynamics can yield stasis in such traits on the scale of entire species. We suggest that predator–prey coevolutionary alternation could be modeled using coupled stochastic differential equations (SDEs), which have been used to study correlative and causal connections among time series. SDEs can handle irregular sampling intervals, measurement uncertainty, and feedback loops between time series and can incorporate environmental proxies and time series from multiple geographic locations. We advocate developing this approach further to test the role of coevolutionary alternation in stasis and make recommendations for how SDEs might be used to model the coevolutionary feedback of predator(s) and multiple prey populations evolving in response to one another across space in a constantly changing environment.

Life-history traits such as dispersal affect population attributes like gene flow, which can have consequences for speciation and extinction rates over macroevolutionary timescales. Here we use the Cheilostomatida, a monophyletic order of marine bryozoans, to test whether a life-history trait, larval brooding, affected the origination and extinction rates of genera throughout their fossil record. Cheilostome lineages that brood their larvae have shorter larval dispersal distances than non-brooding lineages, which has led to the hypothesis that the evolution of larval brooding decreased gene flow, increased origination, and drove their Cretaceous diversification. Brooding cheilostomes are far more diverse than non-brooding cheilostomes today, but it remains to be shown that brooding lineages have a higher origination rate than non-brooders. We fit time-varying Pradel seniority capture–mark–recapture models to look at the effect of brooding on origination and extinction rates during the Cretaceous cheilostome diversification, the Cretaceous/Paleogene mass extinction and recovery, and through the Cenozoic. Our results support the hypothesis that brooding affects origination rate, but only in the Cenomanian to Campanian. Extinction rates do not differ between brooding and non-brooding genera, and there is no regime shift specific to the Cretaceous/Paleogene mass extinction. Our work illustrates the importance of using fossil occurrences and time-varying models, which can detect interval-specific diversification differentials.

Genome size (GS) is thought to be a key life-history trait and important for controlling plant distributions and evolutionary dynamics, but a full understanding of GS variation through evolutionary history requires proxy measurements from fossils. Here, we compare two potential GS proxies: guard cell length (GCL) and sporomorph size. We generated GCL and pollen size data from angiosperms growing in the University of Münster Botanical Garden, compiled sporomorph size data from the literature, and related these to GS using phylogenetic regression models. We also fit evolutionary models to the botanical garden data and used a published dataset to validate GCL as a GS proxy. The majority of the analyses conducted revealed a positive relationship between GS and sporomorph size, but in most cases, the explanatory power of the regressions was low. GCL showed a stronger and more consistent relationship with GS, and independent validation of the relationship showed a generally good match between predicted and observed GS. Sporomorph size is not suitable as a cross-taxon GS proxy, but some specific taxa (e.g., Pinus) may contain useful GS information. GCL has much more potential for measuring paleo-GS, but requires further research for us to better understand possible environmental controls on cell size variation.

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.

Although the theory of punctuated equilibria has stood the test of time, critics have sometimes highlighted the lack of a complementary molecular mechanism. The developmental gene hypothesis (DGH) provides just such a mechanism and is reviewed and significantly expanded in the present paper, taking advantage of concepts of active and passive evolvability, genetic drift, and the nearly neutral theory of molecular evolution, and compensatory adaptation in the face of weakly deleterious genetic variation. In addition, with the use of game theory, the author models the behavior of developmental regulatory (DevReg) genes, which are integral to the proposed hypothesis, in order to better understand their roles in stasis and speciation.

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.

Prospective and early-career paleontologists deserve an accurate assessment of employment opportunities in their chosen field of study. Drawing on a wide range of sources, we have produced an admittedly incomplete analysis of the current status and recent trends of permanent academic employment in the discipline. Obtaining more complete longitudinal data on employment trends is a major difficulty; this is a challenge that needs to be addressed. The number of job seekers is far in excess of available positions. There has been a clear erosion in the number of academic paleontologists in the United States, a trend exacerbated in recent years. The decline, in constant dollars, of federal funding for paleontological research has potential strong negative impacts on future hiring. The loss of paleontology positions has also had a deleterious effect on our professional societies, which have seen a loss of regular (professional) membership, although student membership remains strong. These trends also potentially negatively impact efforts to diversify the field. Professional societies need to better coordinate their efforts to address these serious issues. Individual paleontologists also must become more effective advocates for the importance and relevance of our science.