Strata of the Ediacaran Period (635–538.8 Ma) yield the oldest known fossils of complex, macroscopic organisms in the geologic record. These “Ediacaran-type” macrofossils (known as the Ediacaran biota) first appear in mid-Ediacaran strata, experience an apparent decline through the terminal Ediacaran, and directly precede the Cambrian (538.8–485.4 Ma) radiation of animals. Existing hypotheses for the origin and demise of the Ediacaran biota include: changing oceanic redox states, biotic replacement by succeeding Cambrian-type fauna, and mass extinction driven by environmental change. Few studies frame trends in Ediacaran and Cambrian macroevolution from the perspective of the sedimentary rock record, despite well-documented Phanerozoic covariation of macroevolutionary patterns and sedimentary rock quantity. Here we present a quantitative analysis of North American Ediacaran–Cambrian rock and fossil records from Macrostrat and the Paleobiology Database. Marine sedimentary rock quantity increases nearly monotonically and by more than a factor of five from the latest Ediacaran to the late Cambrian. Ediacaran–Cambrian fossil quantities exhibit a comparable trajectory and have strong (rs > 0.8) positive correlations with marine sedimentary area and volume flux at multiple temporal resolutions. Even so, Ediacaran fossil quantities are dramatically reduced in comparison to the Cambrian when normalized by the quantity of preserved marine rock. Although aspects of these results are consistent with the expectations of a simple fossil preservation–induced sampling bias, together they suggest that transgression–regression and a large expansion of marine shelf environments coincided with the diversification of animals during a dramatic transition that is starkly evident in both the sedimentary rock and fossil records.

Continuous morphological traits play a crucial role in phylogenetic inference, yet they are often discretized due to limited software support and challenges of handling missing data efficiently. We present a new implementation of the Brownian motion model for continuous trait evolution in the Bayesian phylogenetics software MrBayes. Our approach efficiently accommodates any proportion of missing data and supports evolutionary rate variation across characters and data partitions. It is compatible with both non-clock and relaxed clock models. We validate the implementation through simulations and apply it to empirical datasets of pterosaurs and ancient humans, demonstrating that continuous traits can improve phylogenetic resolution. This development expands the methodological tool kit for morphological and total-evidence phylogenetics and is applicable across diverse taxonomic groups.

The fossil record is spatiotemporally heterogeneous: taxon occurrence data have patchy spatial distributions, and this patchiness varies through time. Large-scale quantitative paleobiology studies that fail to account for heterogeneous sampling coverage will generate uninformative inferences at best and confidently draw wrong conclusions at worst. Explicitly spatial methods of standardization are necessary for analyses of large-scale fossil datasets, because nonspatial sample standardization, such as diversity rarefaction, is insufficient to reduce the signal of varying spatial coverage through time or between environments and clades. Spatial standardization should control both geographic area and dispersion (spread) of fossil localities. In addition to standardizing the spatial distribution of data, other factors may be standardized, including environmental heterogeneity or the number of publications or field collecting units that report taxon occurrences. Using a case study of published global Paleobiology Database occurrences, we demonstrate strong signals of sampling; without spatial standardization, these sampling signatures could be misattributed to biological processes. We discuss practical issues of implementing spatial standardization via subsampling and present the new R package divvy to improve the accessibility of spatial analysis. The software provides three spatial subsampling approaches, as well as related tools to quantify spatial coverage. After reviewing the theory, practice, and history of equalizing spatial coverage between data comparison groups, we outline priority areas to improve related data collection, analysis, and reporting practices in paleobiology.

Over the past 10 million years, coastal-marine settings along the Peruvian Margin have undergone profound geographic and oceanographic transformations, resulting in extensive changes in coastal-marine communities. While mollusk taxonomy research is slowly being integrated into ecosystem-wide analyses, which have historically centered on vertebrates, a long-term chronostratigraphically controlled analysis of molluscan diversity and compositional changes has not been undertaken for this region. We compiled a database covering 152 species, 97 genera, and 51 families of mollusk fossils from the Peruvian Margin (13–16°S) to assess long-term diversification patterns and faunal turnover from the late Miocene to the present. We identified two distinctive molluscan assemblages. The first, dating to the late Miocene (10–6 Ma), underwent a substantial shift during the Mio-Pliocene transition (6–4 Ma), culminating in a second assemblage more akin to modern counterparts. This shift resulted in an increase in diversity, with the younger assemblage (6–0 Ma) exhibiting greater genus richness than the former late Miocene assemblage. The turnover at 6–4 Ma was driven by peaks in bivalve origination (6–5 Ma) along with elevated extinction rates for gastropods (6–5 Ma) and bivalves (5–4 Ma). Ecological analyses revealed that no single ecological trait consistently changed during this interval, indicating that the turnover resulted from a broad reorganization of ecological strategies. We propose that the major molluscan turnover during the late Miocene–early Pliocene is associated with geomorphological changes related to the Andean uplift, the disappearance of semi-embayments, and a sea-level rise.

We know the fossil record is incomplete, but just how much biodiversity does it miss? We produce the first geographically controlled estimate by comparing the geographic ranges of 34,266 modern tetrapods with a map of the world's sedimentary basins. By modeling which tetrapods live within sedimentary basins, we produce a first-order estimate of what might be found in the fossil record of the future. In this record, nearly 30% of tetrapod species have almost no chance of fossilizing, and more stringent criteria for fossilization exclude far more diversity. This geographically structured fossil record preserves disparate patterns of taxonomic and phylogenetic diversity in different tetrapod groups and underpreserves projected extinctions. For the globally threatened amphibians, the magnitude of the extinction of all endangered species would be underestimated by 66–98% in our future record. These results raise profound questions about the structure of the fossil record. Is it capable of recording major origination and extinction events on land? Have swaths of terrestrial diversity gone unrecorded based on geography alone? There are chapters of Earth history that paleontologists can never hope to know, but what is missing, and why?

The Fezouata Shale Formation has dramatically impacted our understanding of Early Ordovician marine ecosystems before the great Ordovician biodiversification event (GOBE), thanks to the abundance and quality of exceptionally preserved animals within it. Systematic work has noted that the shelly fossil subassemblages of the Fezouata Shale biota are typical of open-marine deposits from the Lower Ordovician, but no studies have tested the quantitative validity of this statement. We extracted 491 occurrences of recalcitrant fossil genera from the Paleobiology Database to reconstruct 31 subassemblages to explore the paleoecology of the Fezouata Shale and other contemporary, high-latitude (66°S–90°S) deposits from the Lower Ordovician (485.4–470 Ma) and test the interpretation that the Fezouata Shale biota is typical for an Ordovician open-marine environment. Sørensen's dissimilarity metrics and Wilcoxon tests indicate that the subassemblages of the Tremadocian-aged lower Fezouata Shale are approximately 20% more heterogenous than the Floian-aged upper Fezouata Shale. Dissimilarity metrics and visualization suggest that while the lower Fezouata and upper Fezouata share faunal components, the two sections have distinct faunas. We find that the faunal composition of the lower Fezouata Shale is comparable with other Tremadocian-aged subassemblages from high latitudes, suggesting that it is typical for an Early Ordovician open-marine environment. We also find differences in faunal composition between Tremadocian- and Floian-aged deposits. Our results corroborate previous field-based and qualitative systematic studies that concluded that the shelly assemblages of the Fezouata Shale are comparable with those of other Lower Ordovician deposits from high latitudes. This establishes the first quantitative baseline for examining the composition and variability within the assemblages of the Fezouata Shale and will be key to future studies attempting to discern the degree to which it can inform our understanding of marine ecosystems just before the start of the GOBE.

The Lilliput Effect, wherein assemblages decrease in mean individual body size after mass extinctions, has not been documented at a wide geographic scale in any of the Late Devonian mass extinction pulses in invertebrate taxa. Based on a dataset of 800 scolecodonts (polychaete jaw elements) from the literature, museum collections, and newly presented data from the Appalachian Basin, we find that scolecodont size distribution per temporal bin decreases across the Frasnian/Famennian Kellwasser Events from a median length of 500 μm before the Kellwasser Events to a median length of 196 μm during the Kellwasser Events. The majority of the small scolecodonts documented during the extinction interval are newly measured specimens from the Kellwasser Events of the Appalachian Basin, although this size change is not unique to the Appalachian Basin. We interpret the reduction in body size as a hypoxia-driven occurrence of the Lilliput Effect because of the susceptibility of benthic invertebrates to hypoxia and the association of this extinction event with hypoxia. While previous studies have shown that polychaete community biomass decreases in response to oxygen stress, our study provides fossil evidence of individual size reduction, plausibly due to oxygen stress.

In the last 50 years, the field of paleobiology has undergone a computational revolution that opened multiple new avenues for recording, storing, and analyzing vital data on the history of life on Earth. With these advances, the amount of data available for research has grown, but so too has our responsibility to ensure that our data tools and infrastructures continue to innovate in order to best serve our diverse community. This review focuses on data equity in paleobiology, an aspirational goal, wherein data in all forms are collected, stored, shared and analyzed in a responsible, equitable, and sustainable manner. While there have been many advancements across the last five decades, inequities persist. Our most significant challenges relate to several interconnected factors, including ethical data collection, sustainable infrastructure, socioeconomic biases, and global inequalities. We highlight the ways in which data equity is critical for paleobiology and stress the need for collaborative efforts across the paleobiological community to urgently address these data equity challenges. We also provide recommendations for actions from individuals, teams, academic publishers, and academic societies in order to continue enhancing data equity and ensuring an equitable and sustainable future for our field.

A traditional typological approach to taxonomy often does not adequately account for intraspecific variation and can result in taxonomic oversplitting. For many groups, including ammonoids of the Placenticeras genus, intraspecific variation documented in recent studies (e.g., ontogenetic changes, sexual dimorphism, polymorphism) challenges the historic proliferation of species names. Here, we used a population approach to taxonomy and quantitatively evaluated morphometric variation in a sample of Late Cretaceous (Santonian–Campanian) Placenticeras from Alabama and adjacent counties.

We used linear mixed models (LMMs) to characterize how morphological variables scale with conch size across the sample, exploiting mixed longitudinal data to evaluate individual variation in growth and inform interpretations of multivariate analyses. Extended LMMs incorporating geological formation evaluated morphological changes through time. Principal component analysis and clustering analysis were then used to evaluate the number of distinct clusters that emerged in multivariate morphospace independent of previous taxon name assignments.

Discontinuous scaling relationships and distinct clusters in multivariate space suggest sexual dimorphism characterized by differences in adult size and, secondarily, shape. Previous Stantonoceras and Placenticeras assignments broadly overlap in our morphospace, failing to justify this historic distinction (as sexual dimorphs or as genera or subgenera). Placenticeras conch morphology and ornament placement changed through time, suggesting a potential utility for coarse (stage-level) biostratigraphy. However, temporal changes were not associated with distinct clusters in morphospace, and our data fail to support the plethora of reported species names. As few as one or two (successive) species may be present in our sample (representing 130 years of collection effort). In addition to highlighting the need for a significant taxonomic revision of the Placenticeras genus, this study demonstrates the utility of LMMs for distinguishing between different sources of morphological variation, improving interpretations of morphospace under a population approach to taxonomy, and maximizing the amount of ontogenetic information that can be obtained nondestructively.

An important question in evolutionary biology and macroecology is whether taxa show systematic trajectories in occupancy, the proportion of geographic area occupied, over macroevolutionary timescales. Past studies have used fossils to document these trajectories, showing a symmetric rise and fall. In this study, I focus on several biases in the analyses of fossil occupancy trajectories that have been unaccounted for. First, better sampling of boundary bins in a taxon’s stratigraphic range, paradoxically, results in lower mean occupancy of taxa in those bins. This is because better sampling allowed more taxa with low occupancies to be included in the mean occupancies of those bins compared with intermediate bins. Second, the possibility that taxa may have incomplete durations within boundary bins could also lower occupancies in those bins. Finally, a bias can also exist when the number of sampled sites is not constant throughout a taxon’s stratigraphic range. I use simulations to show that the first bias can be corrected by conditioning these boundary bins to be sampled in the same way as intermediate bins. To mitigate the second bias, I use higher-resolution time bins to constrain the intervals over which taxa’s occupancies are measured so that they are comparable between boundary and intermediate time bins. I also present an approach that can correct for the last bias by subsampling geographic sites, testing its impact in a simulation. Considering these factors, the occupancy trajectories of marine animal genera look to be a relatively gradual rise post-origination with a sudden decline before extinction.

Sexual dimorphism, a widespread phenomenon, has been extensively researched in extant and fossil crustaceans. However, identifying sexual dimorphism in phyllocarid fossils preserved as isolated parts is often challenging, except in cases where the specimens are exceptionally well preserved, including those with soft tissues. This study proposes a novel approach by introducing the use of geometric morphometric techniques to identify sexual dimorphism in phyllocarid fossils based on carapace morphology. It presents a comprehensive re-analysis of Soomicaris ordosensis Liu et al., 2023a, carapaces from the Upper Ordovician in North China and Tarim Plates. Elliptic Fourier analysis was applied to quantify the size and shape variation in nearly 100 specimens. The results demonstrate the presence of significant sexual dimorphism in the length and shape of the S. ordosensis carapace. The carapace shape exhibited variation between the sexes: the posterodorsal margin of one group of carapaces gradually extends backward to form a posterodorsal spine; the carapaces of the other group have a convex posterior margin and lack a posterodorsal spine. Additionally, both types manifest an overall allometric growth pattern, albeit with distinct growth coefficients. Furthermore, the observed approximately 1:1 ratio between the two forms suggests that the population of S. ordosensis may have exhibited a dioecious mating system. Geometric morphometrics are a highly effective method for elucidating the subtle variations in the carapace morphology of S. ordosensis, thereby underscoring the cryptic dimorphism characteristics of fossil animals. This finding offers the first indirect evidence for egg-brooding behavior within the extinct order Archaeostraca.

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.

Digenean trematodes are parasites with a complex life cycle that often infest shell-bearing mollusks and produce distinct traces on the host skeleton that are recognizable in the fossil record. Here, three bivalve species (Transennella conradina, Abra segmentum, and Chamelea gallina) from Pleistocene and Holocene deposits of Florida and Italy were used to evaluate the hypothesis that trematode infestation affects shell morphology. The morphological effects of infestation were evaluated using geometric morphometrics and the pallial sinus index (PSI = pallial sinus length/shell length). For all three host species: (1) large size classes possess higher trematode prevalence (i.e., proportion of specimens possessing trematode-induced pits within a population) and higher per-specimen frequency of trematode-induced scars when compared with smaller size classes, suggesting ontogenetic accumulation of parasites; and (2) infested and non-infested specimens significantly differ in shell landmark-based morphology. Geometric morphometric analyses indicate that in two out of three species (Transennella conradina, Abra segmentum): (1) PSI and thin-plate spline analyses suggest significant pallial sinus reduction in infested specimens relative to non-infested; and (2) overall morphospace range, estimated by sample-standardized principal component (PC) hypervolume, was inflated with the inclusion of infested specimens. Consistent with previous studies, results indicate that trematode-induced morphological changes may influence the burrowing capabilities of the studied bivalves, affecting their ecological functioning and fitness. Changes in morphospace induced by trematode parasites hamper species delineation and confound morphometric and disparity patterns in the fossil record of infestation-prone species. Excluding fossil specimens with trematode traces can mitigate those confounding effects. Conversely, comparative morphometric analyses of infested and non-infested host specimens may allow us to investigate host responses to parasites over evolutionary timescales.

Body mass is an important facet of reconstructing the paleobiology of fossil species and has, historically, been estimated from individual skeletal measurements. This paper demonstrates the potential advantages of estimating body mass using 3D geometric morphometrics on limb bones, which allows size to be explicitly contextualized within the functional morphology of the animal. Geometric morphometrics of the humerus and femur is used to estimate body mass in domestic dogs and wild canids, and the resulting estimates are compared with estimates made using limb bone dimensions and centroid size. In both groups, 3D methods produced more accurate estimates of body mass than linear dimensions. Additionally, centroid size was a poor predictor of body mass and should not be preferred over linear measurements. The use of 3D methods also reveals specific aspects of shape that are associated with different sizes. In general, relatively heavier individuals were associated with more robust bones and wider articulation sites, as well as larger attachment sites for muscles related to flexion and extension of the shoulder and hip joints. The body-mass equations constructed based on dogs were further evaluated on wild canids, as a test of their potential efficacy on fossil canids. With some adjustments, the body-mass estimation equations made for domestic dogs were able to reliably predict the mass of wild canids. These equations were then used to estimate body mass for a selection of fossil canids: Canis latrans, 16 kg; Aenocyon dirus, 67 kg; Phlaocyon multicuspus, 8 kg; and Hesperocyon gregarius, 2.5 kg.

The Paleozoic represents a key time interval in the origins and early diversification of chondrichthyans (cartilaginous fishes), but their diversity and macroevolution are largely obscured by heterogenous spatial and temporal sampling. The predominantly cartilaginous skeletons of chondrichthyans pose an additional limitation on their preservation potential and hence on the quality of their fossil record. Here, we use a newly compiled genus-level dataset and the application of sampling standardization methods to analyze global total-chondrichthyan diversity dynamics through time from their first appearance in the Ordovician through to the end of the Permian. Subsampled estimates of chondrichthyan genus richness were initially low in the Ordovician and Silurian but increased substantially in the Early Devonian. Richness reached its maximum in the middle Carboniferous before dropping across the Carboniferous/Permian boundary and gradually decreasing throughout the Permian. Sampling is higher in both the Devonian and Carboniferous compared with the Silurian and most of the Permian stages. Shark-like scales from the Ordovician are too limited to allow for some of the subsampling techniques. Our results detect two Paleozoic radiations in chondrichthyan diversity: the first in the earliest Devonian, led by acanthodians (stem-group chondrichthyans), which then decline rapidly by the Late Devonian, and the second in the earliest Carboniferous, led by holocephalans, which increase greatly in richness across the Devonian/Carboniferous boundary. Dispersal of chondrichthyans, specifically holocephalans, into deeper-water environments may reflect a niche expansion following the faunal displacement in the aftermath of the Hangenberg extinction event at the end of the Devonian.

Orthoceratoid cephalopods had straight or slightly curved shells that often contained enigmatic calcareous structures in their chambers. These cameral deposits have been interpreted as counterweights, allowing these cephalopods to assume postures other than a default, downward-facing orientation. These animals must have balanced the proportions of their soft body, cameral deposits, and air-filled chambers to maintain a condition near neutral buoyancy. Lower body chamber ratios (BCRs) allow more mass to be dedicated to cameral deposits, increasing their influence over the total mass distribution. Using 43 computer reconstructions, we calculated the proportion of chamber contents that satisfy a neutrally buoyant condition across different BCRs. Furthermore, we explored the limits of cameral deposit distributions inside the shell to better understand their influence over orientation, stability, and maneuverability. Cephalopods with 40% BCR cannot accommodate any deposits and assume stable, downward-facing orientations. Cephalopods with 30% BCR allow some cameral deposits, which negligibly reduce stability. A slight reduction in BCR to 25% can considerably improve maneuverability, allowing these cephalopods to assume a wider range of postures while swimming. While our results are most relevant to some subset of orthocone cephalopods (Pseudorthoceratida), we also highlight similar constraints faced by broader orthocone groups. Swimming capabilities are extremely sensitive to BCR, which likely constrains the life habits and ecology of these animals. Our results add context to (1) the physical constraints of orthocone cephalopods, (2) their functional complexity in Paleozoic ecosystems, and (3) how these early swimmers navigated physical trade-offs between stability and maneuverability.

The evolution of mammalian innovations like elevated growth rates, endothermy, and live birth has been the subject of paleobiological work for decades. Bone histology provides one of the best lines of evidence for assessing growth rates and life-history traits in the fossil record. However, little ontogenetic information is available for nonmammalian cynodonts, the stock lineage that eventually gave rise to mammals. Here, I report the bone histology of two traversodontid cynodonts from the Triassic Manda Formation of Tanzania. Using two femoral size series, I correlate bone tissue composition and limb size in Scalenodon angustifrons and Luangwa drysdalli. Fifteen individuals were analyzed from seven penecontemporaneous localities to assess intraspecific histovariation within traversodontid ontogenetic development for the first time. My results show that Scalenodon and Luangwa have disparate life histories despite being similarly sized contemporaries. Luangwa is characterized by parallel-fibered bone that transitions to woven-parallel bone early in ontogeny, interpreted as a growth spurt. This increase in growth rate is seen in small- and middle-sized individuals but is resorbed and remodeled in the largest, skeletally mature individual. By contrast, Scalenodon is characterized by woven-parallel tissue in early ontogeny. However, femur size is not correlated with changes in bone tissue composition, as multiple individuals show peripheral slower-growing tissue regardless of size, interpreted as highly developmentally plastic growth. Together, these results demonstrate that coeval members of Traversodontidae show disparate life histories. The underlying mechanisms to explain different life histories in these taxa are likely due to (1) intrinsic differences in growth rates and (2) varying degrees of developmentally flexible growth. The implication of this work is that intraspecific variation in growth dynamics may be more widespread than currently understood in cynodonts and that size is not a good indicator of maturity for some species.

Crocodylomorpha is a large and diverse clade with a long evolutionary history now restricted to modern crocodilians. Tethysuchia is a less-inclusive clade of semi-amphibious taxa that crossed two biological crises: the second Oceanic Anoxic Event (OAE 2) and the Cretaceous/Paleogene (K/Pg) crisis. Numerous studies have sought to find the driving factors explaining crocodylomorph evolution, producing contradictory conclusions. Studies of included groups may be useful. Here, we study factors driving tethysuchian evolution using phylogenetically informed statistical analyses. First, we tested the phylogenetic structure of tethysuchian extinction at the OAE 2 and K/Pg crises. We then used phylogenetic comparative methods to test the influence of intrinsic (body size, snout proportion) and extrinsic (temperature, paleolatitude) factors on the evolution of tethysuchian diversity at the OAE 2 and the K/Pg crises. Finally, we tested whether temperature influenced the evolution of body size. We conclude that (1) extinction was not random in regard to phylogeny for Tethysuchia at the OAE 2 and K/Pg crises; (2) while an important tethysuchian turnover follows OAE 2, the K/Pg crisis was followed by an explosion in diversity of tethysuchians, probably linked to the colonization of emptied ecological niches; (3) tethysuchians lived in warmer environments after the OAE 2 crisis, possibly because of both global warming and latitudinal distribution shifts; (4) there is a significant change of snout proportion after the OAE 2 and the K/Pg crises, likely caused by niche partitioning; and (5) there is a positive correlation between body size and temperature, possibly because of a longer growth season.

Bones preserved in fluvial sediments make up the majority of the terrestrial vertebrate fossil record, and unsteady flows (overbank floods, levee breaches, debris flows, etc.) are often invoked as agents of bone transport and burial. Experiments exploring transport of mammal bones under steady-state flow led to the development of Voorhies Groups, which are used as indicators of winnowing and transport at fossil sites. Some studies have raised concerns about the use of transport groups beyond the scope of the original experiments, especially regarding untested taxa and flow conditions. Here we investigate transport of hadrosauroid dinosaur bone models and modern sheep bones in experimental sheet floods. We find that evolving flow dynamics in unsteady flows can influence bone mobility behaviors. Factors such as bedforms and interactions with other bones caused shorter transport distances than might be expected in some elements, which would be heightened in real flooding situations where trapping mechanisms are common. Our hadrosauroid bones sorted into two statistically significant groups and one overlapping intermediate group based on transport distance. However, those groups could not be identified among sheep bones. Distributions of transport distances in both taxa do not fully match predictions based on Voorhies Groups. Our results indicate that Voorhies Groups do not quantitatively apply to all potential fluvial settings and taxa, and we thus advise caution in interpretations of fossil site taphonomic history based on Voorhies Groups. Further exploration of variables underlying bone transport and burial may allow for more broadly comparative examinations of fluvial biostratinomy.

A central goal in ecology is investigating the impact of major perturbations, such as invasion, on the structure of biological communities. One promising line of inquiry is using co-occurrence analyses to examine how species’ traits mediate coexistence and how major ecological, climatic, and environmental disturbances can affect this relationship and underlying mechanisms. However, present communities are heavily influenced by anthropogenic behaviors and may exhibit greater or lesser resistance to invasion than communities that existed before human arrival. Therefore, to disentangle the impact of individual disturbances on mammalian communities, it is important to examine community dynamics before humans. Here, we use the North American fossil record to evaluate the co-occurrence structure of mammals across the Great American Biotic Interchange. We compiled 126 paleocommunities from the late Pliocene (4–2.5 Ma) and early Pleistocene (2.5–1 Ma). Genus-level co-occurrence was calculated to identify significantly aggregated (co-occur more than expected) and segregated (co-occur less than expected) genus pairs. A functional diversity analysis was used to calculate functional distance between genus pairs to evaluate the relationship between pair association strength and functional role. We found that the strength distribution of aggregating and segregating genus pairs does not significantly change from the late Pliocene to the early Pleistocene, even with different mammals forming the pairs, including immigrant mammals from South America. However, we did find that significant pairs, both aggregations and segregations, became more similar in their functional roles following the Plio-Pleistocene transition. Due to different mammals and ecological roles forming significant associations and the stability of co-occurrence structure across this interval, our study suggests that mammals have fundamental ways of assembling that may have been altered by humans in the present.