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Theropods are obligate bipedal dinosaurs that appeared 230 Ma and are still extant as birds. Their history is characterized by extreme variations in body mass, with gigantism evolving convergently between many lineages. However, no quantification of hindlimb functional morphology has shown whether these body mass increases led to similar specializations between distinct lineages. Here we studied femoral shape variation across 41 species of theropods (n = 68 specimens) using a high-density 3D geometric morphometric approach. We demonstrated that the heaviest theropods evolved wider epiphyses and a more distally located fourth trochanter, as previously demonstrated in early archosaurs, along with an upturned femoral head and a mediodistal crest that extended proximally along the shaft. Phylogenetically informed analyses highlighted that these traits evolved convergently within six major theropod lineages, regardless of their maximum body mass. Conversely, the most gracile femora were distinct from the rest of the dataset, which we interpret as a femoral specialization to “miniaturization” evolving close to Avialae (bird lineage). Our results support a gradual evolution of known “avian” features, such as the fusion between lesser and greater trochanters and a reduction of the epiphyseal offset, independent from body mass variations, which may relate to a more “avian” type of locomotion (more knee than hip driven). The distinction between body mass variations and a more “avian” locomotion is represented by a decoupling in the mediodistal crest morphology, whose biomechanical nature should be studied to better understand the importance of its functional role in gigantism, miniaturization, and higher parasagittal abilities.
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.
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.
Electron backscatter diffraction (EBSD) has been widely used in recent studies of eggshells for its convenience in collecting in situ crystallographic information. China has a wide variety of dinosaur eggshells, although nearly none have been studied with this technique. Elongatoolithid eggs include many oogenera, although the microstructural differences of some were not highly appreciated, leading to several parataxonomic problems. In this paper, we surveyed seven elongatoolithid oogenera in China using EBSD in order to acquire more information about their microstructural variation. It is shown in this paper that in some elongatoolithid eggshells, scaly calcite grains that form the squamatic ultrastructure are not the only form of calcite in the continuous layer. Large columnar grains separated by high-angled grain boundaries and slender subgrains separated by radially arranged low-angled grain boundaries could exist in certain areas of the eggshells such as Macroolithus and Macroelongatoolithus. This paper discusses the criteria for identifying squamatic ultrastructure and proposes type I (rich in rugged high-angled grain boundaries) and type II (rich in both rugged high- and low-angled grain boundaries) squamatic ultrastructures. A pathological layer is found in Undulatoolithus pengi. An external zone is identified in the eggshell of Heishanoolithus changii, which does not support its position within the oofamily Elongatoolithidae. We argue that Paraelongatoolithus no longer belongs to Elongatoolithidae based on a combination of reticulated ornamentation, columnar continuous layer, and acicular mammillae. The high structural variation in elongatoolithid eggshells also implies that it may be inappropriate to relate all previous elongatoolithid eggshells to oviraptorosaurs and assume they are non-monophyletic.
The fossil record is subject to multiple biases that can distort macroevolutionary and paleoecological inferences. Although temporal and spatial sampling biases have received substantial attention, other sources of fossil sampling heterogeneity remain less well quantified. Using the Triton database of planktonic foraminifera, we assess the influence of geographic, ecological, morphological, and methodological factors on fossil recovery rates. We first apply a temporal subsampling method to standardize fossil occurrences over geologic time, validating this approach against an expert-curated lineage-through-time trajectory. After subsampling, the occurrences remain unevenly distributed throughout species’ lifetimes and inhomogeneously distributed across species, reflecting biological signal and/or persistent sampling biases.
We then investigate this residual sampling heterogeneity with a generalized additive model incorporating relevant predictors from Triton. Our results reveal that, after correcting for temporal biases, geographic predictors (paleolatitude, paleolongitude, longitudinal spread) explain nearly a third of sampling variation. Species-specific ecological and morphological attributes contribute an additional fraction, among which mean relative abundance emerges as the main factor. Additional predictors of fossil sampling rates include age-calculation methods and biostratigraphic sampling biases. Despite accounting for multiple sources of variation, 37% of the deviance remains unexplained, suggesting unmodeled biological, stratigraphic, diagenetic, or taxonomic drivers of sampling heterogeneity.
Overall, observed recovery rates question the validity of the homogeneous-sampling assumption used in most diversification models, and this heterogeneity cannot be reduced to a single dominant factor. This conclusion reinforces the need for integrated subsampling approaches and process-based models that explicitly account for heterogeneous fossilization rates to improve the reliability of macroevolutionary analyses.
During the late Miocene and early Pliocene (latest Hemphillian, Hh4 interval, 5.7 to 4.75 Ma) a distinctive suite of four species of extinct horses (Family Equidae) were widespread in North America. This includes Nannippus aztecus, Neohipparion eurystyle, Astrohippus stocki, and Dinohippus mexicanus. In Florida, two additional equid species, Pseudhipparion simpsoni and Cormohipparion emsliei, are also typically found at Hh4 localities. Here we compare horses from four Hh4 Florida fossil sites, including three from the Bone Valley mines and a fourth from the recently discovered Montbrook site. Two of these sites have all six expected species, one has five species, and one has only four species present. To explain these differences, we used species counts from research databases and rarefaction simulation to clarify the relative abundances, species richness, and occurrences of these horses from these four sites. The Palmetto Mine (Agrico) site, with five equid species, appears to lack the sixth species owing to ecological reasons. This is different from Montbrook, the site with only four of the six species. Results indicate that Montbrook is likely lacking two missing equid species for a couple of reasons: sampling bias and biological/ecological causes. Our results demonstrate that sampling biases can account for observed equid species richness when the overall abundance of certain equid species is low. Nevertheless, other factors, including ecology and with sufficient resolution, perhaps also time, may also explain the distribution and occurrences of individual species at these and other fossil sites. In a broader perspective, analyses such as this example provide an opportunity to address a persistent challenge in paleontology, that is, how do we explain absences of extinct taxa from the fossil record?
Over the last 50 years, access to new data and analytical tools has expanded the study of analytical paleobiology, contributing to innovative analyses of biodiversity dynamics over Earth's history. Despite—or even spurred by—this growing availability of resources, analytical paleobiology faces deep-rooted obstacles that stem from the need for more equitable access to data and best practices to guide analyses of the fossil record. Recent progress has been accelerated by a collective push toward more collaborative, interdisciplinary, and open science, especially by early-career researchers. Here, we survey four challenges facing analytical paleobiology from an early-career perspective: (1) accounting for biases when interpreting the fossil record; (2) integrating fossil and modern biodiversity data; (3) building data science skills; and (4) increasing data accessibility and equity. We discuss recent efforts to address each challenge, highlight persisting barriers, and identify tools that have advanced analytical work. Given the inherent linkages between these challenges, we encourage discourse across disciplines to find common solutions. We also affirm the need for systemic changes that reevaluate how we conduct and share paleobiological research.
Traditional approaches to studying paleontological biodiversity have focused on richness, that is, the number of taxa at a given level in the Linnaean hierarchy. While valuable, this approach does not capture the phylogenetic dimension of biodiversity as reflected by its taxonomic structure. This paper introduces a novel perspective in the study of past biodiversity by applying taxonomic distinctness (TD) metrics, average taxonomic distinctness (AvTD) and variation in taxonomic distinctness (VarTD), to investigate the rediversification of marine bivalves following the Permian–Triassic mass extinction. Using a dataset of 59 fossil faunas, we explore the dynamics of the taxonomic tree during the Triassic and along paleolatitudinal gradients. Subtropical assemblages show higher AvTD and lower VarTD than tropical ones, suggesting ecological and biogeographic influences on the taxonomic structure. In the temporal dimension, our data reveal an Early Triassic decrease in AvTD and an increase in VarTD, followed by a long-term trend of increasing AvTD and decreasing VarTD that persisted until the Norian. This pattern suggests that vacant ecospace after the end-Permian mass extinction did not stimulate the immediate appearance of a disproportional number of supraspecific taxa, possibly due to the unusual biotic and abiotic environment of the early postextinction time. In contrast, the long-term increase in AvTD that followed this Early Triassic lag phase indicates a low but steady surplus of supraspecific taxa relative to newly evolved species. This observation supports macroevolutionary scenarios in which evolutionary rates correlate with ecological opportunity. However, the long duration of this phase is unexpected, particularly when compared with conventional indicators for “complete” recovery. Based on TD metrics, recovery from the greatest Phanerozoic mass extinction went on at least until the end of the Norian, nearly 50 Ma after the crisis.
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.
Phylogenetic inferences using combined datasets of both extant and extinct species have grown increasingly popular, in part thanks to the development of the fossilized birth–death (FBD) process. The FBD process provides a powerful model for the evolution of past and present lineages and can be used for both inference and simulation. Simulations in particular are very helpful for new users to gain better understanding of the model and its different components. In this work, we present FossilSimShiny, a visual application for simulating phylogenies, fossil samples, and fossil taxonomies under the FBD process. The app integrates a wide range of simulation models and presents the simulation results in clear, customizable figures. As a teaching tool, FossilSimShiny allows lecturers to create illustration plots and students to directly experiment with the model. For research applications, the app can help researchers save time and effort by testing and calibrating simulation setups before running them on a large scale.
Ontogenetic change is a major source of phenotypic variation among members of a species and is often of greater magnitude than the anatomical differences that distinguish closely related species. Ontogeny has therefore become a problematic confounding variable in vertebrate paleontology, especially in study systems distant from extant crown clades, rendering taxonomic hypothesis testing (a fundamental process in evolutionary biology) rife with difficulty. Paleontologists have adopted quantitative methods to compensate for the perception that juvenile specimens lack diagnostic apomorphies seen in their adult conspecifics. Here, I critically evaluate these methods and the assumptions that guide their interpretation using a μCT dataset comprising growth series of American and Chinese alligator. I find that several widespread assumptions are scientifically unjustifiable and that two popular methods—geometric morphometrics and cladistic analysis of ontogeny—have unacceptably high rates of type II error and present numerous procedural difficulties. However, I also identify a suite of ontogenetically invariant characters that differentiate the living species of Alligator throughout ontogeny. These characters overwhelmingly correspond to anatomical systems that develop before (and play a signaling role in) the development of the cranial skeleton itself, suggesting that their ontogenetic invariance is a consequence of the widely conserved vertebrate developmental program. These observations suggest that the architecture of the cranium is fixed early in embryonic development and that ontogenetic remodeling does not alter the topological relationships of the cranial bones or the soft tissue structures they house. I propose a general model for future taxonomic hypothesis tests in the fossil record, in which the hypothesis that two specimens are different ontogenetic stages of a single species can be falsified by the discovery of character differences that cannot be attributed plausibly to ontogenetic variation.
The present-day climate crisis is transforming coral reef communities, potentially undermining ecosystem functioning. Evolutionary trade-offs between species traits result in diverse life-history strategies, enabling corals to survive disturbance events through specific adaptive mechanisms. Trait–trait relationship networks offer insights into trait turnover and changing life-history strategies during environmental changes. Paleoecological insights from the fossil records can further illustrate how species adapt to environmental shifts, highlighting resilience traits.
We highlight coral traits that promote resilience in the Caribbean based on fossil occurrences and morphological traits, examining biological determinants of species and trait turnover across the Cenozoic. We use traits that underpin the survival of corals during disturbances, for example, corallite diameter, colony growth form, corallite integration, and budding type. We analyzed species turnover and extinctions with a bipartite network and explored trait turnover with trait–trait co-occurrence networks based on 4268 species records at 421 sites over ~40 Myr.
Our findings support existing evidence that species turnover coincided with major environmental and biogeographic changes across the Cenozoic. Additionally, our results provide new insight into functional changes throughout the Cenozoic. Past cooler climates favored corals with a fast growing and reproducing (competitive) life-history strategy, which boosts short-term success, but also increases susceptibility to diseases and thermal stress. Cenozoic species and trait turnover occurred during environmental change, corroborating expectations of such turnover in the future. We found trait co-occurrence modules associated with competitive and stress-tolerant life-history strategies. The transition from the “greenhouse” (Paleogene) to the “icehouse” (Neogene) climate over ~40 Myr favored competitive traits, which supported fast-growing, shallow reefs. With rising temperatures and declining Acropora in the Caribbean, future reefs may resemble Eocene reefs: dominated by stress-tolerant, slow-growing corals adapted to marginal environments.
The trend of global cooling across the Cenozoic transformed the North American landscape from closed forest to more open grasslands, resulting in dietary adaptations in herbivores in response to shifting resources. In contrast, the material properties of the predator food source (muscle, skin, and bone) have remained constant over this transition, suggesting a corresponding lack of change in predator dietary adaptations. We investigated the North American mammalian predator fossil record using a tooth-shape metric and body mass, predicting that the former would exhibit stability. Instead, we found that mean molar morphology became more blade-like, with our tooth-shape metric sharply increasing in the late Eocene and remaining high from the Oligocene onward. Subsequent tests in extant carnivorans reveal taxa with more bladelike teeth are prevalent in more open environments. Our results reveal an unexpected functional shift among North American predators in response to large-scale environmental changes across the Cenozoic.
Paleo-ecological niche modeling (paleoENM) estimates the niches and distributions of extinct species using fossil paleo-coordinates and local environmental data. While general circulation models (GCMs) have been used to estimate climate conditions in deep time, primarily for terrestrial vertebrates, variations in paleo-elevation models used in GCM construction can influence paleoENM outcomes. This study (1) examines the impact of the Cretaceous–Paleogene (K-Pg) mass extinction on the niche dimensions of the marine invertebrate group Turritellinae (Cerithoidea: Turritellidae) and (2) compares two paleo-elevation models’ effects on GCM-based species’ distribution predictions. Fossil occurrence data from the Maastrichtian and Danian periods were collected from the Paleobiology Database (PBDB), museum collections, and published literature. Environmental data were extracted from HadCM3L GCM simulations using Scotese- and Getech-based paleogeographic and pCO2 boundary conditions. We estimated the niche dimensions of turritellines using maximum entropy (MaxEnt) and performed ordination analysis using kernel density estimation. MaxEnt model metrics showed that the Getech-based GCM outperformed the Scotese-based GCM. Geographic projections revealed minor differences in suitable habitat between the Maastrichtian and Danian in the Getech-based GCM, but overinflated predictions in the Scotese-based GCM. Niche overlap between the Maastrichtian and Danian was high, with both GCMs supporting niche similarity and equivalency. Our results suggest that differences in elevation model boundary conditions affected predicted distribution and niche patterns. This study offers a novel approach to understanding ecological persistence in invertebrates after mass extinction events, examines the robustness of GCM boundary conditions in paleoENM studies, and provides a framework for future paleoecological research on fossil invertebrates.
The mammalian skull plays important roles in sensing, communication, and obtaining and processing food. These diverse functions make the skull a natural target for investigating how developmental processes and selective pressures related to ecology interact to produce patterns of morphological evolution. Here, we investigate the evolution of the equid skull through the lens of craniofacial evolutionary allometry (CREA), a pattern of relative facial elongation in larger mammals that has been hypothesized to result from similar patterns of facial elongation over ontogeny. Using 3D geometric morphometrics and linear measurements, we describe the major axes of shape variation in the equid skull, test whether equids follow or deviate from the CREA pattern over the clade’s evolutionary history, and assess whether the evolution of high-crowned teeth (hypsodonty) is related to relative facial proportions, all in an explicitly phylogenetic context. We find that equids deviate from the CREA pattern and that the evolution of hypsodont dentition did not significantly influence facial proportions in the group. Importantly, these results are only apparent when using statistics appropriate for phylogenetic data. Comparison with previously published data on facial proportions from modern equids indicates that ontogenetic patterns of facial elongation do not scale to produce patterns of facial proportions observed at the intraspecific and evolutionary levels. Taken together, our results complicate the historic narrative that a single set of selective factors drove patterns of morphological evolution within the group.
Caryocaridids are a unique representative of pelagic arthropods from the Ordovician period. They are typically found as flattened carapaces in mudstones and shales. This study reports on a species of caryocaridids, Soomicaris cedarbergensis, discovered in the Lower Ordovician of northwestern Xinjiang, NW China. The species shows the rare enrolled carapaces with a preserved cuticular ultrastructure. These specimens of caryocaridids from Xinjiang are the first reported in the Yili Block, and provide the substantial evidence that the paleogeographic distribution of caryocaridid phyllocarids could extend to the Central Asian Orogenic Belt. This species existed from the late Tremadocian until the end of the Ordovician (Hirnantian), making it the longest-ranging known species of caryocaridids. The carapace cuticle of S. cedarbergensis is composed of carbonate-fluorapatite and can be divided into three mineralized lamellae: outer, middle, and inner. The outer and inner lamellae each consist of three layers that correspond to the epicuticle, exocuticle, and endocuticle of extant crustacean carapaces. Moreover, the polygonal reticulation structure of the carapace in archaeostracans appears to be similar in shape and size to the hemolymph sinuses of leptostracans. This unique ultrastructure of the carapace cuticle in caryocaridids is believed to be better suited for a pelagic lifestyle.
Sauropodomorphs were the largest terrestrial animals to have ever existed on Earth.
While previous studies using skeletal remains and digital 3D models suggested that manus became more circular shape and relatively larger over time, this study uses trackways to test these hypotheses with a much larger sample size. In this study, we used the sauropodomorph trackway record from the Early Jurassic to the Late Cretaceous to analyze temporal changes in manus and pes shapes and their relationship with body size. We analyzed footprint measurements from 690 trackways and, for the first time, applied elliptic Fourier descriptors (EFDs) to sauropodomorph footprint outlines to quantitatively assess their morphology. Analyses revealed temporal and size-related changes in manus morphology, whereas no changes were observed in pedes. Specifically, manus shape became anteroposteriorly longer over time and in larger individuals. The relative manus area compared with pes area (the heteropody) also increased chronologically; however, manus areas showed negative allometry relative to pes areas. These results indicate that the temporal change toward circular or horseshoe-shaped manus morphology in sauropodomorphs represents an adaptation to gigantism, while the chronological increase in manus size relative to the pes reflects the radiation of a clade (Titanosauria) with proportionally large manus.
Fossils from the deep-sea Ediacaran biotas of Newfoundland are among the oldest architecturally complex soft-bodied macroorganisms on Earth. Most organisms in the Mistaken Point–type biotas of Avalonia—particularly the fractal-branching frondose Rangeomorpha— have been traditionally interpreted as living erect within the water column during life. However, due to the scarcity of documented physical sedimentological proxies associated with fossiliferous beds, Ediacaran paleocurrents have been inferred in some instances from the preferential orientation of fronds. This calls into question the relationship between frond orientation and paleocurrents. In this study, we present an integrated approach from a newly described fossiliferous surface (the “Melrose Surface” in the Fermeuse Formation at Melrose, on the southern portion of the Catalina Dome in the Discovery UNESCO Global Geopark) combining: (1) physical sedimentological evidence for paleocurrent direction in the form of climbing ripple cross-lamination and (2) a series of statistical analyses based on modified polythetic and monothetic clustering techniques reflecting the circular nature of the recorded orientation of Fractofusus misrai specimens. This study demonstrates the reclining rheotropic mode of life of the Ediacaran rangeomorph taxon Fractofusus misrai and presents preliminary inferences suggesting a similar mode of life for Bradgatia sp. and Pectinifrons abyssalis based on qualitative evidence. These results advocate for the consideration of an alternative conceptual hypothesis for position of life of Ediacaran organisms in which they are interpreted as having lived reclined on the seafloor, in the position that they are preserved.
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.
Tribrachidium heraldicum is an Ediacaran body fossil characterized by triradial symmetry. Previous work has suggested that the anatomy of Tribrachidium was conducive to passive suspension feeding; however, these analyses used an inaccurate model and a relatively simple set of simulations. Using computational fluid dynamics, we explore the functional morphology of Tribrachidium in unprecedented detail by gauging how the presence or absence of distinctive anatomical features (e.g., apical pits and arms) affects flow patterns. Additionally, we map particle pathways, quantify deposition rates at proposed feeding sites, and assess gregarious feeding habits to more fully reconstruct the lifestyle of this enigmatic taxon. Our results provide strong support for interpreting Tribrachidium as a macroscopic suspension feeder, with the apical pits representing loci of particle collection (and possibly ingestion) and the triradial arms representing morphological adaptations for interrupting flow and inducing settling. More speculatively, we suggest that the radial grooves may represent ciliated pathways through which food particles accumulating in the wake of the organism were transported toward the apical pits. Finally, our results allow us to generate new functional hypotheses for other Ediacaran taxa with a triradial body plan. This work refines our understanding of the appearance of suspension feeding in shallow-water paleoenvironments, with implications for the radiation of Metazoa across the Ediacaran/Cambrian boundary.