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bioRxiv2019

Constructing a Timescale of Biotic Recovery across the Cretaceous Paleogene Boundary, Corral Bluffs, Denver Basin, Colorado

ABSTRACTThe Cretaceous–Paleogene (K–Pg) boundary interval represents one of the most significant mass extinctions and ensuing biotic recoveries in Earth history. Earliest Paleocene fossil mammal faunas corresponding to the Puercan North American Land Mammal Age (NALMA) are thought to be highly endemic and potentially diachronous, necessitating precise chronostratigraphic controls at key fossil localities to constrain recovery dynamics in continental biotas following the K–Pg mass extinction. The Laramide synorgenic sedimentary deposits within the Denver Basin preserve one of the most continuous and fossiliferous records of the K–Pg boundary interval in North America. However, poor exposure in much of the Denver Basin makes it difficult to correlate between outcrops. In order to constrain fossil localities across the basin, previous studies have relied upon chronostratigraphic methods such as magnetostratigraphy. Here we present a new high-resolution magnetostratigraphy of 10 lithostratigraphic sections spanning the K–Pg boundary interval at Corral Bluffs located east of Colorado Springs in the southern part of the Denver Basin. Fossil localities from Corral Bluffs have yielded limited dinosaur remains, mammal fossils assigned to the Puercan NALMA, and numerous fossil leaf localities. Palynological analysis identifying the K–Pg boundary in three sections and two independent but nearly identical 206Pb/238U age estimates for the same volcanic ash, provide key temporal calibration points. Our paleomagnetic analysis has identified clear polarity reversal boundaries from Chron C30n to Chron C28r across the sections. It is now possible to place the fossil localities at Corral Bluffs within the broader basin-wide chronostratigraphic framework and evaluate them in the context of K–Pg boundary extinction and recovery.View Full Text

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bioRxiv2019

Saving Old Bones: a non-destructive method for bone collagen prescreening

Bone collagen is an important material for radiocarbon, paleodietary, and paleoproteomic analyses, but it degrades over time. Various methods have been employed to prescreen bone for collagen preservation, but these are often destructive and/or require exportation for analysis. Here we show that near-infrared spectroscopy can be used to determine bone collagen content quickly and non-destructively on site.rView Full Text

paleontology↗

bioRxiv2019

Fossil amber reveals springtails’ longstanding dispersal by social insects

Dispersal is essential for terrestrial organisms living in disjunct habitats and constitutes a significant challenge for the evolution of wingless taxa. Springtails (Collembola), the sister-group of all insects (with dipluran), are reported since the Lower Devonian and thought to have originally been subterranean. The order Symphypleona is reported since the early Cretaceous with genera distributed on every continent, implying an ability to disperse over oceans although never reported in marine water contrary to other springtail orders. Despite being highly widespread, modern springtails are generally rarely reported in any kind of biotic association. Interestingly, the fossil record has provided occasional occurrences of Symphypleona attached by the antennae onto the bodies of larger arthropods. Here, we document the case of a ~16 Ma old fossil association: a winged termite and ant displaying not some, but 25 springtails attached or closely connected to the body. The collembola exhibit rare features for fossils, reflecting their courtship and phoretic behaviors. By observing the modes of attachment of springtails on different arthropods, the sex representation and ratios in springtail antennal anatomies in new and previously reported cases, we infer a likely mechanism for dispersal in Symphypleona. By revealing hidden evidence of modern springtail associations with other invertebrates such as ants and termites, new compelling assemblages of fossil springtails and the drastic increase of eusocial insects’ abundance over Cenozoic (ants/termites comprising more than the third of insects in Miocene amber), we stress that attachment with winged casts of ants and termites may have been a mechanism for the worldwide dispersal of this significant springtail lineage. Moreover, by comparing the general constraints applying to the other wingless soil-dwelling arthropods known to disperse through phoresy, we suggest biases in the collection and observation of phoretic Symphypleona related to their reflexive detachment and infer that this behavior continues today. The specific case of tree resin entrapment represents the (so far) only condition uncovering the actual dispersal mechanism of springtails - one of the oldest terrestrial arthropod lineages living today. Associations with soil-dwelling social insects over time would have been at the origin of this behavioural specialization.View Full Text

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bioRxiv2019

Epidemiology of human culture: parietal rock art requires high population density

Demographic models of human cultural evolution have high explanatory potential but weak empirical support. Here we use a global dataset of rock art sites and climate and genetics-based estimates of ancient population densities to test a new model based on epidemiological principles. The model focuses on the process whereby a cultural innovation becomes endemic in a population. It predicts that this cannot occur unless population density exceeds a critical value. Analysis of the data, using a Bayesian statistical framework, shows that the model has stronger empirical support than a null model, where rock art detection rates and population density are independent, or a proportional model where detection is directly proportional to population density. Comparisons between results for different geographical areas and periods yield qualitatively similar results, supporting the robustness of the model. Re-analysis of the rock art data, using a second set of independent population estimates, yields similar results. We conclude that population density above a critical threshold is a necessary condition for the maintenance of rock art as a stable part of a populations cultural repertoire. Methods similar to those described can be used to test the model for other classes of archaeological artifact and to compare it against other models.

paleontology↗

bioRxiv2019

SYSTEMATIC REVISION AND REDEFINITION OF THE GENUS SCIRROTHERIUM EDMUND & THEODOR, 1997 (CINGULATA, PAMPATHERIIDAE): IMPLICATIONS FOR THE ORIGIN OF PAMPATHERIIDS AND THE EVOLUTION OF THE SOUTH AMERICAN LINEAGE INCLUDING HOLMESINA

The intrageneric relationships of the pampatheriid genus Scirrotherium and its affinities with supposedly related genera, i.e., Kraglievichia and Holmesina, are revised through parsimony phylogenetic analyses and new comparative morphological descriptions. For this work, unpublished material of pampatheriids (numerous osteoderms, one partial skull and a few postcranial bones) from Neogene formations of Colombia was analyzed. The results show that Scirrotherium is paraphyletic if we include all its referred species, i.e., Scirrotherium hondaensis, S. carinatum and S. antelucanus. The species S. carinatum is closer to Kraglievichia paranensis than to S. hondaensis or S. antelucanus, therefore the new name K. carinatum comb. nov. is proposed. The relationship among S. hondaensis and S. antelucanus could not be resolved, so these species should be designated in aphyly. In spite of failing to recover S. hondaensis and S. antelucanus as one single clade, here is preferred to maintain the generic name Scirrotherium in both species based on diagnostic evidence. New emended diagnoses for Scirrotherium, S. hondaensis and Kraglievichia are provided. The genus Holmesina was found to be monophyletic and positioned as the sister clade of Scirrotherium + Kraglievichia. The evolutionary and biogeographic implications of the new phylogeny and taxonomic re-arrangements are discussed. A possible geographic origin of the family Pampatheriidae and Scirrotherium in low latitudes of South America as early as Early Miocene times is claimed. The South American ancestor or sister taxon of Holmesina is predicted to be morphologically more similar to Scirrotherium than to Kraglievichia.View Full Text

paleontology↗

bioRxiv2019

Crocodylomorph cranial shape evolution and its relationship with body size and ecology

Crocodylomorpha, which includes living crocodylians and their extinct relatives, has a rich fossil record, extending back for more than 200 million years. Unlike modern semi-aquatic crocodylians, extinct crocodylomorphs exhibited more varied lifestyles, ranging from marine to fully terrestrial forms. This ecological diversity was mirrored by a remarkable morphological disparity, particularly in terms of cranial morphology, which seems to be closely associated with ecological roles in the group. Here, I use geometric morphometrics to comprehensively investigate cranial shape variation and disparity in Crocodylomorpha. I quantitatively assess the relationship between cranial shape and ecology (i.e. terrestrial, aquatic, and semi-aquatic lifestyles), as well as possible allometric shape changes. I also characterise patterns of cranial shape evolution and identify regime shifts. I found a strong link between shape and size, and a significant influence of ecology on the observed shape variation. Terrestrial taxa, particularly notosuchians, have significantly higher disparity, and shifts to more longirostrine regimes are associated with large-bodied aquatic or semi-aquatic species. This demonstrates an intricate relationship between cranial shape, body size and lifestyle in crocodylomorph evolutionary history. Additionally, disparity-through-time analyses were highly sensitive to different phylogenetic hypotheses, suggesting the description of overall patterns among distinct trees. For crocodylomorphs, most results agree in an early peak during the Early Jurassic and another in the middle of the Cretaceous, followed by nearly continuous decline until today. Since only crown-group members survived through the Cenozoic, this decrease in disparity was likely the result of habitat loss, which narrowed down the range of crocodylomorph lifestyles.View Full Text

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bioRxiv2019

Early Tetrapodomorph Biogeography: Controlling for Fossil Record Bias in Macroevolutionary Analyses

ABSTRACTThe fossil record provides direct empirical data for understanding macroevolutionary patterns and processes. Inherent biases in the fossil record are well known to confound analyses of this data. Sampling bias proxies have been used as covariates in regression models to test for such biases. Proxies, such as formation count, are associated with paleobiodiversity, but are insufficient for explaining species dispersal owing to a lack of geographic context. Here, we develop a sampling bias proxy that incorporates geographic information and test it with a case study on early tetrapodomorph biogeography. We use recently-developed Bayesian phylogeographic models and a new supertree of early tetrapodomorphs to estimate dispersal rates and ancestral habitat locations. We find strong evidence that geographic sampling bias explains supposed radiations in dispersal rate (potential adaptive radiations). Our study highlights the necessity of accounting for geographic sampling bias in macroevolutionary and phylogenetic analyses and provides an approach to test for its effect.View Full Text

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bioRxiv2019

The oldest freshwater crabs: claws on dinosaur bones

With approximately 1,500 extant species, freshwater crabs (Decapoda: Brachyura) are among the most diverse decapod crustaceans. Nevertheless, their fossil record is extremely limited: only Potamidae, Potamonautidae and Trichodactylidae are reported up to the Eocene of the Neotropics so far. This work documents unusually large decapod claws from the Upper Cretaceous (Campanian) continental deposits of Velaux and vicinity (southern France), in close association with large vertebrate remains. In addition to (1) the systematic assignment of these claws, the study addresses (2) the salinity trends in the deposit environment from its faunal assemblage and the elementary chemical patterns of fossils, and (3) the likely scenario for their auto/allochtony in the Velaux fluvial system. These claws belong to a new taxon, Dinocarcinus velauciensis n. gen. n. sp., referred to as Portunoidea sensu lato, a group of “true” crabs nowadays linked to marine systems. However, the faunal assemblage, the claw taphonomy and the carbonates Y/Ho signatures support their ancient freshwater/terrestrial ecology, making them the oldest reported continental brachyurans and extending the presence of crabs in freshwater environments by 40 Ma. Either as primary or as secondary freshwater crabs, the occurrence of these portunoids in Velaux is an evidence for the independent colonizations of continental environments by multiple brachyuran clades over time, as early as the Campanian.View Full Text

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bioRxiv2019

Defining variation in pre-human ecosystems can guide conservation: An example from a Caribbean coral reef

There is a consensus that Caribbean coral reefs are a pale shadow of what they once were, yet a reef’s pre-human state is typically assumed or estimated using space-for-time substitution approaches. These approaches may fail to account for past variation before human impact which could mislead conservation priorities and actions. In this study we use a suite of fossilised mid-Holocene (7.2-5.6 ka) fringing reefs in Caribbean Panama to define the Historical Range of Variation (HRV) in coral community structure before human-impact to provide context for the states of modern reefs in the same area. Using the abundances of coral taxa to quantify communities, we found that most of the modern coral communities exist in novel ecosystem states with no fossil precedence. We do however identify one modern reef that is indistinguishable in coral community structure from the mid-Holocene reefs. Reef-matrix cores show that the community on this reef has remained in a stable state for over 760 years, suggesting long-term resistance to the region-wide shift to novel states. Without historical context this robust and stable reef would be overlooked since it does not fulfil expectations of what a “pristine” coral reef should look like. This example illustrates how defining past variation using the fossil record can place modern degradation in historical context and improve conservation recommendations.View Full Text

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bioRxiv2019

Dispersal ability predicts evolutionary success among mammalian carnivores

Understanding why some clades contain more species than others is a major challenge in evolutionary biology, and variation in dispersal ability and its connection to diversification rate may be part of the explanation. Several studies have suggested a negative relationship between dispersal capacity and diversification rate among living mammals. However, this pattern may differ when also considering extinct species, given known extinction biases. The colonization of new areas by various lineages may be associated with both diversity increases in those colonising lineages and declines in the lineages already present. Past diversity declines are, however, effectively impossible to infer based on phylogenies of extant taxa, and the underlying process may, therefore, be difficult to determine. Here we produce a novel species-level phylogeny of all known extant and extinct species of the order Carnivora and related extinct groups (1,723 species in total) to show that there is instead a positive relationship between dispersal rate and diversification rate when all extinct species are included. Species that disperse between continents leave more descendant species than non-dispersers, and dispersing species belong to lineages that at the time of dispersal were diversifying faster than the average non-disperser. Our study showcases the importance of combining fossils and phylogenies to better understand evolutionary and biogeographic patterns.View Full Text

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bioRxiv2019

Hydraulic diversity of El Cien Formation (Baja California Sur, Mexico) and the consequences of functional diversity in paleoclimate estimation using fossil wood

ABSTRACTCommunity assembly processes, environmental filtering and limiting similarity, determine functional traits values within communities. Because environment influences the number of viable functional strategies species might take, a strong effect of environmental filter often results in communities having species with similar trait values and narrow functional niches. On the other hand, limiting similarity lead to communities with broader functional spaces. The degree to community assembly processes influence wood trait variation has important implications for paleoclimate estimation using fossil wood since the main tenet of the approach is environmental driven trait convergence, and assumes a central role of environmental filtering. We used functional diversity (FD) to determine how three wood anatomical traits vary in 14 extant communities (272 species) growing under different climate regimes, and inferred the prevalence of environmental filtering/limiting similarity. We also calculated FD metrics for the El Cien Formation fossil woods and discussed the results in light of the current knowledge of the flora. We found lower anatomical diversity in communities growing in dry/cool places (smaller functional spaces and lower abundance of trait combinations), suggesting that strong wood anatomical trait convergence could be the result of stronger habitat filtering in these communities. A lower strength of environmental filter in warm/wet environments, likely results in an amplification of the role of other drivers that promote higher number of hydraulic strategies through niche partition in highly structured communities. More complex ecological structures in mild tropical places likely lead to a higher spread of wood trait values. This asymmetry in the strength of environmental filter along climate gradients, suggest that the imbalances in strength of the trait-climate convergence, should be incorporated in paleoclimate prediction models. FD approach can be used to recognize promising traits with narrow niches along climate gradients, and therefore a constant effect of environmental filter.View Full Text

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bioRxiv2019

The fast and the frugal: Divergent locomotory strategies drive limb lengthening in theropod dinosaurs

Limb length, cursoriality and speed have long been areas of significant interest in theropod paleobiology as locomotory capacity, especially running ability, is critical in not just in prey pursuit but also to avoid become prey oneself. One aspect that is traditionally overlooked is the impact of allometry on running ability and the limiting effect of large body size. Since several different non-avian theropod lineages have each independently evolved body sizes greater than any known terrestrial carnivorous mammal, ∼1000kg or more, the effect that such larger mass has on movement ability and energetics is an area with significant implications for Mesozoic paleoecology. Here using expansive datasets, incorporating several different metrics to estimate body size, limb length and running speed, to calculate the effects of allometry running We test both on traditional metrics used to evaluate cursoriality in non-avian theropods such as distal limb length, relative hindlimb length as well as comparing the energetic cost savings of relative hindlimb elongation between members of the Tyrannosauridae and more basal megacarnivores such as Allosauroids or Ceratosauridae. We find that once the limiting effects of body size increase is incorporated, no commonly used metric including the newly suggested distal limb index (Tibia + Metatarsus/ Femur length) shows a significant correlation to top speed. The data also shows a significant split between large and small bodied theropods in terms of maximizing running potential suggesting two distinct strategies for promoting limb elongation based on the organisms’ size. For small and medium sized theropods increased leg length seems to correlate with a desire to increase top speed while amongst larger taxa it corresponds more closely to energetic efficiency and reducing foraging costs. We also find, using 3D volumetric mass estimates, that the Tyrannosauridae show significant cost of transport savings compared to more basal clades, indicating reduced energy expenditures during foraging and likely reduced need for hunting forays. This suggests that amongst theropods while no one strategy dictated hindlimb evolution. Amongst smaller bodied taxa the competing pressures of being both a predator and a prey item dominant while larger ones, freed from predation pressure, seek to maximize foraging ability. We also discuss the implications both for interactions amongst specific clades and Mesozoic paleobiology and paleoecological reconstructions as a whole.View Full Text

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bioRxiv2019

Reptile-like physiology in Early Jurassic stem-mammals

There is uncertainty regarding the timing and fossil species in which mammalian endothermy arose, with few studies of stem-mammals on key aspects of endothermy such as basal or maximum metabolic rates, or placing them in the context of living vertebrate metabolic ranges. Synchrotron X-ray imaging of incremental tooth cementum shows two Early Jurassic stem-mammals, Morganucodon and Kuehneotherium, had lifespans (a basal metabolic rate proxy) considerably longer than comparably sized living mammals, but similar to reptiles, and that Morganucodon had femoral blood flow rates (a maximum metabolic rate proxy) intermediate between living mammals and reptiles. This shows maximum metabolic rates increased evolutionarily before basal rates, and that contrary to previous suggestions of a Triassic origin, Early Jurassic stem-mammals lacked the endothermic metabolism of living mammals.One Sentence Summary Surprisingly long lifespans and low femoral blood flow suggest reptile-like physiology in key Early Jurassic stem-mammals.View Full Text

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bioRxiv2019

Evolution of crab eye structures and the utility of ommatidia morphology in resolving phylogeny

ABSTRACTImage-forming compound eyes are such a valuable adaptation that similar visual systems have evolved independently across crustaceans. But if different compound eye types have evolved independently multiple times, how useful are eye structures and ommatidia morphology for resolving phylogenetic relationships? Crabs are ideal study organisms to explore these questions because they have a good fossil record extending back into the Jurassic, they possess a great variety of optical designs, and details of eye form can be compared between extant and fossil groups. True crabs, or Brachyura, have been traditionally divided into two groups based on the position of the sexual openings in males and females: the so-called ‘Podotremata’ (females bearing their sexual openings on the legs), and the Eubrachyura, or ‘higher’ true crabs (females bearing their sexual openings on the thorax). Although Eubrachyura appears to be monophyletic, the monophyly of podotreme crabs remains controversial and therefore requires exploration of new character systems. The earliest podotremous lineages share the plesiomorphic condition of ‘mirror’ reflecting superposition eyes with most shrimp, lobsters, and anomurans (false crabs and allies). The optical mechanisms of fossil and extant podotreme groups more closely related to Eubrachyura, however, are still poorly investigated. To better judge the phylogenetic utility of compound eye form, we investigated the distribution of eye types in fossil and extant podotreme crabs. Our findings suggest the plesiomorphic ‘mirror’ eyes—seen in most decapod crustaceans including the earliest true crabs—has been lost in several ‘higher’ podotremes and in eubrachyurans. We conclude that the secondary retention of larval apposition eyes has existed in eubrachyurans and some podotremes since at least the Early Cretaceous, and that the distribution of eye types among true crabs supports a paraphyletic podotreme grade, as suggested by recent molecular and morphological phylogenetic studies. We also review photoreceptor structure and visual pigment evolution, currently known in crabs exclusively from eubrachyuran representatives. These topics are critical for future expansion of research on podotremes to deeply investigate the homology of eye types across crabs.View Full Text

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bioRxiv2019

Modeling Dragons: Using linked mechanistic physiological and microclimate models to explore environmental, physiological, and morphological constraints on the early evolution of dinosaurs

We employed the widely-tested biophysiological modeling software, Niche Mapper™ to investigate the metabolic function of Late Triassic dinosaurs Plateosaurus and Coelophysis during global greenhouse conditions. We tested them under a variety of assumptions about resting metabolic rate, evaluated within six microclimate models that bound paleoenvironmental conditions at 12° N paleolatitude, as determined by sedimentological and isotopic proxies for climate within the Chinle Formation of the southwestern United States. Sensitivity testing of metabolic variables and simulated “metabolic chamber” analyses support elevated “ratite-like” metabolic rates and intermediate “monotreme-like” core temperature ranges in these species of early saurischian dinosaur. Our results suggest small theropods may have needed partial to full epidermal insulation in temperate environments, while fully grown prosauropods would have likely been heat stressed in open, hot environments and should have been restricted to cooler microclimates such as dense forests (under any vegitative cover) or those seen at higher latitudes and elevations. This is in agreement with the Late Triassic fossil record and may have contributed to the latitudinal gap in the Triassic prosauropod record.View Full Text

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bioRxiv2019

First fossil skull of an anteater (Vermilingua, Myrmecophagidae) from northern South America, a taxonomic reassessment of Neotamandua and a discussion of the myrmecophagid diversification

The evolutionary history of the South American anteaters, Vermilingua, is incompletely known as consequence of the fragmentary and geographically biased nature of the fossil record of this group. Neotamandua borealis is the only recorded extinct species from northern South America, specifically from the Middle Miocene of La Venta area, southwestern Colombia. A new genus and species of myrmecophagid for La Venta, Gen. et sp. nov., is here described based on a new partial skull. Additionally, given that the co-occurrent species of Gen. et sp. nov., N. borealis, was originally referred to as Neotamandua, the taxonomic status of this genus is revised. The morphological and taxonomic analyses of these taxa indicate that Gen. et sp. nov. may be related to Tamandua and that the justification of the generic assignments of the species referred to as Neotamandua is weak or insufficient. Two species previously referred to as Neotamandua (N. magna and N.? australis) were designated as species inquirendae and new diagnostic information for the redefined genus and its type species, N. conspicua, is provided. Together, these results suggest that the diversification of Myrmecophagidae was taxonomically and biogeographically more complex than what has been proposed so far. Considering the new evidence, it is proposed a synthetic model on the diversification of these xenartrans during the late Cenozoic based on the probable relationships between their intrinsic ecological constraints and some major abiotic changes in the Americas.

paleontology↗

bioRxiv2019

Classification of down-core foraminifera image sets using convolutional neural networks.

Manual identification of foraminifera species or morphotypes under stereoscopic microscopes is time-consuming for the taxonomist, and a long-time goal has been automating this process to improve efficiency and repeatability. Recent advances in computation hardware have seen deep convolutional neural networks emerge as the state-of-the-art technique for image-based automated classification. Here, we describe a method for classifying large down-core foraminifera image set using convolutional neural networks. Construction of the classifier is demonstrated on the publically available endless forams image set with an best accuracy of approximately 90%. A complete down-core analysis is performed for benthic species in the Holocene period for core MD02-2518 from the North Eastern Pacific, and the relative abundances compare favourably with manual counting, showing the same signal dynamics. Using our workflow opens the way to automated paleo-reconstruction based on computer image analysis, and can be employed using our labelling and classification software ParticleTrieur.View Full Text

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bioRxiv2019

A unique predator in a unique ecosystem: modelling the apex predator from the Late Cretaceous crocodyliform-dominated fauna in Brazil.

ABSTRACTTheropod dinosaurs were relatively scarce in the Late Cretaceous ecosystems of southeast Brazil. Instead, hypercarnivorous crocodyliforms known as baurusuchids were abundant and probably occupied the ecological role of apex predators. Baurusuchids exhibited a series of morphological adaptations hypothesised to be associated with this ecological role, but quantitative biomechanical analyses of their morphology have so far been lacking. Here, we employ a biomechanical modelling approach, applying finite element analysis (FEA) to models of the skull and mandibles of a baurusuchid specimen. This allowed us to characterise the craniomandibular apparatus of baurusuchids, as well as to compare the functional morphology of the group to that of other archosaurian carnivores, such as theropods and crocodylians. Our results support the ecological role of baurusuchids as specialised apex predators in the continental Late Cretaceous ecosystems of South America. With a relatively weak bite force (∼600 N), baurusuchids’ predation strategies likely relied on other morphological specializations, such as ziphodont dentition and strong cervical musculature. Comparative assessments of the stress distribution and magnitude of scaled models of other predators (the theropod Allosaurus fragilis and the living crocodylian Alligator mississippiensis) consistently show different responses to loadings under the same functional scenarios, suggesting distinct predatory behaviours for these animals. The unique selective pressures in the arid to semi-arid Late Cretaceous ecosystems of southeast Brazil, which were dominated by crocodyliforms, possibly drove the emergence and evolution of the biomechanical features seen in baurusuchids, which are distinct from those previously reported for other predatory taxa.View Full Text

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