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The turtle shell is a remarkable structure that has intrigued not only evolutionary biologists but also engineering and material scientists because of its multi-scale complexity and various functions. Although protection is its most apparent role, the carapace and plastron are also related to many physiological functions and their shape influences hydrodynamics and self-righting ability. As such, analysing the functional morphology of the shell could help understanding the ecology of Triassic stem-turtles, which will contribute to the century-long debate on the evolutionary origins of turtles. Here, we used 3D imaging techniques to digitize the shells of two of the earliest stem-turtle taxa, Proganochelys and Proterochersis, and submitted their models to biomechanical and shape analyses. We analysed the strength performance under five predation scenarios and tested the function of two morphological traits found in stem-turtles, the epiplastral processes and an attached pelvic girdle. The latter, also present in the crown-lineage of side-necked turtles, has been suggested to increase load-bearing capacity of the shell or to improve swimming in pleurodires. Our results do not confirm the shell-strengthening hypothesis and, together with the results of our shape analyses, suggest that at least one of the first stem-turtles (Proterochersis) was an aquatic animal.

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Body size is of fundamental importance to our understanding of extinct organisms. Physiology, ecology and life history are all strongly influenced by body size and shape, which ultimately determine how a species interacts with its environment. Reconstruction of body size and form in extinct animals provides insight into the dynamics underlying community composition and faunal turnover in past ecosystems and broad macroevolutionary trends. Many extinct animals are known only from incomplete remains, necessitating the use of anatomical proxies to reconstruct body size and form. Numerous limitations affecting the appropriateness of these proxies are often overlooked, leading to controversy and downstream inaccuracies in studies for which reconstructions represent key input data. In this perspective, we discuss four prominent case studies (Dunkleosteus, Helicoprion, Megalodon and Perucetus) in which proxy taxa have been used to estimate body size and shape from fragmentary remains. We synthesise the results of these and other studies to discuss nuances affecting the validity of taxon selection when reconstructing extinct organisms, as well as mitigation measures that can ensure the selection of the most appropriate proxy. We argue that these precautionary measures are necessary to maximise the robustness of reconstructions in extinct taxa for better evolutionary and ecological inferences.

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The Longxiang tracksite (lower Upper Cretaceous, Shanghang Basin) includes twelve didactyl deinonychosaur tracks that fall into two morphologies, differentiated by both size and form. The smaller tracks (∼11 cm long) are referable to the ichnogenus Velociraptorichnus. The larger tracks (∼36 cm long) establish the ichnotaxon Fujianipus yingliangi. Based on the size of the tracks, F. yingliangi has an estimated hip height of over 1.8 m, a size comparable to that of the largest known deinonychosaurs, i.e., Austroraptor and Utahraptor. The reduced form of digit IV, relative to digit III, indicates that F. yingliangi is a probable troodontid. Gigantism evidently evolved independently at least four times within the Deinonychosauria and within at least three major lineages: the Eudromaeosauria, Unenlagiidae, and Troodontidae. In the mid-Cretaceous of Asia, the evolution of F. yingliangi overlapped with that of early large-bodied tyrannosauroids and with previously established large allosaurids (although the latter may have been in decline).

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Vertebrate-mediated seed dispersal is a common attribute of many living plants, and variation in the size and abundance of fleshy diaspores is influenced by regional climate and by the nature of vertebrate seed dispersers among present-day floras. However, potential drivers of large-scale variation in the abundance and size distributions of fleshy diaspores through geological time, and the importance of geographic variation, are incompletely known. This knowledge gap is important because fleshy diaspores are a key mechanism of energy transfer from photosynthesis to animals and may in part explain the diversification of major groups within birds and mammals. Various hypotheses have been proposed to explain variation in the abundance and size distribution of fleshy diaspores through time, including plant-frugivore co-evolution, angiosperm diversification, and changes in vegetational structure and climate. We present a new data set of more than 800 georeferenced fossil diaspore occurrences spanning the Triassic-Oligocene, across low to mid- to high palaeolatitudes. We use this to quantify patterns of long-term change in fleshy diaspores, examining the timing and geographical context of important shifts as a test of the potential evolutionary and climatic explanations. We find that the fleshy fruit sizes of angiosperms increased for much of the Cretaceous, during the early diversification of angiosperms from herbaceous ancestors with small fruits. Nevertheless, this did not cause a substantial net change in the fleshy diaspore size distributions across seed plants, because gymnosperms had achieved a similar size distribution by at least the Late Triassic. Furthermore, gymnosperm-dominated Mesozoic ecosystems were mostly open, and harboured low proportions of specialised frugivores until the latest Cretaceous, suggesting that changes in vegetation structure and plant-frugivore co-evolution were probably not important drivers of fleshy diaspore size distributions over long timescales. Instead, fleshy diaspore size distributions may be largely constrained by physical or life-history limits that are shared among groups and diversify as a plant group expands into different growth forms/sizes, habitats, and climate regimes. Mesozoic gymnosperm floras had a low abundance of fleshy diaspores (<50% fleshy diaspore taxa), that was surpassed by some low-latitude angiosperm floras in the Cretaceous. Eocene angiosperm floras show a mid- to high latitude peak in fleshy fruit abundance, with very high proportions of fleshy fruits that even exceed those seen at low latitudes both in the Eocene and today. Mid- to high latitude proportions of fleshy fruits declined substantially over the Eocene-Oligocene transition, resulting in a shift to more modern-like geographic distributions with the highest proportion of fleshy fruits occurring in low-latitude tropical assemblages. This shift was coincident with global cooling and the onset of Southern Hemisphere glaciation, suggesting that rapid cooling at mid- and high latitudes caused a decrease in availability of the climate conditions most favourable for fleshy fruits in angiosperms. Future research could be focused on examining the environmental niches of modern fleshy fruits, and the potential effects of climate change on fleshy fruit and frugivore diversity.

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The white shark, Carcharodon carcharias, is the main top predator of the present-day Mediterranean Sea. The deep past of C. carcharias in the Mediterranean is witnessed by a rather conspicuous, mostly Pliocene fossil record. Here, we provide a synthesis of the palaeobiology and palaeoecology of the Mediterranean white sharks. Phenetically modern white shark teeth first appeared around the Miocene-Pliocene transition in the Pacific, and soon after in the Mediterranean. Molecular phylogenetic analyses support an origin of the Mediterranean white shark population from the dispersal of Australian/Pacific palaeopopulations, which may have occurred through the Central American Seaway. Tooth dimensions suggest that the Mediterranean white sharks could have grown up to about 7 m total length during the Pliocene. A richer-than-today marine mammal fauna was likely pivotal in supporting the Mediterranean white sharks through the Pliocene and most of the Quaternary. White sharks have seemingly become more common as other macropredators declined and disappeared, notwithstanding the concurrent demise of many potential prey items in the context of the latest Pliocene and Quaternary climatic and environmental perturbations of the Mediterranean region. The overall generalist trophic habits of C. carcharias were likely crucial for securing ecological success in the highly variable Mediterranean scenario by allowing the transition to a mostly piscivorous diet as the regional marine mammal fauna shrank.

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Extant amniotes show remarkable postural diversity. Broadly speaking, limbs with erect (strongly adducted, more vertically oriented) posture are found in mammals that are particularly heavy (graviportal) or show good running skills (cursorial), while crouched (highly flexed) limbs are found in taxa with more generalized locomotion. In Reptilia, crocodylians have a "semi-erect" (somewhat adducted) posture, birds have more crouched limbs and lepidosaurs have sprawling (well-abducted) limbs. Both synapsids and reptiles underwent a postural transition from sprawling to more erect limbs during the Mesozoic Era. In Reptilia, this postural change is prominent among archosauriforms in the Triassic Period. However, limb posture in many key Triassic taxa remains poorly known. In Synapsida, the chronology of this transition is less clear, and competing hypotheses exist. On land, the limb bones are subject to various stresses related to body support that partly shape their external and internal morphology. Indeed, bone trabeculae (lattice-like bony struts that form the spongy bone tissue) tend to orient themselves along lines of force. Here, we study the link between femoral posture and the femoral trabecular architecture using phylogenetic generalized least squares. We show that microanatomical parameters measured on bone cubes extracted from the femoral head of a sample of amniote femora depend strongly on body mass, but not on femoral posture or lifestyle. We reconstruct ancestral states of femoral posture and various microanatomical parameters to study the "sprawling-to-erect" transition in reptiles and synapsids, and obtain conflicting results. We tentatively infer femoral posture in several hypothetical ancestors using phylogenetic flexible discriminant analysis from maximum likelihood estimates of the microanatomical parameters. In general, the trabecular network of the femoral head is not a good indicator of femoral posture. However, ancestral state reconstruction methods hold great promise for advancing our understanding of the evolution of posture in amniotes.

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Deuterostomes are characterized by some of the most widely divergent body plans in the animal kingdom. These striking morphological differences have hindered efforts to predict ancestral characters, with the origin and earliest evolution of the group remaining ambiguous. Several iconic Cambrian fossils have been suggested to be early deuterostomes and hence could help elucidate ancestral character states. However, their phylogenetic relationships are controversial. Here, we describe new, exceptionally preserved specimens of the discoidal metazoan Rotadiscus grandis from the early Cambrian Chengjiang biota of China. These reveal a previously unknown double spiral structure, which we interpret as a chordate-like covering to a coelomopore, located adjacent to a horseshoe-shaped tentacle complex. The tentacles differ in key aspects from those seen in lophophorates and are instead more similar to the tentacular systems of extant pterobranchs and echinoderms. Thus, Rotadiscus exhibits a chimeric combination of ambulacrarian and chordate characters. Phylogenetic analyses recover Rotadiscus and closely related fossil taxa as stem ambulacrarians, filling a significant morphological gap in the deuterostome tree of life. These results allow us to reconstruct the ancestral body plans of major clades of deuterostomes, revealing that key traits of extant forms, such as a post-anal region, gill bars, and a U-shaped gut, evolved through convergence.

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The water-to-land transition by the first tetrapod vertebrates represents a key stage in their evolution. Selection pressures exerted by this new environment on animals led to the emergence of new locomotor and postural strategies that favoured access to different ecological niches and contributed to their evolutionary success. Today, amniotes show great locomotor and postural diversity, particularly among Reptilia, whose extant representatives include parasagittally locomoting erect and crouched bipeds (birds), sub-parasagittal 'semi-erect' quadrupeds (crocodylians) and sprawling quadrupeds (squamates and turtles). But the different steps leading to such diversity remain enigmatic and the type of locomotion adopted by many extinct species raises questions. This is notably the case of certain Triassic taxa such as Euparkeria and Marasuchus. The exploration of the bone microanatomy in reptiles could help to overcome these uncertainties. Indeed, this locomotor and postural diversity is accompanied by great microanatomical disparity. On land, the bones of the appendicular skeleton support the weight of the body and are subject to multiple constraints that partly shape their external and internal morphology. Here we show how microanatomical parameters measured in cross-section, such as bone compactness or the position of the medullocortical transition, can be related to locomotion. We hypothesised that this could be due to variations in cortical thickness. Using statistical methods that take phylogeny into account (phylogenetic flexible discriminant analyses), we develop different models of locomotion from a sample of femur cross-sections from 51 reptile species. We use these models to infer locomotion and posture in 7 extinct reptile taxa for which they remain debated or not fully clear. Our models produced reliable inferences for taxa that preceded and followed the quadruped/biped and sprawling/erect transitions, notably within the Captorhinidae and Dinosauria. For taxa contemporary with these transitions, such as Terrestrisuchus and Marasuchus, the inferences are more questionable. We use linear models to investigate the effect of body mass and functional ecology on our inference models. We show that body mass seems to significantly impact our model predictions in most cases, unlike the functional ecology. Finally, we illustrate how taphonomic processes can impact certain microanatomical parameters, especially the eccentricity of the section, while addressing some other potential limitations of our methods. Our study provides insight into the evolution of enigmatic locomotion in various early reptiles. Our models and methods could be used by palaeontologists to infer the locomotion and posture in other extinct reptile taxa, especially when considered in combination with other lines of evidence.

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The study of development is critical for revealing the evolution of major vertebrate lineages. Coelacanths have one of the longest evolutionary histories among osteichthyans, but despite access to extant representatives, the onset of their weakly ossified endoskeleton is still poorly understood. Here we present the first palaeohistological and skeletochronological study of Miguashaia bureaui from the Upper Devonian of Canada, pivotal for exploring the palaeobiology and early evolution of osteogenesis in coelacanths. Cross sections of the caudal fin bones show that the cortex is made of layers of primary bone separated by lines of arrested growth, indicative of a cyclical growth. The medullary cavity displays remnants of calcified cartilage associated with bony trabeculae, characteristic of endochondral ossification. A skeletochronological analysis indicates that rapid growth during a short juvenile period was followed by slower growth in adulthood. Our new analysis highlights the life history and palaeoecology of Miguashaia bureaui and reveals that, despite differences in size and habitat, the poor endoskeletal ossification known in the extant Latimeria chalumnae can be traced back at least 375 million years ago.

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This review asks some hard questions about what the enigmatic graphoglyptid trace fossils are, documents some of their early fossil record from the Ediacaran-Cambrian transition and explores the idea that they may not have been fossils at all. Most researchers have considered the Graphoglyptida to have had a microbial-farming mode of life similar to that proposed for the fractal Ediacaran Rangeomorpha. This begs the question "What are the Graphoglyptida if not the Rangeomorpha persevering" and if so then "What if…?". This provocative idea has at its roots some fundamental questions about how to distinguish burrows sensu-stricto from the external molds of endobenthic sediment displacive organisms.

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Dinosaur bonebeds with amber content, yet scarce, offer a superior wealth and quality of data on ancient terrestrial ecosystems. However, the preserved palaeodiversity and/or taphonomic characteristics of these exceptional localities had hitherto limited their palaeobiological potential. Here, we describe the amber from the Lower Cretaceous dinosaur bonebed of Ariño (Teruel, Spain) using a multidisciplinary approach. Amber is found in both a root layer with amber strictly in situ and a litter layer mainly composed of aerial pieces unusually rich in bioinclusions, encompassing 11 insect orders, arachnids, and a few plant and vertebrate remains, including a feather. Additional palaeontological data-charophytes, palynomorphs, ostracods- are provided. Ariño arguably represents the most prolific and palaeobiologically diverse locality in which fossiliferous amber and a dinosaur bonebed have been found in association, and the only one known where the vast majority of the palaeontological assemblage suffered no or low-grade pre-burial transport. This has unlocked unprecedentedly complete and reliable palaeoecological data out of two complementary windows of preservation-the bonebed and the amber-from the same site.

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BACKGROUND: The radiation of ecdysozoans (moulting animals) during the Cambrian gave rise to panarthropods and various groups of worms including scalidophorans, which played an important role in the elaboration of early marine ecosystems. Although most scalidophorans were infaunal burrowers travelling through soft sediment at the bottom of the sea, Selkirkia lived inside a tube. RESULTS: We explore the palaeobiology of these tubicolous worms, and more generally the origin and evolutionary significance of tube-dwelling in early animals, based on exceptionally preserved fossils from the early Cambrian Chengjiang Lagerstätte (Stage 3, China) including a new species, Selkirkia transita sp. nov. We find that the best phylogenetic model resolves Selkirkia as a stem-group priapulid. Selkirkia secreted a protective cuticular thickening, the tube, inside which it was able to move during at least part of its life. Partly based on measured growth patterns, we construe that this tube was separated from the trunk during a moulting process that has no direct equivalent in other scalidophorans. Although the ontogeny of Selkirkia is currently unknown, we hypothesize that its conical tube might have had the same ecological function and possibly even deep development origin as the lorica, a protective cuticular thickening found in larval priapulids and adult loriciferans. Selkirkia is seen as a semi-sedentary animal capable of very shallow incursions below the water/sediment interface, possibly for feeding or during the tube-secreting phase. Brachiopod epibionts previously reported from the Xiaoshiba Lagerstätte (ca. 514 Ma) also presumably occur in Selkirkia sinica from Chengjiang (ca. 518 Ma). CONCLUSIONS: Our critical and model-based approach provides a new phylogenetic framework for Scalidophora, upon which to improve in order to study the evolution of morphological characters in this group. Tube-dwelling is likely to have offered Selkirkia better protection and anchoring to sediment and has developed simultaneously in other Cambrian animals such as hemichordates, annelids or panarthropods. Often lost in modern representatives in favour of active infaunal lifestyles, tube-dwelling can be regarded as an early evolutionary response of various metazoans to increasing environmental and biological pressure in Cambrian marine ecosystems.

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The health of reef-building corals has declined due to climate change and pollution. However, less is known about whether giant clams, reef-dwelling bivalves with a photosymbiotic partnership similar to that found in reef-building corals, are also threatened by environmental degradation. To compare giant clam health against a prehistoric baseline, we collected fossil and modern Tridacna shells from the Gulf of Aqaba, Northern Red Sea. After calibrating daily/twice-daily growth lines from the outer shell layer, we determined that modern individuals of all three species (Tridacna maxima, T. squamosa and T. squamosina) grew faster than Holocene and Pleistocene specimens. Modern specimens also show median shell organic δ15N values 4.2‰ lower than fossil specimens, which we propose is most likely due to increased deposition of isotopically light nitrate aerosols in the modern era. Nitrate fertilization accelerates growth in cultured Tridacna, so nitrate aerosol deposition may contribute to faster growth in modern wild populations. Furthermore, colder winter temperatures and past summer monsoons may have depressed fossil giant clam growth. Giant clams can serve as sentinels of reef environmental change, both to determine their individual health and the health of the reefs they inhabit.

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Extinction events in the geological past are similar to the present-day biodiversity crisis in that they have a pronounced biogeography, producing dramatic changes in the spatial distributions of species. Reconstructing palaeobiogeographic patterns from fossils therefore allows us to examine the long-term processes governing the formation of regional biotas, and potentially helps build spatially explicit models for future biodiversity loss. However, the extent to which biogeographic patterns can be preserved in the fossil record is not well understood. Here, we perform a suite of simulations based on the present-day distribution of North American mammals, aimed at quantifying the preservation potential of beta diversity and spatial richness patterns over extinction events of varying intensities, and after applying a stepped series of taphonomic filters. We show that taphonomic biases related to body size are the biggest barrier to reconstructing biogeographic patterns over extinction events, but that these may be compensated for by both the small mammal record preserved in bird castings, as well as range expansion in surviving species. Overall, our results suggest that the preservation potential of biogeographic patterns is surprisingly high, and thus that the fossil record represents an invaluable dataset recording the changing spatial distribution of biota over key intervals in Earth History.

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BACKGROUND: The two main primate groups recorded throughout the European Miocene, hominoids and pliopithecoids, seldom co-occur. Due to both their rarity and insufficiently understood palaeoecology, it is currently unclear whether the infrequent co-occurrence of these groups is due to sampling bias or reflects different ecological preferences. Here we rely on the densely sampled primate-bearing sequence of Abocador de Can Mata (ACM) in Spain to test whether turnovers in primate assemblages are correlated with palaeoenvironmental changes. We reconstruct dietary evolution through time (ca. 12.6-11.4 Ma), and hence climate and habitat, using tooth-wear patterns and carbon and oxygen isotope compositions of enamel of the ubiquitous musk-deer Micromeryx. RESULTS: Our results reveal that primate species composition is strongly correlated with distinct environmental phases. Large-bodied hominoids (dryopithecines) are recorded in humid, densely-forested environments on the lowermost portion of the ACM sequence. In contrast, pliopithecoids inhabited less humid, patchy ecosystems, being replaced by dryopithecines and the small-bodied Pliobates toward the top of the series in gallery forests embedded in mosaic environments. CONCLUSIONS: These results support the view that pliopithecoid primates preferred less humid habitats than hominoids, and reveal that differences in behavioural ecology were the main factor underpinning their rare co-occurrence during the European Miocene. Our findings further support that ACM hominoids, like Miocene apes as a whole, inhabited more seasonal environments than extant apes. Finally, this study highlights the importance of high-resolution, local investigations to complement larger-scale analyses and illustrates that continuous and densely sampled fossiliferous sequences are essential for deciphering the complex interplay between biotic and abiotic factors that shaped past diversity.

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Heteromorphs are ammonoids forming a conch with detached whorls (open coiling) or non-planispiral coiling. Such aberrant forms appeared convergently four times within this extinct group of cephalopods. Since Wiedmann's seminal paper in this journal, the palaeobiology of heteromorphs has advanced substantially. Combining direct evidence from their fossil record, indirect insights from phylogenetic bracketing, and physical as well as virtual models, we reach an improved understanding of heteromorph ammonoid palaeobiology. Their anatomy, buoyancy, locomotion, predators, diet, palaeoecology, and extinction are discussed. Based on phylogenetic bracketing with nautiloids and coleoids, heteromorphs like other ammonoids had 10 arms, a well-developed brain, lens eyes, a buccal mass with a radula and a smaller upper as well as a larger lower jaw, and ammonia in their soft tissue. Heteromorphs likely lacked arm suckers, hooks, tentacles, a hood, and an ink sac. All Cretaceous heteromorphs share an aptychus-type lower jaw with a lamellar calcitic covering. Differences in radular tooth morphology and size in heteromorphs suggest a microphagous diet. Stomach contents of heteromorphs comprise planktic crustaceans, gastropods, and crinoids, suggesting a zooplanktic diet. Forms with a U-shaped body chamber (ancylocone) are regarded as suspension feeders, whereas orthoconic forms additionally might have consumed benthic prey. Heteromorphs could achieve near-neutral buoyancy regardless of conch shape or ontogeny. Orthoconic heteromorphs likely had a vertical orientation, whereas ancylocone heteromorphs had a near-horizontal aperture pointing upwards. Heteromorphs with a U-shaped body chamber are more stable hydrodynamically than modern Nautilus and were unable substantially to modify their orientation by active locomotion, i.e. they had no or limited access to benthic prey at adulthood. Pathologies reported for heteromorphs were likely inflicted by crustaceans, fish, marine reptiles, and other cephalopods. Pathologies on Ptychoceras corroborates an external shell and rejects the endocochleate hypothesis. Devonian, Triassic, and Jurassic heteromorphs had a preference for deep-subtidal to offshore facies but are rare in shallow-subtidal, slope, and bathyal facies. Early Cretaceous heteromorphs preferred deep-subtidal to bathyal facies. Late Cretaceous heteromorphs are common in shallow-subtidal to offshore facies. Oxygen isotope data suggest rapid growth and a demersal habitat for adult Discoscaphites and Baculites. A benthic embryonic stage, planktic hatchlings, and a habitat change after one whorl is proposed for Hoploscaphites. Carbon isotope data indicate that some Baculites lived throughout their lives at cold seeps. Adaptation to a planktic life habit potentially drove selection towards smaller hatchlings, implying high fecundity and an ecological role of the hatchlings as micro- and mesoplankton. The Chicxulub impact at the Cretaceous/Paleogene (K/Pg) boundary 66 million years ago is the likely trigger for the extinction of ammonoids. Ammonoids likely persisted after this event for 40-500 thousand years and are exclusively represented by heteromorphs. The ammonoid extinction is linked to their small hatchling sizes, planktotrophic diets, and higher metabolic rates than in nautilids, which survived the K/Pg mass extinction event.

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BACKGROUND: Ecdysozoa are the moulting protostomes, including arthropods, tardigrades, and nematodes. Both the molecular and fossil records indicate that Ecdysozoa is an ancient group originating in the terminal Proterozoic, and exceptional fossil biotas show their dominance and diversity at the beginning of the Phanerozoic. However, the nature of the ecdysozoan common ancestor has been difficult to ascertain due to the extreme morphological diversity of extant Ecdysozoa, and the lack of early diverging taxa in ancient fossil biotas. RESULTS: Here we re-describe Acosmia maotiania from the early Cambrian Chengjiang Biota of Yunnan Province, China and assign it to stem group Ecdysozoa. Acosmia features a two-part body, with an anterior proboscis bearing a terminal mouth and muscular pharynx, and a posterior annulated trunk with a through gut. Morphological phylogenetic analyses of the protostomes using parsimony, maximum likelihood and Bayesian inference, with coding informed by published experimental decay studies, each placed Acosmia as sister taxon to Cycloneuralia + Panarthropoda-i.e. stem group Ecdysozoa. Ancestral state probabilities were calculated for key ecdysozoan nodes, in order to test characters inferred from fossils to be ancestral for Ecdysozoa. Results support an ancestor of crown group ecdysozoans sharing an annulated vermiform body with a terminal mouth like Acosmia, but also possessing the pharyngeal armature and circumoral structures characteristic of Cambrian cycloneuralians and lobopodians. CONCLUSIONS: Acosmia is the first taxon placed in the ecdysozoan stem group and provides a constraint to test hypotheses on the early evolution of Ecdysozoa. Our study suggests acquisition of pharyngeal armature, and therefore a change in feeding strategy (e.g. predation), may have characterised the origin and radiation of crown group ecdysozoans from Acosmia-like ancestors.

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BACKGROUND: Saber-toothed mammals, now all extinct, were cats or "cat-like" forms with enlarged, blade-like upper canines, proposed as specialists in taking large prey. During the last 66 Ma, the saber-tooth ecomorph has evolved convergently at least in five different mammalian lineages across both marsupials and placentals. Indeed, Thylacosmilus atrox, the so-called "marsupial saber-tooth," is often considered as a classic example of convergence with placental saber-tooth cats such as Smilodon fatalis. However, despite its superficial similarity to saber-toothed placentals, T. atrox lacks many of the critical anatomical features related to their inferred predatory behavior-that of employing their enlarged canines in a killing head strike. METHODS: Here we follow a multi-proxy approach using canonical correspondence analysis of discrete traits, biomechanical models of skull function using Finite Element Analysis, and 3D dental microwear texture analysis of upper and lower postcanine teeth, to investigate the degree of evolutionary convergence between T. atrox and placental saber-tooths, including S. fatalis. RESULTS: Correspondence analysis shows that the craniodental features of T. atrox are divergent from those of placental saber-tooths. Biomechanical analyses indicate a superior ability of T. atrox to placental saber-tooths in pulling back with the canines, with the unique lateral ridge of the canines adding strength to this function. The dental microwear of T. atrox indicates a soft diet, resembling that of the meat-specializing cheetah, but its blunted gross dental wear is not indicative of shearing meat. CONCLUSIONS: Our results indicate that despite its impressive canines, the "marsupial saber-tooth" was not the ecological analogue of placental saber-tooths, and likely did not use its canines to dispatch its prey. This oft-cited example of convergence requires reconsideration, and T. atrox may have had a unique type of ecology among mammals.

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Analyses of morphological disparity have been used to characterize and investigate the evolution of variation in the anatomy, function and ecology of organisms since the 1980s. While a diversity of methods have been employed, it is unclear whether they provide equivalent insights. Here, we review the most commonly used approaches for characterizing and analysing morphological disparity, all of which have associated limitations that, if ignored, can lead to misinterpretation. We propose best practice guidelines for disparity analyses, while noting that there can be no 'one-size-fits-all' approach. The available tools should always be used in the context of a specific biological question that will determine data and method selection at every stage of the analysis.

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Reconstructing trophic interactions in ancient ecosystems is an important and fascinating branch of palaeontological research. Here we describe small bioerosional traces that are preserved on sauropod bone from the early Late Jurassic Qigu Formation (Oxfordian) of Liuhuanggou gorge in the southern Junggar Basin (Xinjiang Province, northwestern China). The most likely producers of these traces are tiny Mesozoic mammals as evinced by the small size of the traces as well as by their paired and opposed arrangement. The feeding traces are only superficially preserved on the bone surface and most likely were inflicted unintentionally during feeding. The occurrence of the bite marks along small ridges and the "gnawed" appearance of the bone surface points to selective feeding on the remaining soft tissues of the dinosaur carcass. The traces represent the oldest direct evidence for mammalian feeding behaviour in the fossil record. Additionally, these traces expand the known range of the early mammalian feeding repertoire significantly and shed light on the palaeobiology and palaeoecology of early mammals, a field that has remained evasive for a long time.

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