Estimation of the forces exerted on the limb long bones of a white rhinoceros (Ceratotherium simum) using musculoskeletal modelling and simulation.

Heavy animals incur large forces on their limb bones, due to the transmission of body weight and ground reaction forces, and the contractions of the various muscles of the limbs. This is particularly true for rhinoceroses, the heaviest extant animals capable of galloping. Several studies have examined their musculoskeletal system and the forces their bones incur, but no detailed quantification has ever been attempted. Such quantification could help understand better the link between form and function in giant land animals. Here we constructed three-dimensional musculoskeletal models of the forelimb and hindlimb of Ceratotherium simum, the heaviest extant rhino species, and used static optimisation (inverse) simulations to estimate the forces applied on the bones when standing at rest, including magnitudes and directions. Overall, unsurprisingly, the most active muscles were antigravity muscles, which generate moments opposing body weight (thereby incurring the ground reaction force), and thus keep the joints extended, avoiding joint collapse via flexion. Some muscles have an antigravity action around several joints, and thus were found to be highly active, likely specialised in body weight support (ulnaris lateralis; digital flexors). The humerus was subjected to the greatest amount of forces in terms of total magnitude; forces on the humerus furthermore came from a great variety of directions. The radius was mainly subject to high-magnitude compressive joint reaction forces, but to little muscular tension, whereas the opposite pattern was observed for the ulna. The femur had a pattern similar to that of the humerus, and the tibia's pattern was intermediate, being subject to great compression in its caudal side but to great tension in its cranial side (i.e. bending). The fibula was subject to by far the lowest force magnitude. Overall, the forces estimated were consistent with the documented morphofunctional adaptations of C. simum's long bones, which have larger insertion areas for several muscles and a greater robusticity overall than those of lighter rhinos, likely reflecting the intense forces we estimated here. Our estimates of muscle and bone (joint) loading regimes for this giant tetrapod improve the understanding of the links between form and function in supportive tissues and could be extended to other aspects of bone morphology, such as microanatomy.

The evolution of femoral morphology in giant non-avian theropod dinosaurs.

Theropods are obligate bipedal dinosaurs that appeared 230 million years ago 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 if 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 epiphyses' offset, independently 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.

Shape variation in the limb long bones of modern elephants reveals adaptations to body mass and habitat.

During evolution, several vertebrate lineages have shown trends towards an increase in mass. Such a trend is associated with physiological and musculoskeletal changes necessary to carry and move an increasingly heavy body. Due to their prominent role in the support and movement of the body, limb long bones are highly affected by these shifts in body mass. Elephants are the heaviest living terrestrial mammals, displaying unique features allowing them to withstand their massive weight, such as the columnarity of their limbs, and as such are crucial to understand the evolution towards high body mass in land mammals. In this study, we investigate the shape variation of the six limb long bones among the modern elephants, Elephas maximus and Loxodonta africana, to understand the effect of body mass and habitat on the external anatomy of the bones. To do so, we use three-dimensional geometric morphometrics (GMMs) and qualitative comparisons to describe the shape variation, at both the intraspecific and interspecific levels. Our results reveal that the two species share similar negative ontogenetic allometric patterns (i.e. becoming stouter with increased length) in their humerus and femur, but not in the other bones: the proximal epiphyses of the stylopod bones develop considerably during growth, while the distal epiphyses, which are involved in load distribution in the elbow and knee joints, are already massive in juveniles. We attribute this pattern to a weight-bearing adaptation already present in young specimens. Among adults of the same species, bone robustness increases with body mass, so that heavier specimens display stouter bones allowing for a better mechanical load distribution. While this robustness variation is significant for the humerus only, all the other bones appear to follow the same pattern. This is particularly visible in the ulna and tibia, but less so in the femur, which suggests that the forelimb and hindlimb adapted differently to high body mass support. Robustness analyses, while significant for the humerus only, suggest more robust long bones in Asian elephants than in African savanna elephants. More specifically, GMMs and qualitative comparisons indicate that three bones are clearly distinct when comparing the two species: in E. maximus the humerus, the ulna and the tibia display enlarged areas of muscular insertions for muscles involved in joint and limb stabilization, as well as in limb rotation. These results suggest a higher limb compliance in Asian elephants, associated with a higher dexterity, which could be linked to their habitat and foraging habits.

How does bone microanatomy and musculature covary? An investigation in the forelimb of two species of martens (Martes foina, Martes martes).

The long bones and associated musculature play a prominent role in the support and movement of the body and are expected to reflect the associated mechanical demands. But in addition to the functional response to adaptive changes, the conjoined effects of phylogenetic, structural and developmental constraints also shape the animal's body. In order to minimise the effect of the aforementioned constraints and to reveal the biomechanical adaptations in the musculoskeletal system to locomotor mode, we here study the forelimb of two closely related martens: the arboreal pine marten (Martes martes) and the more terrestrial stone marten (Martes foina), focusing on their forelimb muscle anatomy and long bone microanatomy; and, especially, on their covariation. To do so, we quantified muscle data and bone microanatomical parameters and created 3D and 2D maps of the cortical thickness distribution for the three long bones of the forelimb. We then analysed the covariation of muscle and bone data, both qualitatively and quantitatively. Our results reveal that species-specific muscular adaptations are not clearly reflected in the microanatomy of the bones. Yet, we observe a global thickening of the bone cortex in the radius and ulna of the more arboreal pine marten, as well a stronger flexor muscle inserting on its elbow. We attribute these differences to variation in their locomotor modes. Analyses of our 2D maps revealed a shift of cortical thickness distribution pattern linked to ontogeny, rather than species-specific patterns. We found that although intraspecific variation is not negligible, species distinction was possible when taking muscular and bone microanatomical data into consideration. Results of our covariation analyses suggest that the muscle-bone correlation is linked to ontogeny rather than to muscular strength at zones of insertion. Indeed, if we find a correlation between cortical thickness distribution and the strength of some muscles in the humerus, that is not the case for the others and in the radius and ulna. Cortical thickness distribution appears rather linked to bone contact zones and ligament insertions in the radius and ulna, and to some extent in the humerus. We conclude that inference on muscle from bone microanatomy is possible only for certain muscles in the humerus.

Sesamoid bones also show functional adaptation in their microanatomy-The example of the patella in Perissodactyla.

The patella is the largest sesamoid bone of the skeleton. It is strongly involved in the knee, improving output force and velocity of the knee extensors, and thus plays a major role in locomotion and limb stability. However, the relationships between its structure and functional constraints, that would enable a better understanding of limb bone functional adaptations, are poorly known. This contribution proposes a comparative analysis, both qualitative and quantitative, of the microanatomy of the whole patella in perissodactyls, which show a wide range of morphologies, masses, and locomotor abilities, in order to investigate how the microanatomy of the patella adapts to evolutionary constraints. The inner structure of the patella consists of a spongiosa surrounded by a compact cortex. Contrary to our expectations, there is no increase in compactness with bone size, and thus body size and weight, but only an increase in the tightness of the spongiosa. No particular thickening of the cortex associated with muscle insertions is noticed but a strong thickening is observed anteriorly at about mid-length, where the strong intermediate patellar ligament inserts. The trabeculae are mainly oriented perpendicularly to the posterior articular surface, which highlights that the main stress is anteroposteriorly directed, maintaining the patella against the femoral trochlea. Conversely, anteriorly, trabeculae are rather circumferentially oriented, following the insertion of the patellar ligament and, possibly also, of the quadriceps tendon. A strong variation is observed among perissodactyl families but also intraspecifically, which is in accordance with previous studies suggesting a higher variability in sesamoid bones. Clear trends are nevertheless observed between the three families. Equids have a much thinner cortex than ceratomorphs. Rhinos and equids, both characterized by a development of the medial border, show an increase in trabecular density laterally suggesting stronger stresses laterally. The inner structure in tapirs is more homogeneous despite the absence of medial development of the medial border with no "compensation" of the inner structure, which suggests different stresses on their knees associated with a different morphology of their patellofemoral joint.

A path to gigantism: Three-dimensional study of the sauropodomorph limb long bone shape variation in the context of the emergence of the sauropod bauplan.

Sauropodomorph dinosaurs include the largest terrestrial animals that ever lived on Earth. The early representatives of this clade were, however, relatively small and partially to totally bipedal, conversely to the gigantic and quadrupedal sauropods. Although the sauropod bauplan is well defined, notably by the acquisition of columnar limbs, the evolutionary sequence leading to its emergence remains debated. Here, we aim to tackle this evolutionary episode by investigating shape variation in the six limb long bones for the first time using three-dimensional geometric morphometrics. The morphological features of the forelimb zeugopod bones related to the sauropod bauplan tend to appear abruptly, whereas the pattern is more gradual for the hindlimb zeugopod bones. The stylopod bones tend to show the same pattern as their respective zeugopods. The abrupt emergence of the sauropod forelimb questions the locomotor abilities of non-sauropodan sauropodomorphs inferred as quadrupeds. Features characterizing sauropods tend to corroborate a view of their locomotion mainly based on stylopod retraction. An allometric investigation of the shape variation in accordance with size highlight differences in hindlimb bone allometries between the sauropods and the non-sauropodan sauropodomorphs. These differences notably correspond to an unexpected robustness decrease trend in the sauropod hindlimb zeugopod. In addition to forelimb bones that appear to be proportionally more gracile than in non-sauropodan sauropodomorphs, sauropods may have relied on limb architecture and features related to the size increase, rather than general robustness, to deal with the role of weight-bearing.

Femoral specializations to locomotor habits in early archosauriforms.

The evolutionary history of archosaurs and their closest relatives is characterized by a wide diversity of locomotor modes, which has even been suggested as a pivotal aspect underlying the evolutionary success of dinosaurs vs. pseudosuchians across the Triassic-Jurassic transition. This locomotor diversity (e.g., more sprawling/erect; crouched/upright; quadrupedal/bipedal) led to several morphofunctional specializations of archosauriform limb bones that have been studied qualitatively as well as quantitatively through various linear morphometric studies. However, differences in locomotor habits have never been studied across the Triassic-Jurassic transition using 3D geometric morphometrics, which can relate how morphological features vary according to biological factors such as locomotor habit and body mass. Herein, we investigate morphological variation across a dataset of 72 femora from 36 different species of archosauriforms. First, we identify femoral head rotation, distal slope of the fourth trochanter, femoral curvature, and the angle between the lateral condyle and crista tibiofibularis as the main features varying between bipedal and quadrupedal taxa, all of these traits having a stronger locomotor signal than the lesser trochanter's proximal extent. We show a significant association between locomotor mode and phylogeny, but with the locomotor signal being stronger than the phylogenetic signal. This enables us to predict locomotor modes of some of the more ambiguous early archosauriforms without relying on the relationships between hindlimb and forelimb linear bone dimensions as in prior studies. Second, we highlight that the most important morphological variation is linked to the increase of body size, which impacts the width of the epiphyses and the roundness and proximodistal position of the fourth trochanter. Furthermore, we show that bipedal and quadrupedal archosauriforms have different allometric trajectories along the morphological variation in relation to body size. Finally, we demonstrate a covariation between locomotor mode and body size, with variations in femoral bowing (anteroposterior curvature) being more distinct among robust femora than gracile ones. We also identify a decoupling in fourth trochanter variation between locomotor mode (symmetrical to semi-pendant) and body size (sharp to rounded). Our results indicate a similar level of morphological disparity linked to a clear convergence in femoral robusticity between the two clades of archosauriforms (Pseudosuchia and Avemetatarsalia), emphasizing the importance of accounting for body size when studying their evolutionary history, as well as when studying the functional morphology of appendicular features. Determining how early archosauriform skeletal features were impacted by locomotor habits and body size also enables us to discuss the potential homoplasy of some phylogenetic characters used previously in cladistic analyses as well as when bipedalism evolved in the avemetatarsalian lineage. This study illuminates how the evolution of femoral morphology in early archosauriforms was functionally constrained by locomotor habit and body size, which should aid ongoing discussions about the early evolution of dinosaurs and the nature of their evolutionary "success" over pseudosuchians.

Effect of mass and habitat on the shape of limb long bones: A morpho-functional investigation on Bovidae (Mammalia: Cetartiodactyla).

Limb long bones are essential to an animal's locomotion, and are thus expected to be heavily influenced by factors such as mass or habitat. Because they are often the only organs preserved in the fossil record, understanding their adaptive trends is key to reconstructing the paleobiology of fossil taxa. In this regard, the Bovidae has always been a prized group of study. This family is extremely diverse in terms of both mass and habitat, and it is expected that their bones will possess adaptations to both factors. Here, we present the first 3D geometric morphometric study focusing on bovid limb long bones. We used anatomical landmarks as well as curve and surface sliding semi-landmarks to accurately describe the stylopod and zeugopod bones. We included 50 species from ten of the twelve currently recognized tribes of bovids, ranging from 4.6 to 725 kg, and living in open plains, forests, mountains, or anywhere in-between. Shape data were correlated with the mean mass of the species and its habitat, even when taking into account the phylogenetic history of our sample. Bones pertaining to heavy species are more robust, adapted for a better repartition of stronger forces. Articulations are especially affected, being proportionally much larger in heavier species. Muscle insertion areas are unevenly affected. Insertion areas of muscles implied in body support and propulsion show a strong increase in their robustness when compared to insertion areas of muscles acting on the limb mostly when it is off the ground. Habitat influences the shape of the humerus, the radius-ulna, and the femur, but not of the tibia, whether the phylogeny is taken into account or not. Specific habitats tend to be associated with particular features on the bones. Articulations are proportionally wider in open-habitat species, and the insertion areas of muscles involved in limb extension and propulsion are wider, reflecting the fact that open habitat species are more cursorial and rely on fast running to avoid predators. Forest and mountain species generally present similar adaptations for increased manoeuvrability, such as a round femoral head, and generally have more gracile bones.

What about limb long bone nutrient canal(s)? - a 3D investigation in mammals.

The nutrient arteries, located in the long bone diaphysis, are the major blood supply to long bones, especially during the early phases of growth and ossification. Their intersection with the central axis of the medullary area corresponds to the ossification center, and their opening on the outer bone surface to the nutrient foramen. Nutrient arteries/foramen have essentially been analyzed in humans, and only to a much lesser extent in a few mammals. Some studies have taken measurements of the nutrient foramen; others have investigated the shape and orientation of the nutrient canals, although only partially. No studies have analyzed the nutrient canal in three dimensions inside the bone and the relationships between nutrient foramen, nutrient canal, growth, and physiology require further investigation. The current study proposes to investigate in three dimensions the shape of the nutrient canal in stylopod bones of various mammals. Qualitative and quantitative parameters are defined to discuss the diversity in, for example, morphology, orientation, and diameter encountered, resorting to two different datasets to maximize differences within mammals and then analyze variation within morphologically and phylogenetically closer taxa. This study highlights a strong intraspecific variation for various parameters, with limited biological signal, but also shows trends. It notably provides evidence that canals are generally more numerous and relatively thinner in less elongated bones. Moreover, it shows that the growth center is located distally in the humerus and proximally in the femur, and that the canals are essentially oriented towards the faster growing end, so that the nutrient foramen does not indicate the location of the growth center. This result seems general in mammals but cannot be generalized outside of Mammalia. Further analyses of the features of nutrient arteries in reptiles are required to make comparisons with the trends observed in mammals.

A first glimpse at the influence of body mass in the morphological integration of the limb long bones: an investigation in modern rhinoceroses.

The appendicular skeleton of tetrapods is a particularly integrated structure due to the shared developmental origin or similar functional constraints exerted on its elements. Among these constraints, body mass is considered strongly to influence its integration but its effect on shape covariation has rarely been addressed in mammals, especially in heavy taxa. Here, we propose to explore the covariation patterns of the long bones in heavy animals and their link to body mass. We investigate the five modern rhinoceros species, which display an important range of bodyweight. We used a 3D geometric morphometric approach to describe the shape covariation of the six bones composing the stylopodium and zeugopodium both among and within species. Our results indicate that the appendicular skeleton of modern rhinos is a strongly integrated structure. At the interspecific level, the shape covariation is roughly similar between all pairs of bones and mainly concerns the muscular insertions related to powerful flexion and extension movements. The forelimb integration appears higher and more related to body mass than that of the hind limb, suggesting a specialization for weight support. The integration of the stylopodium elements does not seem to relate to body mass in our sample, which suggests a greater effect of shared developmental factors. Conversely, the covariation of the zeugopodium bones seems more associated with body mass, particularly for the radius-ulna pair. The fibula appears poorly integrated with other bones, especially within non-Rhinoceros species, which may represent a case of parcellation due to a functional dissociation between the hind limb bones. The exploration of the integration patterns at the intraspecific level also highlights a more prominent effect of age over individual body mass on shape covariation within C. simum. This study lends support to previous hypotheses indicating a link between high body mass and high integration level.

New hominoid fossils from Moroto II, Uganda and their bearing on the taxonomic and adaptive status of Morotopithecus bishopi.

The early Miocene site of Moroto II, Uganda has yielded some of the oldest known hominoid fossils. A new partial mandible (UMP MORII 03'551) is notable for its long tooth row and large, narrow M2 with well-developed cristids - a morphological combination previously unknown for large bodied catarrhines of the Early Miocene and suggesting folivory. The tooth proportions are compatible with belonging to the same taxon as the maxilla UMP 62-11, the holotype of Morotopithecus bishopi; likewise, the long tooth row and vertical planum of UMP MORII 03'551 suggest that it may represent the same taxon as mandible(s) UMP 66-01 and UMP 62-10. Canine size strongly suggests UMP MORII 03'551 is a female. Comparisons of the tooth crown morphology and tooth row proportions, relative enamel thickness, enamel-dentine junction morphology, long-period line periodicity, and dental wear patterns support significant morphological, developmental, and inferred dietary differentiation, and therefore generic-level distinctiveness, among Afropithecus, Morotopithecus and the Proconsul clade. An isolated M1 (UMP MORII 03'559) is morphologically dissimilar, and much smaller than the actual or inferred size of molars in UMP MORII 03'551, UMP 66-01 and UMP 62-10, supporting the presence of two hominoid taxa at Moroto II, M. bishopi and a smaller bodied proconsulid. Given the high level of body mass dimorphism inferred for Morotopithecus and other early Miocene catarrhines, the known postcrania from Moroto II could be attributable to either taxon. However, UMP MORII 03'551 and the femora UMP MORII 94'80 derive from the same stratigraphic interval, while the isolated M1 was deposited later, increasing the likelihood that the mandible and femora are from the same individual. These new fossils expand our understanding of the taxonomic and adaptive diversity of early Miocene catarrhines.

3D quantitative comparative analysis of long bone diaphysis variations in microanatomy and cross-sectional geometry.

Long bone inner structure and cross-sectional geometry display a strong functional signal, leading to convergences, and are widely analyzed in comparative anatomy at small and large taxonomic scales. Long bone microanatomical studies have essentially been conducted on transverse sections but also on a few longitudinal ones. Recent studies highlighted the interest in analyzing variations of the inner structure along the diaphysis using a qualitative as well as a quantitative approach. With the development of microtomography, it has become possible to study three-dimensional (3D) bone microanatomy and, in more detail, the form-function relationships of these features. This study focused on the selection of quantitative parameters to describe in detail the cross-sectional shape changes and distribution of the osseous tissue along the diaphysis. Two-dimensional (2D) virtual transverse sections were also performed in the two usual reference planes and results were compared with those obtained based on the whole diaphysis analysis. The sample consisted in 14 humeri and 14 femora of various mammalian taxa that are essentially terrestrial. Comparative quantitative analyses between different datasets made it possible to highlight the parameters that are strongly impacted by size and phylogeny and the redundant ones, and thus to estimate their relevance for use in form-function analyses. The analysis illustrated that results based on 2D transverse sections are similar for both sectional planes; thus if a strong bias exists when mixing sections from the two reference planes in the same analysis, it would not problematic to use either one plane or the other in comparative studies. However, this may no longer hold for taxa showing a much stronger variation in bone microstructure along the diaphysis. Finally, the analysis demonstrated the significant contribution of the parameters describing variations along the diaphysis, and thus the interest in performing 3D analyses; this should be even more fruitful for heterogeneous diaphyses. In addition, covariation analyses showed that there is a strong interest in removing the size effect to access the differences in the microstructure of the humerus and femur. This methodological study provides a reference for future quantitative analyses on long bone inner structure and should make it possible, through a detailed knowledge of each descriptive parameter, to better interpret results from the multivariate analyses associated with these studies. This will have direct implications for studies in vertebrate anatomy, but also in paleontology and anthropology.

Comparative morphology of snake (Squamata) endocasts: evidence of phylogenetic and ecological signals.

Brain endocasts obtained from computed tomography (CT) are now widely used in the field of comparative neuroanatomy. They provide an overview of the morphology of the brain and associated tissues located in the cranial cavity. Through anatomical comparisons between species, insights on the senses, the behavior, and the lifestyle can be gained. Although there are many studies dealing with mammal and bird endocasts, those performed on the brain endocasts of squamates are comparatively rare, thus limiting our understanding of their morphological variability and interpretations. Here, we provide the first comparative study of snake brain endocasts in order to bring new information about the morphology of these structures. Additionally, we test if the snake brain endocast encompasses a phylogenetic and/or an ecological signal. For this purpose, the digital endocasts of 45 snake specimens, including a wide diversity in terms of phylogeny and ecology, were digitized using CT, and compared both qualitatively and quantitatively. Snake endocasts exhibit a great variability. The different methods performed from descriptive characters, linear measurements and the outline curves provided complementary information. All these methods have shown that the shape of the snake brain endocast contains, as in mammals and birds, a phylogenetic signal but also an ecological one. Although phylogenetically related taxa share several similarities between each other, the brain endocast morphology reflects some notable ecological trends: e.g. (i) fossorial species possess both reduced optic tectum and pituitary gland; (ii) both fossorial and marine species have cerebral hemispheres poorly developed laterally; (iii) cerebral hemispheres and optic tectum are more developed in arboreal and terrestrial species.

How does the shape of the wing and hindlimb bones of aquatic birds relate to their locomotor abilities?

Aquatic birds represent diverse ecologies and locomotion types. Some became flightless or lost the ability for effective terrestrial locomotion, yet, certain species excel in water, on land, and in air, despite differing physical characteristics associated with each medium. In this exploratory study, we intend to quantitatively analyze the morphological variety of multiple limb bones of aquatic birds using 3D geometric morphometrics. Morphological variation is mainly driven by phylogeny, which also affects size and locomotion. However, the shape of the ulna, including the proportion and orientation of the epiphyses is influenced by size and aquatic propulsive techniques even when phylogeny is taken into consideration. Certain trends, possibly linked to functions, can be observed too in other bones, notably in cases where phylogenetic and functional signals are probably mixed when some taxa only englobe species with similar functional requirements: penguins exhibit the most distinctive wing bone morphologies, highly adapted to wing-propulsion; advanced foot-propellers exhibit femur morphology that reduces proximal mobility but supports stability; knee structures, like cnemial crests of varied sizes and orientations, are crucial for muscle attachments and efficient movement in water and on land; taxa relying on their feet in water but retaining terrestrial abilities share features enabling swimming and walking postures. Size-linked changes distinguish the wing bones of non-wing-propelled taxa. For hindlimbs, larger size relates to robust bones probably linked to terrestrial abilities, but robustness in femora can be connected to foot-propulsion. These results help us better understand birds' skeletal adaptation and can be useful inferring extinct species' ecology.

Jack-of-all-trades master of all? Snake vertebrae have a generalist inner organization.

Snakes are a very speciose group of squamates that adapted to various habitats and ecological niches. Their ecological diversity is of particular interest and functional demands associated with their various styles of locomotion are expected to result in anatomical specializations. In order to explore the potential adaptation of snakes to their environment we here analyze variation in vertebral structure at the microanatomical level in species with different locomotor adaptations. Vertebrae, being a major element of the snake body, are expected to display adaptations to the physical constraints associated with the different locomotor modes and environments. Our results revealed a rather homogenous vertebral microanatomy in contrast to what has been observed for other squamates and amniotes more generally. We here suggest that the near-absence of microanatomical specializations in snake vertebrae might be correlated to their rather homogeneous overall morphology and reduced range of morphological diversity, as compared to lizards. Thus, snakes appear to retain a generalist inner morphology that allows them to move efficiently in different environments. Only a few ecologically highly specialized taxa appear to display some microanatomical specializations that remain to be studied in greater detail.

Femora from an exceptionally large population of coeval ornithomimosaurs yield evidence of sexual dimorphism in extinct theropod dinosaurs.

Sexual dimorphism is challenging to detect among fossils due to a lack of statistical representativeness. The Angeac-Charente Lagerstätte (France) represents a remarkable 'snapshot' from a Berriasian (Early Cretaceous) ecosystem and offers a unique opportunity to study intraspecific variation among a herd of at least 61 coeval ornithomimosaurs. Herein, we investigated the hindlimb variation across the best-preserved specimens from the herd through 3D Geometric Morphometrics and Gaussian Mixture Modeling. Our results based on complete and fragmented femora evidenced a dimorphism characterized by variations in the shaft curvature and the distal epiphysis width. Since the same features vary between sexes among modern avian dinosaurs, crocodilians, and more distant amniotes, we attributed this bimodal variation to sexual dimorphism based on the extant phylogenetic bracketing approach. Documenting sexual dimorphism in fossil dinosaurs allows a better characterization and accounting of intraspecific variations, which is particularly relevant to address ongoing taxonomical and ecological questions relative to dinosaur evolution.

Evolution: Back to heavy bones in salty seas.

In amniote vertebrates that transitioned from land to sea, bone mass typically increases and later decreases as active swimming evolves. A new study now has found that heavy bones re-evolved in some fossil marine mammals, suggesting that this trajectory can be reversed.

Early Triassic ichthyopterygian fossils from the Russian Far East.

Ichthyopterygia is a major clade of reptiles that colonized the ocean after the end-Permian mass extinction, with the oldest fossil records found in early Spathian substage (late Olenekian, late Early Triassic) strata in the western USA. Here, we describe reptilian remains found in situ in the early Spathian Neocolumbites insignis ammonoid zone of South Primorye in the Russian Far East. Specimen NSM PV 23854 comprises fragmentary axial elements exhibiting a combination of morphological characteristics typical of Ichthyopterygia. The cylindrical centra suggest that the specimen represents a basal ichthyopterygian, and its size is comparable to that of Utatsusaurus. Specimen NSM PV 24995 is represented by a single limb bone, which is tentatively identified as an ichthyopterygian humerus. With a body length of approximately 5 m estimated from the humeral length, NSM PV 24995 represents one of the largest specimens of early Spathian marine reptiles known to date. Such size variation among the earliest ichthyopterygians might suggest an explosive diversification in size immediately after the end-Permian mass extinction. Both vertebrae and humerus specimens exhibit an extremely cancellous inner structure, suggesting a high degree of aquatic adaptation in ichthyopterygians, despite their short history of evolution in the ocean.

Limb myology and muscle architecture of the Indian rhinoceros Rhinoceros unicornis and the white rhinoceros Ceratotherium simum (Mammalia: Rhinocerotidae).

Land mammals support and move their body using their musculoskeletal system. Their musculature usually presents varying adaptations with body mass or mode of locomotion. Rhinocerotidae is an interesting clade in this regard, as they are heavy animals potentially reaching three tons but are still capable of adopting a galloping gait. However, their musculature has been poorly studied. Here we report the dissection of both forelimb and hindlimb of one neonate and one adult each for two species of rhinoceroses, the Indian rhinoceros (Rhinoceros unicornis) and the white rhinoceros (Ceratotherium simum). We show that their muscular organisation is similar to that of their relatives, equids and tapirs, and that few evolutionary convergences with other heavy mammals (e.g. elephants and hippopotamuses) are present. Nevertheless, they show clear adaptations to their large body mass, such as more distal insertions for the protractor and adductor muscles of the limbs, giving them longer lever arms. The quantitative architecture of rhino muscles is again reminiscent of that of horses and tapirs, although contrary to horses, the forelimb is much stronger than the hindlimb, which is likely due to its great role in body mass support. Muscles involved mainly in counteracting gravity (e.g. serratus ventralis thoracis, infraspinatus, gastrocnemius, flexores digitorum) are usually highly pennate with short fascicles facilitating strong joint extension. Muscles involved in propulsion (e.g. gluteal muscles, gluteobiceps, quadriceps femoris) seem to represent a compromise between a high maximal isometric force and long fascicles, allowing a reasonably fast and wide working range. Neonates present higher normalized maximal isometric force than the adults for almost every muscle, except sometimes for the extensor and propulsor muscles, which presumably acquire their great force-generating capacity during the growth of the animal. Our study clarifies the way the muscles of animals of cursorial ancestry can adapt to support a greater body mass and calls for further investigations in other clades of large body mass.

Vertebral microanatomy in squamates: structure, growth and ecological correlates.

The histological study of vertebrae in extant squamates shows that the internal vertebral structure in this group differs from that of other tetrapods. Squamate vertebrae are lightly built and basically composed of two roughly concentric osseous tubes--one surrounding the neural canal and the other constituting the peripheral cortex of the vertebra--connected by few thin trabeculae. This structure, which characteristically evokes that of a tubular bone, results from a peculiar remodelling process characterised by an imbalance between local bone resorption and redeposition; in both periosteal and endosteo-endochondral territories, bone is extensively resorbed but not reconstructed in the same proportion by secondary deposits. This process is particularly intense in the deep region of the centrum, where originally compact cortices are made cancellous, and where the endochondral spongiosa is very loose. This remodelling process starts at an early stage of development and remains active throughout subsequent growth. The growth of squamate centra is also strongly asymmetrical, with the posterior (condylar) part growing much faster than the anterior (cotylar) part. Preliminary analyses testing for associations between vertebral structure and habitat use suggest that vertebrae of fossorial taxa are denser than those of terrestrial taxa, those in aquatic taxa being of intermediate density. However, phylogenetically informed analyses do not corroborate these findings, thus suggesting a strong phylogenetic signal in the data. As our analyses demonstrate that vertebrae in snakes are generally denser than those of lizards sensu stricto, this may drive the presence of a phylogenetic signal in the data. More comprehensive sampling of fossorial and aquatic lizards is clearly needed to more rigorously evaluate these patterns.