Unexpectedly uneven distribution of functional trade-offs explains cranial morphological diversity in carnivores.

Functional trade-offs can affect patterns of morphological and ecological evolution as well as the magnitude of morphological changes through evolutionary time. Using morpho-functional landscape modelling on the cranium of 132 carnivore species, we focused on the macroevolutionary effects of the trade-off between bite force and bite velocity. Here, we show that rates of evolution in form (morphology) are decoupled from rates of evolution in function. Further, we found theoretical morphologies optimising for velocity to be more diverse, while a much smaller phenotypic space was occupied by shapes optimising force. This pattern of differential representation of different functions in theoretical morphological space was highly correlated with patterns of actual morphological disparity. We hypothesise that many-to-one mapping of cranium shape on function may prevent the detection of direct relationships between form and function. As comparatively only few morphologies optimise bite force, species optimising this function may be less abundant because they are less likely to evolve. This, in turn, may explain why certain clades are less variable than others. Given the ubiquity of functional trade-offs in biological systems, these patterns may be general and may help to explain the unevenness of morphological and functional diversity across the tree of life.

Raptorial appendages of the Cambrian apex predator Anomalocaris canadensis are built for soft prey and speed.

The stem-group euarthropod Anomalocaris canadensis is one of the largest Cambrian animals and is often considered the quintessential apex predator of its time. This radiodont is commonly interpreted as a demersal hunter, responsible for inflicting injuries seen in benthic trilobites. However, controversy surrounds the ability of A. canadensis to use its spinose frontal appendages to masticate or even manipulate biomineralized prey. Here, we apply a new integrative computational approach, combining three-dimensional digital modelling, kinematics, finite-element analysis (FEA) and computational fluid dynamics (CFD) to rigorously analyse an A. canadensis feeding appendage and test its morphofunctional limits. These models corroborate a raptorial function, but expose inconsistencies with a capacity for durophagy. In particular, FEA results show that certain parts of the appendage would have experienced high degrees of plastic deformation, especially at the endites, the points of impact with prey. The CFD results demonstrate that outstretched appendages produced low drag and hence represented the optimal orientation for speed, permitting acceleration bursts to capture prey. These data, when combined with evidence regarding the functional morphology of its oral cone, eyes, body flaps and tail fan, suggest that A. canadensis was an agile nektonic predator that fed on soft-bodied animals swimming in a well-lit water column above the benthos. The lifestyle of A. canadensis and that of other radiodonts, including plausible durophages, suggests that niche partitioning across this clade influenced the dynamics of Cambrian food webs, impacting on a diverse array of organisms at different sizes, tiers and trophic levels.

Homo sapiens and Neanderthals share high cerebral cortex integration into adulthood.

There is controversy around the mechanisms that guided the change in brain shape during the evolution of modern humans. It has long been held that different cortical areas evolved independently from each other to develop their unique functional specializations. However, some recent studies suggest that high integration between different cortical areas could facilitate the emergence of equally extreme, highly specialized brain functions. Here, we analyse the evolution of brain shape in primates using three-dimensional geometric morphometrics of endocasts. We aim to determine, firstly, whether modern humans present unique developmental patterns of covariation between brain cortical areas; and secondly, whether hominins experienced unusually high rates of evolution in brain covariation as compared to other primates. On the basis of analyses including modern humans and other extant great apes at different developmental stages, we first demonstrate that, unlike our closest living relatives, Homo sapiens retain high levels of covariation between cortical areas into adulthood. Among the other great apes, high levels of covariation are only found in immature individuals. Secondly, at the macro-evolutionary level, our analysis of 400 endocasts, representing 148 extant primate species and 6 fossil hominins, shows that strong covariation between different areas of the brain in H. sapiens and Homo neanderthalensis evolved under distinctly higher evolutionary rates than in any other primate, suggesting that natural selection favoured a greatly integrated brain in both species. These results hold when extinct species are excluded and allometric effects are accounted for. Our findings demonstrate that high covariation in the brain may have played a critical role in the evolution of unique cognitive capacities and complex behaviours in both modern humans and Neanderthals.

Biomechanical analyses of pterygotid sea scorpion chelicerae uncover predatory specialisation within eurypterids.

Eurypterids (sea scorpions) are extinct aquatic chelicerates. Within this group, members of Pterygotidae represent some of the largest known marine arthropods. Representatives of this family all have hypertrophied, anteriorly-directed chelicerae and are commonly considered Silurian and Devonian apex predators. Despite a long history of research interest in these appendages, pterygotids have been subject to limited biomechanical investigation. Here, we present finite element analysis (FEA) models of four different pterygotid chelicerae-those of Acutiramus bohemicus, Erettopterus bilobus, Jaekelopterus rhenaniae, and Pterygotus anglicus-informed through muscle data and finite element models (FEMs) of chelae from 16 extant scorpion taxa. We find that Er. bilobus and Pt. anglicus have comparable stress patterns to modern scorpions, suggesting a generalised diet that probably included other eurypterids and, in the Devonian species, armoured fishes, as indicated by co-occurring fauna. Acutiramus bohemicus is markedly different, with the stress being concentrated in the proximal free ramus and the serrated denticles. This indicates a morphology better suited for targeting softer prey. Jaekelopterus rhenaniae exhibits much lower stress across the entire model. This, combined with an extremely large body size, suggests that the species likely fed on larger and harder prey, including heavily armoured fishes. The range of cheliceral morphologies and stress patterns within Pterygotidae demonstrate that members of this family had variable diets, with only the most derived species likely to feed on armoured prey, such as placoderms. Indeed, increased sizes of these forms throughout the mid-Palaeozoic may represent an 'arms race' between eurypterids and armoured fishes, with Devonian pterygotids adapting to the rapid diversification of placoderms.

Earliest evidence for fruit consumption and potential seed dispersal by birds.

The Early Cretaceous diversification of birds was a major event in the history of terrestrial ecosystems, occurring during the earliest phase of the Cretaceous Terrestrial Revolution, long before the origin of the bird crown-group. Frugivorous birds play an important role in seed dispersal today. However, evidence of fruit consumption in early birds from outside the crown-group has been lacking. Jeholornis is one of the earliest-diverging birds, only slightly more crownward than Archaeopteryx, but its cranial anatomy has been poorly understood, limiting trophic information which may be gleaned from the skull. Originally hypothesised to be granivorous based on seeds preserved as gut contents, this interpretation has become controversial. We conducted high-resolution synchrotron tomography on an exquisitely preserved new skull of Jeholornis, revealing remarkable cranial plesiomorphies combined with a specialised rostrum. We use this to provide a near-complete cranial reconstruction of Jeholornis, and exclude the possibility that Jeholornis was granivorous, based on morphometric analyses of the mandible (3D) and cranium (2D), and comparisons with the 3D alimentary contents of extant birds. We show that Jeholornis provides the earliest evidence for fruit consumption in birds, and indicates that birds may have been recruited for seed dispersal during the earliest stages of the avian radiation. As mobile seed dispersers, early frugivorous birds could have expanded the scope for biotic dispersal in plants, and might therefore explain, at least in part, the subsequent evolutionary expansion of fruits, indicating a potential role of bird-plant interactions in the Cretaceous Terrestrial Revolution.

Birds and plants have a close relationship that has developed over millions of years. Birds became diverse and abundant around 135 million years ago. Shortly after, plants started developing new and different kinds of fruits. Today, fruit-eating birds help plants to reproduce by spreading seeds in their droppings. This suggests that birds and plants have coevolved, changing together over time. But it is not clear exactly how their relationship started. One species that might hold the answers is an early bird species known as Jeholornis. It lived in China in the Early Cretaceous, around 120 million years ago. Palaeontologists have discovered preserved seeds inside its fossilised remains. The question is, how did they get there? Some birds eat seeds directly, cracking them open or grinding them up in the stomach to extract the nutrients inside. Other birds swallow seeds when they are eating fruit. If Jeholornis belonged to this second group, it could represent one of the early steps in plant-bird coevolution. Hu et al. scanned and reconstructed a preserved Jeholornis skull and compared it to the skulls, especially the mandibles, of modern birds, including species that grind seeds, species that crack seeds and species that eat fruits, leaving the seeds whole. The analyses ruled out seed cracking. But it could not distinguish between seed grinding and fruit eating. Hu et al. therefore compared the seed remains found inside Jeholornis fossils to seeds eaten by modern birds. The fossilised seeds were intact and showed no evidence of grinding. This suggests that Jeholornis ate whole fruits for at least part of the year. At around the time Jeholornis was alive, the world was entering a phase called the Cretaceous Terrestrial Revolution, which was characterized by an explosion of new species and an expansion of both flowering plants and birds. This finding opens new avenues for scientists to explore how plant and birds might have evolved together. Similar analyses could unlock new information about how other species interacted with their environments.

The extinct shark Otodus megalodon was a transoceanic superpredator: Inferences from 3D modeling.

Although shark teeth are abundant in the fossil record, their bodies are rarely preserved. Thus, our understanding of the anatomy of the extinct Otodus megalodon remains rudimentary. We used an exceptionally well-preserved fossil to create the first three-dimensional model of the body of this giant shark and used it to infer its movement and feeding ecology. We estimate that an adult O. megalodon could cruise at faster absolute speeds than any shark species today and fully consume prey the size of modern apex predators. A dietary preference for large prey potentially enabled O. megalodon to minimize competition and provided a constant source of energy to fuel prolonged migrations without further feeding. Together, our results suggest that O. megalodon played an important ecological role as a transoceanic superpredator. Hence, its extinction likely had large impacts on global nutrient transfer and trophic food webs.

More Challenging Diets Sustain Feeding Performance: Applications Toward the Captive Rearing of Wildlife.

The rescue and rehabilitation of young fauna is of substantial importance to conservation. However, it has been suggested that incongruous diets offered in captive environments may alter craniofacial morphology and hinder the success of reintroduced animals. Despite these claims, to what extent dietary variation throughout ontogeny impacts intrapopulation cranial biomechanics has not yet been tested. Here, finite element models were generated from the adult crania of 40 rats (n = 10 per group) that were reared on 4 different diet regimes and stress magnitudes compared during incisor bite simulations. The diets consisted of (1) exclusively hard pellets from weaning, (2) exclusively soft ground pellet meal from weaning, (3) a juvenile switch from pellets to meal, and (4) a juvenile switch from meal to pellets. We hypothesized that a diet of exclusively soft meal would result in the weakest adult skulls, represented by significantly greater stress magnitudes at the muzzle, palate, and zygomatic arch. Our hypothesis was supported at the muzzle and palate, indicating that a diet limited to soft food inhibits bone deposition throughout ontogeny. This finding presents a strong case for a more variable and challenging diet during development. However, rather than the "soft" diet group resulting in the weakest zygomatic arch as predicted, this region instead showed the highest stress among rats that switched as juveniles from hard pellets to soft meal. We attribute this to a potential reduction in number and activity of osteoblasts, as demonstrated in studies of sudden and prolonged disuse of bone. A shift to softer foods in captivity, during rehabilitation after injury in the wild for example, can therefore be detrimental to healthy development of the skull in some growing animals, potentially increasing the risk of injury and impacting the ability to access full ranges of wild foods upon release. We suggest captive diet plans consider not just nutritional requirements but also food mechanical properties when rearing wildlife to adulthood for reintroduction.

Biomechanical analyses of Cambrian euarthropod limbs reveal their effectiveness in mastication and durophagy.

Durophagy arose in the Cambrian and greatly influenced the diversification of biomineralized defensive structures throughout the Phanerozoic. Spinose gnathobases on protopodites of Cambrian euarthropod limbs are considered key innovations for shell-crushing, yet few studies have demonstrated their effectiveness with biomechanical models. Here we present finite-element analysis models of two Cambrian trilobites with prominent gnathobases-Redlichia rex and Olenoides serratus-and compare these to the protopodites of the Cambrian euarthropod Sidneyia inexpectans and the modern American horseshoe crab, Limulus polyphemus. Results show that L. polyphemus, S. inexpectans and R. rex have broadly similar microstrain patterns, reflecting effective durophagous abilities. Conversely, low microstrain values across the O. serratus protopodite suggest that the elongate gnathobasic spines transferred minimal strain, implying that this species was less well-adapted to masticate hard prey. These results confirm that Cambrian euarthropods with transversely elongate protopodites bearing short, robust gnathobasic spines were likely durophages. Comparatively, taxa with shorter protopodites armed with long spines, such as O. serratus, were more likely restricted to a soft food diet. The prevalence of Cambrian gnathobase-bearing euarthropods and their various feeding specializations may have accelerated the development of complex trophic relationships within early animal ecosystems, especially the 'arms race' between predators and biomineralized prey.

Global elongation and high shape flexibility as an evolutionary hypothesis of accommodating mammalian brains into skulls.

Little is known about how the large brains of mammals are accommodated into the dazzling diversity of their skulls. It has been suggested that brain shape is influenced by relative brain size, that it evolves or develops according to extrinsic or intrinsic mechanical constraints, and that its shape can provide insights into its proportions and function. Here, we characterize the shape variation among 84 marsupial cranial endocasts of 57 species including fossils, using three-dimensional geometric morphometrics and virtual dissections. Statistical shape analysis revealed four main patterns: over half of endocast shape variation ranges from elongate and straight to globular and inclined; little allometric variation with respect to centroid size, and none for relative volume; no association between locomotion and endocast shape; limited association between endocast shape and previously published histological cortex volumes. Fossil species tend to have smaller cerebral hemispheres. We find divergent endocast shapes in closely related species and within species, and diverse morphologies superimposed over the main variation. An evolutionarily and individually malleable brain with a fundamental tendency to arrange into a spectrum of elongate-to-globular shapes-possibly mostly independent of brain function-may explain the accommodation of brains within the enormous diversity of mammalian skull form.

Quantitatively assessing mekosuchine crocodile locomotion by geometric morphometric and finite element analysis of the forelimb.

Morphological shifts observed in the fossil record of a lineage potentially indicate concomitant shifts in ecology of that lineage. Mekosuchine crocodiles of Cenozoic Australia display departures from the typical eusuchian body-plan both in the cranium and postcranium. Previous qualitative studies have suggested that these crocodiles had a more terrestrial habitus than extant crocodylians, yet the capacity of mekosuchine locomotion remains to be tested. Limb bone shape, such as diaphyseal cross-section and curvature, has been related to habitual use and locomotory function across a wide variety of taxa. Available specimens of mekosuchine limbs, primarily humeri, are distinctly columnar compared with those of extant crocodylians. Here we apply a quantitative approach to biomechanics in mekosuchine taxa using both geomorphic morphometric and finite element methods to measure bone shape and estimate locomotory stresses in a comparative context. Our results show mekosuchines appear to diverge from extant semi-aquatic saltwater and freshwater crocodiles in cross-sectional geometry of the diaphysis and generate different structural stresses between models that simulate sprawling and high-walk gaits. The extant crocodylians display generally rounded cross-sectional diaphyseal outlines, which may provide preliminary indication of resistance to torsional loads that predominate during sprawling gait, whereas mekosuchine humeri appear to vary between a series of elliptical outlines. Mekosuchine structural stresses are comparatively lower than those of the extant crocodylians and reduce under high-walk gait in some instances. This appears to be a function of bending moments induced by differing configurations of diaphyseal curvature. Additionally, the neutral axis of structural stresses is differently oriented in mekosuchines. This suggests a shift in the focus of biomechanical optimisation, from torsional to axial loadings. Our results lend quantitative support to the terrestrial habitus hypothesis in so far as they suggest that mekosuchine humeri occupied a different morphospace than that associated with the semi-aquatic habit. The exact adaptational trajectory of mekosuchines, however, remains to be fully quantified. Novel forms appear to emerge among mekosuchines during the late Cenozoic. Their adaptational function is considered here; possible applications include navigation of uneven terrain and burrowing.

Morphometric analysis of the hominin talus: Evolutionary and functional implications.

The adoption of bipedalism is a key benchmark in human evolution that has impacted talar morphology. Here, we investigate talar morphological variability in extinct and extant hominins using a 3D geometric morphometric approach. The evolutionary timing and appearance of modern human-like features and their contributions to bipedal locomotion were evaluated on the talus as a whole, each articular facet separately, and multiple combinations of facets. Distinctive suites of features are consistently present in all fossil hominins, despite the presence of substantial interspecific variation, suggesting a potential connection of these suites to bipedal gait. A modern human-like condition evolved in navicular and lateral malleolar facets early in the hominin lineage compared with other facets, which demonstrate more complex morphological variation within Homininae. Interestingly, navicular facet morphology of Australopithecus afarensis is derived in the direction of Homo, whereas more recent hominin species such as Australopithecus africanus and Australopithecus sediba retain more primitive states in this facet. Combining the navicular facet with the trochlea and the posterior calcaneal facet as a functional suite, however, distinguishes Australopithecus from Homo in that the medial longitudinal arch had not fully developed in the former. Our results suggest that a more everted foot and stiffer medial midtarsal region are adaptations that coincide with the emergence of bipedalism, whereas a high medial longitudinal arch emerges later in time, within Homo. This study provides novel insights into the emergence of talar morphological traits linked to bipedalism and its transition from a facultative to an obligate condition.

The influence of mobility strategy on the modern human talus.

OBJECTIVES: The primate talus is known to have a shape that varies according to differences in locomotion and substrate use. While the modern human talus is morphologically specialized for bipedal walking, relatively little is known on how its morphology varies in relation to cultural and environmental differences across time. Here we compare tali of modern human populations with different subsistence economies and lifestyles to explore how cultural practices and environmental factors influence external talar shape. MATERIALS AND METHODS: The sample consists of digital models of 142 tali from 11 archaeological and post-industrial modern human groups. Talar morphology was investigated through 3D (semi)landmark based geometric morphometric methods. RESULTS: Our results show distinct differences between highly mobile hunter-gatherers and more sedentary groups belonging to a mixed post-agricultural/industrial background. Hunter-gatherers exhibit a more "flexible" talar shape, everted posture, and a more robust and medially oriented talar neck/head, which we interpret as reflecting long-distance walking strictly performed barefoot, or wearing minimalistic footwear, along uneven ground. The talus of the post-industrial population exhibits a "stable" profile, neutral posture, and a less robust and orthogonally oriented talar neck/head, which we interpret as a consequence of sedentary lifestyle and use of stiff footwear. DISCUSSION: We suggest that talar morphological variation is related to the adoption of constraining footwear in post-industrial society, which reduces ankle range of motion. This contrasts with hunter-gatherers, where talar shape shows a more flexible profile, likely resulting from a lack of footwear while traversing uneven terrain. We conclude that modern human tali vary with differences in locomotor and cultural behavior.

Head to head: the case for fighting behaviour in Megaloceros giganteus using finite-element analysis.

The largest antlers of any known deer species belonged to the extinct giant deer Megaloceros giganteus. It has been argued that their antlers were too large for use in fighting, instead being used only in ritualized displays to attract mates. Here, we used finite-element analysis to test whether the antlers of M. giganteus could have withstood forces generated during fighting. We compared the mechanical performance of antlers in M. giganteus with three extant deer species: red deer (Cervus elaphus), fallow deer (Dama dama) and elk (Alces alces). Von Mises stress results suggest that M. giganteus was capable of withstanding some fighting loads, provided that their antlers interlocked proximally, and that their antlers were best adapted for withstanding loads from twisting rather than pushing actions, as are other deer with palmate antlers. We conclude that fighting in M. giganteus was probably more constrained and predictable than in extant deer.

Evolution of the vomer and its implications for cranial kinesis in Paraves.

Most living birds exhibit cranial kinesis-movement between the rostrum and braincase-in which force is transferred through the palatal and jugal bars. The palate alone distinguishes the Paleognathae from the Neognathae, with cranial kinesis more developed in neognaths. Most previous palatal studies were based on 2D data and rarely incorporated data from stem birds despite great interest in their kinetic abilities. Here we reconstruct the vomer of the Early Cretaceous stem bird Sapeornis and the troodontid Sinovenator, taxa spanning the dinosaur-bird transition. A 3D shape analysis including these paravians and an extensive sampling of neornithines reveals their strong similarity to paleognaths and indicates that morphological differences in the vomer between paleognaths and neognaths are intimately related to their different kinetic abilities. These results suggest the skull of Mesozoic paravians lacked the kinetic abilities observed in neognaths, a conclusion also supported by our identification of an ectopterygoid in Sapeornis here. We conclude that cranial kinesis evolved relatively late, likely an innovation of the Neognathae, and is linked to the transformation of the vomer. This transformation increased palatal mobility, enabling the evolution of a diversity of kinetic mechanisms and ultimately contributing to the extraordinary evolutionary success of this clade.

Impact of transition to a subterranean lifestyle on morphological disparity and integration in talpid moles (Mammalia, Talpidae).

BACKGROUND: Understanding the mechanisms promoting or constraining morphological diversification within clades is a central topic in evolutionary biology. Ecological transitions are of particular interest because of their influence upon the selective forces and factors involved in phenotypic evolution. Here we focused on the humerus and mandibles of talpid moles to test whether the transition to the subterranean lifestyle impacted morphological disparity and phenotypic traits covariation between these two structures. RESULTS: Our results indicate non-subterranean species occupy a significantly larger portion of the talpid moles morphospace. However, there is no difference between subterranean and non-subterranean moles in terms of the strength and direction of phenotypic integration. CONCLUSIONS: Our study shows that the transition to a subterranean lifestyle significantly reduced morphological variability in talpid moles. However, this reduced disparity was not accompanied by changes in the pattern of traits covariation between the humerus and the mandible, suggesting the presence of strong phylogenetic conservatism within this pattern.

Morphological integration affects the evolution of midline cranial base, lateral basicranium, and face across primates.

OBJECTIVES: The basicranium and face are two integrated bony structures displaying great morphological diversity across primates. Previous studies in hominids determined that the basicranium is composed of two independent modules: the midline basicranium, mostly influenced by brain size, and the lateral basicranium, predominantly associated with facial shape. To better assess how morphological integration impacts the evolution of primate cranial shape diversity, we test to determine whether the relationships found in hominids are retained across the order. MATERIALS AND METHODS: Three-dimensional landmarks (29) were placed on 143 computed tomography scans of six major clades of extant primate crania. We assessed the covariation between midline basicranium, lateral basicranium, face, and endocranial volume using phylogenetically informed partial least squares analyses and phylogenetic generalized least squares models. RESULTS: We found significant integration between lateral basicranium and face and between midline basicranium and face. We also described a significant correlation between midline basicranium and endocranial volume but not between lateral basicranium and endocranial volume. DISCUSSION: Our findings demonstrate a significant and pervasive integration in the craniofacial structures across primates, differing from previous results in hominids. The uniqueness of module organization in hominids may explain this distinction. We found that endocranial volume is significantly integrated to the midline basicranium but not to the lateral basicranium. This finding underlines the significant effect of brain size on the shape of the midline structures of the cranial base in primates. With the covariations linking the studied features defined here, we suggest that future studies should focus on determining the causal links between them.

Elemental signatures of Australopithecus africanus teeth reveal seasonal dietary stress.

Reconstructing the detailed dietary behaviour of extinct hominins is challenging1-particularly for a species such as Australopithecus africanus, which has a highly variable dental morphology that suggests a broad diet2,3. The dietary responses of extinct hominins to seasonal fluctuations in food availability are poorly understood, and nursing behaviours even less so; most of the direct information currently available has been obtained from high-resolution trace-element geochemical analysis of Homo sapiens (both modern and fossil), Homo neanderthalensis4 and living apes5. Here we apply high-resolution trace-element analysis to two A. africanus specimens from Sterkfontein Member 4 (South Africa), dated to 2.6-2.1 million years ago. Elemental signals indicate that A. africanus infants predominantly consumed breast milk for the first year after birth. A cyclical elemental pattern observed following the nursing sequence-comparable to the seasonal dietary signal that is seen in contemporary wild primates and other mammals-indicates irregular food availability. These results are supported by isotopic evidence for a geographical range that was dominated by nutritionally depauperate areas. Cyclical accumulation of lithium in A. africanus teeth also corroborates the idea that their range was characterized by fluctuating resources, and that they possessed physiological adaptations to this instability. This study provides insights into the dietary cycles and ecological behaviours of A. africanus in response to food availability, including the potential cyclical resurgence of milk intake during times of nutritional challenge (as observed in modern wild orangutans5). The geochemical findings for these teeth reinforce the unique place of A. africanus in the fossil record, and indicate dietary stress in specimens that date to shortly before the extinction of Australopithecus in South Africa about two million years ago.

Raptor talon shape and biomechanical performance are controlled by relative prey size but not by allometry.

Most birds of prey (raptors), rely heavily on their talons for capturing prey. However, the relationship between talon shape and the ability to take prey is poorly understood. In this study we investigate whether raptor talons have evolved primarily in response to adaptive pressures exerted by different dietary demands, or if talon morphology is largely constrained by allometric or phylogenetic factors. We focus on the hallux talon and include 21 species in total varying greatly in body mass and feeding ecology, ranging from active predation on relatively large prey to obligate scavenging. To quantify the variation in talon shape and biomechanical performance within a phylogenetic framework, we combined three dimensional (3D) geometric morphometrics, finite element modelling and phylogenetic comparative methods. Our results indicate that relative prey size plays a key role in shaping the raptorial talon. Species that hunt larger prey are characterised by both distinct talon shape and mechanical performance when compared to species that predate smaller prey, even when accounting for phylogeny. In contrast to previous results of skull-based analysis, allometry had no significant effect. In conclusion, we found that raptor talon evolution has been strongly influenced by relative prey size, but not allometry and, that talon shape and mechanical performance are good indicators of feeding ecology.

Comparative Three-Dimensional Moment Arm Analysis of the Sauropod Forelimb: Implications for the Transition to a Wide-Gauge Stance in Titanosaurs.

The evolution of extraordinarily large size among Sauropoda was associated with a number of biomechanical adaptations. Changes in muscle moment arms undoubtedly accompanied these adaptations, but since muscles rarely fossilize, our ability to understand them has been restricted. Here, we use three-dimensional (3D) musculoskeletal modeling to reconstruct and quantitatively assess leverage of forelimb muscles in the transition from the narrow-gauge stance of basal sauropods to a wide-gauge stance in titanosaurs. A comparative analysis is conducted on three neosauropods: the narrow-gauge diplodocid Apatosaurus louisae, the intermediate-gauge titanosariform Giraffatitan brancai, and the wide-gauge titanosaur Diamantinasaurus matildae. In this study, moment arm magnitudes and corresponding morphological evidence indicates multiple changes across the narrow-gauge to wide-gauge transition in sauropods. High shoulder adduction was found in Diamantinasaurus, suggesting functional changes for supporting a wider stance and a limb less aligned with ground reaction force. High leverage in shoulder extension of Diamantinasaurus and Giraffatitan is possibly related to the increased use of the forelimb in forward propulsion with an anterior shift in center of mass. In addition, the prominence of the olecranon process in Diamantinasaurus produced high moment arm leverage in elbow flexion and extension, suggesting titanosaurs might have maintained a more flexed forelimb posture and displayed an increased degree of maneuverability. Other results are more variable between taxa but still indicate smaller scale changes. A sensitivity analysis was also conducted to measure the reliability of our models and test specific uncertainties within the modeling process, as well as other uncertainties uncovered during analysis. Anat Rec, 2018. © 2018 Wiley Periodicals, Inc. Anat Rec, 302:794-817, 2019. © 2018 Wiley Periodicals, Inc.

Three-Dimensional Musculoskeletal Modeling of the Sauropodomorph Hind Limb: The Effect of Postural Change on Muscle Leverage.

The biomechanical constraints for life at massive size can be directly observed in the evolutionary history of sauropodomorph dinosaurs. Members of this lineage underwent a number of major postural transitions as they increased in size from relatively small bipedal dinosaurs to massive titanosaurs that include the largest terrestrial animals of all time. To better understand the impact of gigantic size on the biomechanics of sauropods, we used three-dimensional musculoskeletal modeling to investigate how hind limb musculature was affected, first by the development of a quadrupedal stance from a bipedal one, and later in the transition from a narrow-gauge to a wide-gauge stance. Muscle moment arms were measured in four sauropodomorph taxa: the bipedal basal sauropodomorph Plateosaurus engelhardti, the narrow-gauge diplodocid Diplodocus carnegii, the titanosauriform Giraffatitan brancai, and the wide-gauge titanosaur Diamantinasaurus matildae. In Plateosaurus, low moment arm leverage in the hip extensors and knee flexors and extensors was observed suggesting high-velocity movement for fast locomotion. A reduction in hip extensor leverage in Diamantinasaurus was found which suggests a reduced role for the hind limb in forward propulsion in titanosaurs. An increase in overall hip adductor leverage and leverage of adductors 1 and 2 in Diamantinasaurus, compared with other taxa studied, might relate to the development of a wide-gauge stance. High knee flexor-extensor leverage in Giraffatitan but not Diamantinasaurus partially refutes the idea that broader femoral condyles in titanosauriforms increased knee torque production capabilities. Sauropodomorph postural changes clearly had an impact on the function and leverage of hind limb muscles. Anat Rec, 301:2145-2163, 2018. © 2018 Wiley Periodicals, Inc.