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The last common ancestor of birds and crocodylians plus all of its descendants (clade Archosauria) dominated terrestrial Mesozoic ecosystems, giving rise to disparate body plans, sizes, and modes of locomotion. As in the fields of vertebrate morphology and paleontology more generally, studies of archosaur skeletal structure have come to depend on tools for acquiring, measuring, and exploring three-dimensional (3-D) digital models. Such models, in turn, form the basis for many analyses of musculoskeletal function. A set of shared conventions for describing 3-D pose (joint or limb configuration) and 3-D kinematics (change in pose through time) is essential for fostering comparison of posture/movement among such varied species, as well as for maximizing communication among scientists. Following researchers in human biomechanics, we propose a standard methodological approach for measuring the relative position and orientation of the major segments of the archosaur pelvis and hindlimb in 3-D. We describe the construction of anatomical and joint coordinate systems using the extant guineafowl and alligator as examples. Our new standards are then applied to three extinct taxa sampled from the wider range of morphological, postural, and kinematic variation that has arisen across250 million years of archosaur evolution. These proposed conventions, and the founding principles upon which they are based, can also serve as starting points for measuring poses between elements within a hindlimb segment, for establishing coordinate systems in the forelimb and axial skeleton, or for applying our archosaurian system more broadly to different vertebrate clades.

The intersection of paleontology and biomechanics can be reciprocally illuminating, helping to improve paleobiological knowledge of extinct species and furthering our understanding of the generality of biomechanical principles derived from study of extant species. However, working with data gleaned primarily from the fossil record has its challenges. Building on decades of prior research, we outline and critically discuss a complete workflow for biomechanical analysis of extinct species, using locomotor biomechanics in the Triassic theropod dinosaurCoelophysisas a case study. We progress from the digital capture of fossil bone morphology to creating rigged skeletal models, to reconstructing musculature and soft tissue volumes, to the development of computational musculoskeletal models, and finally to the execution of biomechanical simulations. Using a three-dimensional musculoskeletal model comprising 33 muscles, a static inverse simulation of the mid-stance of running shows thatCoelophysisprobably used more upright (extended) hindlimb postures and was likely capable of withstanding a vertical ground reaction force of magnitude more than 2.5 times body weight. We identify muscle force-generating capacity as a key source of uncertainty in the simulations, highlighting the need for more refined methods of estimating intrinsic muscle parameters such as fiber length. Our approach emphasizes the explicit application of quantitative techniques and physics-based principles, which helps maximize results robustness and reproducibility. Although we focus on one specific taxon and question, many of the techniques and philosophies explored here have much generality to them, so they can be applied in biomechanical investigation of other extinct organisms.

Randau, Marcela, Goswami, Anjali, Hutchinson, John R., Cuff, Andrew R., Pierce, Stephanie E. (2016): Cryptic complexity in felid vertebral evolution: shape differentiation and allometry of the axial skeleton. Zoological Journal of the Linnean Society 178 (1): 183-202, DOI: 10.1111/zoj.12403, URL: http://dx.doi.org/10.1111/zoj.12403

The first finite element (FE) validation of a complete avian cranium was performed on an extant palaeognath, the ostrich (Struthio camelus). Ex-vivo strains were collected fromthe cranial bone and rhamphotheca. These experimental strains were then compared to convergence tested, specimen-specific finite element (FE) models. The FE models contained segmented cortical and trabecular bone, sutures and the keratinous rhamphotheca as identified frommicro-CT scan data. Each of these individual materials was assigned isotropic material properties either fromthe literature or fromnanoindentation, and the FE models compared to the ex-vivo results. The FE models generally replicate the location of peak strains and reflect the correct mode of deformation in the rostral region. The models are too stiff in regions of experimentally recorded high strain and too elastic in regions of low experimentally recorded low strain. The mode of deformation in the low strain neurocranial region is not replicated by the FE models, and although the models replicate strain orientations to within 10? in some regions, in most regions the correlation is not strong. Cranial sutures, as has previously been found in other taxa, are important for modifying both strain magnitude and strain patterns across the entire skull, but especially between opposing the sutural junctions. Experimentally, we find that the strains on the surface of the rhamphotheca are much lower than those found on nearby bone. The FE models producemuch higher principal strains despite similar strain ratios across the entirety of the rhamphotheca. This study emphasises the importance of attempting to validate FE models, modelling sutures and rhamphothecae in birds, and shows that whilst location of peak strain and patterns of deformation can be modelled, replicating experimental data in digital models of avian crania remains problematic.

We developed a three‐dimensional, computational biomechanical model of a juvenile Nile crocodile (Crocodylus niloticus) pelvis and hindlimb, composed of 47 pelvic limb muscles, to investigate muscle function. We tested whether crocodiles, which are known to use a variety of limb postures during movement, use limb orientations (joint angles) that optimise the moment arms (leverages) or moment‐generating capacities of their muscles during different limb postures ranging from a high walk to a sprawling motion. We also describe the three‐dimensional (3D) kinematics of the crocodylian hindlimb during terrestrial locomotion across an instrumented walkway and a treadmill captured via X‐ray Reconstruction of Moving Morphology (biplanar fluoroscopy; ‘XROMM’). We reconstructed the 3D positions and orientations of each of the hindlimb bones and used dissection data for muscle lines of action to reconstruct a focal, subject‐specific 3D musculoskeletal model. Motion data for different styles of walking (a high, crouched, bended and two types of sprawling motion) were fed into the 3D model to identify whether any joints adopted near‐optimal poses for leverage across each of the behaviours. We found that (1) the hip adductors and knee extensors had their largest leverages during sprawling postures and (2) more erect postures typically involved greater peak moment arms about the hip (flexion‐extension), knee (flexion) and metatarsophalangeal (flexion) joints. The results did not fully support the hypothesis that optimal poses are present during different locomotory behaviours because the peak capacities were not always reached around mid‐stance phase. Furthermore, we obtained few clear trends for isometric moment‐generating capacities. Therefore, perhaps peak muscular leverage in Nile crocodiles is instead reached either in early/late stance or possibly during swing phase or other locomotory behaviours that were not studied here, such as non‐terrestrial movement. Alternatively, our findings could reflect a trade‐off between having to execute different postures, meaning that hindlimb muscle leverage is not optimised for any singular posture or behaviour. Our model, however, provides a comprehensive set of 3D estimates of muscle actions in extant crocodiles which can form a basis for investigating muscle function in extinct archosaurs., We created an open‐source computational 3D biomechanical model of the Nile crocodile hindlimb containing 47 digitally dissected muscles, permitting, for the first time, a comprehensive investigation into muscular leverage in each of the locomotory behaviours/postures used by crocodiles. Here we integrate data from three‐dimensional anatomy, 3D locomotor kinematics in vivo and 3D biomechanical modelling to quantify how Nile crocodiles move. We found that our results did not fully support the hypothesis that biomechanically optimal poses (in terms of leverage) are adopted during different locomotory behaviours in Nile crocodiles. Therefore, we infer that the wide range of locomotor behaviours used by crocodylians on land prohibits simple optimisation of muscle leverage to any single particular behaviour.