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Fossil footprints (i.e., tracks) were believed to document arch anatomical evolution, although our recent work has shown that track arches record foot kinematics instead. Analyses of track arches can thereby inform the evolution of human locomotion, although quantifying this 3-D aspect of track morphology is difficult. Here, we present a volumetric method for measuring the arches of 3-D models of human tracks and feet, using both Autodesk Maya and Blender software. The method involves generation of a 3-D object that represents the space beneath the longitudinal arch, and measurement of that arch object's geometry and spatial orientation. We provide relevant tools and guidance for users to apply this technique to their own data. We present three case studies to demonstrate potential applications. These include, (1) measuring the arches of static and dynamic human feet, (2) comparing the arches of human tracks with the arches of the feet that made them, and (3) direct comparisons of human track and foot arch morphology throughout simulated track formation. The volumetric measurement tool proved robust for measuring 3-D models of human tracks and feet, in static and dynamic contexts. This tool enables researchers to quantitatively compare arches of fossil hominin tracks, in order to derive biomechanical interpretations from them, and/or offers a different approach for quantifying foot morphology in living humans., (© 2024 Wiley Periodicals LLC.)

Our knowledge of vertebrate functional evolution depends on inferences about joint function in extinct taxa. Without rigorous criteria for evaluating joint articulation, however, such analyses risk misleading reconstructions of vertebrate animal motion. Here we propose an approach for synthesizing raycast-based measurements of 3-D articular overlap, symmetry, and congruence into a quantitative "articulation score" for any non-interpenetrating six-degree-of-freedom joint configuration. We apply our methodology to bicondylar hindlimb joints of two extant dinosaurs (guineafowl, emu) and, through comparison with in vivo kinematics, find that locomotor joint poses consistently have high articulation scores. We then exploit this relationship to constrain reconstruction of a pedal walking stride cycle for the extinct dinosaur Deinonychus antirrhopus, demonstrating the utility of our approach. As joint articulation is investigated in more living animals, the framework we establish here can be expanded to accommodate additional joints and clades, facilitating improved understanding of vertebrate animal motion and its evolution., (© 2024. The Author(s).)

Major transformations in the locomotor system of archosaurs (a major clade of reptiles including birds, crocodiles, dinosaurs, and pterosaurs) were accompanied by significant modifications to ankle anatomy. How the evolution of such a complex multi-joint structure is related to shifts in ankle function and locomotor diversity across this clade remains unclear and weakly grounded in extant experimental data. Here, we used X-ray Reconstruction of Moving Morphology to reconstruct skeletal motion and quantify the sources of three-dimensional ankle mobility in the American alligator, a species that retains the ancestral archosaur ankle structure. We then applied the observed relationships between joint excursion and locomotor behaviors to predict ankle function in extinct archosaurs. High-resolution reconstructions of Alligator skeletal movement revealed previously unseen regionalized coordination among joints responsible for overall ankle rotation. Differences in joint contributions between maneuvers and steady walking parallel transitions in mobility inferred from the ankle structure of fossil taxa in lineages with more erect hind limb postures. Key ankle structures related to ankle mobility were identified in the alligator, which permitted the characterization of ancestral archosaur ankle function. Modifications of these structures provide morphological evidence for functional convergence among sublineages of bird-line and crocodylian-line archosaurs. Using the dynamic insight into the internal sources of Alligator ankle mobility and trends among locomotor modes, we trace anatomical shifts and propose a mechanistic hypothesis for the evolution of ankle structure and function across Archosauria., (© 2022 The Authors. Journal of Anatomy published by John Wiley & Sons Ltd on behalf of Anatomical Society.)

The longitudinal arch of the human foot is viewed as a pivotal adaptation for bipedal walking and running. Fossil footprints from Laetoli, Tanzania, and Ileret, Kenya, are believed to provide direct evidence of longitudinally arched feet in hominins from the Pliocene and Pleistocene, respectively. We studied the dynamics of track formation using biplanar X-ray, three-dimensional animation and discrete element particle simulation. Here, we demonstrate that longitudinally arched footprints are false indicators of foot anatomy; instead they are generated through a specific pattern of foot kinematics that is characteristic of human walking. Analyses of fossil hominin tracks from Laetoli show only partial evidence of this walking style, with a similar heel strike but a different pattern of propulsion. The earliest known evidence for fully modern human-like bipedal kinematics comes from the early Pleistocene Ileret tracks, which were presumably made by members of the genus Homo. This result signals important differences in the foot kinematics recorded at Laetoli and Ileret and underscores an emerging picture of locomotor diversity within the hominin clade., (© 2023. The Author(s), under exclusive licence to Springer Nature Limited.)

Living birds (Aves) have bodies substantially modified from the ancestral reptilian condition. The avian pelvis in particular experienced major changes during the transition from early archosaurs to living birds 1,2 . This stepwise transformation is well documented by an excellent fossil record 2-4 ; however, the ontogenetic alterations that underly it are less well understood. We used embryological imaging techniques to examine the morphogenesis of avian pelvic tissues in three dimensions, allowing direct comparison with the fossil record. Many ancestral dinosaurian features 2 (for example, a forward-facing pubis, short ilium and pubic 'boot') are transiently present in the early morphogenesis of birds and arrive at their typical 'avian' form after transitioning through a prenatal developmental sequence that mirrors the phylogenetic sequence of character acquisition. We demonstrate quantitatively that avian pelvic ontogeny parallels the non-avian dinosaur-to-bird transition and provide evidence for phenotypic covariance within the pelvis that is conserved across Archosauria. The presence of ancestral states in avian embryos may stem from this conserved covariant relationship. In sum, our data provide evidence that the avian pelvis, whose early development has been little studied 5-7 , evolved through terminal addition-a mechanism 8-10 whereby new apomorphic states are added to the end of a developmental sequence, resulting in expression 8,11 of ancestral character states earlier in that sequence. The phenotypic integration we detected suggests a previously unrecognized mechanism for terminal addition and hints that retention of ancestral states in development is common during evolutionary transitions., (© 2022. The Author(s), under exclusive licence to Springer Nature Limited.)

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 across >250 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., (© 2022 The Authors. Journal of Anatomy published by John Wiley & Sons Ltd on behalf of Anatomical Society.)

The stance phase of walking is when forces are applied to the environment to support, propel, and maneuver the body. Unlike solid surfaces, deformable substrates yield under load, allowing the foot to sink to varying degrees. For bipedal birds and their dinosaurian ancestors, a shared response to walking on these substrates has been identified in the looping path the digits follow underground. Because a volume of substrate preserves a 3-D record of stance phase in the form of footprints or tracks, understanding how the bipedal stride cycle relates to this looping motion is critical for building a track-based framework for the study of walking in extinct taxa. Here we used biplanar X-ray imaging to record and analyze 161 stance phases from 81 trials of three Helmeted Guineafowl (Numida meleagris) walking on radiolucent substrates of different consistency (solid, dry granular, firm to semi-liquid muds). Across all substrates, the feet sank to a range of depths up to 78% of hip height. With increasing substrate hydration, the majority of foot motion shifted from above to below ground. Walking kinematics sampled across all stride cycles revealed six sequential gait-based events originating from both feet, conserved throughout the spectrum of substrate consistencies during normal alternating walking. On all substrates that yielded, five sub-phases of gait were drawn out in space and formed a loop of varying shape. We describe the two-footed coordination and weight distribution that likely contributed to the observed looping patterns of an individual foot. Given such complex subsurface foot motion during normal alternating walking and some atypical walking behaviors, we discuss the definition of "stance phase" on deformable substrates. We also discuss implications of the gait-based origins of subsurface looping on the interpretation of locomotory information preserved in fossil dinosaur tracks., (© The Author(s) 2022. Published by Oxford University Press on behalf of the Society for Integrative and Comparative Biology.)

Paleobiological reconstructions of joint mobility are an essential component of functional analyses of extinct animals. Over the past half-decade, the methods underlying mobility studies have advanced rapidly in three main areas: increasing complexity of virtual joint manipulation, formalizing pose viability criteria, and constructing more rigorous quantitative frameworks. Here we contextualize and review the recent history of this field, and call attention to remaining challenges and potential future directions. Additionally, we make available and describe a set of user-friendly scripts for the animation software Autodesk Maya. In doing so, we aim to make many of the latest approaches for virtual mobility reconstruction more easily accessible to other researchers, encouraging their broader adoption and collaborative improvement., (© The Author(s) 2022. Published by Oxford University Press on behalf of the Society for Integrative and Comparative Biology.)

Virtual and augmented reality (VR/AR) are new technologies with the power to revolutionize the study of morphology. Modern imaging approaches such as computed tomography, laser scanning, and photogrammetry have opened up a new digital world, enabling researchers to share and analyze morphological data electronically and in great detail. Because this digital data exists on a computer screen, however, it can remain difficult to understand and unintuitive to interact with. VR/AR technologies bridge the analog-to-digital divide by presenting 3D data to users in a very similar way to how they would interact with actual anatomy, while also providing a more immersive experience and greater possibilities for exploration. This manuscript describes VR/AR hardware, software, and techniques, and is designed to give practicing morphologists and educators a primer on using these technologies in their research, pedagogy, and communication to a wide variety of audiences. We also include a series of case studies from the presentations and workshop given at the 2019 International Congress of Vertebrate Morphology, and suggest best practices for the use of VR/AR in comparative morphology., (© 2021 Wiley Periodicals LLC.)

Paleobiologists typically exclude impossible joint poses from reconstructions of extinct animals by estimating the rotational range of motion (ROM) of fossil joints. However, this ubiquitous practice carries the assumption that osteological estimates of ROM consistently overestimate true joint mobility. Because studies founded on ROM-based exclusion have contributed substantially to our understanding of functional and locomotor evolution, it is critical that this assumption be tested. Here, we evaluate whether ROM-based exclusion is, as currently implemented, a reliable strategy. We measured the true mobilities of five intact cadaveric joints using marker-based X-ray Reconstruction of Moving Morphology and compared them to virtual osteological estimates of ROM made allowing (a) only all three rotational, (b) all three rotational and one translational, and (c) all three rotational and all three translational degrees of freedom. We found that allowing combinations of motions in all six degrees of freedom is necessary to ensure that true mobility is always successfully captured. In other words, failing to include joint translations in ROM analyses results in the erroneous exclusion of many joint poses that are possible in life. We therefore suggest that the functional and evolutionary conclusions of existing paleobiological reconstructions may be weakened or even overturned when all six degrees of freedom are considered. We offer an expanded methodological framework for virtual ROM estimation including joint translations and outline recommendations for future ROM-based exclusion studies., (© 2021 Anatomical Society.)

The emergence of bipedalism had profound effects on human evolutionary history, but the evolution of locomotor patterns within the hominin clade remains poorly understood. Fossil tracks record in vivo behaviours of extinct hominins, and they offer great potential to reveal locomotor patterns at various times and places across the human fossil record. However, there is no consensus on how to interpret anatomical or biomechanical patterns from tracks due to limited knowledge of the complex foot-substrate interactions through which they are produced. Here, we implement engineering-based methods to understand human track formation with the ultimate goal of unlocking invaluable information on hominin locomotion from fossil tracks. We first developed biplanar X-ray and three-dimensional animation techniques that permit visualization of subsurface foot motion as tracks are produced, and that allow for direct comparisons of foot kinematics to final track morphology. We then applied the discrete element method to accurately simulate the process of human track formation, allowing for direct study of human track ontogeny. This window lets us observe how specific anatomical and/or kinematic variables shape human track morphology, and it offers a new avenue for robust hypothesis testing in order to infer patterns of foot anatomy and motion from fossil hominin tracks., (© 2021 The Author(s).)

Feet must mediate substrate interactions across an animal's entire range of limb poses used in life. Metatarsals, the 'bones of the sole', are the dominant pedal skeletal elements for most tetrapods. In plantigrade species that walk on the entirety of their sole, such as living crocodylians, intermetatarsal mobility offers the potential for a continuum of reconfiguration within the foot itself. Alligator hindlimbs are capable of postural extremes from a belly sprawl to a high walk to sharp turns - how does the foot morphology dynamically accommodate these diverse demands? We implemented a hybrid combination of marker-based and markerless X-ray reconstruction of moving morphology (XROMM) to measure 3D metatarsal kinematics in three juvenile American alligators (Alligator mississippiensis) across their locomotor and maneuvering repertoire on a motorized treadmill and flat-surfaced arena. We found that alligators adaptively conformed their metatarsals to the ground, maintaining plantigrade contact throughout a spectrum of limb placements with non-planar feet. Deformation of the metatarsus as a whole occurred through variable abduction (twofold range of spread) and differential metatarsal pitching (45 deg arc of skew). Internally, metatarsals also underwent up to 65 deg of long-axis rotation. Such reorientation, which correlated with skew, was constrained by the overlapping arrangement of the obliquely expanded metatarsal bases. Such a proximally overlapping metatarsal morphology is shared by fossil archosaurs and archosaur relatives. In these extinct taxa, we suggest that intermetatarsal mobility likely played a significant role in maintaining ground contact across plantigrade postural extremes., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2021. Published by The Company of Biologists Ltd.)

Reconstructions of movement in extinct animals are critical to our understanding of major transformations in vertebrate locomotor evolution. Estimates of joint range of motion (ROM) have long been used to exclude anatomically impossible joint poses from hypothesized gait cycles. Here we demonstrate how comparative ROM data can be harnessed in a different way to better constrain locomotor reconstructions. As a case study, we measured nearly 600,000 poses from the hindlimb joints of the Helmeted Guineafowl and American alligator, which represent an extant phylogenetic bracket for the archosaurian ancestor and its pseudosuchian (crocodilian line) and ornithodiran (bird line) descendants. We then used joint mobility mapping to search for a consistent relationship between full potential joint mobility and the subset of joint poses used during locomotion. We found that walking and running poses are predictably located within full mobility, revealing additional constraints for reconstructions of extinct archosaurs. The inferential framework that we develop here can be expanded to identify ROM-based constraints for other animals and, in turn, will help to unravel the history of vertebrate locomotor evolution., Competing Interests: The authors declare no competing interest., (Copyright © 2021 the Author(s). Published by PNAS.)

Three-dimensional studies of range of motion currently plot joint poses in a 'Euler space' whose axes are angles measured in the joint's three rotational degrees of freedom. Researchers then compute the volume of a pose cloud to measure rotational mobility. However, pairs of poses that are equally different from one another in orientation are not always plotted equally far apart in Euler space. This distortion causes a single joint's mobility to change when measured based on different joint coordinate systems and precludes fair comparison among joints. Here, we present two alternative spaces inspired by a 16th century map projection - cosine-corrected and sine-corrected Euler spaces - that allow coordinate-system-independent comparison of joint rotational mobility. When tested with data from a bird hip joint, cosine-corrected Euler space demonstrated a 10-fold reduction in variation among mobilities measured from three joint coordinate systems. This new quantitative framework enables previously intractable, comparative studies of articular function., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2020. Published by The Company of Biologists Ltd.)

The feet of ground-dwelling birds retain many features of their dinosaurian ancestry. Experiments with living species offer insights into the complex interplay among anatomy, kinematics and substrate during the formation of Mesozoic footprints. However, a key aspect of the track-making process, sub-surface foot movement, is hindered by substrate opacity. Here, we use biplanar X-rays to image guineafowl walking through radiolucent substrates of different consistency (solid, dry granular, firm to semi-liquid muds). Despite substantial kinematic variation, the foot consistently moves in a looping pattern below ground. As the foot sinks and then withdraws, the claws of the three main toes create entry and exit paths in different locations. Sampling these paths at incremental horizons captures two-dimensional features just as fossil tracks do, allowing depth-based zones to be characterized by the presence and relative position of digit impressions. Examination of deep, penetrative tracks from the Early Jurassic confirms that bipeds had an equivalent looping response to soft substrates approximately 200 Ma. Our integration of extant and extinct evidence demonstrates the influence of substrate properties on sinking depth and sub-surface foot motion, both of which are significant sources of track variation in the fossil record of dinosaurs.

Extant archosaurs exhibit highly divergent articular soft tissue anatomies between avian and crocodilian lineages. However, the general lack of understanding of the dynamic interactions among archosaur joint soft tissues has hampered further inferences about the function and evolution of these joints. Here we use contrast-enhanced computed tomography to generate 3D surface models of the pelvis, femora, and hip joint soft tissues in an extant archosaur, the American alligator. The hip joints were then animated using marker-based X-Ray Reconstruction of Moving Morphology (XROMM) to visualize soft tissue articulation during forward terrestrial locomotion. We found that the anatomical femoral head of the alligator travels beyond the cranial extent of the bony acetabulum and does not act as a central pivot, as has been suggested for some extinct archosaurs. Additionally, the fibrocartilaginous surfaces of the alligator's antitrochanter and femoral neck remain engaged during hip flexion and extension, similar to the articulation between homologous structures in birds. Moreover, the femoral insertion of the ligamentum capitis moves dorsoventrally against the membrane-bound portion of the medial acetabular wall, suggesting that the inner acetabular foramen constrains the excursion of this ligament as it undergoes cyclical stretching during the step cycle. Finally, the articular surface of the femoral cartilage model interpenetrates with those of the acetabular labrum and antitrochanter menisci; we interpret such interpenetration as evidence of compressive deformation of the labrum and of sliding movement of the menisci. Our data illustrate the utility of XROMM for studying in vivo articular soft tissue interactions. These results also allow us to propose functional hypotheses for crocodilian hip joint soft tissues, expanding our knowledge of vertebrate connective tissue biology and the role of joint soft tissues in locomotor behavior., (© 2019 Anatomical Society.)

Recent discoveries have made hominin tracks an increasingly prevalent component of the human fossil record, and these data have the capacity to inform long-standing debates regarding the biomechanics of hominin locomotion. However, there is currently no consensus on how to decipher biomechanical variables from hominin tracks. These debates can be linked to our generally limited understanding of the complex interactions between anatomy, motion, and substrate that give rise to track morphology. These interactions are difficult to study because direct visualization of the track formation process is impeded by foot and substrate opacity. To address these obstacles, we developed biplanar X-ray and computer animation methods, derived from X-ray Reconstruction of Moving Morphology (XROMM), to analyze the 3D dynamics of three human subjects' feet as they walked across four substrates (three deformable muds and rigid composite panel). By imaging and reconstructing 3D positions of external markers, we quantified the 3D dynamics at the foot-substrate interface. Foot shape, specifically heel and medial longitudinal arch deformation, was significantly affected by substrate rigidity. In deformable muds, we found that depths measured across tracks did not directly reflect the motions of the corresponding regions of the foot, and that track outlines were not perfectly representative of foot size. These results highlight the complex, dynamic nature of track formation, and the experimental methods presented here offer a promising avenue for developing and refining methods for accurately inferring foot anatomy and gait biomechanics from fossil hominin tracks., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

Measuring range of motion (ROM) is a valuable technique that can link bone morphology to joint function in both extant and extinct taxa. ROM results are commonly presented as tables or graphs of maxima and minima for each rotational degree of freedom. We investigate the interactions among three degrees of freedom using X-ray reconstruction of moving morphology (XROMM) to measure ROM of the main hind limb joints of Helmeted Guineafowl (Numida meleagris). By plotting each rotation on an axis, we generate three-dimensional ROM volumes or envelopes composed of hundreds of extreme joint positions for the hip, knee, and intertarsal joints. We find that the shapes of ROM volumes can be quite complex, and that the contribution of long-axis rotation is often substantial. Plotting in vivo poses from individual birds executing different behaviors shows varying use of potential rotational combinations within their ROM envelopes. XROMM can provide unprecedented high-resolution data on the spatial relationship of skeletal elements and thereby illuminate/elucidate the complex ways in which soft and hard tissues interact to produce functional joints. In joints with three rotational degrees of freedom, two-dimensional representations of ROM (flexion/extension and abduction/adduction) are incomplete., (© 2017 Anatomical Society.)

Human patellae (kneecaps) are thought to act as gears, altering the mechanical advantage of knee extensor muscles during running. Similar sesamoids have evolved in the knee extensor tendon independently in birds, but it is unknown if these also affect the mechanical advantage of knee extensors. Here, we examine the mechanics of the patellofemoral joint in the helmeted guineafowl Numida meleagris using a method based on muscle and tendon moment arms taken about the patella's rotation centre around the distal femur. Moment arms were estimated from a computer model representing hindlimb anatomy, using hip, knee and patellar kinematics acquired via marker-based biplanar fluoroscopy from a subject running at 1.6 ms -1 on a treadmill. Our results support the inference that the patella of Numida does alter knee extensor leverage during running, but with a mechanical advantage generally greater than that seen in humans, implying relatively greater extension force but relatively lesser extension velocity.

Marker-based XROMM requires software tools for: (1) correcting fluoroscope distortion; (2) calibrating X-ray cameras; (3) tracking radio-opaque markers; and (4) calculating rigid body motion. In this paper we describe and validate XMALab, a new open-source software package for marker-based XROMM (C++ source and compiled versions on Bitbucket). Most marker-based XROMM studies to date have used XrayProject in MATLAB. XrayProject can produce results with excellent accuracy and precision, but it is somewhat cumbersome to use and requires a MATLAB license. We have designed XMALab to accelerate the XROMM process and to make it more accessible to new users. Features include the four XROMM steps (listed above) in one cohesive user interface, real-time plot windows for detecting errors, and integration with an online data management system, XMAPortal. Accuracy and precision of XMALab when tracking markers in a machined object are ±0.010 and ±0.043 mm, respectively. Mean precision for nine users tracking markers in a tutorial dataset of minipig feeding was ±0.062 mm in XMALab and ±0.14 mm in XrayProject. Reproducibility of 3D point locations across nine users was 10-fold greater in XMALab than in XrayProject, and six degree-of-freedom bone motions calculated with a joint coordinate system were 3- to 6-fold more reproducible in XMALab. XMALab is also suitable for tracking white or black markers in standard light videos with optional checkerboard calibration. We expect XMALab to increase both the quality and quantity of animal motion data available for comparative biomechanics research., (© 2016. Published by The Company of Biologists Ltd.)

Horse racing is a multi-billion-dollar industry that has raised welfare concerns due to injured and euthanized animals. Whilst the cause of musculoskeletal injuries that lead to horse morbidity and mortality is multifactorial, pre-existing pathologies, increased speeds and substrate of the racecourse are likely contributors to foot disease. Horse hooves have the ability to naturally deform during locomotion and dissipate locomotor stresses, yet farriery approaches are utilised to increase performance and protect hooves from wear. Previous studies have assessed the effect of different shoe designs on locomotor performance; however, no biomechanical study has hitherto measured the effect of horseshoes on the stresses of the foot skeleton in vivo. This preliminary study introduces a novel methodology combining three-dimensional data from biplanar radiography with inverse dynamics methods and finite element analysis (FEA) to evaluate the effect of a stainless steel shoe on the function of a Thoroughbred horse's forefoot during walking. Our preliminary results suggest that the stainless steel shoe shifts craniocaudal, mediolateral and vertical GRFs at mid-stance. We document a similar pattern of flexion-extension in the PIP (pastern) and DIP (coffin) joints between the unshod and shod conditions, with slight variation in rotation angles throughout the stance phase. For both conditions, the PIP and DIP joints begin in a flexed posture and extend over the entire stance phase. At mid-stance, small differences in joint angle are observed in the PIP joint, with the shod condition being more extended than the unshod horse, whereas the DIP joint is extended more in the unshod than the shod condition. We also document that the DIP joint extends more than the PIP after mid-stance and until the end of the stance in both conditions. Our FEA analysis, conducted solely on the bones, shows increased von Mises and Maximum principal stresses on the forefoot phalanges in the shod condition at mid-stance, consistent with the tentative conclusion that a steel shoe might increase mechanical loading. However, because of our limited sample size none of these apparent differences have been tested for statistical significance. Our preliminary study illustrates how the shoe may influence the dynamics and mechanics of a Thoroughbred horse's forefoot during slow walking, but more research is needed to quantify the effect of the shoe on the equine forefoot during the whole stance phase, at faster speeds/gaits and with more individuals as well as with a similar focus on the hind feet. We anticipate that our preliminary analysis using advanced methodological approaches will pave the way for new directions in research on the form/function relationship of the equine foot, with the ultimate goal to minimise foot injuries and improve animal health and welfare.

As one of the earliest-known mammaliaforms, Haramiyavia clemmenseni from the Rhaetic (Late Triassic) of East Greenland has held an important place in understanding the timing of the earliest radiation of the group. Reanalysis of the type specimen using high-resolution computed tomography (CT) has revealed new details, such as the presence of the dentary condyle of the mammalian jaw hinge and the postdentary trough for mandibular attachment of the middle ear-a transitional condition of the predecessors to crown Mammalia. Our tests of competing phylogenetic hypotheses with these new data show that Late Triassic haramiyids are a separate clade from multituberculate mammals and are excluded from the Mammalia. Consequently, hypotheses of a Late Triassic diversification of the Mammalia that depend on multituberculate affinities of haramiyidans are rejected. Scanning electron microscopy study of tooth-wear facets and kinematic functional simulation of occlusion with virtual 3D models from CT scans confirm that Haramiyavia had a major orthal occlusion with the tallest lingual cusp of the lower molars occluding into the lingual embrasure of the upper molars, followed by a short palinal movement along the cusp rows alternating between upper and lower molars. This movement differs from the minimal orthal but extensive palinal occlusal movement of multituberculate mammals, which previously were regarded as relatives of haramiyidans. The disparity of tooth morphology and the diversity of dental functions of haramiyids and their contemporary mammaliaforms suggest that dietary diversification is a major factor in the earliest mammaliaform evolution.

In avian bipeds performing steady locomotion, right and left limbs are typically assumed to act out of phase, but with little kinematic disparity. However, outwardly appearing steadiness may harbor previously unrecognized asymmetries. Here, we present marker-based XROMM data showing that guineafowl on a treadmill routinely yaw away from their direction of travel using asymmetrical limb kinematics. Variation is most strongly reflected at the hip joints, where patterns of femoral long-axis rotation closely correlate to degree of yaw divergence. As yaw deviations increase, hip long-axis rotation angles undergo larger excursions and shift from biphasic to monophasic patterns. At large yaw angles, the alternately striding limbs exhibit synchronous external and internal femoral rotations of substantial magnitude. Hip coordination patterns resembling those used during sidestep maneuvers allow birds to asymmetrically modulate their mediolateral limb trajectories and thereby advance using a range of body orientations., (© 2015. Published by The Company of Biologists Ltd.)

Locomotion over deformable substrates is a common occurrence in nature. Footprints represent sedimentary distortions that provide anatomical, functional, and behavioral insights into trackmaker biology. The interpretation of such evidence can be challenging, however, particularly for fossil tracks recovered at bedding planes below the originally exposed surface. Even in living animals, the complex dynamics that give rise to footprint morphology are obscured by both foot and sediment opacity, which conceals animal-substrate and substrate-substrate interactions. We used X-ray reconstruction of moving morphology (XROMM) to image and animate the hind limb skeleton of a chicken-like bird traversing a dry, granular material. Foot movement differed significantly from walking on solid ground; the longest toe penetrated to a depth of ∼5 cm, reaching an angle of 30° below horizontal before slipping backward on withdrawal. The 3D kinematic data were integrated into a validated substrate simulation using the discrete element method (DEM) to create a quantitative model of limb-induced substrate deformation. Simulation revealed that despite sediment collapse yielding poor quality tracks at the air-substrate interface, subsurface displacements maintain a high level of organization owing to grain-grain support. Splitting the substrate volume along "virtual bedding planes" exposed prints that more closely resembled the foot and could easily be mistaken for shallow tracks. DEM data elucidate how highly localized deformations associated with foot entry and exit generate specific features in the final tracks, a temporal sequence that we term "track ontogeny." This combination of methodologies fosters a synthesis between the surface/layer-based perspective prevalent in paleontology and the particle/volume-based perspective essential for a mechanistic understanding of sediment redistribution during track formation.

Ground-dwelling birds are typically characterized as erect bipeds having hind limbs that operate parasagittally. Consequently, most previous research has emphasized flexion/extension angles and moments as calculated from a lateral perspective. Three-dimensional (3D) motion analyses have documented non-planar limb movements, but the skeletal kinematics underlying changes in foot orientation and transverse position remain unclear. In particular, long-axis rotation of the proximal limb segments is extremely difficult to measure with topical markers. Here, we present six degree of freedom skeletal kinematic data from maneuvering guineafowl acquired by marker-based XROMM (X-ray Reconstruction of Moving Morphology). Translations and rotations of the hips, knees, ankles and pelvis were derived from animated bone models using explicit joint coordinate systems. We distinguished sidesteps, sidestep yaws, crossover yaws, sidestep turns and crossover turns, but birds often performed a sequence of blended partial maneuvers. Long-axis rotation of the femur (up to 38 deg) modulated the foot's transverse position. Long-axis rotation of the tibiotarsus (up to 65 deg) also affected medio-lateral positioning, but primarily served to either re-orient a swing phase foot or yaw the body about a stance phase foot. Tarsometatarsal long-axis rotation was minimal, as was hip, knee and ankle abduction/adduction. Despite having superficially hinge-like joints, birds coordinate substantial long-axis rotations of the hips and knees to execute complex 3D maneuvers while striking a diversity of non-planar poses., (© 2014. Published by The Company of Biologists Ltd.)

Crocodylians occupy a key phylogenetic position for investigations of archosaur locomotor evolution. Compared to the well-studied hindlimb, relatively little is known about the skeletal movements and mechanics of the forelimb. In this study, we employed manual markerless XROMM (X-ray Reconstruction Of Moving Morphology) to measure detailed 3-D kinematics of the shoulder girdle and forelimb bones of American alligators (Alligator mississippiensis) walking on a treadmill. Digital models of the interclavicle, scapulocoracoid, humerus, radius and ulna were created using a 3-D laser scanner. Models were articulated and aligned to simultaneously recorded frames of fluoroscopic and standard light video to reconstruct and measure joint motion. Joint coordinate systems were established for the coracosternal, glenohumeral and elbow joints. Our analysis revealed that the limb joints only account for about half of fore/aft limb excursion; the remaining excursion results from shoulder girdle movements and lateral bending of the vertebral column. Considerable motion of each scapulocoracoid relative to the vertebral column is consistent with coracosternal mobility. The hemisellar design of the glenohumeral joint permits some additional translation, or sliding in the fore-aft plane, but this movement does not have much of an effect on the distal excursion of the bone., (© 2013 Anatomical Society.)

Theropod dinosaurs show striking morphological and functional tail variation; e.g., a long, robust, basal theropod tail used for counterbalance, or a short, modern avian tail used as an aerodynamic surface. We used a quantitative morphological and functional analysis to reconstruct intervertebral joint stiffness in the tail along the theropod lineage to extant birds. This provides new details of the tail's morphological transformation, and for the first time quantitatively evaluates its biomechanical consequences. We observe that both dorsoventral and lateral joint stiffness decreased along the non-avian theropod lineage (between nodes Theropoda and Paraves). Our results show how the tail structure of non-avian theropods was mechanically appropriate for holding itself up against gravity and maintaining passive balance. However, as dorsoventral and lateral joint stiffness decreased, the tail may have become more effective for dynamically maintaining balance. This supports our hypothesis of a reduction of dorsoventral and lateral joint stiffness in shorter tails. Along the avian theropod lineage (Avialae to crown group birds), dorsoventral and lateral joint stiffness increased overall, which appears to contradict our null expectation. We infer that this departure in joint stiffness is specific to the tail's aerodynamic role and the functional constraints imposed by it. Increased dorsoventral and lateral joint stiffness may have facilitated a gradually improved capacity to lift, depress, and swing the tail. The associated morphological changes should have resulted in a tail capable of producing larger muscular forces to utilise larger lift forces in flight. Improved joint mobility in neornithine birds potentially permitted an increase in the range of lift force vector orientations, which might have improved flight proficiency and manoeuvrability. The tail morphology of modern birds with tail fanning capabilities originated in early ornithuromorph birds. Hence, these capabilities should have been present in the early Cretaceous, with incipient tail-fanning capacity in the earliest pygostylian birds.

Past studies have shown that birds use their wings not only for flight, but also when ascending steep inclines. Uphill flap-running or wing-assisted incline running (WAIR) is used by both flight-incapable fledglings and flight-capable adults to retreat to an elevated refuge. Despite the broadly varying direction of travel during WAIR, level, and descending flight, recent studies have found that the basic wing path remains relatively invariant with reference to gravity. If so, joints undergo disparate motions to maintain a consistent wing path during those specific flapping modes. The underlying skeletal motions, however, are masked by feathers and skin. To improve our understanding of the form-functional relationship of the skeletal apparatus and joint morphology with a corresponding locomotor behavior, we used XROMM (X-ray Reconstruction of Moving Morphology) to quantify 3-D skeletal kinematics in chukars (Alectoris chukar) during WAIR (ascending with legs and wings) and ascending flight (AF, ascending with wings only) along comparable trajectories. Evidence here from the wing joints demonstrates that the glenohumeral joint controls the vast majority of wing movements. More distal joints are primarily involved in modifying wing shape. All bones are in relatively similar orientations at the top of upstroke during both behaviors, but then diverge through downstroke. Total excursion of the wing is much smaller during WAIR and the tip of the manus follows a more vertical path. The WAIR stroke appears "truncated" relative to ascending flight, primarily stemming from ca. 50% reduction in humeral depression. Additionally, the elbow and wrist exhibit reduced ranges of angular excursions during WAIR. The glenohumeral joint moves in a pattern congruent with being constrained by the acrocoracohumeral ligament. Finally, we found pronounced lateral bending of the furcula during the wingbeat cycle during ascending flight only, though the phasic pattern in chukars is opposite of that observed in starlings (Sturnus vulgaris).

Tetrapod limbs exhibit diverse postures and movements during terrestrial locomotion. As with morphological traits, the history of kinematic evolution should be accessible to reconstruction through analysis of limb motion patterns in a phylogenetic framework. However, the angular data comprising most kinematic descriptions appear to suffer from limitations that preclude meaningful comparison among disparate species. Using simple planar models, we discuss how geometric constraints render joint and elevation angles independent of neither morphology, degree of crouch, nor one another during the stance phase of locomotion. The implicit null hypothesis of potential similarity is invalidated because angular data are not viably transferable among limbs of dissimilar proportion and/or degree of crouch. Overlooking or dismissing the effect of constraints on angular parameterization hampers efforts to quantitatively elucidate the evolution of locomotion. We advocate a search for alternative methods of measuring limb movement that can decouple intersegmental coordination from morphology and posture., (Copyright © 2011 Elsevier Ltd. All rights reserved.)

Three-dimensional skeletal movement is often impossible to accurately quantify from external markers. X-ray imaging more directly visualizes moving bones, but extracting 3-D kinematic data is notoriously difficult from a single perspective. Stereophotogrammetry is extremely powerful if bi-planar fluoroscopy is available, yet implantation of three radio-opaque markers in each segment of interest may be impractical. Herein we introduce scientific rotoscoping (SR), a new method of motion analysis that uses articulated bone models to simultaneously animate and quantify moving skeletons without markers. The three-step process is described using examples from our work on pigeon flight and alligator walking. First, the experimental scene is reconstructed in 3-D using commercial animation software so that frames of undistorted fluoroscopic and standard video can be viewed in their correct spatial context through calibrated virtual cameras. Second, polygonal models of relevant bones are created from CT or laser scans and rearticulated into a hierarchical marionette controlled by virtual joints. Third, the marionette is registered to video images by adjusting each of its degrees of freedom over a sequence of frames. SR outputs high-resolution 3-D kinematic data for multiple, unmarked bones and anatomically accurate animations that can be rendered from any perspective. Rather than generating moving stick figures abstracted from the coordinates of independent surface points, SR is a morphology-based method of motion analysis deeply rooted in osteological and arthrological data.

X-Ray Reconstruction of Moving Morphology (XROMM) comprises a set of 3D X-ray motion analysis techniques that merge motion data from in vivo X-ray videos with skeletal morphology data from bone scans into precise and accurate animations of 3D bones moving in 3D space. XROMM methods include: (1) manual alignment (registration) of bone models to video sequences, i.e., Scientific Rotoscoping; (2) computer vision-based autoregistration of bone models to biplanar X-ray videos; and (3) marker-based registration of bone models to biplanar X-ray videos. Here, we describe a novel set of X-ray hardware, software, and workflows for marker-based XROMM. Refurbished C-arm fluoroscopes retrofitted with high-speed video cameras offer a relatively inexpensive X-ray hardware solution for comparative biomechanics research. Precision for our biplanar C-arm hardware and analysis software, measured as the standard deviation of pairwise distances between 1 mm tantalum markers embedded in rigid objects, was found to be +/-0.046 mm under optimal conditions and +/-0.084 mm under actual in vivo recording conditions. Mean error in measurement of a known distance between two beads was within the 0.01 mm fabrication tolerance of the test object, and mean absolute error was 0.037 mm. Animating 3D bone models from sets of marker positions (XROMM animation) makes it possible to study skeletal kinematics in the context of detailed bone morphology. The biplanar fluoroscopy hardware and computational methods described here should make XROMM an accessible and useful addition to the available technologies for studying the form, function, and evolution of vertebrate animals.

Despite recent advances in aerodynamic, neuromuscular and kinematic aspects of avian flight and dozens of relevant fossil discoveries, the origin of aerial locomotion and the transition from limbs to wings continue to be debated. Interpreting this transition depends on understanding the mechanical interplay of forces in living birds, particularly at the shoulder where most wing motion takes place. Shoulder function depends on a balance of forces from muscles, ligaments and articular cartilages, as well as inertial, gravitational and aerodynamic loads on the wing. Here we show that the force balance system of the shoulder evolved from a primarily muscular mechanism to one in which the acrocoracohumeral ligament has a critical role. Features of the shoulder of Mesozoic birds and closely related theropod dinosaurs indicate that the evolution of flight preceded the acquisition of the ligament-based force balance system and that some basal birds are intermediate in shoulder morphology.

Avian striding bipedalism was studied in the helmeted guineafowl, Numida meleagris. High-speed cineradiographs, light films, and videos were used to record hind limb movements across a wide range of speeds. In particular, direct visualization of the skeleton in X-ray images allowed changes in pelvic and femoral position to be quantified with great accuracy for the first time. With the exception of limb protraction angle, all stride parameters are speed-dependent. During the stance phase, guineafowl primarily employ knee flexion at very low speeds. At higher speeds, the magnitudes of hip and knee extension in the second half of stance progressively increase. Pelvic rotations are relatively small, but birds gradually pitch further forward with speed. An aerial phase is not present at speeds less than 2.0 m/sec, but discontinuities in the relationship of some parameters to speed indicate a gait transition near 0.9 m/sec. Birds are considered to be flying theropod dinosaurs, making characterization of bipedalism in living birds essential to understanding the evolution of theropod locomotion. Data from guineafowl, including the kinematic effects of speed, are informative about several aspects of locomotion in extinct theropods. However, many details of avian bipedalism evolved only within a subset of Theropoda, and are therefore not directly applicable to all members of the clade. J. Morphol. 240:115-125, 1999. © 1999 Wiley-Liss, Inc., (Copyright © 1999 Wiley-Liss, Inc.)

The neuromuscular control of the hind limb of helmeted guineafowl (Numida meleagris) locomoting on a treadmill at 1.0 m/sec was analyzed using simultaneous electromyography (EMG) and cineradiography. Activity from 16 heads representing 14 hip and knee muscles was recorded and correlated with limb movement and myological data to help discern muscle function. The first half of the stance phase is characterized by activity in many hip extensors, which counteract a flexor moment of the ground reaction force to yield hip stability. Simultaneously, medial rotators of the femur mediate pelvic roll and coactive antagonists about the knee control knee flexion of ca. 60°. Later in stance, hip extensors pull the hip through an arc of ca. 25°; knee extension occurs in some strides. N. meleagris hind limb motor patterns were compared to those of their homologs in representative lizards and crocodilians. Using a cladogram of living saurians, motor patterns were reconstructed in hypothetical ancestors. Although data are limited, lizards appear to have very conservative muscle activity similar to that of the ancestral saurian. The extant crocodilian Alligator mississippiensis resembles the reconstructed ancestor of Archosauria in at least 9 of 11 hind limb motor patterns. In contrast, N. meleagris differs from this same ancestor in at least four muscles. Most novelties in extant saurian motor patterns arose on the line to living birds. J. Morphol. 240:127-142, 1999. © 1999 Wiley-Liss, Inc., (Copyright © 1999 Wiley-Liss, Inc.)

The neuromuscular control of the hindlimb of American alligators (Alligator mississippiensis) walking on a treadmill was analyzed using simultaneous electromyography (EMG) and cineradiography. EMG and kinematic data were integrated with myological information to discern the interplay of muscles mediating hip and knee movement during the high walk. Twelve muscles, subdivided into 23 individual heads, cross the hip joint of Alligator. Activity patterns of 12 heads of 11 hip muscles and one knee muscle were recorded and quantified. An additional five heads from four muscles were recorded in single individuals. During the stance phase, the caudofemoralis longus prevents hip flexion and actively shortens to retract the femur through an arc of 60-80°. At the same time, the adductor femoris 1 and pubo-ischio-tibialis control femoral abduction. The knee is extended 30-40° during stance by contraction of the femoro-tibialis internus. These stance phase muscles often produce discontinuous, periodic EMG signals within their normal burst profile. In late stance and early swing, the ilio-fibularis and the pubo-ischio-tibialis are responsible for flexing the knee. The limb is protracted by the pubo-ischio-femoralis internus 2 and pubo-ischio-femoralis externus 2, which flex the hip. The ilio-femoralis abducts the limb during swing to suspend it above the tread. The role of the ambiens 1, which is active in midswing, is unclear. The ilio-tibialis 2, flexor-tibialis externus and flexor-tibialis internus 2 yield sporadic, low amplitude EMGs; these muscles are recruited at a very low level, if at all, during the slow high walk. Although EMGs do not conclusively delineate muscle function, activity patterns are particularly helpful in elucidating the complex interaction of muscular heads in this system. J. Morphol. 234:197-212, 1997. © 1997 Wiley-Liss, Inc., (Copyright © 1997 Wiley-Liss, Inc.)

Isolated teeth referred to the family Haramiyidae are among the earliest known fossil evidence of mammals. First discovered in European Late Triassic deposits a century and a half ago, haramiyids have been interpreted as related to multituberculates, a diverse and widespread lineage that occupied a rodent-like niche from the Late Jurassic to the Early Tertiary. Nonetheless, haramiyid relationships have remained enigmatic because the orientation and position of the teeth in the upper or lower jaw could not be determined with certainty; even their mammalian status has been questioned. The discovery of haramiyid dentaries, a maxilla and other skeletal remains in the Upper Triassic of East Greenland reveals haramiyids as highly specialized mammals with a novel pattern of puncture-crushing occlusion that differs dramatically from the grinding or shearing mechanisms of other Early Mesozoic mammals.

Modern birds have extremely short tail skeletons relative to Archaeopteryx and nonavialian theropod dinosaurs. Long- and short-tailed birds also differ in the conformation of main tail feathers making up the flight surface: frond shaped in Archaeopteryx and fan shaped in extant fliers. Mechanisms of tail fanning were evaluated by electromyography in freely flying pigeons and turkeys and by electrical stimulation of caudal muscles in anesthetized birds. Results from these experiments reveal that the pygostyle, rectrices, rectricial bulbs, and bulbi rectricium musculature form a specialized fanning mechanism. Contrary to previous models, our data support the interpretation that the bulbi rectricium independently controls tail fanning; other muscles are neither capable of nor necessary for significant rectricial abduction. This bulb mechanism permits rapid changes in tail span, thereby allowing the exploitation of a wide range of lift forces. Isolation of the bulbs on the pygostyle effectively decouples tail fanning from fan movement, which is governed by the remaining caudal muscles. The tail of Archaeopteryx, however, differs from this arrangement in several important respects. Archaeopteryx probably had a limited range of lift forces and tight coupling between vertebral and rectricial movement. This would have made the tail of this primitive flier better suited to stabilization than maneuverability. The capacity to significantly alter lift and manipulate the flight surface without distortion may have been two factors favoring tail shortening and pygostyle development during avian evolution., (© 1996 The Society for the Study of Evolution.)

The evolution of avian flight can be interpreted by analyzing the sequence of modifications of the primitive tetrapod locomotor system through time. Herein, we introduce the term "locomotor module" to identify anatomical subregions of the musculoskeletal system that are highly integrated and act as functional units during locomotion. The first tetrapods, which employed lateral undulations of the entire body and appendages, had one large locomotor module. Basal dinosaurs and theropods were bipedal and possessed a smaller locomotor module consisting of the hind limb and tail. Bird flight evolved as the superimposition of a second (aerial) locomotor capability onto the ancestral (terrestrial) theropod body plan. Although the origin of the wing module was the primary innovation, alterations in the terrestrial system were also significant. We propose that the primitive theropod locomotor module was functionally and anatomically subdivided into separate pelvic and caudal locomotor modules. This decoupling freed the tail to attain a new and intimate affiliation with the forelimb during flight, a configuration unique to birds. Thus, the evolution of flight can be viewed as the origin and novel association of locomotor modules. Differential elaboration of these modules in various lineages has produced the diverse locomotor abilities of modern birds., (© 1996 The Society for the Study of Evolution.)

The living members of the clade Archosauria, crocodilians and birds, differ markedly in the morphology of their deep dorsal thigh muscles. To investigate whether this diversity is accompanied by differences in motor pattern and muscle function, the hindlimbs of representative archosaurs were studied by electromyography and cineradiography during terrestrial locomotion. In a crocodilian, Alligator, the iliofemoralis and pubo-ischio-femoralis internus part 2 are both active during the swing phase of the stride cycle. This appears to be the primitive motor pattern for archosaurs. There are four avian homologues of these muscles in the helmeted guineafowl, Numida. These are primarily active in the propulsive phase (iliotrochantericus caudalis and iliotrochantericus medius), the swing phase (iliotrochantericus cranialis) and a speed-dependent combination of the propulsive and/or swing phases (iliofemoralis externus). Differences between Alligator and Numida in the number and attachment of deep dorsal muscles are associated with dissimilar motor patterns and functions. Evolutionary modifications of neuromuscular control must be recognized when evaluating avian locomotor history, but are rarely considered by paleontologists. Even within the deep dorsal thigh muscles of Numida, developmentally and anatomically similar muscles are active out-of-phase. Therefore, although the actions of two adjacent muscles appear equivalent, their functions may differ dramatically. The diversity of deep dorsal thigh muscles in modern birds may be a good model for studying the relationship between activity pattern and peripheral morphology.

In adult rats, each neuromuscular compartment of the lateral gastrocnemius muscle (LG) is exclusively innervated by a primary branch of the LG nerve. In neonates, however, a small percentage of LG cells receives inputs from more than one primary nerve branch; these inputs are known as cross-compartmental. Cross-compartmental inputs are normally lost from the medial compartment of LG (LGm) by the 8th postnatal day. To investigate the mechanisms involved in the elimination of cross-compartmental inputs, muscle fibers in the LGm compartment were denervated by cutting the LGm nerve branch in 1-4 day old rat pups and in adult rats. We then assessed the degree of cross-compartmental innervation within the "denervated" compartment using intracellular recordings from neonatal muscle fibers or immunohistochemical staining for nerve cell adhesion molecule (N-CAM) and neurofilament protein in adult muscles. Following LGm axotomy in neonates, cross-compartmental innervation is more extensive than in controls and is present as late as 20 days after birth. Thus, in the absence of "native" LGm axons, neonatal cross-compartmental inputs proliferate by axonal sprouting and the formation of new synapses on vacant LGm fibers. In contrast, axotomized adults do not form new cross-compartmental inputs over the same time period. The differential response of neonates and adults to muscle nerve branch denervation is evidence for the existence of some form of compartment-specific recognition. We propose that compartmental identity either arises or becomes relatively more potent during ontogeny and normally acts selectively to eliminate foreign axons and deter the formation of new cross-compartmental inputs.

An analysis of hind limb skeletal elements of non-avian theropods and ground-dwelling birds was performed to reveal patterns of change in shape and proportion with size. When femora of equal length are compared, birds exhibit a significantly larger midshaft diameter than non-avian theropods. As total limb length increases, avian femora become relatively shorter (negative allometry), while those of non-avian theropods become relatively longer (positive allometry). Avian femoral/tibiotarsal ratios are all below 0.8 and decrease with limb size, whereas ratios of non-avian theropods are well above 0.8 and tend to increase with limb size. In addition, avian femora exhibit a unique diameter/length relationship not seen in other theropod hind limb bones. Several studies have shown that within the avian limb, the short, robust femur resists bending to a far greater degree than the relatively longer, slimmer tibiotarsus. This is to be expected, as analyses of running birds show that the femur is oriented relatively perpendicular to the ground reaction force throughout the stride, which would subject it to high bending moments. When compared to birds, non-avian theropods have relatively long, slender femora that do not seem to be built to withstand the forces associated with such an orientation. Reconstructing all non-avian theropods in avian-like poses (subhorizontal femur, knee well flexed) with avian locomotor kinematics (relatively little hip extension at most speeds) ignores major differences in scaling between these groups of organisms., (Copyright © 1991 Wiley-Liss, Inc.)