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Common hippopotamuses (hippos) are among the largest extant land mammals. They thus offer potential further insight into how giant body size on land influences locomotor patterns and abilities. Furthermore, as they have semi-aquatic habits and unusual morphology, they prompt important questions about how locomotion evolved in Hippopotamidae. However, basic information about how hippos move is limited and sometimes contradictory. We aimed to test if hippos trot at all speeds and if they ever use an aerial (suspended) phase, and to quantify how their locomotor patterns (footfalls and stride parameters) change with approximate speed. We surveyed videos available online and collected new video data from two zoo hippos in order to calculate the data needed to achieve our aims; gathering a sample of 169 strides from 32 hippos. No hippos studied used other than trotting (or near-trotting) footfall patterns, but at the fastest relative speeds hippos used brief aerial phases, apparently a new discovery. Hippos exhibit relatively greater athletic capacity than elephants in several ways, but perhaps not greater than rhinoceroses. Our data help form a baseline for assessing if other hippos use normal locomotion; relevant to clinical veterinary assessments of lameness; and for reconstructing the evolutionary biomechanics of hippo lineages., Competing Interests: John Hutchinson is an Academic Editor for PeerJ., (© 2024 Hutchinson and Pringle.)

Here, we review the modern interface of three-dimensional (3D) empirical (e.g. motion capture) and theoretical (e.g. modelling and simulation) approaches to the study of terrestrial locomotion using appendages in tetrapod vertebrates. These tools span a spectrum from more empirical approaches such as XROMM, to potentially more intermediate approaches such as finite element analysis, to more theoretical approaches such as dynamic musculoskeletal simulations or conceptual models. These methods have much in common beyond the importance of 3D digital technologies, and are powerfully synergistic when integrated, opening a wide range of hypotheses that can be tested. We discuss the pitfalls and challenges of these 3D methods, leading to consideration of the problems and potential in their current and future usage. The tools (hardware and software) and approaches (e.g. methods for using hardware and software) in the 3D analysis of tetrapod locomotion have matured to the point where now we can use this integration to answer questions we could never have tackled 20 years ago, and apply insights gleaned from them to other fields., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2023. Published by The Company of Biologists Ltd.)

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., (© 2021 The Authors. Journal of Anatomy published by John Wiley & Sons Ltd on behalf of Anatomical Society.)

The arrangement and physiology of muscle fibres can strongly influence musculoskeletal function and whole-organismal performance. However, experimental investigation of muscle function during in vivo activity is typically limited to relatively few muscles in a given system. Computational models and simulations of the musculoskeletal system can partly overcome these limitations, by exploring the dynamics of muscles, tendons and other tissues in a robust and quantitative fashion. Here, a high-fidelity, 26-degree-of-freedom musculoskeletal model was developed of the hindlimb of a small ground bird, the elegant-crested tinamou (Eudromia elegans, ~550 g), including all the major muscles of the limb (36 actuators per leg). The model was integrated with biplanar fluoroscopy (XROMM) and forceplate data for walking and running, where dynamic optimization was used to estimate muscle excitations and fibre length changes throughout both gaits. Following this, a series of static simulations over the total range of physiological limb postures were performed, to circumscribe the bounds of possible variation in fibre length. During gait, fibre lengths for all muscles remained between 0.5 to 1.21 times optimal fibre length, but operated mostly on the ascending limb and plateau of the active force-length curve, a result that parallels previous experimental findings for birds, humans and other species. However, the ranges of fibre length varied considerably among individual muscles, especially when considered across the total possible range of joint excursion. Net length change of muscle-tendon units was mostly less than optimal fibre length, sometimes markedly so, suggesting that approaches that use muscle-tendon length change to estimate optimal fibre length in extinct species are likely underestimating this important parameter for many muscles. The results of this study clarify and broaden understanding of muscle function in extant animals, and can help refine approaches used to study extinct species., Competing Interests: The authors have declared that no competing interests exist.

Extant Crocodylia are exceptional because they employ almost the full range of quadrupedal footfall patterns ("gaits") used by mammals; including asymmetrical gaits such as galloping and bounding. Perhaps this capacity evolved in stem Crocodylomorpha, during the Triassic when taxa were smaller, terrestrial, and long-legged. However, confusion about which Crocodylia use asymmetrical gaits and why persists, impeding reconstructions of locomotor evolution. Our experimental gait analysis of locomotor kinematics across 42 individuals from 15 species of Crocodylia obtained 184 data points for a wide velocity range (0.15-4.35 ms -1 ). Our results suggest either that asymmetrical gaits are ancestral for Crocodylia and lost in the alligator lineage, or that asymmetrical gaits evolved within Crocodylia at the base of the crocodile line. Regardless, we recorded usage of asymmetrical gaits in 7 species of Crocodyloidea (crocodiles); including novel documentation of these behaviours in 5 species (3 critically endangered). Larger Crocodylia use relatively less extreme gait kinematics consistent with steeply decreasing athletic ability with size. We found differences between asymmetrical and symmetrical gaits in Crocodylia: asymmetrical gaits involved greater size-normalized stride frequencies and smaller duty factors (relative ground contact times), consistent with increased mechanical demands. Remarkably, these gaits did not differ in maximal velocities obtained: whether in Alligatoroidea or Crocodyloidea, trotting or bounding achieved similar velocities, revealing that the alligator lineage is capable of hitherto unappreciated extreme locomotor performance despite a lack of asymmetrical gait usage. Hence asymmetrical gaits have benefits other than velocity capacity that explain their prevalence in Crocodyloidea and absence in Alligatoroidea-and their broader evolution.

Ontogenetic information is crucial to understand life histories and represents a true challenge in dinosaurs due to the scarcity of growth series available. Mussaurus patagonicus was a sauropodomorph dinosaur close to the origin of Sauropoda known from hatchling, juvenile and mature specimens, providing a sufficiently complete ontogenetic series to reconstruct general patterns of ontogeny. Here, in order to quantify how body shape and its relationship with locomotor stance (quadruped/biped) changed in ontogeny, hatchling, juvenile (~1 year old) and adult (8+ years old) individuals were studied using digital models. Our results show that Mussaurus rapidly grew from about 60 g at hatching to ~7 kg at one year old, reaching >1000 kg at adulthood. During this time, the body's centre of mass moved from a position in the mid-thorax to a more caudal position nearer to the pelvis. We infer that these changes of body shape and centre of mass reflect a shift from quadrupedalism to bipedalism occurred early in ontogeny in Mussaurus. Our study indicates that relative development of the tail and neck was more influential in determining the locomotor stance in Sauropodomorpha during ontogeny, challenging previous studies, which have emphasized the influence of hindlimb vs. forelimb lengths on sauropodomorph stance.

Background: Morphological diversity of limb bone lengths, diameters, and proportions in mammals is known to vary strongly with locomotor habit. It remains less well known how different locomotor habits are correlated with cross-sectional traits of the limb skeleton, such as cross-sectional area (CSA), second moments of area (SMA), and section modulus (MOD) and whether these traits have evolved adaptively. CSA and SMA represent the bone's resistance to axial compression and bending, respectively, whereas MOD represents bone structural strength related to shape. Sampling 28 species of mustelids, a carnivoran lineage with diverse locomotor habits, we tested for differences in humeral, radial, and ulnar cross-sectional traits among specialists for climbing, digging, and swimming, in addition to generalists. Given that the limbs of digging specialists function in the dense substance of soil, and that swimming specialists need to counteract buoyancy, we predicted that these mustelids with these specializations should have the greatest values of cross-sectional traits., Results: We analyzed cross-sectional traits (calculated via μCT scanning and rendered dimensionless) in 5% increments along a bone's length and found significant differences among locomotor habits, though differences in ulnar cross-sectional traits were fewer than those for the humerus and radius. Swimming specialists had the greatest values of cross-sectional traits, followed by digging specialists. Climbing specialists had the lowest values of cross-sectional traits. However, phylogenetic affinity underlies these results. Fitting models of trait evolution to CSA and SMA revealed that a multi-rate Brownian motion model and a multi-optima Ornstein-Uhlenbeck model are the best-fitting models of evolution for these traits. However, inspection of α-values uncovered that many of the OU models did not differ from a Brownian motion model., Conclusions: Within Mustelidae, differences in limb function and locomotor habit influence cross-sectional traits in ways that produce patterns that may diverge from adaptive patterns exhibited by external traits (e.g., bone lengths) of the mammalian limb skeleton. These results suggest that not all the traits of a single organ evolve under a single evolutionary process and that models of trait evolution should be fit to a range of traits for a better understanding of the evolution of the mammalian locomotor system.

Giraffes ( Giraffa camelopardalis ) possess specialised anatomy. Their disproportionately elongate limbs and neck confer recognised feeding advantages, but little is known about how their morphology affects locomotor function. In this study, we examined the stride parameters and ground reaction forces from three adult giraffes in a zoological park, across a range of walking speeds. The patterns of GRFs during walking indicate that giraffes, similar to other mammalian quadrupeds, maintain a forelimb-biased weight distribution. The angular excursion of the neck has functional links with locomotor dynamics in giraffes, and was exaggerated at faster speeds. The horizontal accelerations of the neck and trunk were out of phase compared with the vertical accelerations, which were intermediate between in and out of phase. Despite possessing specialised morphology, giraffes' stride parameters were broadly predicted from dynamic similarity, facilitating the use of other quadrupedal locomotion models to generate testable hypotheses in giraffes., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2019. Published by The Company of Biologists Ltd.)

Reconstructing the locomotion of extinct vertebrates offers insights into their palaeobiology and helps to conceptualize major transitions in vertebrate evolution 1-4 . However, estimating the locomotor behaviour of a fossil species remains a challenge because of the limited information preserved and the lack of a direct correspondence between form and function 5,6 . The evolution of advanced locomotion on land-that is, locomotion that is more erect, balanced and mechanically power-saving than is assumed of anamniote early tetrapods-has previously been linked to the terrestrialization and diversification of amniote lineages 7 . To our knowledge, no reconstructions of the locomotor characteristics of stem amniotes based on multiple quantitative methods have previously been attempted: previous methods have relied on anatomical features alone, ambiguous locomotor information preserved in ichnofossils or unspecific modelling of locomotor dynamics. Here we quantitatively examine plausible gaits of the stem amniote Orobates pabsti, a species that is known from a complete body fossil preserved in association with trackways 8 . We reconstruct likely gaits that match the footprints, and investigate whether Orobates exhibited locomotor characteristics that have previously been linked to the diversification of crown amniotes. Our integrative methodology uses constraints derived from biomechanically relevant metrics, which also apply to extant tetrapods. The framework uses in vivo assessment of locomotor mechanics in four extant species to guide an anatomically informed kinematic simulation of Orobates, as well as dynamic simulations and robotics to filter the parameter space for plausible gaits. The approach was validated using two extant species that have different morphologies, gaits and footprints. Our metrics indicate that Orobates exhibited more advanced locomotion than has previously been assumed for earlier tetrapods 7,9 , which suggests that advanced terrestrial locomotion preceded the diversification of crown amniotes. We provide an accompanying website for the exploration of the filters that constrain our simulations, which will allow revision of our approach using new data, assumptions or methods.

Brittle stars (Phylum Echinodermata, Class Ophiuroidea) have evolved rapid locomotion employing muscle and skeletal elements within their (usually) five arms to apply forces in a manner analogous to that of vertebrates. Inferring the inner workings of the arm has been difficult as the skeleton is internal and many of the ossicles are sub-millimeter in size. Advances in 3D visualization and technology have made the study of movement in ophiuroids possible. We developed six virtual 3D skeletal models to demonstrate the potential range of motion of the main arm ossicles, known as vertebrae, and six virtual 3D skeletal models of non-vertebral ossicles. These models revealed the joint center and relative position of the arm ossicles during near-maximal range of motion. The models also provide a platform for the comparative evaluation of functional capabilities between disparate ophiuroid arm morphologies. We made observations on specimens of Ophioderma brevispina and Ophiothrix angulata. As these two taxa exemplify two major morphological categories of ophiuroid vertebrae, they provide a basis for an initial assessment of the functional consequences of these disparate vertebral morphologies. These models suggest potential differences in the structure of the intervertebral articulations in these two species, implying disparities in arm flexion mechanics. We also evaluated the differences in the range of motion between segments in the proximal and distal halves of the arm length in a specimen of O. brevispina, and found that the morphology of vertebrae in the distal portion of the arm allows for higher mobility than in the proximal portion. Our models of non-vertebral ossicles show that they rotate further in the direction of movement than the vertebrae themselves in order to accommodate arm flexion. These findings raise doubts over previous hypotheses regarding the functional consequences of ophiuroid arm disparity. Our study demonstrates the value of integrating experimental data and visualization of articulated structures when making functional interpretations instead of relying on observations of vertebral or segmental morphology alone. This methodological framework can be applied to other ophiuroid taxa to enable comparative functional analyses. It will also facilitate biomechanical analyses of other invertebrate groups to illuminate how appendage or locomotor function evolved., (© 2018 Anatomical Society.)

Broiler chickens are increasingly at the forefront of global meat production but the consequences of fast growth and selection for an increase in body mass on bird health are an ongoing concern for industry and consumers. To better understand the implications of selection we evaluated energetics and behaviour over the 6-week hatch-to-slaughter developmental period in a commercial broiler. The effect of posture on resting metabolic rate becomes increasingly significant as broilers grow, as standing became more energetically expensive than sitting. The proportion of overall metabolic rate accounted for by locomotor behaviour decreased over development, corresponding to declining activity levels, mean and peak walking speeds. These data are consistent with the inference that broilers allocate energy to activity within a constrained metabolic budget and that there is a reducing metabolic scope for exercise throughout their development. Comparison with similarly sized galliforms reveals that locomotion is relatively energetically expensive in broilers.

Centre of mass is a fundamental anatomical and biomechanical parameter. Knowledge of centre of mass is essential to inform studies investigating locomotion and other behaviours, through its implications for segment movements, and on whole body factors such as posture. Previous studies have estimated centre of mass position for a range of organisms, using various methodologies. However, few studies assess the accuracy of the methods that they employ, and often provide only brief details on their methodologies. As such, no rigorous, detailed comparisons of accuracy and repeatability within and between methods currently exist. This paper therefore seeks to apply three methods common in the literature (suspension, scales and digital modelling) to three 'calibration objects' in the form of bricks, as well as three birds to determine centre of mass position. Application to bricks enables conclusions to be drawn on the absolute accuracy of each method, in addition to comparing these results to assess the relative value of these methodologies. Application to birds provided insights into the logistical challenges of applying these methods to biological specimens. For bricks, we found that, provided appropriate repeats were conducted, the scales method yielded the most accurate predictions of centre of mass (within 1.49 mm), closely followed by digital modelling (within 2.39 mm), with results from suspension being the most distant (within 38.5 mm). Scales and digital methods both also displayed low variability between centre of mass estimates, suggesting they can accurately and consistently predict centre of mass position. Our suspension method resulted not only in high margins of error, but also substantial variability, highlighting problems with this method., (© 2017 The Authors. Journal of Anatomy published by John Wiley & Sons Ltd on behalf of Anatomical Society.)

Despite their semi-aquatic mode of life, modern crocodylians use a wide range of terrestrial locomotor behaviours, including asymmetrical gaits otherwise only found in mammals. The key to these diverse abilities may lie in the axial skeleton. Correlations between vertebral morphology and both intervertebral joint stiffness and locomotor behaviour have been found in other animals, but the vertebral mechanics of crocodylians have not yet been experimentally and quantitatively tested. We measured the passive mechanics and morphology of the thoracolumbar vertebral column in Crocodylus niloticus in order to validate a method to infer intervertebral joint stiffness based on morphology. Passive stiffness of eight thoracic and lumbar joints was tested in dorsal extension, ventral flexion and mediolateral flexion using cadaveric specimens. Fifteen measurements that we deemed to be potential correlates of stiffness were taken from each vertebra and statistically tested for correlation with joint stiffness. We found that the vertebral column of C. niloticus is stiffer in dorsoventral flexion than in lateral flexion and, in contrast to that of many mammals, shows an increase in joint stiffness in the lumbar region. Our findings suggest that the role of the axial column in crocodylian locomotion may be functionally different from that in mammals, even during analogous gaits. A moderate proportion of variation in joint stiffness (R(2)=0.279-0.520) was predicted by centrum width and height, neural spine angle and lamina width. These results support the possible utility of some vertebral morphometrics in predicting mechanical properties of the vertebral column in crocodiles, which also should be useful for forming functional hypotheses of axial motion during locomotion in extinct archosaurs.

Over the past century, various modern analogs have been used to infer the evolution of locomotor performance in stem tetrapods and their fish ancestors, with varying success. Here, we conduct a phylogenetic review of these modern analogs, from chondrichthyans to mammals, highlighting the broad spectrum of vertebrate clades and locomotor behaviors. The pros and cons behind utilizing modern analogs for the early stages of the transition from water to land also are discussed. In particular, it is noted that any hypothesis about locomotion not only must be supported by evidence from living animals but must also be consistent with character transformations in the fossil record. A "total-evidence" approach that emphasizes what extinct taxa could not do, rather than focusing on the specifics of how they functioned, is thus recommended. An example of this approach, which investigates mobility of the limb joints in modern semi-aquatic animals and in the Devonian stem tetrapod Ichthyostega, is detailed. We propose that various locomotion behaviors of modern quadrupeds can be ruled out for Ichthyostega, but that forelimb "crutching" motions, as seen in living mudskippers, may have been possible. The potential for movement in other known Devonian stem tetrapods is assessed through an anatomical comparison of limb joint morphology-and associated mobility-with Ichthyostega, and deemed to have been quite similar.

Locomotion in living birds (Neornithes) has two remarkable features: feather-assisted flight, and the use of unusually crouched hindlimbs for bipedal support and movement. When and how these defining functional traits evolved remains controversial. However, the advent of computer modelling approaches and the discoveries of exceptionally preserved key specimens now make it possible to use quantitative data on whole-body morphology to address the biomechanics underlying this issue. Here we use digital body reconstructions to quantify evolutionary trends in locomotor biomechanics (whole-body proportions and centre-of-mass position) across the clade Archosauria. We use three-dimensional digital reconstruction to estimate body shape from skeletal dimensions for 17 archosaurs along the ancestral bird line, including the exceptionally preserved, feathered taxa Microraptor, Archaeopteryx, Pengornis and Yixianornis, which represent key stages in the evolution of the avian body plan. Rather than a discrete transition from more-upright postures in the basal-most birds (Avialae) and their immediate outgroup deinonychosauria, our results support hypotheses of a gradual, stepwise acquisition of more-crouched limb postures across much of theropod evolution, although we find evidence of an accelerated change within the clade Maniraptora (birds and their closest relatives, such as deinonychosaurs). In addition, whereas reduction of the tail is widely accepted to be the primary morphological factor correlated with centre-of-mass position and, hence, evolution of hindlimb posture, we instead find that enlargement of the pectoral limb and several associated trends have a much stronger influence. Intriguingly, our support for the onset of accelerated morpho-functional trends within Maniraptora is closely correlated with the evolution of flight. Because we find that the evolution of enlarged forelimbs is strongly linked, via whole-body centre of mass, to hindlimb function during terrestrial locomotion, we suggest that the evolution of avian flight is linked to anatomical novelties in the pelvic limb as well as the pectoral.

Crocodylians exhibit a fascinating diversity of terrestrial gaits and limb motions that remain poorly described and are of great importance to understanding their natural history and evolution. Their musculoskeletal anatomy is pivotal to this diversity and yet only qualitative studies of muscle-tendon unit anatomy exist. The relative masses and internal architecture (fascicle lengths and physiological cross-sectional areas) of muscles of the pectoral and pelvic limbs of American alligators (Alligator mississippiensis Daudin 1801) were recorded for an ontogenetic series of wild specimens (n = 15, body masses from 0.5 to 60 kg). The data were analysed by reduced major axis regression to determine scaling relationships with body mass. Physiological cross-sectional areas and therefore muscle force-generating capacity were found to be greater in the extensor (anti-gravity) muscles of the pelvic limb than in the pectoral limb, reflecting how crocodylians differ from mammals in having greater loading of the hindlimbs than the forelimbs. Muscle masses and architecture were generally found to scale isometrically with body mass, suggesting an ontogenetic decrease in terrestrial athleticism. This concurs with the findings of previous studies showing ontogenetic decreases in limb bone length and the general scaling principle of a decline of strength : weight ratios with increasing size in animals. Exceptions to isometric scaling found included positive allometry in fascicle length for extensor musculature of both limbs, suggesting an ontogenetic increase in working range interpreted as increasing postural variability - in particular the major hip extensors - the interpretation of which is complicated by previous described ontogenetic increase of moment arms for these muscles.

As the largest extant terrestrial animals, elephants do not trot or gallop but can move smoothly to faster speeds without markedly changing their kinematics, yet with a shift from vaulting to bouncing kinetics. To understand this unusual mechanism, we quantified the forelimb and hindlimb motions of eight Asian elephants (Elephas maximus) and seven African elephants (Loxodonta africana). We used 240 Hz motion analysis (tracking 10 joint markers) to measure the flexion/extension angles and angular velocities of the limb segments and joints for 288 strides across an eightfold range of speeds (0.6-4.9 m s(-1)) and a sevenfold range of body mass (521-3684 kg). We show that the columnar limb orientation that elephants supposedly exemplify is an oversimplification--few segments or joints are extremely vertical during weight support (especially at faster speeds), and joint flexion during the swing phase is considerable. The 'inflexible' ankle is shown to have potentially spring-like motion, unlike the highly flexible wrist, which ironically is more static during support. Elephants use approximately 31-77% of their maximal joint ranges of motion during rapid locomotion, with this fraction increasing distally in the limbs, a trend observed in some other running animals. All angular velocities decrease with increasing size, whereas smaller elephant limbs are not markedly more flexed than adults. We find no major quantitative differences between African and Asian elephant locomotion but show that elephant limb motions are more similar to those of smaller animals, including humans and horses, than commonly recognized. Such similarities have been obscured by the reliance on the term ;columnar' to differentiate elephant limb posture from that of other animals. Our database will be helpful for identifying elephants with unusual limb movements, facilitating early recognition of musculoskeletal pathology.

For centuries, elephant locomotion has been a contentious and confusing challenge for locomotion scientists to understand, not only because of technical difficulties but also because elephant locomotion is in some ways atypical of more familiar quadrupedal gaits. We analyzed the locomotor kinematics of over 2400 strides from 14 African and 48 Asian elephant individuals (body mass 116-4632 kg) freely moving over ground at a 17-fold range of speeds, from slow walking at 0.40 m s(-1) to the fastest reliably recorded speed for elephants, 6.8 m s(-1). These data reveal that African and Asian elephants have some subtle differences in how size-independent kinematic parameters change with speed. Although elephants use a lateral sequence footfall pattern, like many other quadrupeds, they maintain this footfall pattern at all speeds, shifting toward a 25% phase offset between limbs (singlefoot) as they increase speed. The duty factors of elephants are greater for the forelimbs than for the hindlimbs, so an aerial phase for the hindquarters is reached at slower speeds than for the forequarters. This aerial phase occurs at a Froude number of around 1, matching theoretical predictions. At faster speeds, stance and swing phase durations approach asymptotes, with the duty factor beginning to level off, concurrent with an increase in limb compliance that likely keeps peak forces relatively low. This increase of limb compliance is reflected by increased compression of the hindlimbs. Like other tetrapods, smaller elephants are relatively more athletic than larger ones, but still move very similarly to adults even at <500 kg. At any particular speed they adopt greater relative stride frequencies and relative stride lengths compared to larger elephants. This extends to near-maximal locomotor performance as well - smaller elephants reach greater Froude numbers and smaller duty factors, hence likely reach relatively greater peak loads on their limbs and produce this force more rapidly. A variety of lines of kinematic evidence support the inference that elephants change their mechanics near a Froude number of 1 (if not at slower speeds), at least to using more compliant limbs, if not spring-like whole-body kinetics. In some ways, elephants move similarly to many other quadrupeds, such as increasing speed mainly by increasing stride frequency (except at fast speeds), and they match scaling predictions for many stride parameters. The main difference from most other animals is that elephants never change their footfall pattern to a gait that uses a whole-body aerial phase. Our large dataset establishes what the normal kinematics of elephant locomotion are, and can also be applied to identify gait abnormalities that may signal musculoskeletal pathologies, a matter of great importance to keepers of captive elephants.

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

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

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

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

Saurischian dinosaurs evolved seven orders of magnitude in body mass, as well as a wide diversity of hip joint morphology and locomotor postures. The very largest saurischians possess incongruent bony hip joints, suggesting that large volumes of soft tissues mediated hip articulation. To understand the evolutionary trends and functional relationships between body size and hip anatomy of saurischians, we tested the relationships among discrete and continuous morphological characters using phylogenetically corrected regression. Giant theropods and sauropods convergently evolved highly cartilaginous hip joints by reducing supraacetabular ossifications, a condition unlike that in early dinosauromorphs. However, transitions in femoral and acetabular soft tissues indicate that large sauropods and theropods built their hip joints in fundamentally different ways. In sauropods, the femoral head possesses irregularly rugose subchondral surfaces for thick hyaline cartilage. Hip articulation was achieved primarily using the highly cartilaginous femoral head and the supraacetabular labrum on the acetabular ceiling. In contrast, theropods covered their femoral head and neck with thinner hyaline cartilage and maintained extensive articulation between the fibrocartilaginous femoral neck and the antitrochanter. These findings suggest that the hip joints of giant sauropods were built to sustain large compressive loads, whereas those of giant theropods experienced compression and shear forces. [ABSTRACT FROM AUTHOR]

The surface area of feet in contact with the ground is a key morphological feature that influences animal locomotion. Underfoot pressures (and consequently stresses experienced by the foot), as well as stability of an animal during locomotion, depend on the size and shape of this area. Here we tested whether the area of a skeletal foot could predict in vivo soft tissue foot surface area. Computed tomography scans of 29 extant tetrapods (covering mammals, reptiles, birds and amphibians) were used to produce models of both the soft tissues and the bones of their feet. Soft tissue models were oriented to a horizontal plane, and their outlines projected onto a surface to produce two‐dimensional silhouettes. Silhouettes of skeletal models were generated either from bones in CT pose or with all autopodial bones aligned to the horizontal plane. Areas of these projections were calculated using alpha shapes (mathematical tight‐fitting outline). Underfoot area of soft tissue was approximately 1.67 times that of skeletal tissue area (~ 2 times for manus, ~ 1.6 times for pes, if analysed separately). This relationship between skeletal foot area and soft tissue area, while variable in some of our study taxa, could provide information about the size of the organisms responsible for fossil trackways, suggest what size of tracks might be expected from potential trackmakers known only from skeletal remains, and aid in soft tissue reconstruction of skeletal remains for biomechanical modelling. [ABSTRACT FROM AUTHOR]

White rhinoceroses (Ceratotherium simum) are odd-toed ungulates that belong to the group Perissodactyla. Being second only to elephants in terms of large body mass amongst extant tetrapods, rhinoceroses make fascinating subjects for the study of how large land animals support and move themselves. Rhinoceroses often are kept in captivity for protection from ivory poachers and for educational/touristic purposes, yet a detrimental side effect of captivity can be foot disease (i.e., enthesopathies and osteoarthritis around the phalanges). Foot diseases in large mammals are multifactorial, but locomotor biomechanics (e.g., pressures routinely experienced by the feet) surely can be a contributing factor. However, due to a lack of in vivo experimental data on rhinoceros foot pressures, our knowledge of locomotor performance and its links to foot disease is limited. The overall aim of this study was to characterize peak pressures and center of pressure trajectories in white rhinoceroses during walking. We asked two major questions. First, are peak locomotor pressures the lowest around the fat pad and its lobes (as in the case of elephants)? Second, are peak locomotor pressures concentrated around the areas with the highest reported incidence of pathologies? Our results show a reduction of pressures around the fat pad and its lobes, which is potentially due to the material properties of the fat pad or a tendency to avoid or limit “heel” contact at impact. We also found an even and gradual concentration of foot pressures across all digits, which may be a by-product of the more horizontal foot roll-off during the stance phase. While our exploratory, descriptive sample precluded hypothesis testing, our study provides important new data on rhinoceros locomotion for future studies to build on, and thus impetus for improved implementation in the care of captive/managed rhinoceroses. [ABSTRACT FROM AUTHOR]

This paper is the second of a three-part series that investigates the architecture of cancellous bone in the main hindlimb bones of theropod dinosaurs, and uses cancellous bone architectural patterns to infer locomotor biomechanics in extinct non-avian species. Cancellous bone is widely known to be highly sensitive to its mechanical environment, and therefore has the potential to provide insight into locomotor biomechanics in extinct tetrapod vertebrates such as dinosaurs. Here in Part II, a new biomechanical modelling approach is outlined, one which mechanistically links cancellous bone architectural patterns with three-dimensional musculoskeletal and finite element modelling of the hindlimb. In particular, the architecture of cancellous bone is used to derive a single 'characteristic posture' for a given species--one in which bone continuum-level principal stresses best align with cancellous bone fabric--and thereby clarify hindlimb locomotor biomechanics. The quasi-static approach was validated for an extant theropod, the chicken, and is shown to provide a good estimate of limb posture at around mid-stance. It also provides reasonable predictions of bone loading mechanics, especially for the proximal hindlimb, and also provides a broadly accurate assessment of muscle recruitment insofar as limb stabilization is concerned. In addition to being useful for better understanding locomotor biomechanics in extant species, the approach hence provides a new avenue by which to analyse, test and refine palaeobiomechanical hypotheses, not just for extinct theropods, but potentially many other extinct tetrapod groups as well. [ABSTRACT FROM AUTHOR]

This paper is the last of a three-part series that investigates the architecture of cancellous bone in the main hindlimb bones of theropod dinosaurs, and uses cancellous bone architectural patterns to infer locomotor biomechanics in extinct non-avian species. Cancellous bone is highly sensitive to its prevailing mechanical environment, and may therefore help further understanding of locomotor biomechanics in extinct tetrapod vertebrates such as dinosaurs. Here in Part III, the biomechanical modelling approach derived previously was applied to two species of extinct, non-avian theropods, Daspletosaurus torosus and Troodon formosus. Observed cancellous bone architectural patterns were linked with quasi-static, three-dimensional musculoskeletal and finite element models of the hindlimb of both species, and used to derive characteristic postures that best aligned continuum-level principal stresses with cancellous bone fabric. The posture identified for Daspletosaurus was largely upright, with a subvertical femoral orientation, whilst that identified for Troodon was more crouched, but not to the degree observed in extant birds. In addition to providing new insight on posture and limb articulation, this study also tested previous hypotheses of limb bone loading mechanics and muscular control strategies in non-avian theropods, and how these aspects evolved on the line to birds. The results support the hypothesis that an upright femoral posture is correlated with bending-dominant bone loading and abduction-based muscular support of the hip, whereas a crouched femoral posture is correlated with torsion-dominant bone loading and long-axis rotation-based muscular support. Moreover, the results of this study also support the inference that hindlimb posture, bone loading mechanics and muscular support strategies evolved in a gradual fashion along the line to extant birds. [ABSTRACT FROM AUTHOR]

This paper is the first of a three-part series that investigates the architecture of cancellous ('spongy') bone in the main hindlimb bones of theropod dinosaurs, and uses cancellous bone architectural patterns to infer locomotor biomechanics in extinct non-avian species. Cancellous bone is widely known to be highly sensitive to its mechanical environment, and has previously been used to infer locomotor biomechanics in extinct tetrapod vertebrates, especially primates. Despite great promise, cancellous bone architecture has remained little utilized for investigating locomotion in many other extinct vertebrate groups, such as dinosaurs. Documentation and quantification of architectural patterns across a whole bone, and across multiple bones, can provide much information on cancellous bone architectural patterns and variation across species. Additionally, this also lends itself to analysis of the musculoskeletal biomechanical factors involved in a direct, mechanistic fashion. On this premise, computed tomographic and image analysis techniques were used to describe and analyse the three-dimensional architecture of cancellous bone in the main hindlimb bones of theropod dinosaurs for the first time. A comprehensive survey across many extant and extinct species is produced, identifying several patterns of similarity and contrast between groups. For instance, more stemward non-avian theropods (e.g. ceratosaurs and tyrannosaurids) exhibit cancellous bone architectures more comparable to that present in humans, whereas species more closely related to birds (e.g. paravians) exhibit architectural patterns bearing greater similarity to those of extant birds. Many of the observed patterns may be linked to particular aspects of locomotor biomechanics, such as the degree of hip or knee flexion during stance and gait. A further important observation is the abundance of markedly oblique trabeculae in the diaphyses of the femur and tibia of birds, which in large species produces spiralling patterns along the endosteal surface. Not only do these observations provide new insight into theropod anatomy and behaviour, they also provide the foundation for mechanistic testing of locomotor hypotheses via musculoskeletal biomechanical modelling. [ABSTRACT FROM AUTHOR]

Members of the mammalian family Felidae (extant and extinct cats) are grossly phenotypically similar, but display a 300-fold range in body size, from less than 1 kg to more than 300 kg. In addition to differences in body mass, felid species show dietary and locomotory specializations that correlate to skull and limb osteological measurements, such as shape or cross-sectional area. However, ecological correlates to the axial skeleton are yet untested. Here, we build on previous studies of the biomechanical and morphological evolution of the felid appendicular skeleton by conducting a quantitative analysis of morphology and allometry in the presacral vertebral column across extant cats. Our results demonstrate that vertebral columns of arboreal, scansorial and terrestrial felids significantly differ in morphology, specifically in the lumbar region, while no distinction based on dietary specialization was found. Body size significantly influences vertebral morphology, with clear regionalization of allometry along the vertebral column, suggesting that anterior (cervicals and thoracics) and posterior (lumbar) vertebrae may be independently subjected to distinct selection pressures. [ABSTRACT FROM AUTHOR]

In quadrupeds the musculature of the hindlimbs is expected to be responsible for generating most of the propulsive locomotory forces, as well as contributing to body support by generating vertical forces. In supporting the body, postural changes from crouched to upright limbs are often associated with an increase of body mass in terrestrial tetrapods. However, felids do not change their crouched limb posture despite undergoing a 300-fold size increase between the smallest and largest extant species. Here, we test how changes in the muscle architecture (masses and lengths of components of the muscle-tendon units) of the hindlimbs and lumbosacral region are related to body mass, to assess whether there are muscular compensations for the maintenance of a crouched limb posture at larger body sizes. We use regression and principal component analyses to detect allometries in muscle architecture, with and without phylogenetic correction. Of the muscle lengths that scale allometrically, all scale with negative allometry (i.e. relative shortening with increasing body mass), whereas all tendon lengths scale isometrically. Only two muscles' belly masses and two tendons' masses scale with positive allometry (i.e. relatively more massive with increasing body mass). Of the muscles that scale allometrically for physiological cross-sectional area, all scale positively (i.e. relatively greater area with increasing body mass). These muscles are mostly linked to control of hip and thigh movements. When the architecture data are phylogenetically corrected, there are few significant results, and only the strongest signals remain. None of the vertebral muscles scaled significantly differently from isometry. Principal component analysis and manovas showed that neither body size nor locomotor mode separate the felid species in morphospace. Our results support the inference that, despite some positively allometric trends in muscle areas related to thigh movement, larger cats have relatively weaker hindlimb and lumbosacral muscles in general. This decrease in power may be reflected in relative decreases in running speeds and is consistent with prevailing evidence that behavioural changes may be the primary mode of compensation for a consistently crouched limb posture in larger cats. [ABSTRACT FROM AUTHOR]

Crocodiles and their kin (Crocodylidae) use asymmetrical (bounding and galloping) gaits when moving rapidly. Despite being morphologically and ecologically similar, it seems alligators and their kin (Alligatoridae) do not. To investigate a possible anatomical basis for this apparent major difference in locomotor capabilities, we measured relative masses and internal architecture (fascicle lengths and physiological cross-sectional areas) of muscles of the pectoral and pelvic limbs of 40 individuals from six representative species of Crocodylidae and Alligatoridae. We found that, relative to body mass, Crocodylidae have significantly longer muscle fascicles (increased working range), particularly in the pectoral limb, and generally smaller muscle physiological cross-sectional areas (decreased force-exerting capability) than Alligatoridae. We therefore hypothesise that the ability of some crocodylians to use asymmetrical gaits may be limited more by the ability to make large, rapid limb motions (especially in the pectoral limb) than the ability to exert large limb forces. Furthermore, analysis of scaling patterns in muscle properties shows that limb anatomy in the two clades becomes more divergent during ontogeny. Limb muscle masses, fascicle lengths and physiological cross-sectional areas scale with significantly larger coefficients in Crocodylidae than Alligatoridae. This combination of factors suggests that inter-clade disparity in maximal limb power is highest in adult animals. Therefore, despite their apparent morphological similarities, both mean values and scaling patterns suggest that considerable diversity exists in the locomotor apparatus of extant Crocodylia. [ABSTRACT FROM AUTHOR]

One of the most extraordinary results of selective breeding is the modern broiler chicken, whose phenotypic attributes reflect its genetic success. Unfortunately, leg health issues and poor walking ability are prevalent in the broiler population, with the exact aetiopathogenesis unknown. Here we present a biomechanical analysis of the gait dynamics of the modern broiler and its two pureline commercial broiler breeder lines (A and B) in order to clarify how changes in basic morphology are associated with the way these chickens walk. We collected force plate and kinematic data from 25 chickens (market age), over a range of walking speeds, to quantify the three-dimensional dynamics of the centre of mass (CoM) and determine how these birds modulate the force and mechanical work of locomotion. Common features of their gait include extremely slow walking speeds, a wide base of support and large lateral motions of the CoM, which primarily reflect changes to cope with their apparent instability and large body mass. These features allowed the chickens to keep their peak vertical forces low, but resulted in high mediolateral forces, which exceeded fore-aft forces. Gait differences directly related to morphological characteristics also exist. This was particularly evident in Pureline B birds, which have a more crouched limb posture. Mechanical costs of transport were still similar across all lines and were not exceptional when compared with more wild-type ground-running birds. Broiler chickens seem to have an awkward gait, but some aspects of their dynamics show rather surprising similarities to other avian bipeds. [ABSTRACT FROM AUTHOR]

Foot pressure distributions during locomotion have causal links with the anatomical and structural configurations of the foot tissues and the mechanics of locomotion. Elephant feet have five toes bound in a flexible pad of fibrous tissue (digital cushion). Does this specialized foot design control peak foot pressures in such giant animals? And how does body size, such as during ontogenetic growth, influence foot pressures? We addressed these questions by studying foot pressure distributions in elephant feet and their correlation with body mass and centre of pressure trajectories, using statistical parametric mapping (SPM), a neuro-imaging technology. Our results show a positive correlation between body mass and peak pressures, with the highest pressures dominated by the distal ends of the lateral toes (digits 3, 4 and 5). We also demonstrate that pressure reduction in the elephant digital cushion is a complex interaction of its viscoelastic tissue structure and its centre of pressure trajectories, because there is a tendency to avoid rear 'heel' contact as an elephant grows. Using SPM, we present a complete map of pressure distributions in elephant feet during ontogeny by performing statistical analysis at the pixel level across the entire plantar/palmar surface. We hope that our study will build confidence in the potential clinical and scaling applications of mammalian foot pressures, given our findings in support of a link between regional peak pressures and pathogenesis in elephant feet. [ABSTRACT FROM AUTHOR]

The structure and motion of elephant limbs are unusual compared with those of other animals. Elephants stand and move with straighter limbs (at least when walking), and have limited speed and gait. We devised novel experiments to examine how the limbs of elephants support and propel their mass and to explore the factors that may constrain locomotor performance in these largest of living land animals. We demonstrate that elephant limbs are remarkably compliant even in walking, which maintains low peak forces. Dogma defines elephant limbs as extremely "columnar" for effective weight support, but we demonstrate that limb effective mechanical advantage (EMA) is roughly one-third of that predicted for their size. EMA in elephants is actually smaller than that in horses, which are only one-tenth their mass; it is comparable to human limb values. EMA drops sharply with speed in elephants, as it does in humans. Muscle forces therefore must increase as the limbs become more flexed, and we show how this flexion translates to greater volumes of muscle recruited for locomotion and hence metabolic cost. Surprisingly, elephants use their forelimbs and hindlimbs in similar braking and propulsive roles, not dividing these functions among limbs as was previously assumed or as in other quadrupeds. Thus, their limb function is analogous to four-wheel-drive vehicles. To achieve the observed limb compliance and low peak forces, elephants synchronize their limb dynamics in the vertical direction, but incur considerable mechanical costs from limbs working against each other horizontally. [ABSTRACT FROM AUTHOR]

Abstract: Archosaurian reptiles evolved a more erect posture and parasagittal gait early on. Early dinosaurs were the first habitual striding bipeds, a trait retained by living birds. Yet there is much more to archosaur locomotor evolution than these two transitions. I review our understanding of the pattern of locomotor evolution from the first archosaurs to Crocodylia and Neornithes, outlining where transitions of locomotor function evolved. I evaluate current research approaches, advocating more experimental work on extant animals to establish rigorous form–function relationships, and more biomechanical research that is bolstered by validation and sensitivity analysis of its assumptions, methods, and results. To cite this article: J.R. Hutchinson, C. R. Palevol 5 (2006) . [Copyright &y& Elsevier]

The anatomy and functions of muscle–tendon complexes and their bony attachments in birds and their outgroups show how the major pelvic limb muscle groups evolved. Fossils reveal that most changes evolved after the divergence of archosaurs in the Triassic, particularly in the dinosaurian precursors to birds. Three-dimensional limb control became concentrated at the hip joint; more distal joints and muscles were restricted to flexion or extension early in dinosaur evolution. Hip extensors expanded even though the primary femoral retractor M. caudofemoralis longus was reduced. Hip flexors and two-joint ‘hamstring’ muscles were simplified to a few large heads. Knee extensors increased their sizes and moment arms early in bipedal dinosaurs, but the patella and cranial cnemial crest evolved later in birds. Lower limb muscles expanded as ossifications such as the hypotarsus increased their moment arms. The ossification of lower limb tendons, particularly in extensors, is a recent novelty of birds. Muscles and tendons that develop large forces, stresses, and moments to stabilize or move the limbs became increasingly prominent on the line to birds. Locomotion evolved in a stepwise pattern that only recently produced the derived limb control mechanisms of crown-group birds, such as the strongly flexed hip and knee joints. [Copyright &y& Elsevier]