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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., (© 2019 Anatomical Society.)

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.

The number of precaudal vertebrae in all extant crocodylians is remarkably conservative, with nine cervicals, 15 dorsals and two sacrals, a pattern present also in their closest extinct relatives. The consistent vertebral count indicates a tight control of axial patterning by Hox genes during development. Here we report on a deviation from this pattern based on an associated skeleton of the giant caimanine Purussaurus , a member of crown Crocodylia, and several other specimens from the Neogene of the northern neotropics. P. mirandai is the first crown-crocodylian to have three sacrals, two true sacral vertebrae and one non-pathological and functional dorsosacral, to articulate with the ilium (pelvis). The giant body size of this caiman relates to locomotory and postural changes. The iliosacral configuration, a more vertically oriented pectoral girdle, and low torsion of the femoral head relative to the condyles are hypothesized specializations for more upright limb orientation or weight support., Competing Interests: TS, JH, OS, MD, JC, RS, MS No competing interests declared, (© 2019, Scheyer et al.)

Dromaeosaurids were a clade of bird-like, carnivorous dinosaurs that are well known for their characteristic morphology of pedal digit II, which bore an enlarged, sickle-shaped claw and permitted an extreme range of flexion-extension. Proposed functions for the claw often revolve around predation, but the exact manner of use varies widely. Musculoskeletal modelling provides an avenue to quantitatively investigate the biomechanics of this enigmatic system, and thereby test different behavioural hypotheses. Here, a musculoskeletal model of the hindlimb and pes of Deinonychus was developed, and mathematical optimization was used to assess the factors that maximize production of force at the claw tip. Optimization revealed that more crouched hindlimb postures (i.e., more flexed knees and ankles) and larger flexor muscle volumes consistently increased claw forces, although the optimal degree of digit flexion or extension depended on assumptions of muscle activity and fibre operating range. Interestingly, the magnitude of force capable of being produced at the claw tip was relatively small, arguing against regular transmission of a large proportion of body weight into a substrate principally via the claw tip. Such transmission would therefore likely have needed to occur via more proximal parts of the foot. Collectively, the results best support a grasping function for digit II (e.g., restraint of prey smaller than the dromaeosaurid's own body size), although other behaviours involving flexed hindlimbs cannot be excluded., Competing Interests: The author declares there are no competing interests.

Complex structures, like the vertebrate skull, are composed of numerous elements or traits that must develop and evolve in a coordinated manner to achieve multiple functions. The strength of association among phenotypic traits (i.e., integration), and their organization into highly-correlated, semi-independent subunits termed modules, is a result of the pleiotropic and genetic correlations that generate traits. As such, patterns of integration and modularity are thought to be key factors constraining or facilitating the evolution of phenotypic disparity by influencing the patterns of variation upon which selection can act. It is often hypothesized that selection can reshape patterns of integration, parceling single structures into multiple modules or merging ancestrally semi-independent traits into a strongly correlated unit. However, evolutionary shifts in patterns of trait integration are seldom assessed in a unified quantitative framework. Here, we quantify patterns of evolutionary integration among regions of the archosaur skull to investigate whether patterns of cranial integration are conserved or variable across this diverse group. Using high-dimensional geometric morphometric data from 3D surface scans and computed tomography scans of modern birds (n = 352), fossil non-avian dinosaurs (n = 27), and modern and fossil mesoeucrocodylians (n = 38), we demonstrate that some aspects of cranial integration are conserved across these taxonomic groups, despite their major differences in cranial form, function, and development. All three groups are highly modular and consistently exhibit high integration within the occipital region. However, there are also substantial divergences in correlation patterns. Birds uniquely exhibit high correlation between the pterygoid and quadrate, components of the cranial kinesis apparatus, whereas the non-avian dinosaur quadrate is more closely associated with the jugal and quadratojugal. Mesoeucrocodylians exhibit a slightly more integrated facial skeleton overall than the other grades. Overall, patterns of trait integration are shown to be stable among archosaurs, which is surprising given the cranial diversity exhibited by the clade. At the same time, evolutionary innovations such as cranial kinesis that reorganize the structure and function of complex traits can result in modifications of trait correlations and modularity., (© The Author(s) 2019. Published by Oxford University Press on behalf of the Society for Integrative and Comparative Biology. All rights reserved. For permissions please email: journals.permissions@oup.com.)

Active muscle performs various mechanical functions during locomotion: work output during shortening, work absorption when resisting (but not preventing) lengthening, and impulse (force-time integral) whenever there is active force. The energetic costs of these functions are important components in the energy budget during locomotion. We investigated how the pattern of stimulation and movement affects the mechanics and energetics of muscle fibre bundles isolated from wild rabbits ( Oryctolagus cuniculus ). The fibres were from muscles consisting of mainly fast-twitch, type 2 fibres. Fibre length was held constant (isometric) or a sinusoidal pattern of movement was imposed at a frequency similar to the stride frequency of running wild rabbits. Duty cycle (stimulation duration×movement frequency) and phase (timing of stimulation relative to movement) were varied. Work and impulse were measured as well as energy produced as heat. The sum of net work (work output-work input) and heat was taken as a measure of energetic cost. Maximum work output was produced with a long duty cycle and stimulation starting slightly before shortening, and was produced quite efficiently. However, efficiency was even higher with other stimulation patterns that produced less work. The highest impulse (considerably higher than isometric impulse) was produced when stimulation started while the muscle fibres were being lengthened. High impulse was produced very economically because of the low cost of producing force during lengthening. Thus, locomotion demanding high work, high impulse or economical work output or impulse requires a distinct pattern of stimulation and movement., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2019. Published by The Company of Biologists Ltd.)

Gravity may influence multiple aspects of legged locomotion, from the periods of limbs moving as pendulums to the muscle forces required to support the body. We present a system for exposing mice to hypergravity using a centrifuge and studying their locomotion and activity during exposure. Centrifuge-induced hypergravity has the advantages that it both allows animals to move freely, and it affects both body and limbs. The centrifuge can impose two levels of hypergravity concurrently, using two sets of arms of different lengths, each carrying a mouse cage outfitted with a force and speed measuring exercise wheel and an infrared high-speed camera; both triggered automatically when a mouse begins running on the wheel. Welfare is monitored using infrared cameras. As well as detailing the design of the centrifuge and instrumentation, we present example data from mice exposed to multiple levels of hypergravity and details of how they acclimatized to hypergravity., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2019. Published by The Company of Biologists Ltd.)

Background: The current study applies both traditional and Diffusible Iodine Contrast Enhanced computed microtomography (DICE µCT) techniques to reveal the musculoskeletal anatomy of Phlyctimantis maculatus . DICE µCT has emerged as a powerful tool to visualise intricate musculoskeletal anatomy. By generating 3D digital models, anatomical analyses can be conducted non-destructively, preserving the in situ 3D topography of the system, therefore eliminating some of the drawbacks associated with traditional methods. We aim to describe the musculature of the spine, pelvis, and hindlimb, compare the musculoskeletal anatomy and pelvic morphology of P. maculatus with functionally diverse frogs, and produce 3D digital anatomy reference data., Method: An adult frog was stained using an aqueous Lugol's solution and scanned in a SkyScan1176 in vivo µCT scanner. Scan images were reconstructed, resampled, and digitally segmented to produce a 3D model. A further adult female frog was dissected traditionally for visualisation of tendinous insertions., Results: Our work revealed three main findings: (1) P. maculatus has similar gross muscular anatomy to Rana catesbeiana (bullfrog) but is distinct from those species that exhibit ancestral traits (leopelmids) and those that are highly specialised (pipids), (2) P. maculatus 's pelvic anatomy best fits the description of Emerson's walking/hopping pelvic morphotype IIA, and (3) a split in the semimembranosus and gracilis major muscles is consistent with the reported myology in other anuran species., Discussion: While DICE µCT methods were instrumental in characterising the 3D anatomy, traditional dissection was still required to visualise important structures such as the knee aponeurosis, tendinous insertions, and fasciae. Nonetheless, the anatomical data presented here marks the first detailed digital description of an arboreal and terrestrial frog. Further, our digital model presents P. maculatus as a good frog model system and as such has formed a crucial platform for further functional analysis within the anuran pelvis and hindlimb., Competing Interests: The authors declare there are no competing interests.

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.

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., Competing Interests: John R. Hutchinson is an Academic Editor for PeerJ.

Some frog species, such as Kassina maculata (red-legged running frog), use an asynchronous walking/running gait as their primary locomotor mode. Prior comparative anatomy work has suggested that lateral rotation of the pelvis improves walking performance by increasing hindlimb stride length; however, this hypothesis has never been tested. Using non-invasive methods, experimental high-speed video data collected from eight animals were used to create two three-dimensional kinematic models. These models, each fixed to alternative local anatomical reference frames, were used to investigate the hypothesis that lateral rotation of the mobile ilio-sacral joint in the anuran pelvis plays a propulsive role in walking locomotion by increasing hindlimb stride length. All frogs used a walking gait (duty factor greater than 0.5) despite travelling over a range of speeds (0.04-0.23 m s -1 ). The hindlimb joint motions throughout a single stride were temporally synchronized with lateral rotation of the pelvis. The pelvis itself, on average, underwent an angular excursion of 12.71° (±4.39°) with respect to the body midline during lateral rotation. However, comparison between our two kinematic models demonstrated that lateral rotation of the pelvis only increases the cranio-caudal excursion of the hindlimb modestly. Thus, we propose that pelvic lateral rotation is not a stride length augmenting mechanism in K. maculata ., Competing Interests: There were no competing interests in the current study.

Electromyography (EMG) is used to understand muscle activity patterns in animals. Understanding how much variation exists in muscle activity patterns in homologous muscles across animal clades during similar behaviours is important for evaluating the evolution of muscle functions and neuromuscular control. We compared muscle activity across a range of archosaurian species and appendicular muscles, including how these EMG patterns varied across ontogeny and phylogeny, to reconstruct the evolutionary history of archosaurian muscle activation during locomotion. EMG electrodes were implanted into the muscles of turkeys, pheasants, quail, guineafowl, emus (three age classes), tinamous and juvenile Nile crocodiles across 13 different appendicular muscles. Subjects walked and ran at a range of speeds both overground and on treadmills during EMG recordings. Anatomically similar muscles such as the lateral gastrocnemius exhibited similar EMG patterns at similar relative speeds across all birds. In the crocodiles, the EMG signals closely matched previously published data for alligators. The timing of lateral gastrocnemius activation was relatively later within a stride cycle for crocodiles compared to birds. This difference may relate to the coordinated knee extension and ankle plantarflexion timing across the swing-stance transition in Crocodylia, unlike in birds where there is knee flexion and ankle dorsiflexion across swing-stance. No significant effects were found across the species for ontogeny, or between treadmill and overground locomotion. Our findings strengthen the inference that some muscle EMG patterns remained conservative throughout Archosauria: for example, digital flexors retained similar stance phase activity and M. pectoralis remained an 'anti-gravity' muscle. However, some avian hindlimb muscles evolved divergent activations in tandem with functional changes such as bipedalism and more crouched postures, especially M. iliotrochantericus caudalis switching from swing to stance phase activity and M. iliofibularis adding a novel stance phase burst of activity., (© 2019 Wiley Periodicals, Inc.)

Unmanned Aerial Systems (UAS) are increasingly being used recreationally, commercially and for wildlife research, but very few studies have quantified terrestrial mammalian reactions to UAS approaches. We used two Vertical Take-off and Landing (VTOL) UAS to approach seven herbivore species in the Moremi Game Reserve, Botswana, after securing the relevant permissions. We recorded responses to 103 vertical and 120 horizontal approaches, the latter from three altitudes above ground level (AGL). We ran mixed logistic regressions to identify factors triggering (i) any response and (ii) an evasive response. We included effects of activity, altitude, direction of approach, distance, habitat, herd type, herd size, other species, target species, time, VTOL type and wind strength. Response triggers were linked to altitude, distance, habitat and target species. Elephant (Loxodonta africana), giraffe (Giraffa camelopardalis), wildebeest (Connochaetes taurinus) and zebra (Equus quagga) were most affected by VTOL approach, impala (Aepyceros melampus) and lechwe (Kobus leche) were least responsive, and tsessebe (Damaliscus lunatus) displayed intermediate sensitivity. VTOLs flown lower than 60 m AGL and closer than 100 m horizontal distance from target animals triggered behavioural responses in most species. Enforced regulations on recreational UAS use in wildlife areas are necessary to minimise disturbance to terrestrial mammals.

The study of animal locomotion can be logistically challenging, especially in the case of large or unhandleable animals in uncontrolled environments. Here we demonstrate the utility of a low cost unmanned aerial vehicle (UAV) in measuring two-dimensional running kinematics from free-roaming giraffes ( Giraffa camelopardalis giraffa ) in the Free State Province, South Africa. We collected 120 Hz video of running giraffes, and calibrated each video frame using metatarsal length as a constant object of scale. We tested a number of methods to measure metatarsal length. The method with the least variation used close range photography and a trigonometric equation to spatially calibrate the still image, and derive metatarsal length. In the absence of this option, a spatially calibrated surface model of the study terrain was used to estimate topographical dimensions in video footage of interest. Data for the terrain models were collected using the same equipment, during the same study period. We subsequently validated the accuracy of the UAV method by comparing similar speed measurements of a human subject running on a treadmill, with treadmill speed. At 8 m focal distance we observed an error of 8% between the two measures of speed. This error was greater at a shorter focal distance, and when the subject was not in the central field of view. We recommend that future users maximise the camera focal distance, and keep the subject in the central field of view. The studied giraffes used a grounded rotary gallop with a speed range of 3.4-6.9 ms -1 (never cantering, trotting or pacing), and lower duty factors when compared with other cursorial quadrupeds. As this pattern might result in adverse increases in peak vertical limb forces with speed, it was notable to find that contralateral limbs became more in-phase with speed. Considering the latter pattern and the modest maximal speed of giraffes, we speculate that tissue safety factors are maintained within tolerable bounds this way. Furthermore, the angular kinematics of the neck were frequently isolated from the pitching of the body during running; this may be a result of the large mass of the head and neck. Further field experiments and biomechanical models are needed to robustly test these speculations., Competing Interests: John R. Hutchinson is an Academic Editor for PeerJ.

In recent years, there have been significant advances in the technology used to collect data on the movement and activity patterns of humans and animals. GPS units, which form the primary source of location data, have become cheaper, more accurate, lighter and less power-hungry, and their accuracy has been further improved with the addition of inertial measurement units. The consequence is a glut of geospatial time series data, recorded at rates that range from one position fix every several hours (to maximize system lifetime) to ten fixes per second (in high dynamic situations). Since data of this quality and volume have only recently become available, the analytical methods to extract behavioral information from raw position data are at an early stage of development. An instance of this lies in the analysis of animal movement patterns. When investigating solitary animals, the timing and location of instances of avoidance and association are important behavioral markers. In this paper, a novel analytical method to detect avoidance and association between individuals is proposed; unlike existing methods, assumptions about the shape of the territories or the nature of individual movement are not needed. Simulations demonstrate that false positives (type I error) are rare (1%-3%), which means that the test rarely suggests that there is an association if there is none.

Like other diapsids, Tyrannosaurus rex has two openings in the temporal skull region. In addition, like in other dinosaurs, its snout and lower jaw show large cranial fenestrae. In T. rex, they are thought to decrease skull weight, because, unlike most other amniotes, the skull proportion is immense compared to the body. Understanding morphofunctional complexity of this impressive skull architecture requires a broad scale phylogenetic comparison with skull types different to that of dinosaurs with fundamentally diverging cranial regionalization. Extant fully terrestrial vertebrates (amniotes) provide the best opportunities in that regard, as their skull performance is known from life. We apply for the first time anatomical network analysis to study skull bone integration and modular constructions in tyrannosaur and compare it with five representatives of the major amniote groups in order to get an understanding of the general patterns of amniote skull modularity. Our results reveal that the tyrannosaur has the most modular skull organization among the amniotes included in our study, with an unexpected separation of the snout in upper and lower sub-modules and the presence of a lower adductor chamber module. Independent pathways of bone reduction in opossum and chicken resulted in different degrees of cranial complexity with chicken having a typical sauropsidian pattern. The akinetic skull of opossum, alligator, and leatherback turtle evolved in independent ways mirrored in different patterns of skull modularity. Kinetic forms also show great diversity in modularity. The complex tyrannosaur skull modularity likely represents a refined mosaic of phylogenetic and ecological factors with food processing being probably most important for shaping its skull architecture. Mode of food processing primarily shaped skull integration among amniotes, however, phylogenetic patterns of skull integration are low in our sampling. Our general conclusions on amniote skull integrity are obviously preliminary and should be tested in subsequent studies. As such, this study provides a framework for future research focusing on the evolution of modularity on lower taxonomic levels.

In this work we present a novel, inductance-based system to measure and control the motion of bellows-driven continuum joints in soft robots. The sensing system relies on coils of wire wrapped around the minor diameters of each bellows on the joint. As the bellows extend, these coils of wire become more distant, decreasing their mutual inductance. Measuring this change in mutual inductance allows us to measure the motion of the joint. By dividing the sensing of the joint into two sections and measuring the motion of each section independently, we are able to measure the overall deformation of the joint with a piece-wise constant-curvature approximation. This technique allows us to measure lateral displacements that would be otherwise unobservable. When measuring bending, the inductance sensors measured the joint orientation with an RMS error of 1.1 °. The inductance sensors were also successfully used as feedback to control the orientation of the joint. The sensors proposed and tested in this work provided accurate motion feedback that would be difficult to achieve robustly with other sensors. This sensing system enables the creation of robust, self-sensing soft robots based on bellows-driven continuum joints., Competing Interests: Disclosure of potential conflicts of interest. Authors Wyatt Felt and C. David Remy are listed as inventors on a patent application for inductance sensing on bellows actuators. The other authors are (or were) employed by Pneubotics and/or its parent organization, Otherlab. Each of them has an investment interest with the company, and each is listed as an inventor in patents related to this work.

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.

Objective: To describe the anaesthetic, physiological and side effects of intramuscular (IM) medetomidine and ketamine, followed by inhalational anaesthesia with sevoflurane, in Nile crocodiles (Crocodylus niloticus)., Study Design: Observational trial., Animals: Ten juvenile captive-bred Nile crocodiles undergoing surgical implantation of skeletal beads and muscular electrodes., Methods: During preanaesthetic examination, the following variables were assessed: heart (HR) and respiratory (f R ) rates, and response to palpebral, corneal and toe- and tail-pinch withdrawal reflexes. The crocodiles were injected IM with an initial combination of medetomidine and ketamine and re-evaluated at 5 minute intervals for 20 minutes, or until they appeared unresponsive. If that did not occur, the drugs were redosed according to a decision tree based on the observed effects. The righting, biting and palatal valve reflexes were assessed in the unresponsive crocodiles, and used to confirm anaesthetic induction. Anaesthesia was maintained with sevoflurane in oxygen. At the end of surgery, medetomidine was antagonized with IM atipamezole., Results: The decision tree identified 0.3 mg kg -1 medetomidine and 15 mg kg -1 ketamine as a useful drug combination, which resulted in anaesthetic induction and surgical anaesthesia 16 ± 8 and 16 (25-20) minutes after injection, respectively. Compared to baseline, HR and f R significantly decreased after anaesthetic induction (p < 0.001), but then remained stable throughout surgery. Intraoperatively, cloacal temperature [27 (26-30) °C] did not change over time (p = 0.48). The total dose of atipamezole was 2 (1-3) mg kg -1 and time to recovery was 36 (20-60) minutes. Perioperative complications were not observed., Conclusions: and clinical relevance Medetomidine and ketamine, injected IM and followed by sevoflurane anaesthesia, may be regarded as a useful anaesthetic technique for juvenile Nile crocodiles undergoing minimally invasive experimental surgery., (Copyright © 2018 Association of Veterinary Anaesthetists and American College of Veterinary Anesthesia and Analgesia. Published by Elsevier Ltd. All rights reserved.)

Background: Both pleasure and competition horses regularly exercise on surfaces such as tarmac, gravel and turf during 'hacking'. Despite this, there is limited evidence relating to the effect of these surfaces upon foot-surface interaction., Objectives: To investigate forelimb foot placement, hoof vibration and movement symmetry in pleasure horses on three commonly encountered hacking surfaces., Study Design: Quantitative gait study in a convenience sample., Methods: Six horses regularly partaking in hacking exercise were ridden in walk and trot on all surfaces. Horses were equipped with one hoof-mounted, accelerometer and four body-mounted inertial measurement units (IMUs) to measure foot impact and movement symmetry. High-speed (400 FPS) video footage of foot-placement was acquired (dorsal, palmar, lateral views). Foot-impact and movement symmetry were analysed with a mixed effects model and Bowker symmetry tests for foot-placement analysis., Results: Vibration power and frequency parameters increase as perceived surface firmness increases from grass, to gravel, to tarmac (P≤0.001). Vibration power parameters were consistently greater at trot compared with walk (P≤0.001), but the same was not true for vibration frequency (P≥0.2). Greatest movement asymmetry was recorded during grass surface trotting. No significant difference in foot-placement was detected between the three surfaces., Main Limitations: This was a field study using three commonly encountered hacking surfaces. Surface properties change easily with water content and temperature fluctuations so care must be taken when considering other similar surfaces, especially at different times of the year. Six leisure horses were used so the results may not be representative of horses of all types., Conclusions: Vibration parameters generally increase as perceived surface firmness increases. Increasing speed alters vibration power but not frequency. Further investigations are required to determine the role that this may play in the development of musculoskeletal disease in horses., (© 2018 EVJ Ltd.)

Despite enhancements in the development of robotic systems, the energy economy of today's robots lags far behind that of biological systems. This is in particular critical for untethered legged robot locomotion. To elucidate the current stage of energy efficiency in legged robotic systems, this paper provides an overview on recent advancements in development of such platforms. The covered different perspectives include actuation, leg structure, control and locomotion principles. We review various robotic actuators exploiting compliance in series and in parallel with the drive-train to permit energy recycling during locomotion. We discuss the importance of limb segmentation under efficiency aspects and with respect to design, dynamics analysis and control of legged robots. This paper also reviews a number of control approaches allowing for energy efficient locomotion of robots by exploiting the natural dynamics of the system, and by utilizing optimal control approaches targeting locomotion expenditure. To this end, a set of locomotion principles elaborating on models for energetics, dynamics, and of the systems is studied., (Copyright © 2018 Kashiri, Abate, Abram, Albu-Schaffer, Clary, Daley, Faraji, Furnemont, Garabini, Geyer, Grabowski, Hurst, Malzahn, Mathijssen, Remy, Roozing, Shahbazi, Simha, Song, Smit-Anseeuw, Stramigioli, Vanderborght, Yesilevskiy and Tsagarakis.)

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.)

Functional outcomes, such as joint flexion and gait, are important indicators of efficacy in musculoskeletal research. Current technologies that objectively assess these parameters, including visual tracking systems and force plates, are challenging to deploy in long-term translational and clinical studies. To that end, we developed a wearable device that measures both physical activity and joint flexion using a single integrated sensor and magnet system, and hypothesized that it could evaluate post-operative functional recovery in an unsupervised setting. To demonstrate the feasibility of measuring joint flexion, we first compared knee motion from the wearable device to that acquired from a motion capture system to confirm that knee flexion measurements during normal human gait, predicted via changes in magnetic field strength, closely correlated with data acquired by motion capture. Using this system, we then monitored a porcine cohort after bilateral stifle arthrotomy to investigate longitudinal changes in physical activity and joint flexion. We found that unsupervised activity declined immediately after surgery, with a return to pre-operative activity occurring over a period of 2 weeks. By providing objective, individualized data on locomotion and joint function, this magnet-based system will facilitate the in vivo assessment of novel therapeutics in translational orthopaedic research.

Flying insects impress by their versatility and have been a recurrent source of inspiration for engineering devices. A large body of literature has focused on various aspects of insect flight, with an essential part dedicated to the dynamics of flapping wings and their intrinsically unsteady aerodynamic mechanisms. Insect wings flex during flight and a better understanding of structural mechanics and aeroelasticity is emerging. Most recently, insights from solid and fluid mechanics have been integrated with physiological measurements from visual and mechanosensors in the context of flight control in steady airs and through turbulent conditions. We review the key recent advances concerning flight in unsteady environments and how the multi-body mechanics of the insect structure-wings and body-are at the core of the flight control question. The issues herein should be considered when applying bio-informed design principles to robotic flapping wings., Competing Interests: Declarations of interest: none.

Standing up from a prone position is a critical daily activity for animals: failing to do so effectively may cause an injurious fall or increase predation susceptibility. This sit-to-stand behaviour (StS) is biomechanically interesting because it necessitates transitioning through near-maximal joint motion ranges from a crouched (i.e., poor mechanical advantage) to a more upright posture. Such large joint excursions should require large length changes of muscle-tendon units. Here we integrate experimental and musculoskeletal simulation methods to quantify the joint motions, limb forces, and muscle fibre forces, activations and length changes during StS in an extreme athlete-the greyhound-which has large hindlimb muscles bearing short-fibred distal muscles and long tendons. Study results indicate that hindlimb anti-gravity muscle fibres operate near their ~50% limits of length change during StS; mostly by starting at highly lengthened positions. StS also requires high muscle activations (>50%), in part due to non-sagittal motions. Finally, StS movements require passive non-muscular support in the distal hindlimb where short-fibred muscles are incapable of sustaining StS themselves. Non-locomotor behaviours like StS likely impose important trade-offs between muscle fibre force capacity and length changes, as well as active and passive mechanisms of support, that have been neglected in locomotor biomechanics studies.

Background: Musculoskeletal models are commonly used to quantify joint motions and loads during human motion. Constraining joint kinematics simplifies these models but the implications of the placement and quantity of markers used during data acquisition remains unclear. The purpose of this study was to establish the effects of marker placement and quantity on lower extremity kinematics calculated using a constrained-kinematic model. We hypothesized that a constrained-kinematic model would produce lower-extremity kinematics errors that correlated with the number of tracking markers removed from the thigh and shank., Methods: Healthy-young adults (N = 10) walked on a treadmill at slow, moderate, and fast speeds while skin-mounted markers were tracked using motion capture. Lower extremity kinematics were calculated for 256 combinations of leg and shank markers to establish the implications of marker placement and quantity on joint kinematics. Marker combinations that yielded differences greater than 5 degrees were tested with paired t-tests and the relationship between number of markers and kinematic errors were modeled with polynomials to determine goodness of fit (R 2 )., Results: Sagittal joint and hip coronal kinematics errors were smaller than documented errors caused by soft-tissue artifact, which tends to be approximately 5 degrees, when excluding thigh and shank markers. Joint angle and center kinematic errors negatively correlated with the number of markers included in the analyses (R 2  > 0.97) and typically showed the greatest error reductions when two markers were included on the thigh or shank segments. Further, we demonstrated that a simplified marker set that included markers on the pelvis, lateral knee condyle, lateral malleolus, and shoes produced kinematics that strongly agreed with the traditional marker set that included 3 tracking markers for each segment., Conclusion: Constrained-kinematic models are resilient to marker placement and quantity, which has implications on study design and post-processing workflows.

Large mammals that live in arid and/or desert environments can cope with seasonal and local variations in rainfall, food and climate 1 by moving long distances, often without reliable water or food en route. The capacity of an animal for this long-distance travel is substantially dependent on the rate of energy utilization and thus heat production during locomotion-the cost of transport 2-4 . The terrestrial cost of transport is much higher than for flying (7.5 times) and swimming (20 times) 4 . Terrestrial migrants are usually large 1-3 with anatomical specializations for economical locomotion 5-9 , because the cost of transport reduces with increasing size and limb length 5-7 . Here we used GPS-tracking collars 10 with movement and environmental sensors to show that blue wildebeest (Connochaetes taurinus, 220 kg) that live in a hot arid environment in Northern Botswana walked up to 80 km over five days without drinking. They predominantly travelled during the day and locomotion appeared to be unaffected by temperature and humidity, although some behavioural thermoregulation was apparent. We measured power and efficiency of work production (mechanical work and heat production) during cyclic contractions of intact muscle biopsies from the forelimb flexor carpi ulnaris of wildebeest and domestic cows (Bos taurus, 760 kg), a comparable but relatively sedentary ruminant. The energetic costs of isometric contraction (activation and force generation) in wildebeest and cows were similar to published values for smaller mammals. Wildebeest muscle was substantially more efficient (62.6%) than the same muscle from much larger cows (41.8%) and comparable measurements that were obtained from smaller mammals (mouse (34%) 11 and rabbit (27%)). We used the direct energetic measurements on intact muscle fibres to model the contribution of high working efficiency of wildebeest muscle to minimizing thermoregulatory challenges during their long migrations under hot arid conditions.

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., Competing Interests: John Hutchinson and Andrew Farke are Academic Editors for PeerJ.

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., Competing Interests: John Hutchinson is an Academic Editor for PeerJ.

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., Competing Interests: John Hutchinson and Andrew Farke are Academic Editors for PeerJ.

Bone adaptation is modulated by the timing, direction, rate and magnitude of mechanical loads. To investigate whether frequent slow, or infrequent fast, gaits could dominate bone adaptation to load, we compared scaling of the limb bones from two mammalian herbivore clades that use radically different high-speed gaits, bipedal hopping (suborder Macropodiformes; kangaroos and kin) and quadrupedal galloping (order Artiodactyla; goats, deer and kin). Forelimb and hindlimb bones were collected from 20 artiodactyl and 15 macropod species (body mass M 1.05-1536 kg) and scanned in computed tomography or X-ray microtomography. Second moment of area ( I max ) and bone length ( l ) were measured. Scaling relations ( y = ax b ) were calculated for l versus M for each bone and for I max versus M and I max versus l for every 5% of length. I max versus M scaling relationships were broadly similar between clades despite the macropod forelimb being nearly unloaded, and the hindlimb highly loaded, during bipedal hopping. I max versus l and l versus M scaling were related to locomotor and behavioural specializations. Low-intensity loads may be sufficient to maintain bone mass across a wide range of species. Occasional high-intensity gaits might not break through the load sensitivity saturation engendered by frequent low-intensity gaits., Competing Interests: The authors declare that we have no competing interests.

Flapping flight is the most power-demanding mode of locomotion, associated with a suite of anatomical specializations in extant adult birds. In contrast, many developing birds use their forelimbs to negotiate environments long before acquiring "flight adaptations," recruiting their developing wings to continuously enhance leg performance and, in some cases, fly. How does anatomical development influence these locomotor behaviors? Isolating morphological contributions to wing performance is extremely challenging using purely empirical approaches. However, musculoskeletal modeling and simulation techniques can incorporate empirical data to explicitly examine the functional consequences of changing morphology by manipulating anatomical parameters individually and estimating their effects on locomotion. To assess how ontogenetic changes in anatomy affect locomotor capacity, we combined existing empirical data on muscle morphology, skeletal kinematics, and aerodynamic force production with advanced biomechanical modeling and simulation techniques to analyze the ontogeny of pectoral limb function in a precocial ground bird ( Alectoris chukar ). Simulations of wing-assisted incline running (WAIR) using these newly developed musculoskeletal models collectively suggest that immature birds have excess muscle capacity and are limited more by feather morphology, possibly because feathers grow more quickly and have a different style of growth than bones and muscles. These results provide critical information about the ontogeny and evolution of avian locomotion by (i) establishing how muscular and aerodynamic forces interface with the skeletal system to generate movement in morphing juvenile birds, and (ii) providing a benchmark to inform biomechanical modeling and simulation of other locomotor behaviors, both across extant species and among extinct theropod dinosaurs.

Structure is an important clinical marker of tendon health; however, current standards use qualitative scores that are not strongly reliable. Therefore, the purpose of this study was to establish the reliability of an image-processing technique that quantifies tendon collagen structure using B-mode ultrasound images. Longitudinal images of the Achilles tendon were collected in 12 healthy young adults, and intra- and inter-rater reliability was assessed over multiple image selections and multiple days. Intraclass correlation coefficients were strong (r ≥ 0.71) for all comparisons. These findings demonstrate that quantitative assessments of tendon structure using B-mode ultrasound are reliable., (© 2018 by the American Institute of Ultrasound in Medicine.)

In this paper, we systematically investigate passive gaits that emerge from the natural mechanical dynamics of a bipedal system. We use an energetically conservative model of a simple spring-leg biped that exhibits well-defined swing leg dynamics. Through a targeted continuation of periodic motions of this model, we systematically identify different gaits that emerge from simple bouncing in place. We show that these gaits arise along one-dimensional manifolds that bifurcate into different branches with distinctly different motions. The branching is associated with repeated breaks in symmetry of the motion. Among others, the resulting passive dynamic gaits include walking, running, hopping, skipping and galloping. Our work establishes that the most common bipedal gaits can be obtained as different oscillatory motions (or nonlinear modes) of a single mechanical system with a single set of parameter values. For each of these gaits, the timing of swing leg motion and vertical motion is matched. This work thus supports the notion that different gaits are primarily a manifestation of the underlying natural mechanical dynamics of a legged system. Our results might explain the prevalence of certain gaits in nature, and may provide a blueprint for the design and control of energetically economical legged robots., (© 2018 The Author(s).)

Although many arthropods have the ability to voluntarily lose limbs, how these animals rapidly adapt to such an extreme perturbation remains poorly understood. It is thought that moving with certain gaits can enable efficient, stable locomotion; however, switching gaits requires complex information flow between and coordination of an animal's limbs. We show here that upon losing two legs, spiders can switch to a novel, more statically stable gait, or use temporal adjustments without a gait change. The resulting gaits have higher overall static stability than the gaits that would be imposed by limb loss. By decreasing the time spent in a low-stability configuration - effectively 'limping' over less-stable phases of the stride - spiders increased the overall stability of the less statically stable gait with no observable reduction in speed, as compared with the intact condition. Our results shed light on how voluntary limb loss could have persisted evolutionarily among many animals, and provide bioinspired solutions for robots when they break or lose limbs., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2018. Published by The Company of Biologists Ltd.)

Animals, including humans, have been shown to maintain a gait during locomotion that minimizes the risk of injury and energetic cost. Despite the importance of understanding the mechanisms of gait regulation, ethical and experimental challenges have prevented full exploration of these. Here we present data on the gait response of mice to rapid, precisely timed, spatially confined mechanical perturbations. Our data elucidate that after the mechanical perturbation, the mouse gait response is anisotropic, preferring deviations away from the trot towards bounding, over those towards other gaits, such as walk or pace. We quantified this shift by projecting the observed gait onto the line between trot and bound, in the space of quadrupedal gaits. We call this projection λ. For λ=0, the gait is the ideal trot; for λ=±π, it is the ideal bound. We found that the substrate perturbation caused a significant shift in λ towards bound during the stride in which the perturbation occurred and the following stride (linear mixed effects model: Δλ=0.26±0.07 and Δλ=0.21±0.07, respectively; random effect for animal, p < 0.05 for both strides, n = 8 mice). We hypothesize that this is because the bounding gait is better suited to rapid acceleration or deceleration, and an exploratory analysis of jerk showed that it was significantly correlated with λ (p < 0.05). Understanding how gait is controlled under perturbations can aid in diagnosing gait pathologies and in the design of more agile robots., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

Inspired by biological control synergies, wherein fixed groups of muscles are activated in a coordinated fashion to perform tasks in a stable way, we present an analogous control approach for the stabilization of legged robots and apply it to a model of running. Our approach is based on the step-to-step notion of stability, also known as orbital stability, using an orbital control Lyapunov function. We map both the robot state at a suitably chosen Poincaré section (an instant in the locomotion cycle such as the mid-flight phase) and control actions (e.g., foot placement angle, thrust force, braking force) at the current step, to the robot state at the Poincaré section at the next step. This map is used to find the control action that leads to a steady state (nominal) gait. Next, we define a quadratic Lyapunov function at the Poincaré section. For a range of initial conditions, we find control actions that would minimize an energy metric while ensuring that the Lyapunov function decays exponentially fast between successive steps. For the model of running, we find that the optimization reveals three distinct control synergies depending on the initial conditions: (1) foot placement angle is used when total energy is the same as that of the steady state (nominal) gait; (2) foot placement angle and thrust force are used when total energy is less than the nominal; and (3) foot placement angle and braking force are used when total energy is more than the nominal.

Many medium and large herbivores locomote forwards very slowly and intermittently when grazing. While the footfall order during grazing is the same as for walking, the relative fore-hind timing-phasing-is quite different. Extended periods of static stability are clearly required during grazing; however, stability requirements are insufficient to account for the timing. Aspects of relatively rapid rolling and pitching-toppling due to the resistance of the back to bending and twisting-can be included in a simplifying geometric model to explain the observation that, in grazing livestock, a step forward with a forefoot is consistently and immediately followed by a step forward from the hind; but not vice versa. The same principles and geometry, but applied to the footfall pattern of walking primates, show that toppling would occur at a different point in the gait cycle. This provides a potential account for the distinctive diagonal-sequence footfall pattern of primates, as it prevents the instant of toppling from being at forefoot placement. Careful and controlled hand positioning would thus be facilitated, presumably beneficial to walking on top of branches, despite a slight energetic cost compared with the usual lateral sequence pattern of horses., (© 2018 The Authors.)

Inspired by the locomotive advantages that an articulated spine enables in quadrupedal animals, we explore and quantify the energetic effect that an articulated spine has in legged robots. We compare two model instances of a conceptual planar quadruped: one with a traditional rigid main body and one with an articulated main body with an actuated spinal joint. Both models feature four distinct legs, series elastic actuation, distributed mass in all body segments, and limits on actuator torque and speed. Using optimal control to find the energetically optimal joint trajectories, actuator inputs, and footfall timing, we examine and compare the positive mechanical work cost of transport of both models across multiple gaits and speeds. Our results show that an articulated spine increases the maximum possible speed and improves the locomotor economy at higher velocities, especially for asymmetrical gaits. The driving factors for these improvements are the same mechanistic effects that facilitate asymmetrical gaits in nature: improved leg recirculation, elastic energy storage in the spine, and enlarged stride lengths.

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.

Background: Agreement among experienced clinicians is poor when assessing the presence and severity of ataxia, especially when signs are mild. Consequently, objective gait measurements might be beneficial for assessment of horses with neurological diseases., Objectives: To assess diagnostic criteria using motion capture to measure variability in spatial gait-characteristics and swing duration derived from ataxic and non-ataxic horses, and to assess if variability increases with blindfolding., Study Design: Cross-sectional., Methods: A total of 21 horses underwent measurements in a gait laboratory and live neurological grading by multiple raters. In the gait laboratory, the horses were made to walk across a runway surrounded by a 12-camera motion capture system with a sample frequency of 240 Hz. They were made to walk normally and with a blindfold in at least three trials each. Displacements of reflective markers on head, fetlock, hoof, fourth lumbar vertebra, tuber coxae and sacrum derived from three to four consecutive strides were processed and descriptive statistics, receiver operator characteristics (ROC) to determine the diagnostic sensitivity, specificity and area under the curve (AUC), and correlation between median ataxia grade and gait parameters were determined., Results: For horses with a median ataxia grade ≥2, coefficient of variation for the location of maximum vertical displacement of pelvic and thoracic distal limbs generated good diagnostic yield. The hoofs of the thoracic limbs yielded an AUC of 0.81 with 64% sensitivity and 90% specificity. Blindfolding exacerbated the variation for ataxic horses compared to non-ataxic horses with the hoof marker having an AUC of 0.89 with 82% sensitivity and 90% specificity., Main Limitations: The low number of consecutive strides per horse obtained with motion capture could decrease diagnostic utility., Conclusions: Motion capture can objectively aid the assessment of horses with ataxia. Furthermore, blindfolding increases variation in distal pelvic limb kinematics making it a useful clinical tool., (© 2017 EVJ Ltd.)

How extinct, non-avian theropod dinosaurs moved is a subject of considerable interest and controversy. A better understanding of non-avian theropod locomotion can be achieved by better understanding terrestrial locomotor biomechanics in their modern descendants, birds. Despite much research on the subject, avian terrestrial locomotion remains little explored in regards to how kinematic and kinetic factors vary together with speed and body size. Here, terrestrial locomotion was investigated in twelve species of ground-dwelling bird, spanning a 1,780-fold range in body mass, across almost their entire speed range. Particular attention was devoted to the ground reaction force (GRF), the force that the feet exert upon the ground. Comparable data for the only other extant obligate, striding biped, humans, were also collected and studied. In birds, all kinematic and kinetic parameters examined changed continuously with increasing speed, while in humans all but one of those same parameters changed abruptly at the walk-run transition. This result supports previous studies that show birds to have a highly continuous locomotor repertoire compared to humans, where discrete 'walking' and 'running' gaits are not easily distinguished based on kinematic patterns alone. The influences of speed and body size on kinematic and kinetic factors in birds are developed into a set of predictive relationships that may be applied to extinct, non-avian theropods. The resulting predictive model is able to explain 79-93% of the observed variation in kinematics and 69-83% of the observed variation in GRFs, and also performs well in extrapolation tests. However, this study also found that the location of the whole-body centre of mass may exert an important influence on the nature of the GRF, and hence some caution is warranted, in lieu of further investigation.

Animal movement is fundamental for ecosystem functioning and species survival, yet the effects of the anthropogenic footprint on animal movements have not been estimated across species. Using a unique GPS-tracking database of 803 individuals across 57 species, we found that movements of mammals in areas with a comparatively high human footprint were on average one-half to one-third the extent of their movements in areas with a low human footprint. We attribute this reduction to behavioral changes of individual animals and to the exclusion of species with long-range movements from areas with higher human impact. Global loss of vagility alters a key ecological trait of animals that affects not only population persistence but also ecosystem processes such as predator-prey interactions, nutrient cycling, and disease transmission., (Copyright © 2018, The Authors, some rights reserved; exclusive licensee American Association for the Advancement of Science. No claim to original U.S. Government Works.)

Background: Horses show compensatory head movement in hindlimb lameness and compensatory pelvis movement in forelimb lameness but little is known about the relationship of withers movement symmetry with head and pelvic asymmetry in horses with naturally occurring gait asymmetries., Objectives: To document head, withers and pelvic movement asymmetry and timing differences in horses with naturally occurring gait asymmetries., Study Design: Retrospective analysis of gait data., Methods: Head, withers and pelvic movement asymmetry and timing of displacement minima and maxima were quantified from inertial sensors in 163 Thoroughbreds during trot-ups on hard ground. Horses were divided into 4 subgroups using the direction of head and withers movement asymmetry. Scatter plots of head vs. pelvic movement asymmetry illustrated how the head-withers relationship distinguishes between contralateral and ipsilateral head-pelvic movement asymmetry. Independent t test or Mann-Whitney U test (P<0.05) compared pelvic movement asymmetry and timing differences between groups., Results: The relationship between head and withers asymmetry (i.e. same sided or opposite sided asymmetry) predicts the relationship between head and pelvic asymmetry in 69-77% of horses. Pelvic movement symmetry was significantly different between horses with same sign vs. opposite sign of head-withers asymmetry (P<0.0001). Timing of the maximum head height reached after contralateral ('sound') stance was delayed compared to withers (P = 0.02) and pelvis (P = 0.04) in horses with contralateral head-withers asymmetry., Main Limitations: The clinical lameness status of the horses was not investigated., Conclusion: In the Thoroughbreds with natural gait asymmetries investigated here, the direction of head vs. withers movement asymmetry identifies the majority of horses with ipsilateral and contralateral head and pelvic movement asymmetries. Withers movement should be further investigated for differentiating between forelimb and hindlimb lame horses. Horses with opposite sided head and withers asymmetry significantly delay the upward movement of the head after 'sound' forelimb stance., (© 2017 The Authors. Equine Veterinary Journal published by John Wiley & Sons Ltd on behalf of EVJ Ltd.)

Many of the major locomotor transitions during the evolution of Archosauria, the lineage including crocodiles and birds as well as extinct Dinosauria, were shifts from quadrupedalism to bipedalism (and vice versa). Those occurred within a continuum between more sprawling and erect modes of locomotion and involved drastic changes of limb anatomy and function in several lineages, including sauropodomorph dinosaurs. We present biomechanical computer models of two locomotor extremes within Archosauria in an analysis of joint ranges of motion and the moment arms of the major forelimb muscles in order to quantify biomechanical differences between more sprawling, pseudosuchian (represented the crocodile Crocodylus johnstoni ) and more erect, dinosaurian (represented by the sauropodomorph Mussaurus patagonicus ) modes of forelimb function. We compare these two locomotor extremes in terms of the reconstructed musculoskeletal anatomy, ranges of motion of the forelimb joints and the moment arm patterns of muscles across those ranges of joint motion. We reconstructed the three-dimensional paths of 30 muscles acting around the shoulder, elbow and wrist joints. We explicitly evaluate how forelimb joint mobility and muscle actions may have changed with postural and anatomical alterations from basal archosaurs to early sauropodomorphs. We thus evaluate in which ways forelimb posture was correlated with muscle leverage, and how such differences fit into a broader evolutionary context (i.e. transition from sprawling quadrupedalism to erect bipedalism and then shifting to graviportal quadrupedalism). Our analysis reveals major differences of muscle actions between the more sprawling and erect models at the shoulder joint. These differences are related not only to the articular surfaces but also to the orientation of the scapula, in which extension/flexion movements in Crocodylus (e.g. protraction of the humerus) correspond to elevation/depression in Mussaurus . Muscle action is highly influenced by limb posture, more so than morphology. Habitual quadrupedalism in Mussaurus is not supported by our analysis of joint range of motion, which indicates that glenohumeral protraction was severely restricted. Additionally, some active pronation of the manus may have been possible in Mussaurus , allowing semi-pronation by a rearranging of the whole antebrachium (not the radius against the ulna, as previously thought) via long-axis rotation at the elbow joint. However, the muscles acting around this joint to actively pronate it may have been too weak to drive or maintain such orientations as opposed to a neutral position in between pronation and supination. Regardless, the origin of quadrupedalism in Sauropoda is not only linked to manus pronation but also to multiple shifts of forelimb morphology, allowing greater flexion movements of the glenohumeral joint and a more columnar forelimb posture., Competing Interests: John Hutchinson is an Academic Editor for PeerJ.

Intra-cortical bone remodelling is a cell-driven process that replaces existing bone tissue with new bone tissue in the bone cortex, leaving behind histological features called secondary osteons. While the scaling of bone dimensions on a macroscopic scale is well known, less is known about how the spatial dimensions of secondary osteons vary in relation to the adult body size of the species. We measured the cross-sectional area of individual intact secondary osteons and their central Haversian canals in transverse sections from 40 stylopodal bones of 39 mammalian species (body mass 0.3-21 000 kg). Scaling analysis of our data shows that mean osteonal resorption area (negative allometry, exponent 0.23, R 2  0.54, p <0.005) and Haversian canal area (negative allometry, exponent 0.31, R 2  0.45, p <0.005) are significantly related to body mass, independent of phylogeny. This study is the most comprehensive of its kind to date, and allows us to describe overall trends in the scaling behaviour of secondary osteon dimensions, supporting the inference that the osteonal resorption area may be limited by the need to avoid fracture in smaller mammalian species, but the need to maintain osteocyte viability in larger mammalian species., Competing Interests: The authors declare no competing interests. C.P., H.C. and M.C. are employed by RCS, which oversees the conservation of the Quekett collection; however, this association had no bearing on the analysis or interpretation of the results.